FOOD GRAINS
Second Edition
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ENCYCLOPEDIA OF
FOOD GRAINS
Second Edition
VOLUME 1
THE WORLD OF FOOD GRAINS
EDITORS
COLIN WRIGLEY
University of Queensland, Brisbane, Australia
HAROLD CORKE
University of Hong Kong, Hong Kong, Peoples Republic of China
KOUSHIK SEETHARAMAN{
JON FAUBION
Kansas State University, Manhattan KS, USA
{
deceased
The following articles are US government works in the public domain and are not subject to copyright:
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This book and the individual contributions contained in it are protected under copyright by the Publisher (other than as may be noted
herein).
Notice
Knowledge and best practice in this field are constantly changing. As new research and experience broaden our understanding, changes in
research methods, professional practices, or medical treatment may become necessary.
Practitioners and researchers may always rely on their own experience and knowledge in evaluating and using any information, methods,
compounds, or experiments described herein. In using such information or methods they should be mindful of their own safety and the
safety of others, including parties for whom they have a professional responsibility.
To the fullest extent of the law, neither the Publisher nor the authors, contributors, or editors, assume any liability for any injury and/or
damage to persons or property as a matter of products liability, negligence or otherwise, or from any use or operation of any methods,
products, instructions, or ideas contained in the material herein.
Preface xvii
Tributes to Former Co-Editors xix
Editors Biography xxiii
Editorial Advisory Board xxv
Contributors xxvii
VOLUME 1
The Basics 1
The Grains that Feed the World 1
CW Wrigley, H Corke, and J Faubion
Grain and Plant Morphology of Cereals and how characters can be used to identify varieties 51
J Wyatt
Maize: Overview 99
MP Scott and M Emery
Wheat: An Overview of the Grain That Provides Our Daily Bread 105
CW Wrigley
v
vi Contents
African Rice (Oryza glaberrima): A Brief History and Its Growing Importance in Current
Rice Breeding Efforts 140
JT Manful and S Graham-Acquaah
Barley: An Overview of a Versatile Cereal Grain with Many Food and Feed Uses 147
AM Stanca, A Gianinetti, F Rizza, and V Terzi
Grain Production and Consumption in Oceania: Australia and Pacific Countries 409
CW Wrigley and RJ French
Research Organizations of the World: Asia-Pacific, CentralSouth America, and AfricaMiddle East 444
JH Skerritt
viii Contents
Research Organizations of the World: Global Trends and the Commercial Sector 451
JH Skerritt
VOLUME 2
Celiac Disease 83
P Koehler, H Wieser, and KA Scherf
Proteins 98
The Protein Chemistry of Cereal Grains 98
F Bekes and CW Wrigley
The Enzymes Associated with Food Grains and Their Functions in Processing 138
DJ Mares, K Mrva, and GB Fincher
Carbohydrates 161
Carbohydrate Metabolism 161
RN Chibbar, S Jaiswal, M Gangola, and M Baga
Fats 248
Lipid Chemistry 248
L Day
Mycotoxins 290
MT Fletcher and BJ Blaney
Units of Grain Science and Trade: Equivalence between the US, Chinese, and Metric Units 308
W Huang and CW Wrigley
VOLUME 3
Wheat-Based Foods 1
Cereals: Overview of Uses: Accent on Wheat Grain 1
CF Morris
Breads 8
TR Moore
Noodles: Starch 64
LS Collado and H Corke
Pasta 79
M Sissons
Tortillas 90
LW Rooney and SO Serna-Saldivar
Non-wheat Foods 97
Maize: Foods from Maize 97
SO Serna-Saldivar
Contents xi
Biodiesel 245
Y Yan
The Gluten Proteins of the Wheat Grain in Relation to Flour Quality 375
F Bekes, MC Gianibelli, and CW Wrigley
VOLUME 4
Wheat: Marketing 9
RL Cracknell and RM Williams
The Nature, Causes, and Control of Grain Diseases in the Major Cereal Species 74
RA McIntosh, PM Williamson, and CW Wrigley
Contaminants of Grain 83
J McLean and CW Wrigley
xiv Contents
Appendix 1. Nutrient-Composition Tables for Grains and for Grain-Based Products 433
SF Schakel, N Van Heel, and J Harnack
Appendix 2. Lists of Standardized Testing Methods for the Analysis of Grain and Grain-Based Foods 450
AR Bridges and CW Wrigley
Appendix 3. Grains, Foods, and Ingredients Suiting Gluten-Free Diets for Celiac Disease 460
JM Jones, V Zevallos, and CW Wrigley
Glossary 467
Index 491
PREFACE
The grain-producing plants are essential to life on earth. They are the primary means by which sunlight, the
primary source of the earths energy, is captured via photosynthesis to turn carbon dioxide and water into
carbohydrates and thus into the wider range of the biochemicals needed by all life forms.
The summary is simple:
The suns energy CO2 H2O
#
Grain-bearing plants
which provide:
Food, Fats, Fuel and Fiber for ourselves
Feed, Forage, Fodder for our animals
Food for Fish Farms
As its title indicates, the Encyclopedia of Food Grains concentrates on the food uses of grains, but details are also
provided about the wider roles of grains. Greater detail about the use of grains for animal feeding is already
available via the on-line Feedipedia encyclopedia (http://www.feedipedia.org/).
Worldwide, however, it is our food that is our obsession first and foremost. We see our health (or
otherwise) to be intimately related to our diet. Therefore nutrition is a major accent of this second edition,
occupying the whole of the second volume, following the first volumes description of the basic aspects of the
world of grains. The third and fourth volumes work their ways back up the value-added grain chain to describe
grain processing and production, thence to breeding and genetics.
The sequence of articles in the first edition (the Encyclopedia of Grain Science, 2004) followed an alphabetical
order. In contrast, the distinct ordering of articles in the second edition assembles fundamental concepts in
Volume 1 The World of Food Grains. Moving into the later volumes, greater complexity and erudition are
evident. Thus, unusual terms such as Transcriptomics, Aeciospore and Allelopathy are more likely to be
found in later volumes than in the first. In any case, reference to the glossary and the index should help in
elucidating such terms.
This second edition is a considerable expansion on the content of the first edition, with its three print volumes
and 168 articles. Now, some twelve years later, the four print volumes of the Encyclopedia of Food Grains provide
xvii
xviii Preface
216 articles, many newly written for the new edition. Those articles that have been reprinted from the first
edition have been updated. In all cases, authors and reviewers are world renowned experts in their respective
fields.
This second edition has been more than three years in its development. In the early stages, the editors were
Harold Corke, Chuck Walker and Colin Wrigley (the three editors of the first edition). Sadly in 2012, Chuck
Walker passed away, leaving a gap that was difficult to fill.
Chuck was replaced by Koushik Seetharaman, who worked avidly in developing the concepts for this second
edition, including his contributions to a meeting of the editors and Elsevier staff in Sydney, January, 2013.
Sadly, Koushik suffered a heart attack in June, 2014. However, his contributions to the second edition during
this critical development stage were such that Elsevier decided that his name should remain as a co-editor.
Tributes to Professors Walker and Seetharaman are provided below.
In July, 2014, Professor Jon Faubion was appointed as a co-editor. Dr Faubion is the Charles Singleton
Professor of Baking and Cereal Science in the Department of Grain Science, Kansas State University. He is a
more than worthy replacement under such tragic circumstances.
The production of an encyclopedia involves a wide range of experts with expertise in a diversity of areas of
knowledge. We acknowledge with grateful thanks the groups of contributors:
Members of the Editorial Advisory Board;
Authors who have shared their special knowledge and experience;
Elsevier staff who have provided extensive advice and administrative services, especially Donna de
Weerd-Wilson, Simon Holt and Rachel Gerlis (successive Supervisory Staff) and Gemma Tomalin,
Gemma Taft, Joanne Williams and Mark Harper (successive Content Project Managers).
The Editors
Harold Corke, Jon Faubion, Koushik Seetharaman, and Colin Wrigley
TRIBUTES TO FORMER CO-EDITORS
Chuck Walker was one of the three editors of the first edition of Elseviers Grains
Encyclopedia (the Encyclopedia of Grain Science, 2004). He was again appointed
as an editor of this second edition. He participated in the planning stages for the
second edition until his untimely death on 26 April 2012. At the time of his
death, Chuck was a Professor Emeritus at the Department of Grain Science and
Industry at Kansas State University, Manhattan, KS, USA.
Charles Eugene Walker was born on 17 December 1936, in Winterset, Mad-
ison county, IA, USA, during the great depression. He was the eldest child of
Albert Cecil and Bessie Irene Headley Walker, poor farmers in Winterset, Mad-
ison county, IA, USA. He walked more than a mile to attend rural one-room
grammar schools in Warren county and graduated from high school in New
Virginia, IA, USA.
Better known as Chuck in later life, Dr Walker received a BS degree in
Chemical Engineering from Iowa State University, Ames, IA, USA, in May 1959. At a mixer party in the fall
of 1957, he met Shannon Rose Philp, a pretty freshman student from Loup city, NE, USA. This was just after a
summer spent working alone on a fire-lookout tower for the US Forest Service, in the wilds of Clearwater
National Forest, in Northern Idaho. They were married on 1 June 1958 and she became his wife of 54 years. She
later rewarded him with two sons, Alan Eugene and Tomas Charles, and they adopted a daughter, Bekianne.
Upon graduation, Chuck worked in flour milling research at the General Mills Central Research Laboratories
in the Minneapolis area from 1959 to 1962. He then accepted a fellowship to graduate school at North Dakota
State University in Fargo from 1962 to 1965, majoring in Cereal Chemistry. Chuck received his PhD in
Chemistry in the summer of 1966. For his PhD dissertation, he developed a micro brewery and studied barley
proteins and their influence on beer quality a rather ironic choice of topics, considering he remained a strict
teetotaler all his life.
While finishing his thesis, he joined the faculty of the North Dakota State College, Valley city, ND, USA
(19651974), teaching various physical science and chemistry courses to undergraduate students. Feeling the
pull of industry, he moved south to work for the Fairmont Foods Central Research Laboratory in Omaha, NE,
USA (19741980), where he did research, product development, and technical assistance on baking, pizza,
snack, and dairy products, eventually reaching the position of Associate Director of Research.
Dismemberment and sale of this Fortune-500 company provided him an opportunity to return to academia
at the University of Nebraska, Lincoln, NE, USA (19801987), where he developed a Cereal Technology
Program within the Department of Food Technology. He developed and taught various food science and
technology courses to both undergraduate and graduate students, and developed a vigorous research program.
He became Interim Department Head and Food Processing Center Director.
In December 1987, Chuck joined the Department of Grain Science and Industry at Kansas State University as
the Bakers National Educational Foundations Baking Science Professor. There he maintained a busy program
xix
xx Tributes to Former Co-Editors
in research, teaching, graduate student advising, publishing, traveling and consulting, specializing in baking
technology.
During his career, Chuck published more than 100 technical papers, and advised about 50 MS and PhD
graduate students. He was heavily involved in applied rheology (mixing and flow characteristics) of flour
products. In recognition of his commitment to international graduate students, Mr Roland Temme, TMCO,
Lincoln, NE, USA, established an international graduate student fellowship in his honor.
Chuck entered 50% phased-in retirement at age 65 in 2002 and retired in 2005, only to be hired back 1/
10-time afterward to assist with seminars and the departments centennial recognition in 2010. He continued
to support the departments activities on a volunteer basis.
Chucks research and consulting travels took him to more than a dozen countries, including many extended
trips to Australia, eventually earning permanent resident status in Australia a country which had fascinated
him since childhood. Most of his time in Australia was spent with the Bread Research Institute in Sydney. He
also made several extended trips to China, serving as guest lecturer at ShanDong Agricultural University, TaiAn,
and at Henan University of Technology, Zhengzhou.
Chuck always commented that the best legacy a person could leave is the knowledge he has shared. A
significant way in which his knowledge has been shared is represented by his contributions to the first and
second editions of Elseviers Encyclopedia of Food Grains.
In August 2012, Koushik Seetharaman was appointed as the third editor of this
second edition of Elseviers Grains Encyclopedia, replacing Professor Walker.
Tragically, Koushiks many contributions to the Encyclopedia of Food Grains were
cut short by his untimely death at 48 years in June 2014.
Koushik received his BS at the Gujarat Agricultural University, Anand, Guja-
rat, India. His MS training was at Cornell University, Ithaca, NY, USA, and his
PhD research was carried out at Texas A&M University, College Station, TX, USA.
At the time of Koushiks appointment as encyclopedia editor (August 2012),
he was Associate Professor and Cereals Chair in the Department of Food Science
at the University of Guelph, Canada. Research in his laboratory spanned the
cereal-grain value chain, focusing on grain quality, processing and ingredient
interactions, and on consumer acceptability and health.
His research on grain quality at Guelph involved collaboration with cereal-breeding companies, grain
farmers of Ontario and local processors, by analyzing Ontario wheats for functionality and nutritional
attributes. Special accents were on antioxidant activity, and on fiber and phenolics contents. Interactions
with breeders and processor related to the chemistry, sensory properties and consumer acceptance of whole
grain products, with specific focus on red versus white wheats in different product matrices.
Koushik had a special interest in the behavior of starch in water-poor systems, such as dough and baked
products. His research group had demonstrated that the constituent polymers of wheat starch are a series of
biopolymers ranging from linear to branched, depending on several factors including genetics and growth
environment. Moreover, the proportion of these polymers further defines their functional properties.
Research funding had been provided by the Canadian Agricultural Adaptation Council with contributions
from the Ontario Cereal Industry Research Council, Kellogg, Kraft, Kraft Canada Mill, Dow AgroSciences, C&M
Seeds, Grain Farmers of Ontario and Brabender GmbH.
Several months before his death, Koushik had been appointed as an Associate Professor to the Department
of Food Science and Nutrition at the University of Minnesota, St Paul, MN, USA, there taking the position of
General Mills Cereal Chair. In that capacity, Koushik had again initiated collaborations with breeders and
geneticists, millers and processors, with a focus on what product quality means for the consumer. His research
focused on interactions of grain biopolymers starch and gluten in particular and the impact on processing,
product attributes and consumer health.
Tributes to Former Co-Editors xxi
His group included three postdoctoral research associates, five doctoral students and one MS student. They
were collaborating extensively with research groups around the world including University of Milan, Italy;
University of Copenhagen, Denmark; Lille University, France; ONIRIS, France; Tamilnadu Agricultural Uni-
versity, India; Iowa State University, USA; African Rice Research Institute, Benin; University of Guelph, Canada;
and Memorial University, Newfoundland, Canada.
Koushiks research background and his collaborations placed him well to contribute to the preparation of
the grains encyclopedia, both by his our writing and via his contacts to engage world-authority contributors.
The grain-science community world-wide was shocked by Koushiks sudden passing. Koushik left a wife
Debra Freedman and an eight-year-old son, Samuel.
Koushiks contribution to the encyclopedia was such that Elsevier decided that his name should remain as a
co-editor even though he did not live to see the end result of his contributions.
Colin W. Wrigleys 55 years in grain-science research have earned him international recognition in
the form of several international and Australian research awards. His work is described in about
600 research publications, including several patents, a series of eight books on Australian cereal
varieties, and many edited books. He was Editor-in-Chief of the first edition of the Encyclopedia of
Grain Science (2004).
His research interests have centered on the characterization of cereal-grain proteins in relation to
processing quality. This has involved developing new methods of protein fractionation, including
gel isoelectric focusing and its 2D combination with gel electrophoresis, leading into proteomic
mapping. Other diagnostic methods developed relate to the evaluation of grain quality in wheat
and barley, such as better methods for variety identification and for characterizing quality in starch
and sprouted grain (as co-patentee of the Rapid ViscoAnalyser). Research involvement has also
included elucidation of grain-quality variation due to environmental factors (heat stress, fertilizer
use, CO2 levels and storage conditions).
In 2009, Wrigley was appointed a Member of the Order of Australia (AM) for service to primary
industry, particularly to grain science as a researcher, and to the development of methods for
improving wheat quality.
He is currently an Honorary Professor at the University of Queensland, Brisbane, Australia.
Harold Corke is a professor in the Food and Nutritional Sciences program at The University of
Hong Kong, and a Chutian Scholar Distinguished Foreign Professor in the Glyn O. Phillips
Hydrocolloid Research Center at HUT, Hubei University of Technology, Wuhan, China. In more
than 22 years in Hong Kong, he has had responsibility for teaching a diverse array of food science
courses, including the major courses Grain Production and Utilization, Food Safety and Quality
Management, Food Product Development, and general education courses Feeding the World, and
Food: Technology, Trade and Culture. He is author or co-author of 180 refereed journal articles,
and his 22 PhD graduates have gone on to successful careers in academia and industry around the
world. He is on the editorial boards of Journal of Food Composition and Analysis, LWT Food Science and
Technology and Journal of Cereal Science, and with Colin Wrigley and Chuck Walker was one of the
editors of the first edition of Encyclopedia of Grain Science, published in 2004. Corke is active in
consulting in grain processing and food safety in Asia and Eastern Europe.
Jon Faubion is the Charles Singleton Professor of Baking and Cereal Science in the Department of
Grain Science, Kansas State University. Over his 35 year career, he has had the sole or shared
responsibility for teaching nine different cereal or food science courses at Texas A&M University,
The University of Minnesota and Kansas State. In 2011, he received the Excellence in Teaching
Award from the American Association of Cereal Chemists International. He is a senior editor of
Cereal Chemistry, and a member of the Board of Directors of the American Association of Cereal
Chemists International and an executive editor of Journal of the Science of Food and Agriculture. Prior
to rejoining the department faculty in 2005, he directed the Applied Technology and Sensory
Science Groups for the research and development arm of The Schwan Food Company.
xxiii
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EDITORIAL ADVISORY BOARD
xxv
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CONTRIBUTORS
E Akhunov BJ Blaney
Kansas State University, Manhattan, KS, USA The University of Queensland, St Lucia, Australia
PC Annis JE Bock
CSIRO Entomology, Canberra, ACT, Australia University of Guelph, Guelph, ON, Canada
GA Annor F Bonomi
University of Ghana, Accra, Ghana University of Milan, Milan, Italy
CA Atkins AR Bridges
The University of Western Australia, Crawley, WA, AACC International, St Paul, MN, USA
Australia
H Brinch-Pedersen
W Atwell Aarhus University, Slagelse, Denmark
Bill Atwell Consulting, Champlin, MN, USA
G Bultosa
M Baga Haramaya University, Dire Dawa, Ethiopia; Botswana
University of Saskatchewan, Saskatoon, SK, Canada College of Agriculture, Gaborone, Botswana
JS Bao W Bushuk
Zhejiang University, Hangzhou, PR China University of Manitoba, Winnipeg, MB, Canada
VJ Barthet YZ Cai
Canadian Grain Commission, Winnipeg, MB, Canada China National Seed Group, Beijing, PR China;
SA Barton Huazhong Agricultural University, Wuhan, PR China;
Oxford Agricultural Trials Ltd, Oxford, UK The University of Hong Kong, Hong Kong, PR China;
Hubei University of Technology, Wuhan, PR China
IL Batey
Sunset Cereal Services, Meadowbank, NSW, Australia EF Caldwell
AACC International, Roseville, MN, USA
F Bekes
FBFD PTY LTD, Beecroft, NSW, Australia KJ Carpenter
University of California, Berkeley, CA, USA
AR Belcher
Oregon State University, Corvallis, OR, USA JC Catullo
INTA, Buenos Aires, Argentina
JN BeMiller
Purdue University, West Lafayette, IN, USA SP Cauvain
BakeTran, Witney, OX, UK
T Beta
University of Manitoba, Winnipeg, Manitoba, Canada S Ceccarelli
Via delle Begonie 2, Ascoli Piceno, Italy
JL Black
John L Black Consulting, Warrimoo, NSW, Australia GK Chandi
University of Guelph, Guelph, ON, Canada
MH Blanco
(Retired) Plant Breeder and Geneticist, Encinitas, N Chaudhary
CA, USA G. J. University of Science and Technology, Hisar, India
xxvii
xxviii Contributors
RN Chibbar JE Dexter
University of Saskatchewan, Saskatoon, SK, Canada Canadian Grain Commission, Winnipeg, MB, Canada
M Chisi B Dobraszczyk
Golden Valley Research Station, Chisamba, Zambia Reading Science Centre, Reading, UK
S Cloutier D Domingues
Agriculture and Agri-Food Canada, Ottawa, ON, General Mills Inc., Minneapolis, MN, USA
Canada
C Dowd
LS Collado General Mills Inc., Minneapolis, MN, USA
The University of the Philippines Los Banos, Laguna,
LM Duizer
Philippines
University of Guelph, Guelph, ON, Canada
NC Collins
RW Duncan
School of Agriculture Food and Wine, The University of
University of Manitoba, Winnipeg, MB, Canada
Adelaide, Glen Osmond, SA, Australia
KG Duodu
H Corke
University of Pretoria, Pretoria, South Africa
The University of Hong Kong, Hong Kong, Pokfulam,
PR China; Hubei University of Technology, Wuhan, SR Eckhoff
PR China (Retired) University of Illinois, Urbana, IL, USA
RL Cracknell M Edney
Crackers Consulting, Mount Eliza, VIC, Australia Canadian Grain Commission, Winnipeg, MB, Canada
GB Crosbie M Emery
Crosbie Grain Quality Consulting, East Fremantle, WA, Iowa State University, Ames, IA, USA
Australia
W Erskine
JS Croser University of Western Australia, Crawley, WA, Australia
University of Western Australia, Crawley, WA, Australia
WD Eustace
A Cruickshank (Retired) Kansas State University, Manhattan, KS, USA
DAFF Queensland, Warwick, QLD, Australia
A Evans
A Cuesta-Marcos Tate and Lyle, Hoffman Estates, IL, USA
Oregon State University, Corvallis, OR, USA
E Evans
MG DEgidio The Tassie Beer Dr, Lindisfarne, TAS, Australia
Unita di ricerca per la Valorizzazione Qualitativa dei
JD Faris
Cereali, Rome, Italy
USDA-ARS Cereal Crops Research Unit, Fargo, ND,
D Dan Ramdath USA
Agriculture and Agri-Food Canada, Guelph, ON, Canada
J Faubion
P Dangi Kansas State University, Manhattan, KS, USA
G. J. University of Science and Technology, Hisar, India
AM Fehily
JK Daun Tinuviel Software, Warrington, UK
(Deceased)
T Filichkin
C David Oregon State University, Corvallis, OR, USA
ISARA Lyon, Lyon, France
GB Fincher
L Day Australian Research Council Centre of Excellence in Plant
AgResearch Ltd., Palmerston North, New Zealand Cell Walls, University of Adelaide, Glen Osmond, SA,
Australia
D Deneke
South Dakota State University, Brookings, SD, USA SP Fisk
Oregon State University, Corvallis, OR, USA
R DePauw
Agriculture and AgriFood Canada, Swift Current, SK, MT Fletcher
Canada The University of Queensland, St Lucia, Australia
Contributors xxix
RJ Fletcher A Gunaratne
University of Queensland, Gatton, QLD, Australia The University of Hong Kong, Hong Kong, PR China
F Fleurat-Lessard YN Guragain
INRA, Mycology and Food Safety Research Unit, Kansas State University, Manhattan, KS, USA
Villenave dOrnon, France
RCE Guy
JA Fox Campden and Chorleywood Food Research Association,
Kansas State University, Manhattan, KS, USA Chipping Campden, UK
RJ French C Hall III
The University of Western Australia, Crawley, WA, North Dakota State University, Fargo, ND, USA
Australia
LM Hall
B Friebe University of Alberta, Edmonton, AB, Canada
Kansas State University, Manhattan, KS, USA
EG Hammond
BJ Furman Iowa State University, Ames, IA, USA
San Jose State University, San Jose, CA, USA
R Haraszi
M Gangola Campden BRI, Gloucestershire, UK
University of Saskatchewan, Saskatoon, SK, Canada
J Harnack
P Gelinas University of Minnesota, Minneapolis, MN, USA
Food Research and Development Centre, Agriculture and
D Hauge
Agri-Food Canada, Saint-Hyacinthe, QC, Canada
Grains for Health Foundation, St. Louis Park, MN, USA
SM Ghazani
AMR Hayes
University of Guelph, Guelph, ON, Canada
St. Catherine University, Lafayette, MN, USA
MC Gianibelli
PM Hayes
(Retired)
Oregon State University, Corvallis, OR, USA
A Gianinetti
L Helgerson
Consiglio per la ricerca in agricoltura e lanalisi
Oregon State University, Corvallis, OR, USA
delleconomia agraria, CRA-GPG, Fiorenzuola dArda,
Italy RJ Henry
University of Queensland, Brisbane, QLD, Australia
BS Gill
Kansas State University, Manhattan, KS, USA D Herb
Oregon State University, Corvallis, OR, USA
JS Godber
LSU Agricultural Center, Baton Rouge, LA, USA I Herencia
Universidad Politecnica de Madrid, Madrid, Spain
A Goldstein
University of Minnesota, St. Paul, MN, USA E Hernandez
Texas A&M University, College Station, TX, USA
R Graebner
Oregon State University, Corvallis, OR, USA KB Hobson
New South Wales Department of Primary Industries,
S Graham-Acquaah
Tamworth, NSW, Australia
Africa Rice Center, Cotonou, Republic of Benin
M Hochhalter
RA Graybosch
North Dakota State University, Fargo, ND, USA
USDA-ARS, University of Nebraska Lincoln, Lincoln,
NE, USA RD Horsley
North Dakota State University, Fargo, ND, USA
S Grundas
Bohdan Dobrzanski Institute of Agrophysics of the Polish W Huang
Academy of Sciences, Lublin, Poland Jiangsu Academy of Agricultural Sciences, Nanjing,
PR China; Jiangnan University, Wuxi, Jiangsu, PR China
TJ Gulya
USDA-ARS, Northern Crop Science Laboratory, Fargo, Y Huang
ND, USA China Agricultural University, Beijing, PR China
xxx Contributors
V Humphrey-Taylor P Koehler
(Retired) New Zealand Institute for Crop Food Research Deutsche Forschungsanstalt fur Lebensmittelchemie,
Limited, Christchurch, New Zealand Leibniz Institut, Freising, Germany
T Hymowitz R Korczak
University of Illinois at Champaign-Urbana, Urbana, University of Minnesota, St. Paul, MN, USA
IL, USA
SG Krattinger
S Iametti University of Zurich, Zurich, Switzerland
University of Milan, Milan, Italy
L Krishnamurthy
C Isaak International Crops Research Institute for the Semi-Arid
University of Manitoba, Winnipeg, MB, Canada Tropics, Patancheru, Andhra Pradesh, India
MS Izydorczyk S Kumar
Canadian Grain Commission, Winnipeg, MB, Canada International Center for Agricultural Research in the Dry
Areas (ICARDA), Rabat, Morocco
P Jacques
Tufts University, Boston, MA, USA OR Larroque
CSIRO Agriculture Flagship, Canberra, Australia
S Jaiswal
University of Saskatchewan, Saskatoon, SK, Canada JW Lawton
J Jane (Retired) ARS-USDA
Iowa State University, Ames, IA, USA EA Lee
LA Johnson University of Guelph, Guelph, ON, Canada
Iowa State University, Ames, IA, USA EC Leonard
JM Jones AgriKnowHow, Maitland, SA, Australia
St. Catherine University, Arden Hills, MN, USA JS Li
A Juhasz China Agricultural University, Beijing, PR China
Centre for Agricultural Research of the Hungarian WD Li
Academy of Sciences, Martonvasar, Hungary Shanxi University, Taiyuan, PR China
BO Juliano Z Li
Philippine Rice Research Institute Los Banos, Laguna, CSIRO Food Futures National Research Flagship,
Philippines Canberra, ACT, Australia; CSIRO Plant Industry,
RS Kadan Canberra, ACT, Australia
(Deceased)
R Lindberg
GP Kadkol Minneapolis Heart Institute Foundation, Minneapolis,
NSW Department of Primary Industries, Calala, NSW, MN, USA
Australia
MG Lindhauer
LW Kannenberg Max Rubner-Institut (MRI), Detmold, Germany
University of Guelph, Guelph, ON, Canada
K Liu
TN Khan Agricultural Research Service, US Department of
University of Western Australia, Crawley, WA, Australia Agriculture, Aberdeen, ID, USA
BS Khatkar Q Liu
G. J. University of Science and Technology, Hisar, India Agriculture and Agri-Food Canada, Guelph, ON, Canada
JG Kling S Lu
Oregon State University, Corvallis, OR, USA National Chung Hsing University, Taichung, Taiwan
EJ Knights OM Lukow
New South Wales Department of Primary Industries, Agriculture and Agri-Food Canada, Winnipeg, MB,
Tamworth, NSW, Australia Canada
Contributors xxxi
BL Ma NJ Mendham
Agriculture and Agri-Food Canada, Ottawa, ON, University of Tasmania, Hobart, TAS, Australia
Canada
R Menon
C-Y Ma The General Mills Bell Institute of Health and Nutrition,
The University of Hong Kong, Hong Kong, PR China Minneapolis, MN, USA
A MacLeod TE Michaels
Canadian Grain Commission, Winnipeg, MB, Canada University of Minnesota, St. Paul, MN, USA
L MacLeod DM Miskelly
Dairy Innovation Australia, Werribee, VIC, Australia Westcott Consultants P/L, Goulburn, NSW, Australia
CK Madsen
CS Moffat
Aarhus University, Slagelse, Denmark
Curtin University, Bentley, WA, Australia
RJ Mailer
ES Monyo
Australian Oils Research, Lambton, NSW, Australia
ICRISAT, Nairobi, Kenya
JT Manful
Africa Rice Center, Cotonou, Republic of Benin TR Moore
AIB International, Manhattan, KS, USA
AG Marangoni
University of Guelph, Guelph, ON, Canada MK Morell
International Rice Research Institute, Los Banos,
DJ Mares Philippines
University of Adelaide, Glen Osmond, SA, Australia
CF Morris
L Marquart USDA-ARS Western Wheat Quality Laboratory,
University of Minnesota, St. Paul, MN, USA; Grains for Pullman, WA, USA
Health Foundation, St. Louis Park, MN, USA
LA Morrison
A Marti
Oregon State University, Corvallis, OR, USA
Universita degli Studi di Milano, Milan, Italy
B Maschoff WK Mousa
Grains for Health Foundation, St. Louis Park, MN, USA University of Guelph, Guelph, ON, Canada; Mansoura
University, Mansoura, Egypt
A Mathiowetz
University of Minnesota, St. Paul, MN, USA; Rochester, K Mrva
MN, USA University of Adelaide, Glen Osmond, SA, Australia
JJ Mayko L Munck
Canola Council of Canada, Mundare, AB, Canada University of Copenhagen, Frederiksberg C, Denmark
CM McDonough R Munns
Texas A&M University, College Station, TX, USA CSIRO Agriculture, Canberra, ACT, Australia; The
University of Western Australia, Crawley, WA, Australia
RA McIntosh
The University of Sydney, Cobbitty, NSW, Australia PA Murphy
Iowa State University, Ames, IA, USA
JD McKeehen
Cereal Partners Worldwide, Minneapolis, MN, USA ED Nafziger
J McLean University of Illinois, Urbana, IL, USA
The University of Newcastle, Callaghan, NSW, Australia SC Nelson
PBE McVetty AACC International (AACCI), St. Paul, MN, USA
University of Manitoba, Winnipeg, MB, Canada T Nleya
B Meints South Dakota State University, Brookings, SD, USA
Oregon State University, Corvallis, OR, USA
L OBrien
A Meldrum The University of Sydney, Narrabri, NSW, Australia;
Pulse Australia, Perth, WA, Australia Solheimar Pty Ltd, Narrabri, NSW, Australia
xxxii Contributors
DK OToole I Rajcan
City University of Hong Kong, Hong Kong, PR China University of Guelph, Guelph, ON, Canada
EA Oelke L Ramsden
University of Minnesota, St. Paul, MN, USA University of Hong Kong, Hong Kong, PR China
RP Oliver KD Rausch
Curtin University, Bentley, WA, Australia University of Illinois at UrbanaChampaign, Urbana, IL,
USA
DE Ortiz
W. K. Kellogg Institute, Battle Creek, MI, USA A Regina
MA Pagani CSIRO Food Futures National Research Flagship,
Universita degli Studi di Milano, Milan, Italy Canberra, ACT, Australia; CSIRO Plant Industry,
Canberra, ACT, Australia
TS Palav
Rich Products Corporation, Buffalo, NY, USA MN Riaz
Texas A&M University, College Station, TX, USA
L Pallas
Yanco Agricultural Institute, Yanco, NSW, Australia D Richard-Molard
(Retired) INRA, Nantes, France
GH Palmer
Heriot-Watt University, Edinburgh, UK F Rizza
Consiglio per la ricerca in agricoltura e lanalisi
RG Palmer delleconomia agraria, CRA-GPG, Fiorenzuola dArda,
Iowa State University, Ames, IA, USA Italy
RF Park MJ Robertson
The University of Sydney, Narellan, NSW, Australia CSIRO Sustainable Ecosystems, St. Lucia, QLD,
GM Paulsen Australia
Kansas State University, Manhattan, KS, USA LW Rooney
LC Paulson Texas A&M University, College Station, TX, USA
The Bama Companies, Tulsa, OK, USA WL Rooney
DS Petterson Texas A&M University, College Station, TX, USA
Department of Agriculture, Nedlands, WA, Australia CM Rosell
RA Porter Institute of Agrochemistry and Food Technology (IATA-
University of Minnesota-NCROC, Grand Rapids, CSIC), Paterna, Valencia, Spain
MN, USA AS Ross
L Privalle Oregon State University, Corvallis, OR, USA
Bayer CropScience, Research Triangle Park, Durham, SJ Roy
NC, USA School of Agriculture Food and Wine, The University of
KV Probst Adelaide, Glen Osmond, SA, Australia
Kansas State University, Manhattan, KS, USA; Grain HD Sapirstein
Processing Corporation, Muscatine, IA, USA University of Manitoba, Winnipeg, MB, Canada
KJ Quail AK Sarkar
Grain Growers Limited, North Ryde, NSW, Australia Canadian International Grains Institute, Winnipeg, MB,
RCN Rachaputi Canada
University of Queensland, Kingaroy, QLD, Australia
A Sarker
H Rahman International Center for Agricultural Research in the Dry
University of Alberta, Edmonton, AB, Canada Areas (ICARDA), New Delhi, India
S Rahman SK Sathe
Monash University Malaysia, Bandar Sunway, Malaysia Florida State University, Tallahassee, FL, USA
MN Raizada SF Schakel
University of Guelph, Guelph, ON, Canada University of Minnesota, Minneapolis, MN, USA
Contributors xxxiii
KA Scherf VA Solah
Deutsche Forschungsanstalt fur Lebensmittelchemie, Curtin University, Perth, WA, Australia
Leibniz Institut, Freising, Germany
AM Stanca
MP Scott University of Modena and Reggio Emilia, Reggio Emilia,
Iowa State University, Ames, IA, USA Italy
K Seetharaman FL Stoddard
(Deceased) University of Helsinki, Helsinki, Finland
GJ Seiler K-C Tan
USDA-ARS, Northern Crop Science Laboratory, Fargo, Curtin University, Bentley, WA, Australia
ND, USA
AS Tatham
SO Serna-Saldivar Cardiff Metropolitan University, Cardiff, UK
Centro de Biotecnologa FEMSA, Escuela de Ingeniera y
JRN Taylor
Ciencias, Monterrey, Mexico
University of Pretoria, Hatfield, South Africa
S Shea Miller
V Terzi
Eastern Cereals and Oilseeds Research Centre,
Consiglio per la ricerca in agricoltura e lanalisi
Agriculture and Agri-Food Canada, Ottawa, ON,
delleconomia agraria, CRA-GPG, Fiorenzuola dArda,
Canada
Italy
DB Sheats
K Tiefenbacher
St. Catherine University, St. Paul, MN, USA
(Retired)
PR Shewry
VK Tiwari
Rothamsted Research, Harpenden, UK; University of
Kansas State University, Manhattan, KS, USA
Reading, Reading, UK
SM Tosh
RD Shillito
Agriculture and Agri-Food Canada, Guelph, ON, Canada
Bayer CropScience, Durham, NC, USA
P Tosi
JP Shroyer
University of Reading, Reading, UK
Kansas State University, Manhattan, KS, USA
LC Trugo
KJ Shroyer
Federal University of Rio de Janeiro, Rio de Janeiro, Brazil
Kansas State University, Manhattan, KS, USA
SE Ulrich
Qingyao Shu
Washington State University, Pullman, WA, USA
Zhejiang University, Hangzhou, PR China
EH Unger
KHM Siddique
Eagle Rock Solutions, Inc., Idaho Falls, ID, USA
University of Western Australia, Crawley, WA, Australia
PV Vadlani
J Singla
Kansas State University, Manhattan, KS, USA
University of Zurich, Zurich, Switzerland
SA Valencia-Chamorro
M Sissons
Escuela Politecnica Nacional, Quito, Ecuador
NSW Department of Primary Industries, Calala, NSW,
Australia; Tamworth Agricultural Institute, Tamworth, V Vamadevan
NSW, Australia University of Minnesota, St. Paul, MN, USA; Agriculture
and Agri-Food Canada, Guelph, ON, Canada
JH Skerritt
University of Queensland, St Lucia, QLD, Australia N Van Heel
University of Minnesota, Minneapolis, MN, USA
DJ Skylas
Grain Growers Limited, North Ryde, NSW, A Vandenberg
Australia University of Saskatchewan, Saskatoon, SK, Canada
DL Smith SK Vasal
McGill University, Ste-Anne-de-Bellevue, QC, Canada BISA, Punjab, India
xxxiv Contributors
Contents
The Grains that Feed the World
Course Structures: Based on EFG Articles
The Grain Crops: An Overview
The Grain Chain: The Route from Genes to Grain-Based Products
Taxonomic Classification of Grain Species
Grain: Morphology of Internal Structure
Grain and Plant Morphology of Cereals and how characters can be used to identify varieties
Grains are the source of half of our food and protein Grains are responsible directly or indirectly for most of the
consumption. food we eat. The big three cereal grains (rice, wheat, and
Current world production for all grains equals 1 kg per maize) have roles as staple foods (major daily sources of
person per day. carbohydrate-based energy) and for processing into animal
Grains also contribute to the wider range of foods via feeds and other diverse uses such as fuel bioethanol. There
animals. has been an explosive growth in meat consumption globally,
Major media claims about grains and health are presented as more and more people come out of poverty and enter into a
in this article. growing middle class. This is most apparent in China, where
Some myths about grain-based foods are busted. demand for meat and other animal products (eggs and milk)
Some claims are supported, at least in part. drives the production of maize domestically and production of
soy beans in major exporting countries (the United States,
Argentina, and Brazil).
A billion people worldwide are obese; a third of the food
Learning Objectives produced in industrialized countries is thrown away uneaten.
In this exuberant excess of food production lie many serious
To understand the significance of grains in feeding the problems a billion people worldwide are chronically mal-
growing human population nourished, food production competes with industry and
To adopt a critical attitude to production and nutritional domestic usage for scarce resources of energy and water,
information about grains presented in the media and to human populations continue to increase, and food production
know how to seek and research authentic information impacts on the environment in ways that may not be sustain-
To be motivated to pursue further topics in depth, starting able. People care about what they eat, about their health, about
with investigating relevant articles in the Encyclopedia of longevity, and about fitness.
Food Grains and going back where necessary to primary In the news media, nutrition sells; there is thus a constant
scientific research articles. output of information on topics such as gluten-free diets,
Today, we know that Jan did not control two important the wide range of elements described later in this article, espe-
factors: the sun and the air (CO2), which together produce all cially nitrogen for protein synthesis. When grain is harvested,
plant life via photosynthesis according to the following reactions: these elements are removed from the field, and they must
ultimately be replaced by some form of fertilizer.
CO2 H2 O light ! O2 glucose a simple sugar
However, the other essentials sunlight, CO2, and water are
Glucose ! carbohydrates, lignin, and cellulose readily available for free on an ongoing basis. Of course, that
plant components might not be claimed for water in many dry climates. The free
claim can certainly be made for CO2 given the rising levels of
Jan Baptist lived in an age of the Aristotelian belief in four this component of our atmosphere. In fact, the sequestration of
elements: water, earth, air, and fire. Even considering these CO2 by plants is one of the many solutions for mitigating the
elements, he took into account the first two, but neglected air rising levels of CO2. There needs to be a realization of the
(CO2) and fire (sunlight). essential nature of CO2 as the source of all plant life and thus
Thus, all our food comes from these sources sunlight, of all our food, thereby to reverse the bad press attributed to CO2.
CO2, and water either directly in the cases of grains,
vegetables, and fruit or indirectly in the cases of animals who
eat the grain and plants. In addition, the soil does contribute Will We Run Out of Food in the Foreseeable Future?
Our reliance on the grains of cereal plants (especially wheat,
rice, and corn) is so great that they supply nearly 60% of our
In 1648, a scientist food energy and 50% of the protein consumed on Earth. Thus,
provided water only grains really do stand between us and starvation. Grains offer
to a small tree the great advantages that they can be stored safely for long
... for 5 years periods and thus transported far from their growth sites. Grains
are adaptable with respect to the wide range of foods made
from them. Grains are also important as animal food and for
Thus, he concluded various forms of industrial processing.
that trees are made from But will we run out of food in the foreseeable future? Efforts
water! He neglected the continue to improve methods of breeding, agronomy, harve-
roles of light and CO2 sting, and processing, showing good signs that grain produc-
tion and quality will continue to increase, providing for a
world of hungry people. The production and yield of wheat
have tripled in the past 40 years, as explained in the Wheat
Overview article (The Cereal Grains: Wheat: An Overview of
Figure 2 The experiment of Belgian scientist, Jan Baptist van Helmont, the Grain That Provides Our Daily Bread). Furthermore,
seeking to determine what plants are made from. Figure 3 shows how there is an ongoing increase in the
2400 600
2200 400
2000 200
1800 0
2004/05 2006/07 2008/09 2010/11 2012/13 2014/15
fcast
Figure 3 The past decade of production and utilization of cereal grains. Source: FAOSTAT.
4 THE BASICS | The Grains that Feed the World
370 14.3
350
14.1
ppm 330
CO2 C
310 CO2 C 13.9
290
13.7
270
250 13.5
1000 1100 1200 1300 1400 1500 1600 1700 1800 1900 2000
Year
Figure 4 Increasing temperatures and levels of carbon dioxide promise increasing grain yields due to the carbon-fertilizer effect.
worldwide production of cereal grains, also including maize, stresses of over 35 C during grain filling. However, there are
rice, barley, and other grass-related grains. The future therefore indications that some wheat genotypes are available that
looks hopeful. provide stable grain quality despite such heat-stress condi-
However, we still have great inequalities in food supply in tions, thereby opening up promising research directions. So,
terms of grains. Although current world production for all considerable research effort is concentrated on breeding for
grains is equivalent to 1 kg per person per day, there is still changed growth conditions (see articles Agronomy of Grain
hunger because some of this great volume of grain serves alter- Growing: Implication of Climate Changes; Sustainable Grain
native important nonfood purposes, such as animal feed and Production and Utilization; Wheat: Biotrophic Pathogen Resis-
industrial uses. The greatest problem, however, is that the major tance; Genetics of Grains: Abiotic Stress Genes and Mecha-
sites of production are distant from the major sites of need, and nisms in Wheat; The Cereal Grains: An Overview of the Family
those in need lack the resources to access the excess grain. of Cereal Grains Prominent in World Agriculture).
Increasing levels of carbon dioxide (Figure 4) offer the
promise of increasing grain yields, but with growth conditions
changing, breeders and agronomists need to focus research on Growing Our Grains
adapting to a changing world scene.
Organic-grown grain is superior nutritionally to other
grain. Wrong!
Climate Change and the Future of Grain Production
GMO grain is hazardous to your health. Wrong!
Predictions of global climate change include increased levels of
carbon dioxide in the atmosphere, causing warmer tempera-
Organic Growing
tures (especially the daily minima), changed patterns of rain-
fall, and increased frequency and severity of heat-stress Organic is an adjective used with the whole range of foods and
episodes, together with decreased frost frequency and rising even fibers. The term is associated with a higher price and
sea levels. These trends imply a future of fluctuations in grain presumed higher value. It should also indicate products of
yield and quality, but overall increases in grain yield are pre- such quality and composition as are expected from growing
dicted. Rising sea levels have serious implications for many these products in the absence of chemicals, instead using nat-
low-lying rice-growing areas in Asia. ural means of farm management. Are organically grown grains,
Irrespective of arguments about causes, increases in atmo- for example, any different nutritionally from other sources of
spheric levels in carbon dioxide have been clearly demon- grain? The answer is generally no. Organic grain will look the
strated, rising from ancient levels of 150280 to 400 ppm same as conventionally grown grain and will not likely to have
and beyond, with a predicted doubling in a century or so. more favorable nutritional content.
One consequence is the provision of carbon fertilizer, leading Organic grain is likely to have a protein content similar to
in the future to more efficient biomass production and greater that of other grains grown under similar conditions. With
yields for many grain species. However, the yield increases may respect to chemicals, the levels of pesticides and herbicides of
be largely in carbohydrate, potentially leaving reduced grain- organic grain may be less than those for conventionally grown
protein levels. The fertilizing effects of carbon dioxide are grain, but the latter are still required to have such compounds
more significant at higher temperatures, the other major effect at levels below the minimum residue levels. A report by the UK
predicted to accompany the global climate challenge. For Crop Protection Association found lower levels of pesticide
wheat, the increases in temperature are also expected to affect residues in organic produce when compared to conventional
potential dough-forming properties. Temperatures in the range but these differences are relatively small, for example, a typical
1530 C during grain filling have been shown to provide residue of 0.1 mg/kg is equivalent to a fly on a ten-ton truck.
optimal dough strength, but grain with weaker dough quality The encyclopedias article on organic grain points out that
is produced when plants are subjected to a few days of heat the great advantage of organic farming is the benefit to land
THE BASICS | The Grains that Feed the World 5
use and the environment (see articles Agronomy of Grain Grow- on the actual composition of GM food and the actual effects
ing: Organic Grain Production and Food Processing; Sustainable of GM plant and animal agricultural practices (see articles
Grain Production and Utilization; Food Grains and the Con- Food Grains and the Consumer: Genetically Modified Grains
sumer: Labelling of Grain-Based Foods). and the Consumer; Genetics of Grains: Detection of Geneti-
cally Modified Grains; Development of Genetically Modified
Grains; The Cereal Grains: Maize: Overview; The Oilseeds:
Genetically Modified Organisms Cottonseed: Overview; Canola: Overview; Soybean:
It is well over a century since grain scientists commenced the Overview).
now usual practice of cross pollination to combine the benefits of
one parental line with those of another. Previously, plant
improvements had involved the selection of promising plants Grains, Our Health, and the Media
(by appearance) from the common mix of plants in the field.
Cross pollination greatly extended the genetic variability of popu- What can we believe? We all know that the French queen Marie
lations from which to select promising new genotypes for use in Antoinette (Figure 5) famously declared Let them eat cake
developing new improved varieties. However, this approach to when she was told that her starving subjects were clamoring for
plant breeding is limited because cross pollination is generally bread. Wrong!
restricted to plants of the same genus, perhaps even the same The saying is stuck in historical tradition because it has been
species. Thus, for example, it is not possible to source genetic a convenient precis of the aristocratic arrogance of the times
resistance to a disease from beyond the target grain species. preceding the French revolution (178999). However, the line
In recent decades, this limitation has been removed by the about eating cake was actually written in the mid-1760s when
introduction of genetic engineering techniques, permitting not Marie Antoinette was a young girl living in Austria. It appears in
only very precise inter-genus transfer of genes but also the target- a work entitled Confessions written by Jean-Jacques Rousseau.
ing of single genes or groups of genes for transfer from unrelated Furthermore, much has been lost in translation. The word
genera (which could, e.g., be as distant as a fish and a different cake is a mistranslation of the French brioche that is a pastry
plant). The improved plant or animal that carries functional with tender crumb made with generous inclusion of egg and
genes from an unrelated genus (a gene transfer that could not butter, hardly like our general concept of cake. The linkage to
happen in nature) is termed genetically modified (GM). Many Marie Antoinette has been attributed to Jean-Baptiste Alphonse
improved grain genotypes have been introduced into agricul- Karr (180890), who wrote it 50 years after her death.
tural production on a very large scale. Such improvements may How does it happen that society accepts the authenticity of
include not only agronomic and disease benefits but also a phrase or story even though it is just plain wrong? The Let
improvements in nutritional value. The major crops involved them eat cake story may be no more than a trivial error of
for large-scale production are maize, soybean, canola, and cot- history. More important are the multitude of other errors and
ton. In the case of GMO cotton especially, there has been great misconceptions in the public mind. To the extent that this
environmental benefit of reduced use of insecticides. There are misinformation affects our way of life and our personal well
GMO variations of many other grain crops, such as golden rice,
GM to contain beta-carotene, a source of vitamin A.
There has been political/philosophical/activist and con-
sumer resistance to the benefits offered by GMO technology,
such that the range of GMO grains has been restricted and the
potential benefits have not yet become available. For example,
there is thus no widespread cultivation of GMO wheat, barley,
or rice. Much of the scientific evidence presented on the poten-
tial harm of GM foods is simply spurious, generated with
disregard to normal experimental practices, and with results
predetermined by a bias on the part of the authors. In conse-
quence of consumer resistance, the spread of GMO grains has
been restricted to some countries and regions mainly to the
Americas. In Argentina, for example, many farmers grow two
crops a year wheat and GMO soy. As the soy is herbicide-
resistant, weeds are eliminated during the soy-growth phase,
and the subsequent wheat crop is harvested free of weed seeds.
Much of Europe and Africa has been far more circumspect
about permitting introduction of GM grains. There is a valid
and essential ongoing public debate to be made about the
ownership or intellectual property basis of genetic resources
that underpin GMO production. There is valid unease about
the potentially excess control that a few biotechnology compa-
nies may exert on world agriculture. There are also arguments
to be made about the environmental impacts both positive
and negative of all forms of agriculture, including that
involving GM plants and animals. But these arguments should Figure 5 The peasants of France were starving for lack of bread. Marie
be made on the basis of accurate and apolitical information Antoinette responded: No bread? Then let them eat cake. Did she?
6 THE BASICS | The Grains that Feed the World
Figure 6 This type of corn dolly is not made from corn (maize); they are
usually made from the stalks and heads of wheat, rye, triticale, and
The Encyclopedia of Food Grains: A Source of barley. This apparent misnomer arises from the use of corn to signify any
Authentic Information grain species; this connotation is still prevalent in parts of Britain,
whereas corn is used in North America to signify maize.
This encyclopedia offers authentic information about all
aspects of food grains as they relate to us and about all At least we know that milk and coffee are gluten-free. Not
sorts of grains: chickpeas, rice, maize, barley, quinoa, soya, necessarily.
plus buckwheat, and the real wheat (the last two grains are Corn dollies are made from corn (maize). No, not usually!
unrelated!). The encyclopedias coverage takes us through the (See Figure 6.)
various stages of the breeding of grains, their growth, produc- There are dietary imbalances in the cereal grains. Partly true!
tion, processing, and the retailing of the wide range of grain- Spinach is the best source of iron in our diet; we know that
based foods. All these issues impact us as consumers, situated because we grew up with Popeye the Sailor Man who ate all
as we are at the culmination of the grain chain (see article The his spinach. Wrong!
Basics: The Grain Chain: The Route from Genes to Grain-Based Wild flax growing in New Zealand differs from cultivated
Products). Veracity for the articles is assured due to authorship flax. Correct! (See Figure 7.)
by world authorities, followed by peer review by members of Organic-grown grain is superior nutritionally to other
the editorial advisory board, the editors, and external experts. grain. Wrong!
GMO grain is hazardous to your health. Wrong!
Correcting Misinformation
OTHERS
ABUNDANT ELEMENTS
1.5%
PHOSPHORUS RARE
SULFUR
1.0%
POTASSIUM
SODIUM
CHLORINE
CALCIUM
MAGNESIUM
1.5% VERY RARE
IRON
FLUORINE
NITROGEN SILICON
3.0% ZINC
STRONTIUM
RUBIDIUM
LEAD
HYDROGEN
MANGANESE
10% COPPER
ULTRA RARE
ALUMINUM
CARBON CADMIUM
TIN
18% BARIUM
MERCURY
SELENIUM
IODINE
OXYGEN
MOLYBDENUM
65% NICKEL
BORON
CHROMIUM
ARSENIC
COBALT
VANADIUM
= MINERALS
Figure 8 The proportions of the chemical elements that make up our bodies. Reproduced from Bunpei Y (2012) Wonderful Life with the Elements. San
Francisco, CA: No Starch Press.
badditives. All such ingredients must be approved by the sourced from fish; they are now becoming available in grains
respective government agencies (see articles Food Grains and via new genotypes of oilseeds such as canola (Figure 13) (see
the Consumer: Fortification of Grain-Based Foods; Wheat- articles Breeding of Grains: Canola/Rapeseed: Genetics and
Based Foods: Breads; Cookies, Biscuits and Crackers: Formu- Breeding; Fats: Healthy Fats and Oils; Lipid Chemistry; The
lation, Processing and Characteristics; Flatbreads of the World; Oilseeds: Canola: Overview).
Sour Dough Technology).
Ca K Fe YOGURT
Zn Mg Mn Ca Mo K Mg
BREAD
P P I Co Na
P K Fe
FRUITS
Na CI Ca
K Mg Cu Zn
Mn
P
BUTTER
Ca Mg Na
K Co P
Se Cu
BACON AND Ca Fe K
EGGS
Na P Zn
BACON
P Cl Mg Cu
Na K S EGGS
COFFEE
BLACK PEPPER Se Fe Ca P Cr
K
Cr K S Zn Co K
Figure 9 Sources of the chemical elements in breakfast. The elements carbon, oxygen, hydrogen, and nitrogen are not included because they are in all
these foods. Reproduced from Bunpei Y (2012) Wonderful Life with the Elements. San Francisco, CA: No Starch Press.
Bread, Cereal,
Rice & Pasta
Group
6-11
SERVINGS
Why the Gluten-Free Diet? feel better. However, beyond these cases, gluten-free diet has
become a dietary fad with many adherents. The inferior quality
People who are diagnosed with celiac disease must adhere to a
of much of the gluten-free bread available is a testament to the
gluten-free diet. Celiac disease is a disease affecting all ages; the
value of gluten in bread making. Nevertheless, there are many
only treatment, a gluten-free diet, is a lifelong commitment.
substitutes for the function of gluten in the production of
There are other conditions, less well characterized, for which
gluten-free foods (Figure 14).
benefit has been obtained by the omission of gluten from the
Strict adherence to a gluten-free is difficult given the use of
diet. In these cases, perhaps, a gluten-free diet will make you
wheat, rye, triticale, and barley in many processed foods. In the
process of avoiding gluten, it may be possible that some normal
nutrient sources are also avoided with the consequent risk of
missing out on some essential nutrients. But overall, it is unlikely
that eating gluten-free diets will be damaging to your health (see
articles Appendix 3: Grains, Foods, and Ingredients Suiting
sugars, fats
Acetyl-CoA
malate [cis-aconitate]
fumarate isocitrate
succinate [oxalosuccinate]
succinyl-coA ketoglutarate
Figure 12 The Krebs cycle via which fats and sugars are broken down to CO2 releasing energy that is stored in compounds such as adenosine
triphosphate.
THE BASICS | The Grains that Feed the World 11
Spinach?
In his comic strip, Popeyes superhuman strength was
attributed to the iron of his spinach diet, consumed by the
canful. There is a statue of Popeye in Crystal City, Texas,
commemorating his contribution to increasing the consump-
tion of spinach by 33% at about the time he was invented. In
the meat-scarce years of World War II, this was an encouraging
propaganda, but it was fraudulent. (According to Mould RF
Figure 14 The gluten-free symbol used to indicate foods with
(1984) Moulds Medical Anecdotes. Bristopl, UK: Adam Hilger
<20 ppm of gluten protein from wheat, rye, triticale, and barley. Ltd, p. 33.) The original researchers displaced the decimal
Buckwheat, however, is a misnomer; it is unrelated to wheat and it is point, giving a tenfold overestimate of the iron content for
satisfactory for people who must have a gluten-free diet (see article The spinach. In fact, spinach has about 30 mg of iron per kilogram.
Legumes and Pseudocereals: Buckwheat: Overview). The population would have done better to concentrate on
a grain-based diet, such as wheat germ with an iron content
Gluten-Free Diets for Celiac Disease; Food Grains: Intolerance, of about 100 mg kg1 (see articles Appendix 1: Nutrient-Com-
Allergy and Diseases: Celiac Disease; Cereal Allergens; The Glu- position Tables for Grains and for Grain-Based Products; Grain
ten-Free Diet, the Appendix on eating gluten-free). Composition and Analysis: The Composition of Food Grains
Even milk and coffee may not be gluten-free. Various sub- and Grain-Based Products, the Appendix of Grain
stitutes for dairy milk involve the use of grains. The most Composition).
obvious is soy milk, which is intrinsically gluten-free as the
soya bean is unrelated to wheat. In addition, rice, oats, and
other cereals have been used to make milk substitutes, having In Summary
the dietary advantage of being lactose-free. Rice milk is gluten-
free. Oats is a satisfactory grain for most people with celiac Given that grains comprise such a major part of our diet, it
disease, but intake of this borderline cereal should be limited; would be good to be able to assert that no food grains
the equivalent of less than one cup per day is recommended are poisonous. However, that is not completely true, because
in this encyclopedias appendix (see articles Appendix 3: there is the very unusual exception of the Calabar bean; see
Grains, Foods, and Ingredients Suiting Gluten-Free Diets for Figure 15. With that possible exception, it is certainly true that
Celiac Disease) on gluten-free diets for celiac disease. the wide family of food grains is responsible directly or indi-
There are various roasted-grain beverages marketed as a rectly for most of the food we eat.
caffeine-free alternatives to coffee. These coffee substitutes Given that current world production for all grains equals
may also be known as grain coffee or ersatz coffee. During about a kilogram of grain per person per day, how can the
World War II, roasted wheat and barley were used as a basis worlds nations work toward the elimination of malnutrition?
for coffee. Commercial products provide a powder form to be
reconstituted with hot water, for example, Nestle Caro.
Exercises for Revision
Grains and General Nutrition How is it possible to adopt a critical attitude to assessing
nutritional information about grains presented in the
There are dietary imbalances in the cereal grains. Partly true! media?
Spinach is the best source of iron in our diet, because we What procedures should be followed to seek and research
grew up with Popeye the Sailor Man who ate up all his authentic information?
spinach. Wrong! Select topics, covered in this article, that are relevant to
you personally. Describe what benefit (or otherwise)
that you have obtained from information provided in
Dietary Imbalances?
this article and from relevant articles in the Encyclopedia
The proteins of the major cereal grains can be criticized of Food Grains.
for having a balance of essential amino acids that does Describe how grain-based foods provide the major catego-
not fully conform to our dietary requirements. That is so. ries of nutrients needed in our nutrition. In which cases
On the other hand, the cereal grains provide an excellent might certain grain products be nutritionally deficient?
balance of proteins, carbohydrates, and fats. Some cereals Describe the categories of people who really need gluten-
are partly deficient in some of the essential amino acids, free foods. Who else can be expected to benefit from gluten-
particularly lysine, methionine, isoleucine, and tryptophan. free foods?
12 THE BASICS | The Grains that Feed the World
Further Reading
Harlan JR (1992) Crops and Man. Madison, WI: American Society of Agronomy.
Mangelsdorf PC (1966) Genetic potentials for increasing yields of food crops and
animals. Proceedings of the National Academy of Sciences 56: 370375.
Stoskopf NC (1985) Cereal Grain Crops. Reston, VA: Reston Publishing Company.
Topic Highlights students in big city Hong Kong are surprisingly ignorant about
the basics of cereal and other grain production and usage, even
Suggested guidelines are presented for the use of the Ency- though they live facing the giant agricultural expanses of main-
clopedia of Food Grains as a resource for teaching undergrad- land China. Many will be unclear on the basic formulation of
uate courses in food, nutrition, and agriculture. baked bread (guessing rice instead of wheat), and most will not
The Encyclopedia of Food Grains is a good starting source for know about yeast or other leavening agents, why their favorite
authoritative material for students preparing for further in- rice has the texture it does, or why bioethanol is not going to
depth research on a topic, essays, presentations, and other replace petroleum-based fuels.
project work. In addition, most students will not become professional food
technologists and will hence not be interested in an engineering
unit operations approach to cereal processing. I use the theme
Learning Objectives Food sufficiency and the global future to try to convince the
students that food grains are at the heart of human economic life,
To view Encyclopedia of Food Grains as a resource for under-
from bread and rice to sweeteners, meat (via animal feeding), soy
graduate and graduate coursework.
proteins, and many more. The emphasis in this course is on the
To understand how Encyclopedia of Food Grains chapters can
interactions among technology and markets in producing,
fit additional diverse teaching and learning needs, such as
processing, and distributing grain products.
for student research project-based work and for industry-
The basic principles are interesting for such students as they
focussed training.
come to realize at first how little they know about the products
they eat every day rice, bread, steamed bread, noodles, etc. I tell
Introduction them that I am worried that after taking a course like this, their
relatives (usually a know-it-all uncle) may ask them something
One of the special features of this new Encyclopedia of Food Grains like So you learned about grains at university, are instant noo-
is the effort made to maximize its usefulness as a teaching dles as bad as everyone says? Everybody listen now! The expert is
resource, particularly at the undergraduate level but also for going to speak. The student needs a core of factual knowledge to
graduate-level training and industry-specific technology courses. be able to answer questions like that. Another major part of the
Education in hands-on technology-based fields such as food course concerns the global-scale (with an emphasis on Asia)
science and nutrition is not all about providing suitable material patterns of economic growth and land utilization that determine
for the instructor to use in preparing lectures or for the student the production, import, and export of food. The student should
to read in preparation for examinations. Modern education is be able to think about issues such as If oil costs $40 a barrel, or
less concerned about the acquisition of specific knowledge and $200 a barrel, who will eat and who will starve? Can agriculture
more about the process of learning and inquiry, such that the (bioethanol and biodiesel) provide meaningful contributions to
student will be prepared for lifelong learning and innovation. fuel production? Where will the grains for meat production for
The knowledge content of a subject for example, the nutri- China come from? These types of question are addressed on a
tional potential of a grain may change, but a well-prepared regular basis in the media, at least in the business context. For
student will always be able to seek out new information and students who will go on to become bankers, or personal inves-
new ways to integrate and use the information, if they have a tors, or just consumers, the ability to read such media reports
sound understanding of basic concepts. with a higher level of insight can prove very useful.
An essential part of the course is the requirement for a
A Core Technology Course: Cereal Science/Grain hands-on lab-based group project. The theme will vary from
Science/Grain Processing year to year, but a typical instruction would be
Having taught this course for 20 years, I can explain in more Clearly, the students can use a resource such as Encyclopedia
depth the underlying concepts and ideas. In my experience, of Food Grains to determine how to make their product of
interest, to understand the likely effect of varying formulations, cereals, legumes, their components, and their processing to
and to seek out interesting nutritional problems and feasible foods. Clearly, the subject area coverage overlaps considerably
additives to make an interesting project. The use of small-scale with my course, described in detail earlier, and it would be up
processing equipment has the advantage of allowing students to the individual instructor for any such course to set out the
to make multiple samples of a product bread, steamed bread, precise learning objectives and to select and recommend par-
or noodles and with the repetition, they can remember better ticular sections of Encyclopedia of Food Grains that would be
the basics and develop the intuition to make intelligent adjust- most helpful for the student. At graduate level, even in-depth
ments to the process. By observing the work of their fellow study of particular areas of emphasis, such as genetics and
groups in the class, and by listening to the final project pre- breeding for quality, or international uses of grains, or frac-
sentations of all groups (in a formal half-day final symposium), tionation of grain, or health uses of grains, could easily be
they become well acquainted with a diversity of topics. supported by Encyclopedia of Food Grains.
Minimum expectations at the end of such a course would
be the ability to discuss freely, with good technical knowledge
and comparative insight, such issues as the following: Science Requirement or Reading-Intensive
Introductory Courses: Feeding the World
What are the differences among rice, wheat, maize, and
barley?
Most undergraduate degrees have a requirement for cross-
What climatic factors influence production of the major
disciplinary knowledge, which can be developed by, for exam-
cereal grains?
ple, requiring nonscience students to take science-based
What are the major food uses for each?
courses (Rocks for Jocks being the famous parody of an easy
How many fermented and distilled products are made from
geology course designed for athletes). Another typical require-
grains?
ment is for a first year seminar-style course that imparts a
To what extent are grains used for industrial products?
university-level way of thinking, of using sources, and of inten-
What limits substitution of one for the other?
sive research and reporting. In many cases, this may be the
What are the roles of cereal starch, oil, and protein?
students first experience of small-group discussion and of
What is the nutritional contribution of cereals to the
academic standards for writing and citation. In any case, such
human diet?
courses should be accessible to students with little science
How does meat production depend critically on grain
background yet challenging for those students who do have
production?
strong science training. Food and nutrition provide in many
Who will feed China?
ways ideal material for such courses. The material is intrinsi-
Suitable Encyclopedia of Food Grains coverage that would support cally interesting and accessible to students, and valuable
the teaching elements of this course could include the following: insights can be added from almost any perspective, from chem-
istry to economics, history, or anthropology.
Any overview articles such as those focussing on wheat, I run a freshman-level course as part of the breadth require-
maize, rice, and soy. ment for our degree programs called Feeding the World. In a
Material on grain production and consumption at the sense, this is a world agriculture course, where students learn
global and regional levels. about the key inputs in agriculture (e.g., energy, water,
For selected health topics, individual topics such as Functional chemicals, and knowledge) and examine the scale of world
Foods: Dietary Fiber, Prebiotics, Probiotics, and Synbiotics. production and its sustainability in face of constraints such as
Elements of starch structure and functionality and analysis excessive waste. The course description reads
of quality would help in later study of starch quality-based
products such as noodles. Protein chemistry of cereal grains Continuing human population increases, competition for water
is central to any study of cereal science. supplies, and concern about energy prices have led to profound
For processing wheat, an overview on wheat could be sup- pessimism about long-term food supplies. Already a billion people
go hungry every day. This course will offer an in-depth look at key
plemented with selected topics according to the instructors
issues in global food sufficiency, food production, food distribution,
and students interest, such as, on breads and on noodles. prospects and constraints. It is accessible to students from all fields
Understanding would be strengthened by concept material of study and academic backgrounds. You will develop an integrated
such as on sensory testing and noodle quality. technical, economic and political understanding of the global food
Similarly for rice, basic ideas on processing could be sup- supply crisis. You will be equipped to understand and appreciate
media reports related to this issue in your lives as informed and
ported according to student need along, for example, more
influential citizens. We consider the technological, political, eco-
in-depth genetic material or perhaps additional material on nomic and ethical dimensions of the global food supply enterprise.
eating quality or on the diverse uses of rice. The capacity of modern agriculture to produce food, and the capac-
For maize, it is important for students to understand at least ity of the political system to distribute food globally will be exam-
ined. The idea is to give an academically rigorous course that will
wet-milling uses and to have a general view on animal feed.
cross disciplinary boundaries and help develop your analytical skills
Traditionally, soy bean has not been considered part of and general understanding of an important topic. The course is open
grain in English language materials emphasizing cereal to students of all backgrounds, and hopefully this diversity can
grains. However, soy bean is classified with grain in Chi- enhance the classroom environment. The level of science required
nese, and it is key agricultural product for worldwide trade, is quite low, and none of the assessments (for example class test) will
particularly favor students with greater science background.
animal feed production, and processing into food ingredi-
ents. A student must know about soy processing, and about
The week-by-week topic coverage includes Global
the use of soy concentrates and isolates.
Production, Focus on China, The Green Revolution, Energy,
Kansas State University has a graduate-level course Cereal Sci- Water, Chemical Inputs, Genetic Modification, Biofuels,
ence, with the simple course description The characteristics of Waste, and Sustainability. Clearly, these topics can be aided
THE BASICS | Course Structures: Based on EFG Articles 15
by reference to selected chapters from Encyclopedia of Food Table 1 Core courses for the food science and industry major,
Grains. However, a major part of the course is the student Kansas State University
project, for which typical guidelines might be
FDSCI 101 Foundations in Food Science & Industry Credits: (1)
GENAG 200 Topics in Agriculture (03) College Careers (0)
Identify an important topic within the general theme of Who Will
FDSCI 302 Introduction to Food Science (3)
Feed Africa? [other themes could be Who Will Feed China?, or Who
FDSCI 305 Fundamentals of Food Processing (3)
Can Produce Rice?, or Can Chinese Money Feed the World?]. The
topic should be one that can be investigated using statistical data FDSCI 500 Food Science Seminar (1)
(such as production, import and export data for crops, livestock, or FDSCI 501 Food Chemistry (3)
chemicals; and/or data on human health and nutrition indicators). FDSCI 600 Food Microbiology (2)
Collect and analyze the data (preferably drawing as far as possible on FDSCI 601 Food Microbiology Lab (2)
FAOStat and OECD.Stat databases). Further research the topic using FDSCI 690 Principles of HACCP (2)
any other appropriate sources of information and prepare a written FDSCI 727 Chemical Methods of Food Analysis (2)
report in Powerpoint form. Be prepared to lead a 20 min presentation FDSCI 728 Physical Methods of Food Analysis (2)
GRSC 540 Engineering Applications to Grain/Food Products (3)
The student is then guided to pick a very narrow and GRSC 541 Engineering Applications to Grain/Food Products
specific topic for in-depth research, for example, the potential Laboratory (1)
for Chinese investment in rice production in Mozambique. In Select one
order to make any sense of this, they need to develop a base of ASI 318 Fundamentals of Nutrition (3)
agronomic knowledge, a sense of how transferable the breed- HN 132 Basic Nutrition (3)
Select one
ing or agronomic technologies may be, and an understanding
FDSCI 695 Quality Assurance of Food Products (3)
of rice import/export balances, quality requirements, and uti-
FDSCI 740 Research and Development of Food Products (4)
lization in different regions. The typical Internet search may
provide a lot of fragmented and often technically superficial
information. However, Internet sources for up-to-date infor-
mation, and articles from magazines such as the Economist on
the investment climate for agriculture, seem to fit well with the Develop the outline for a course in Food Product Develop-
solid agricultural, production, and processing information ment, and suggest a health-based theme for the students to
provided by Encyclopedia of Food Grains. follow in developing a team-based new product develop-
ment project. Show how the use of Encyclopedia of Food
The Diversity of Courses in a Major Grains could help in providing a starting point for project
planning.
I have described just two typical types of undergraduate course,
rooted in knowledge of grains and grain products. We can
examine the diversity of courses spanning a typical food
science major (Table 1), albeit one with a stronger-than- Exercises for Readers to Explore the Topic Further
average focus on grains. Literally, all of these courses could
potentially be drawing on Encyclopedia of Food Grains for course Use Encyclopedia of Food Grains to develop teaching mate-
material, whether in agriculture, processing, chemistry, micro- rials for your favorite course, and share the materials online
biology, analysis, engineering, or nutrition. with our website!
A similar listing of courses for a major in agriculture/agron-
omy or nutrition would also reveal substantial overlap of
course coverage with material that could be taken from Ency- See also: Barley, Rice and Maize Processing: Maize: Wet
clopedia of Food Grains. Milling; Rice Processing: Beyond the Farm Gate; Carbohydrates:
Resistant Starch and Health; Starch: Analysis of Quality; Food Grains
and Well-being: Functional Foods: Dietary Fibers, Prebiotics,
Industry Training Probiotics, and Synbiotics; Food-quality Testing: Noodles: Testing
for Quality; The Application of Sensory Science to the Evaluation of
Typical training for industry includes 23-day short courses
Grain-Based Foods; Grains Around the World: Grain Production
on fairly general topics, such as starch quality, Asian foods,
and Consumption: Overview; Non-food Products from Grains:
breakfast cereals, frozen doughs, and functional foods. These
Cereal Grains as Animal Feed; Oilseed and Legume Processing:
are often aimed at technical or sales staff from related areas
Soybean: Processing; Soybean: Soy Concentrates and Isolates;
moving into work on a new topic for the first time. There are
Proteins: The Protein Chemistry of Cereal Grains; The Cereal
also in-house training courses run by big companies to keep
Grains: Maize: Overview; Wheat: An Overview of the Grain That
their staff up-to-date with developments in, for example, grain
Provides Our Daily Bread; The Oilseeds: Soybean: Overview;
processing for healthy ingredients, quality testing, or new
Wheat-Based Foods: Breads; Cereals: Overview of Uses: Accent on
product development. The flexibility and diversity of materials
Wheat Grain; Noodles: Asian Wheat Flour Noodles.
in Encyclopedia of Food Grains provide a good starting point for
instructors building course materials for most such training.
Topic Highlights included species in orders, families, and, for the two most
important families, the Poaceae and the Fabaceae, into tribes.
Definition of a grain Listed crops may have relatives cultivated locally (see ars-grin.
Botanical classification of grain crops gov http://www.ars-grin.gov for additional information).
Utilization of grain crops Common names are the local, regional, or national names
Definition and classification of a pseudocereal for plants. With the multitude of spoken languages in exis-
Systematic listing of grain crop species and their botanical tence, common names for many crop species often are numer-
families ous. In scientific literature, common names for plants should
Classification of primitive grain crops be used only when the scientific name is presented at first
mention of a plant species. The use of common rather than
scientific names can lead to needless confusion. Some plant
Learning Objectives species are known, even within a single language, by a multi-
tude of common names. For example, the grain crop proso
millet (Panicum miliaceum L.) is also known by the English
To understand the definition of grain
common names common millet, French millet, hog millet,
To appreciate the evolutionary diversity of grains and struc-
broomcorn, and broomcorn millet. Also, some common
tures utilized as grains
names have been applied to totally different species. In the
To comprehend the difference between common and sci-
Americas, corn refers to Zea mays L., while in Europe, the term
entific names and understand the need for precision and
the need for scientific names in technical literature has been applied to barley (Hordeum vulgare L.) or to spelt
wheat (Triticum aestivum L. subsp. spelta (L.) Thell.). The use
To have knowledge of the difference between cereals and
of the scientific name would avoid the obvious confusion that
pseudocereals
might result from use of common names.
To appreciate the diversity of grain crops and their
Due to extensive morphological variation, numerous sub-
utilization
specific names have been assigned to some grain crops and
their relatives. Subspecies names are useful as a means of
differentiating various forms of cultivated crops, especially
Introduction when distinct market classes exist for each type. For example,
common or bread wheat is classified as T. aestivum subsp.
Plants considered grain crops are those producing small, hard, aestivum. Two morphologically distinct forms of T. aestivum
dry seed or fruit consumed by man or his domesticated animals L., club wheat (T. aestivum L. subsp. compactum (Host) Mackey)
as a foodstuff or processed for food or industrial purposes. and spelt (T. aestivum L. subsp. spelta (L.) Thell.) are grown and
Grain crops, as a grouping, is, however, largely artificial. marketed for different end uses. The use of the subspecies
Plants producing useful grains have evolved in a number of names, in such cases, is useful as it differentiates the various
plant families, and these families are not always closely related. cultivated forms. Subspecies names also are useful to differen-
Botanically speaking, grains themselves are heterogeneous. The tiate cultivated forms of plant species from wild ones. For
term was derived from the Latin granum meaning seed. Cereal example, durum wheat (T. turgidum L. subsp. durum) has sev-
grains, however, actually are formed of an entire fruit (caryop- eral wild, semiwild, and formerly cultivated relatives including
sis), while those of other grain crops are the dry seeds of various T. turgidum subsp. dicoccoides and T. turgidum subsp. dicoccon.
types of fruits including legumes (pulses), achenes (sunflowers In both examples, the use of the subspecies names serves to
and buckwheats), siliques (canola), and capsules (cotton). differentiate the morphologically different forms while retain-
Grains, therefore, are the result of convergent evolution or the ing an indication of the close genetic relationships between the
development of similar structures (grains) in diverse organ- various types. Such information is useful as an indication of
isms. The actual genes involved in the formation of these the existence of close relatives of grain crops for use as gene
structures might be different in each family of grain crops. donors in crop improvement programs. It also is useful as an
indicator to people with dietary restrictions (e.g., those
afflicted with celiac disease) of the true genetic relationship
The Grain Crops between grain crops.
Plants classified as grain crops are found in both classes of
Grain crops of significance are listed in Table 1 according to the division Anthophyta (Table 1) and include representatives
their common English names and the equivalent botanical of 10 orders and 11 families. The families with the largest
names (as genus and species). When such lists are formulated, number of members are the Poaceae (Gramineae), also
it is valuable to arrange species in a systematic or taxonomic known as the grass or cereal family, and the Fabaceae (Legu-
fashion. This has been accomplished in Table 1 by grouping minosae), the legume or bean family. These two families
Table 1 The grain crops, their family groupings, and botanical names (genus and species), plus common names
Class: Liliopsida
Order: Cyperales
Family: Poaceae (Gramineae)
Subfamily: Pooideae
Tribe Poeae Avena Avena abyssinica Hochst. in Schimper Abyssinian oat
Avena brevis Roth Short oat
Avena nuda L. Naked oat
Avena sativa L. Oats, common oat
Avena strigosa Schreb. Black oat, small oat
Phalaris Phalaris canariensis L. Canary grass
Tribe Triticeae Hordeum Hordeum vulgare L. Barley
Secale Secale cereale L. Rye
Secale derzhavinii Tzvelev Perennial rye
X Triticosecale Triticosecale Wittm. Triticale
Triticum Triticum aestivum L. subsp. aestivum Common wheat, bread wheat
Triticum aestivum L. subsp. compactum (Host) Club wheat
Mackey
Triticum aestivum L. subsp. spelta (L.) Thell. Spelt, farro
Triticum monococcum L. subsp. monococcum Small spelt, einkorn, farro
Triticum timopheevii (Zhuk.) Zhuk. subsp. Zanduri wheat, Georgian wheat
timopheevii
Triticum turgidum L. subsp. durum (Desf.) Husn. Durum, durum wheat
Triticum turgidum L. subsp. carthlicum (Nevski) Persian wheat
A. & D. Love
Triticum turgidum L. subsp. dicoccon (Schrank) Emmer
Thell.
Triticum turgidum L. subsp. polonicum (L.) Thell. Polish wheat
Triticum turgidum L. subsp. turanicum (Jakubc.) Khorasan wheat
A. & D. Love
Triticum turgidum L. subsp. turgidum Cone wheat, pollard wheat, rivet wheat
Tribe Stipeae Achnatherum Achnatherum hymenoides (Roem. & Schult.) Indian ricegrass
Barkworth
Subfamily: Panicoideae
Coix Coix lacryma-jobi L. Coix, Jobs tears
Zea Zea mays L. Maize, Indian corn, corn
Tribe Andropogoneae Sorghum Sorghum bicolor (L.) Moench Sorghum
Tribe Paniceae Pennisetum Pennisetum glaucum (L.) R. Br. Pearl millet
Panicum Panicum miliaceum L. Proso millet, common millet, French millet,
hog millet, broomcorn millet
Panicum hiritcaule J. Presl Sowi millet, sauwi
Panicum sumatrense Roth Little millet, blue panic
Setaria Setaria italica (L.) P. Beauv. Foxtail millet, Hungarian millet, Italian millet
Echinochloa Echinochloa esculenta (A. Braun) H. Scholz Japanese millet, Japanese barnyard millet
Echinochloa frumentacea Link Sawa millet, sawa, billion dollar grass
Paspalum Paspalum scrobiculatum L. Kodo millet
Digitaria Digitaria exilis (Kipp.) Stapf Fonio, white fonio, white fonio millet
Digitaria iburua Stapf Black fonio
Urochloa Urochloa deflexa (Schumach.) H. Scholz Guinea millet, animal fonio
Subfamily: Chloridoideae
Tribe Chlorideae Eleusine Eleusine coracana (L.) Gaertn. Finger millet, birdsfoot
Tribe Eragrosteae Eragrostis Eragrostis tef (Zuccagni) Trotter Teff
Subfamily: Bambusoideae
Tribe Oryzeae Oryza Oryza sativa L. Rice
Oryza glaberrima Steud. African rice
Zizania Zizania palustris L. Northern wild rice
Zizania aquatica L. Wild rice, zizania
Class: Magnoliopsida
Order: Fabales
Family: Fabaceae (Leguminosae)
Tribe Dalbergieae Arachis Arachis hypogaea L. subsp. hypogaea Virginia peanut
Arachis hypogaea L. subsp. fastigiata Waldr. Peanut, Spanish or Valencia types
Tribe Cicereae Cicer Cicer arietinum L. Chickpea
(Continued)
18 THE BASICS | The Grain Crops: An Overview
Table 1 (Continued)
contain over about 70% of the species considered grain crops. (Khorasan wheat), and T. turgidum L. subsp. turgidum (cone,
Five species alone, including four grasses, wheat (T. aestivum pollard, or rivet wheat). Other important members of the tribe
and T. turgidum subsp. durum), maize (Z. mays L.), rice (Oryza Triticeae include barley (H. vulgare L.) and rye (Secale cereale L.).
sativa L.), and barley (H. vulgare L.), and the legume soybean Triticale (X Triticosecale Wittm.) is a man-made crop, grown
(Glycine max (L.) Merr.) also provide over 70% of the worlds primarily not only as forage but also as a grain in some parts of
metric tonnage of food from annual plants. the world. Triticales were developed via hybridizations between
common or durum wheats and rye. Octoploid triticales contain
all the genes of hexaploid wheat and rye, while hexaploid triti-
The Grasses cales contain the genes of tetraploid durum wheat and rye.
Secale derzhavinii Tzvelev, another man-made crop, has been
The grass family (Poaceae) contains the largest and, in terms of developed as a perennial grain crop. Its cultivation is limited.
annual production, most important grain crops, the cereals. The oats, Avena, are more distantly related, being placed
The Poaceae is one of the largest plant families, commonly along with the wheats in the subfamily Pooideae, but in a
divided into subfamilies, each, in turn, containing one or more different tribe, the Poeae. Common oat, Avena sativa L., is the
tribes. Cereal grains (Table 1) are found in the subfamilies most commonly cultivated species. Its importance has dimin-
Pooideae, Bambusoideae, Chloridoideae, and Panicoideae, ished in historical times with the change from animal-driven to
with the Pooideae and Panicoideae containing the largest mechanized agriculture, but oats are still an important grain
numbers of grain crop species. Early systems of grass taxonomy crop for human consumption, especially in the northern hemi-
relied heavily on morphological features, especially spikelet sphere. Other species of Avena have been grown as grain crops,
and inflorescence morphology. However, beginning in the but they either are no longer cultivated or have been reduced to
1950s, information regarding chromosome numbers, embryo the status of relic crops. These included A. abyssinica Hochst. in
structure, leaf anatomy, interspecific hybridizations, and types Schimper (Abyssinian oat), A. brevis Roth (short oat), A. nuda
of photosynthetic systems began to accumulate, and it became L. (naked oat), and A. strigosa Schreb. (black or small oat).
clear that the early arrangement of grass genera into subfam- Phalaris canariensis L., another member of the tribe Poeae,
ilies and tribes was largely artificial. In the 1960s, grass taxon- primarily is grown as a source of feed for caged birds.
omy was completely revised. Many earlier volumes still in wide
use (e.g., Hitchcock and Chases Manual of the Grasses of the Maize, sorghum, and Jobs tears
United States) reflect the original groupings of grass genera. Maize (Z. mays L.) and sorghum (S. bicolor (L.) Moench.) are
More recent references (e.g., Watson and Dallwitzs Grass both members of the subfamily Andropogoneae. Jobs tears
Genera of the World http://delta-intkey.com/grass/), followed (Coix lacryma-jobi L.), rarely cultivated as a grain, but more
herein, should be consulted for proper tribal affiliation of grass frequent now as an ornamental crop, also is a member of this
genera. tribe. Many morphological types of both maize and sorghum
Wheat and other members of the tribe Triticeae (subfamily exist; in both cases, however, only a single species and subspe-
Pooideae) are among the most important grain crops. Com- cies are recognized.
mon or bread wheat, T. aestivum L. subsp. aestivum, and durum
wheat, T. turgidum (L.) subsp. durum (Desf.) Husn., are the two The millets and teff
most widely cultivated. Two additional subspecies of T. aesti- The millets are a taxonomically confusing group. The term
vum, T. aestivum L. subsp. compactum (Host) Mackey (club millet evidently has been applied to any grass with a small
wheat) and T. aestivum L. subsp. spelta (L.) Thell. (spelt or round seed. Thus, the term is used to describe a number of, at
farro), also are cultivated. Club wheats are used to produce times, only distantly related species. Most of the millets are
pastry and other low protein flours, and spelt is grown largely placed in the subfamily Panicoideae, tribe Paniceae. The most
as an heirloom crop, often in organic production systems. important millet in terms of annual production is pearl millet
Some authors consider these to be separate species, but they (Pennisetum glaucum (L.) R.Br.), most commonly grown in sub-
both differ from typical bread wheat only in some aspects of Saharan Africa and in India. The remaining millets often are
spike morphology controlled by one or a few genes, and the termed minor millets. Of these, proso millet (P. miliaceum L.)
forms will all freely hybridize, so recognition as separate spe- is the most widespread, grown as a grain crop throughout Asia
cies is unwarranted. T. monococcum L. subsp. monococcum and into Eastern Europe. In North America, it is largely grown
(small spelt, einkorn, or farro) was once grown as a grain as a feed for caged and domestic birds. Several additional
crop in most states bordering the Mediterranean Sea and millets are grown, primarily in Asia. The following also are
throughout Europe. T. timopheevii (Zhuk.) Zhuk. (Georgian members of the tribe Paniceae: Setaria italica (L.) P. Beauv.
or zanduri wheat) was cultivated in what is now the nation (foxtail millet), Echinochloa esculenta (A. Braun) H. Scholz
of Georgia. Both T. monococcum and T. timopheevii have all but (Japanese millet), E. frumentacea Link. (sawa millet), Paspalum
disappeared from modern cultivation. Several additional sub- scrobiculatum L. (kodo millet), Digitaria exilis (Kipp.) Stapf
species of T. turgidum have been cultivated in historical times, (fonio or white fonio), D. iburua Stapf (black fonio), and
but either their importance has diminished, or they are culti- Urochloa deflexa (Schumach.) H. Scholz (Guinea millet). Eleu-
vated in local, isolated areas as relic crops. These include T. sine coracana (L.) Gaertn, known as finger millet or birdsfoot,
turgidum L. subsp. carthlicum (Nevski) A. & D. Love (Persian actually is a member of the subfamily Chloridoideae, tribe
wheat), T. turgidum L. subsp. dicoccon (Schrank) Thell. Chlorideae, indicating that it is more closely related to teff
(emmer), T. turgidum L. subsp. polonicum (L.) Thell. (Polish than it is to the rest of the millets. Teff (Eragrostis tef (Zuccagni)
wheat), T. turgidum L. subsp. turanicum (Jakubc.) A. & D. Love Trotter), subfamily Chloridoideae, tribe Eragrosteae, is
20 THE BASICS | The Grain Crops: An Overview
cultivated as a grain crop in Ethiopia and as a forage grass include P. sativum L. subsp. asiaticum Govor. (Asiatic pea),
elsewhere. It is the only cultivated grain species from a fairly cultivated in Asia, and P. sativum L. subsp. transcaucasicum
large genus of grasses. Govor., cultivated in the Caucasus mountain region. Legumes
of lesser importance include Cyamopsis tetragonoloba (L.) Taub.
Rice and its relatives (L.) Taub. in Engl. & Prantl (guar), cultivated at times as a grain
Rice (O. sativa L., tribe Oryzeae) is the major food of nearly but now more important as a source of guar gum. Other less
50% of the worlds population. Two forms or races of rice are common grain legumes are three species of the genus Lupinus,
recognized, indica types (race indica), grown in the tropics, L. luteus L. (yellow lupin), L. angustifolius (blue lupin), and L.
and japonica types (races japonica and javonica), the temperate albus (white lupin). The lupins also are grown as ornamental
forms. These types are not recognized as subspecies. African plants. Widely cultivated, though of only local importance as
rice (O. glaberrima Steud.) long has been cultivated in Western grain crops, are two species of the genus Canavalia (jack bean
and Central Africa, but it has largely been replaced by O. sativa. and sword bean), Lablab purpureus (L.) Sweet (hyacinth bean),
Zizania palustris L., northern wild rice, and Z. aquatica L. also Lathyrus sativus L. (chickling pea), Macrotyloma geocarpum
members of the tribe Oryzeae, are cultivated and gathered from (Harms) Marechal & Baudet (Kerstings groundnut), Psophocar-
wild populations in North America. Oryza and Zizania are the pus tetragonolobus (Stickm.) DC. (winged bean), Vicia ervilia
only grain crops assigned to the subfamily Bambusoideae. (L.) Willd. (bitter vetch), Mucuna pruriens (L.) DC. (velvet
Indian ricegrass (Achnatherum hymenoides (Roem. & Schult.) bean), and several species of the genus Vigna, V. angularis
Barkworth), formerly was gathered as a grain crop by Native (Willd.) Ohwi & Ohashi (adzuki bean), V. aconitifolia (Jacq.)
Americans, and there have been some recent attempts to culti- Marechal (moth bean), V. radiata (L.) R. Wilczek (mung bean),
vate and market it. Indian ricegrass is so named for its super- V. mungo (L.) Hepper (black gram), V. subterranea (L.) Verdc.
ficial morphological resemblance to rice, but it actually is a (Bambara groundnut), V. umbellata (Thunb.) Ohwi & Ohashi
member of the subfamily Pooideae. (rice bean), and V. unguiculata (L.) Walp. (black-eyed pea or
cowpea).
The Legumes
Oilseed Crops and Pseudocereals
The Fabaceae (legume or bean family) is second only to the
Poaceae in terms of economic importance to man. This large Several crops of various families produce grains from which
family is divided into three subfamilies, the Papilionoideae, edible or industrial oils are processed. The mustard family
Caesalpinioideae, and Mimosoideae. Some authors treat these (Brassicaceae or Cruciferae) contains several oilseed species,
as separate families. All of the grain legumes are members of most important of which is Brassica napus L. (rapeseed or
the Papilionoideae. Most grain legumes are members of the canola). B. nigra (L.) Koch in Rohling (black mustard) and B.
tribes Phaseoleae and Vicieae, although some also are assigned juncea (L.) Czern. (brown mustard) occasionally are cultivated
to the tribes Dalbergieae, Cicereae, Genisteae, and Indigofereae for oil and to produce mustard. Crambe hispanica L. subsp.
(Table 1). abyssinica (Hochst. ex R. E. Fr.) Prina (crambe) is another
Soybean, or soya (Glycine max L.), is the most important, in member of the mustard family used as a source of oil for
terms of annual metric tonnage of production. G. soja Siebold various industrial purposes. One member of the family Peda-
& Zucc., the wild progenitor of soybean, is a close relative, and liaceae, Sesamum indicum L. (sesame), is cultivated as both a
the two are, at times, treated as one species. Intermediate types spice and a source of sesame oil. Ricinus communis L. (castor
exist and formerly were designated G. gracilis Skvortzov, but bean), of the family Euphorbiaceae, is the source of castor oil,
they now are recognized as representatives of the subspecies G. used in industrial applications. It is also infamous as the source
max subsp. gracilis (Skvortzov) Enken. of the toxin ricin, a seed protein. Linseed oil is derived from
Edible dry beans are assigned to the species Phaseolus vul- Linum usitatissimum L., a member of the Linaceae. L. usitatissi-
garis L. A large number of morphological variants exist for this mum also is the source of flax. Chia (Salvia potus Epling) a
species, some being consumed as pulses, others for the edible member of the mint family (Lamiaceae) is used as a source of
pods. Common names include dry edible beans, kidney bean, edible oil and consumed directly in various ways including a
navy bean, wax bean, green bean, and field bean. All are topping for yogurt and as an ingredient for breakfast cereals.
considered members of the same species. Other species of Chia seed also is used to produce chia pets.
Phaseolus are cultivated, including the tepary bean, P. acutifolius The Compositae (Asteraceae), or sunflower family, proba-
A. Gray, P. coccineus L. (scarlet runner bean), and P. lunatus L. bly is the worlds largest family, in terms of number of species.
(butter bean or lima bean). However, it contains only three members cultivated as grain
Arachis hypogaea L. (peanut) is widely cultivated in both the crops, Helianthus annuus L. (sunflower), Guizotia abyssinica (L.
Old and New Worlds. Two subspecies are grown, A. hypogaea L. f.) Cass. (Niger thistle), and Carthamus tinctorius L. (safflower).
subsp. hypogaea (Virginia peanut) and A. hypogaea L. subsp. Sunflower and safflower primarily are grown as oilseed crops,
fastigiata Waldr. (Spanish or Valencia peanut). Other impor- but sunflower also is used as a grain for human and avian
tant grain legumes include Vicia faba L. (broad bean or fava), consumers. Niger thistle also is grown for birdseed.
Cajanus cajan (L.) Millsp. (pigeon pea), Cicer arietinum L. Cotton species (family Malvaceae, genus Gossypium) inde-
(chickpea), Lens culinaris Medik. (lentil), and Pisum sativum L. pendently were domesticated in both the Old and New
(field or sweet pea). Several subspecies of P. sativum are culti- Worlds. G. herbaceum L. (Arabian or Egyptian cotton) was of
vated, with P. sativum L. subsp. sativum, the common garden Old World origin, while G. hirsutum L. (American upland
pea, being the predominant form. Other less common forms cotton) and G. barbadense L. (American Pima cotton) and
THE BASICS | The Grain Crops: An Overview 21
G. arboreum L. (tree cotton) were domesticated in the New Soybean: Germplasm, Breeding, and Genetics; Wheat Breeding:
World. Over 90% of the worlds cotton production now is Exploiting and Fixing Genetic Variation by Selection and Evaluation;
from G. hirsutum L. While primarily grown as a fiber crop, Genetics of Grains: Maize: Genetics; The Basics: Taxonomic
both oil and protein are processed from cotton seeds. Classification of Grain Species; The Cereal Grains: An Overview of
Pseudocereals are nonmembers of the grass family often the Family of Cereal Grains Prominent in World Agriculture; Cereals:
milled to flour and used to produce flat breads and other Domestication of the Cereal Grains; Rice: Overview; The Legumes
products similar to those derived from true cereals. This is and Pseudocereals: Amaranth: Overview; Beans: Overview;
another artificial grouping, containing members of several Buckwheat: Overview; Chickpea: Overview; Grain Legumes and Their
plant families. Two members of the Chenopodiaceae are culti- Dietary Impact: Overview; Lupin: Overview; Pea: Overview;
vated, Chenopodium quinoa Willd. (quinoa) and C. pallidicaule Pseudocereals: Overview; The Oilseeds: Oilseeds: Overview.
Aellen (canihua). The grain amaranths, Amaranthus hypochon-
driacus L., A. caudatus L., and A. cruentus L., are assigned to the
Amaranthaceae. These five species all were components of the
ancestral cultivated flora of Mesoamerica and South America. Further Reading
Additional pseudocereals include Fagopyrum esculentum Belton PS and Taylor JRN (2002) Pseudocereals and Less Common Cereals: Grain
Moench. (buckwheat) and F. tataricum (L.) Gaertn. (Tartary Properties and Utilization Potential. Berlin: Springer.
buckwheat), both members of the Polygonaceae. Buckwheats Brummitt RK and Powell CE (eds.) (1992) Authors of Plant Names. Kew, UK: Royal
probably were first cultivated in Western China. Botanical Gardens.
Cronquist A (1981) An Integrated System of Classification of Flowering Plants.
New York: Columbia University Press.
Cronquist A (1988) Evolution and Classification of Flowering Plants. New York:
Exercises for Revision New York Botanical Gardens.
Duke JA (1981) Handbook of Legumes of World Economic Importance. New York:
What is the difference between a cereal and a pseudocereal? Plenum Press.
Gould FW (1968) Grass Systematics. New York: McGraw Hill.
Explain the difference between a scientific and a common Hanelt P (ed.) (2001) Mansfelds Encyclopedia of Agricultural and Horticultural Crops.
name? When should scientific names be used? What is the Heidelberg: Springer.
point of using names that are difficult to spell and Harlan JR (1992) Crops and Man, 2nd edn. Madison, WI: American Society of
remember? Agronomy.
Hitchcock AS (1971) Manual of the Grasses of the United States, 2nd edn. New York:
What is the difference between a species and a subspecies?
Dover, revised by A. Chase.
What is a millet? Are all millets derived from the same Lawrence GHM (1951) Taxonomy of Vascular Plants. New York: Macmillan.
species? Mabberley DJ (1997) The Plant-Book. Cambridge: Cambridge University Press.
Which are more closely related wheat and oats or wheat Pohlhill RM and Raven PH (1981) Advances in Legume Systematics. Kew, UK: Royal
and rice? Why? Botanic Gardens.
Smartt J (1990) Grain Legumes. Evolution and Genetic Resources. Cambridge:
Why do you suppose the majority of our grain crops derive Cambridge University Press.
from only two plant families? Vavilov NI (1960) World Resources of Cereals, Leguminous Seed Crops and Flax and
Their Utilization in Plant Breeding. Jerusalem: Israel Program for Scientific
Translations.
Wiersema JH and Blanca L (1999) World Economic Plants: A Standard. Boca Raton, FL:
Exercises for Readers to Explore the Topic Further CRC Press.
Topic Highlights On the other hand, the whole sequence may occur in one
village for a family on subsistence farming, with some grains
The value-addition chain for grain starts with the breeder, being cooked and consumed as is by the family and other
who builds suitable genes into a specific variety so that it grains being processed in a village-owned mill. In place of the
will provide a good yield of quality grain. marketing stage of the chain in Figure 1, there may be the
Farm management is designed to optimize yield and grain interchange of various types of food or labor. Nevertheless,
quality, although some aspects of the growth environment even in this case, there will be the aspect of consumer reaction
may be beyond the growers control. affecting decisions about what species and type of grain to sow
After harvest, grain may be tested and segregated according in the subsequent season.
to its suitability for processing. A circle format (Figure 2) is an alternative representation of
The next stage of processing covers a wide range of possi- the value-addition sequence because it emphasizes the feed-
bilities, given the many grain species and the great variety of back function, primarily the communication from the con-
possible processes. Some examples are given. sumer back to the breeder, who must produce varieties with
The emphasis is on the consumer as the ultimate person to quality traits that suit the specific consumer products. But in
determine product quality by deciding what products to addition, many other feedback functions are needed:
buy.
Thus, there is feedback up the grain chain according to the From the grower to the breeder (I need disease-resistant,
high-yielding varieties)
quality of the product or intermediate provided by the
previous person in the chain. From the rice-mill operator to the grower (I need rice grain
that mills well with a minimum of broken grains)
From the maltster to the grower (I need barley that germi-
nates uniformly)
Learning Objectives
From the hummus maker to the grower or grain trader
To achieve an understanding of all the stages leading to the (I need good chickpea grain for making my hummus)
eventual consumer product, via breeding, grain production, From the miller to the grower (I need clean grain that gives
processing, and marketing a high flour yield)
To gain an appreciation of the need for those involved at From the baker to the miller (I need flour that forms a
each stage to satisfy the requirements of those further down strong extensible dough that makes good bread)
the chain, most importantly the consumer From the retailer to the food processer (I need products
that look attractive, that taste good, and that sell well)
Figure 1 The grain-supply chain extends from the breeding of a variety to the product that reaches the consumer.
Figure 3 Depiction of a breeder at work in the 1890s when crossbreeding was in its infancy. (a) Cross-pollinating. (b) Examining progeny from a cross.
Note the great height of mature wheat plants of that time.
interact with breeders to obtain authentic seed of specific vari- When grain is delivered to the mill, storage silo, or grain
eties for growing under controlled conditions to ensure that elevator, it must be assessed for its quality and variety. This
the resulting grain conforms to the requirements for founda- assessment determines both the price to be paid to the grower
tion or registered seed. and the manner of segregation of the grain delivered. At any
delivery site, there may be several different grades or types of
Grow: Farm Management Must Optimize Grain Yield grain. It is obviously critical that a new delivery of grain should
and Quality be combined with grain of similar quality type in a distinct
storage cell.
In addition to decisions relating to seed for sowing, the grain
grower has decisions about soil preparation, fertilizer use at
sowing and during growth, sowing rate, and the possible use of
irrigation and of herbicides and pesticides. The task of maxi-
mizing financial returns involves on-farm management of
grain quality as well as grain yield, paying attention to the
combination of grain volume harvested, and market value
upon delivery. Even before harvest, the grower may have nego-
tiated marketing of the crop, based on expectations of yield
and quality attributes.
Storage and Transport: Identity Preservation Is Needed Process: Production of the Many Grain-Based Foods
for Grain Type and Quality and Nonfood Items
Further grain transport may involve delivery, probably by rail, It is at the stage of processing that the effectiveness of quality-
to the export terminal for loading into the hold of a ship based segregation is evident and its cost-effectiveness can be
(Figure 11). In any case, some grain is likely to be stored on- evaluated; for any situation, the advantages of better processing
farm for seed use in the next season and for animal feed. outcomes must justify the significant costs involved in the
In situations of subsistence farming, the family will be directly activities of testing and identity preservation. Due to these
dependent on the stored grain for food throughout the commitments, grain-quality assessment is again a priority
coming year. when the grain arrives at the flour mill, malt house, feed mill,
Beyond the initial stage of storage, the value of the segre- or other forms of processing factory. This type of assessment
gated grain must be preserved throughout transport and on determines how a specific grain shipment will be stored,
through further storage so that it still conforms to the original utilized, and managed in the sequence of events leading to
quality type when the grain is delivered for ultimate the product that is offered for final sale to the consumer.
processing. Throughout these stages, the grain is at risk of This sequence may be short: for example, the milling and
deterioration especially due to insect attack. Also, throughout boiling of rice grown by a subsistence farmer, with the family
these stages, commencing even before the farm gate, there is commenting on the eating quality. Even in a commercial situa-
the marketing of the grain at the level of the grain grower, by tion, processing may be minimal involving whole-grain foods
grain merchants, and internationally often at inter- such as rice, couscous, sweet corn, and beans (Figure 12). Min-
governmental levels. imal processing may involve sprouting to provide salad foods
(Figure 13). It may involve the stages of flour milling, leading to
bread and cake baking for sale in a hot-bread shop (Figure 14).
Further processes include the manufacture of breakfast cereals,
the use of cooked rice for sushi (Figure 15), the recovery of oils
Figure 9 Bags of grain, as used for storage and transport prior to bulk
storage. Figure 11 Loading grain into the hold of a ship for international trade.
Figure 10 Grain storage facilities are generally situated beside rail connections to facilitate transport to the mill or export terminal.
THE BASICS | The Grain Chain: The Route from Genes to Grain-Based Products 27
Figure 14 Bread rolls require the further stages of flour milling and
baking.
GENOME
Genes (DNA)
TRANSCRIPTOME PROTEOME
mRNA of expressed genes Polypeptides
ENVIRONMENT FUNCTIONAL PROTEINS
Climate, nutrients
CO2 GRAIN COMPOSITION
PROCESSING
PRODUCTS
Consumer acceptance
Figure 17 The sequence of biochemical events that occur in the grain,
leading to the grain-based products that reach the consumer.
Human
Figure 16 Feed pellets, made from partly ground mixed grains, are
destined for animal feed a further stage of food production beyond the Arabidopsis
stages in Figure 1.
Rice
This feedback loop from the consumer to others up the Wheat
chain may be short (a subsistence farming community) or Cotton
long (involving great distances and long time periods). The
latter case relates to the extent of worldwide grain trade as
the source of the grain involved. The ultimate feedback is to Barley
the grain breeder, for whom a change in breeding direction
involves the many years from an initial cross to the selection of
an elite line ready for registration and sale to growers. Coming Figure 18 Comparative sizes of the genomes of several plant species.
back around the grain circle (clockwise in Figure 2), there is Illustration kindly supplied by Dr. Colin Cavanagh, CSIRO, Canberra,
the further lag factor of the many months that grain may be in Australia.
storage and transport before it is processed. In addition, feed-
back may involve long distances, such as a flour miller in the
Middle East or a legume trader in India reporting grain defects reaching its ultimate destination the consumer. The general
to grain traders or breeders in grain-producing countries such sequence in Figure 17 is true for all grains with respect to their
as the Americas, Australia, or Europe. production and end-use quality. Central to the sequence is the
Consumer preferences also relate to their perception of genome the full DNA complement of genes. The relative sizes
nutritional quality. Government agencies publicize nutritional of several genomes are depicted in Figure 18. Outstandingly
guidelines that generally recommend the consumption of large is the genome of wheat, partly because common (bread)
grain-based foods as a major part of food intake. This emphasis wheat is a hexaploid, having three subgenomes, giving it a
on grains in the diet provides a major boost for the grain- complex genotype (the genetic makeup of the plant).
growing and grain-processing industries of the world. More The first opportunity to examine the effects of growth con-
specific recommendations, such as for greater levels of soluble ditions, according to the scheme in Figure 17, comes by the
fiber, may well target the breeder as the ultimate source of analysis of the transcriptome (mRNA) the transcripts from
significant changes in grain nutrition. Consumer feedback those genes that are active under the environmental conditions
may also target the processor with demands relating to the at the specific stage of grain development. This is the first
incorporation of specific additives in foods as well as targeting indication of what genes are being expressed in the endosperm
government agencies about legislation and food labeling. and also of how much of each gene product will be produced.
The next biological step is the resulting synthesis of the
many polypeptides corresponding to the mRNA transcripts
the proteome. The growth, yield, and suitability of grain for
The Biochemical Grain Chain processing of a grain are determined by the interaction of its
genotype (variety) with the environment (G E). Genes being
In contrast to the supply-chain concepts of Figures 1 and 2, expressed in the transcriptome (mRNA) are the focus of
there is the biochemical concept of Figure 17 that sets out the research attention as candidates for use as markers for specific
events in the context of genome, proteome, and grain compo- attributes, thus being candidates for marker-assisted selection
sition, for the events in the developing grain, and the many in breeding. The corresponding polypeptides in the proteome
changes that occur during processing prior to the product may also serve this purpose if the proteome is used as the target
THE BASICS | The Grain Chain: The Route from Genes to Grain-Based Products 29
for marker selection. In either of these cases, the breeder has composition to permit segregation prior to dumping
the satisfaction of knowing that expressed genes are being (comingling), after which distinct identity is lost. More specific
targeted, rather than genes in the genome that may not even knowledge of the grain quality of specific storage cells offers
be expressed. valuable market intelligence for use in maximizing return on
The transcriptome and the proteome also offer the advan- grain and in providing the processor (next in the grain chain)
tage of reflecting the effects of the growth environment. In with grain of the required quality specifications.
recent decades, we have learned much more about genotype
relationships than about the effects of environmental factors,
the latter being more difficult to study. With respect to wheat- Future Prospects
dough quality, research has concentrated on the genes coding
for the polypeptides that cross-link by disulfide bonds to form The general sequence of grain production and processing in
the polymers of glutenin protein, the critical part of dough. Figures 1 and 2 must continue into the future, but there will be
These polypeptides (subunits) are classed as the high- and changes within each of the stages. The breeding of new and
low-molecular-weight subunits, HMW-GS and LMW-GS, improved genotypes (varieties) is ongoing, so too are develop-
whose many alleles are coded at the Glu-1 and Glu-3 loci, ments in agronomy and farm management. However, these
respectively, on the Group-1 chromosomes of wheat. Informa- strategies will need to change and develop in view of the
tion about these gluten genes is available for many wheat ongoing climate challenges.
varieties from around the world. This allele information can The rising levels of carbon dioxide in the atmosphere con-
be used to predict dough properties as dough strength (Rmax) stitute an effective carbon fertilizer. Field experiments with
and extensibility (Ext) (both parameters from the Brabender ambient and elevated carbon dioxide have clearly shown that
Extensigraph), based on the alleles for both HMW-GS and the carbon fertilizer effect produces greater yields of biomass
LMW-GS, assuming normal grain-filling conditions and aver- and of grain. However, the protein content of the grain is
age protein content. lower, thus affecting grain quality and utilization.
Although we realize that wheat quality is also governed by Grain production will also be affected by predictions of
some other protein factors, this information is valuable to rising temperatures, shifting rainfall patterns, and more fre-
breeders and further down the grain chain for quality-based quent stress situations such as heat shock, storms, and violent
segregation at harvest. However, there are still serious gaps in winds. Furthermore, rising sea levels present the risk that some
our knowledge surrounding the effects of fluctuations in agricultural regions will become unsuitable for cropping.
growth conditions. The grain-production industry worldwide needs to able to
More important, but more poorly understood, is the man- adapt to a changing climate while also ensuring that the nec-
ner by which these glutenin polypeptides are disulfide cross- essary changes are consistent with sustainable production. In
linked into functional glutenin (Figure 17). The glutenin poly- line with these production challenges, markets will require
mers are so large that conventional methods of protein analysis greater uniformity and certainty of grain-quality specifications,
are inadequate. Field-flow fractionation (FFF) has been used to although these discerning markets must be prepared to pay
examine the upper size ranges, which are critical to dough premiums in contrast to commodity-level grain.
strength. FFF has shown, for example, that the dough- Processors and retailers will meet more customer-based
weakening effects of high temperatures during grain filling pressure to supply specific nutritional requirements, while
are accompanied by a reduction in the proportion of very large supermarket companies will exert downward pressure
large glutenin polymers. on suppliers.
However, we are yet to form a valid understanding of the
factors that determine the formation of disulfide bonds
between glutenin subunits and the extent of the ongoing poly- Exercises for Revision
merizations. Models have been suggested for the interactions
between the HMW-GS and LMW-GS and for polymeric glute- What features distinguish the added-value route of one
nin with gliadin proteins, but we do not understand the G E grain species from another? Compare the stages for two or
aspects of this very significant stage of grain-quality three contrasting grain species.
determination. Describe some grain-based products that have shorter and
We are aware that the formation of disulfide bonds within longer chains.
the gluten complex can continue during storage, especially at What are some critical feedback messages that are indicated
temperatures over about 30 C. In addition, disulfide bonds by the grain chains in Figures 1 and 2?
are a target for manipulation during processing (P) the third The shorthand expression G E P is used in relation to
factor that determines end-product quality (G E P). Figure 17. To what extent does it explain the grain chain?
Past the farm gate, there is a marketing opportunity that we
cannot presently grasp fully the possibility of segregating
grain deliveries based on their suitability for end-product man- Exercises for Readers to Explore the Topic Further
ufacture, considering the effects of both G and E. At this stage,
knowledge of variety is of less importance than of quality type, Select some grain-based products and outline the grain
which might be better specified by quantitative analysis of chain that has led to the production of each of these.
glutenin alleles. Some promise is offered for this purpose by Some examples are
on-the-spot equipment for the analysis of glutenin-subunit high-fructose syrups used as sweetening agents,
30 THE BASICS | The Grain Chain: The Route from Genes to Grain-Based Products
margarine from canola or any other oilseed, Marketing; Wheat: Grading and Segregation; Non-wheat Foods:
cornflakes or another breakfast cereal, Maize: Foods from Maize; Rice: Chinese Food Uses; Processing of
ethanol-based biofuels, Grains: Grain Quality Attributes for Cereals Other than Wheat;
oilseed-based diesel fuel. Evaluation of Wheat-Grain Quality Attributes; The Cereal Grains: An
In your region, identify the main grains that are grown and/ Overview of the Family of Cereal Grains Prominent in World
or processed. What forms of quality-based segregation, Agriculture; Wheat: An Overview of the Grain That Provides Our Daily
storage, and transport are used? Bread; The Legumes and Pseudocereals: Grain Legumes and
Investigate the biochemical sequence in Figure 17 for some Their Dietary Impact: Overview; The Oilseeds: Oilseeds: Overview;
grain species other than wheat. Wheat-Based Foods: Cereals: Overview of Uses: Accent on Wheat
Protein isolates are produced from several grains. Investi- Grain.
gate the production and uses of some, such as wheat gluten,
soy-protein isolate, and corn protein (sometimes called
corn gluten).
Examine some current consumer issues related to grain- Further Reading
based foods. In which of the upstream parts of the grain
chain is there the need for more diligent attention to Abdel-Aal E and Peter Wood P (eds.) (2005) Specialty Grains for Food and Feed.
quality? St Paul, MN: AACC International Press.
Arendt EK and Zannini E (eds.) (2013) Cereal Grains for the Food and Beverage
Industries. Oxford: Woodhead Publishing Ltd.
Faridi H and Faubion J (1995) Wheat End Uses Around the World. St. Paul, MN:
See also: Barley, Rice and Maize Processing: Rice Processing: American Association of Cereal Chemists.
Beyond the Farm Gate; Breeding of Grains: Maize: Breeding; Wheat Henry RJ and Kettlewell PS (eds.) (1996) Cereal Grain Quality. London: Chapman and
Breeding: Exploiting and Fixing Genetic Variation by Selection and Hall.
Kent NL and Evers AD (eds.) (1994) Technology of Cereals. An Introduction for
Evaluation; Genetics of Grains: Genomics of Food Grains; Grain Students of Food Science and Agriculture, 4th edn. Oxford: Pergamon Press.
Harvest, Storage and Transport: Wheat: Harvesting, Transport, Wrigley CW and Batey IL (eds.) (2010) Cereal Grains: Assessing and Managing Quality.
and Storage; Grain Marketing and Grading: Barley: Grading and Oxford: Woodhead Publishing Ltd.
Taxonomic Classification of Grain Species
LA Morrison, Oregon State University, Corvallis, OR, USA
CW Wrigley, The University of Queensland, Brisbane, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
Modern taxonomy is a uniform system for assigning scien- Common names can often have multiple or misleading asso-
tific names to plant species, thereby aiding communication ciations. If not qualified, pigweed can collectively refer to
among scientists. members of the genus Amaranthus without distinguishing
Classification is structured as a hierarchical system that between the weedy and grain amaranth forms. The wheat in
follows a concise set of rules for both wild and domesti- buckwheat (Fagopyrum esculentum; Figure 1) and Inca wheat
cated plant species. (Amaranthus caudatus; Figure 2) creates the false impression of
Species names often convey information about a grains a relationship with wheat (Triticum species; Figure 3), when in
function or traits or the activities associated with its use. fact wheat and these two pseudocereal species are far apart
Grains can be divided into two large groups, each with from each other in evolutionary relationships, in biology,
different evolutionary histories: cereals (grass species) and and in grain attributes. A more reliable means of communicat-
pseudocereals and pulses (nongrass species categories). ing is provided by scientific names, which are assigned by a
Taxonomic groupings reflect evolutionary relationships system known as taxonomy.
with interpretations often varying among traditional versus A scientific or taxonomic name consists of a genus and a
phylogenetic classifications. species name that, when combined as a binomial, provide a
For domesticated plants, the terms cultivar and landrace unique identifier for the grain in question. This name is uni-
are not interchangeable. versally recognized in all scientific circles and removes the
Taxonomic classification enables informed decisions for confusion created by common names, as the following exam-
accurately identifying and selecting research material of ple illustrates. Amaranthus caudatus, A. creuntas, and A. hypo-
grain species. chondriacus form the group known as grain amaranths, major
Evolutionary relationships are useful for identifying poten- food staples in the pre-Columbian Americas, which have now
tial allergic cross-reactions to grain species. regained importance and promise as pseudocereal grains in
Using classification to display phylogenetic relationships is international agriculture. Their taxonomic names distinguish
at the center of current controversies over the future direc- these grain amaranths from other weedy Amaranthus species
tion of Linnaean taxonomy. variously known as pigweed, amaranth, or water hemp.
Although some common names have been accepted into
international usage, they can still create problems within a
Learning Objective
scientific context. For example, in large species groups such as
the wheats, scientific names make distinctions where common
To achieve understanding of the role of taxonomy as a scien-
names cannot. The term wheat does not differentiate between
tific system for both classifying domesticated grain species
the wild and domesticated species of Triticum nor between the
and facilitating communication about them among scientists
species of Triticum and its sister genus Aegilops, whose members
are all wild species. The common name farro can collectively
refer to any one of three domesticated wheat species, all hulled
Introduction (non-free-threshing) T. monococcum, a diploid wheat that is a
minor crop species; T. dicoccum, a primitive tetraploid wheat
The common names of grain species often derive from local- also known as emmer; and T. spelta (T. aestivum ssp. spelta), a
ized, figurative terms. They can lead to considerable confusion hexaploid wheat known as spelt that is still grown in Europe and
due to variations in region and language. In English, the prob- Asia and now a specialty wheat in the United States and Canada.
lem is exacerbated by the adoption of common names from Common wheat and bread wheat are equivalent names that can
several languages, either as Anglicized words or direct trans- encompass a complex of domesticated hexaploid wheats when
lations. The grain legume Cicer arietinum offers a very good they are classified under one species name, Triticum aestivum,
example of multiple common names. In languages of Europe thereby including forms that are neither common nor necessar-
and Asia, it is known as cece (Italian), channa dhal (Hindi), ily amenable to bread-making (Figure 3).
garbanzo (Spanish), grao-de-bico (Portuguese: beaked
grain), hummus (Arabic), Kichererbse (German: giggle
peas), and pois chiche (French: chickpea). In English, it is
variously known as chickpea, gram, Bengal gram, chana, Egyp- Taxonomy: A System of Relationships
tian pea, and garbanzo bean. Another problem is the changed
usage of a common name from one region to another. In Inherent in taxonomic groupings is an indication of evolution-
Europe, corn refers to cereal grains generally, whereas in the ary relationships. Thus, the taxonomy of grain species has
Americas, it specifically refers to maize (Zea mays). some bearing on shared traits of plant morphology, grain
Figure 3 Variations in the appearance of heads of wheat species, one of many morphological characteristics used for their taxonomic classification.
From left to right, the wheat species are (including their genome assignments and common names): T. boeoticum (2x: wild einkorn), T. monococcum
(2x: einkorn), T. dicoccoides (4x: wild emmer), T. dicoccum (4x: emmer), T. durum (4x: macaroni wheat), T. carthlicum (4x: Persian wheat),
T. turgidum (4x: rivet wheat, cone wheat), T. polonicum (4x: Polish wheat), T. timopheevii (4x: Timopheevs wheat), T. aestivum (6x: bread wheat),
T. sphaerococcum (6x: shot wheat, Indian dwarf wheat), T. compactum (6x: club wheat), T. spelta (6x: spelt), and T. macha (6x: macha wheat). The
diploid A-genome species, T. urartu, is not shown here. 2x diploid; 4x tetraploid; 6x hexaploid. Adapted from Mangelsdorf PC (1953) Wheat.
Scientific American 189: 5059, 1953 by Scientific American Inc.
Taxonomy: What It Is and How It Started (taxon) or entities (taxa). Naming, or nomenclature, is gov-
erned by a set of rules laid out in the International Code of
Taxonomy provides a system by which plants are classified, Botanical Nomenclature (ICBN) and by the specialized rules
named, and identified. Classification consists of the circum- for domesticated plants found in the International Code of
scription, grouping, and hierarchical ranking of a plant entity Nomenclature for Cultivated Plants (ICNCP). Identification,
34 THE BASICS | Taxonomic Classification of Grain Species
which usually goes hand in hand with nomenclature, is a hails back to an older, no longer accepted, species name
determination based on similarity of a taxons membership T. vulgare, which translates as common wheat. The use and
in an existing group or a new group. derivation of corn and the genus name, Zea, provide another
Linnaeus initiated modern taxonomy with the publication example. The English word corn has its roots in the German
of Species Plantarum in 1753, which classified every plant then word for grain, korn. The Greek word zea, like corn, referred
known to him. The Linnaean system was not an isolated to grain in a general sense. Linnaeus selected Zea as a genus
endeavor. It was built on a long history of scientific attempts, name for the new cereal grain species that had arrived in
beginning with the Greek Theophrastus (third century BC) and Europe following the discovery of America and for which
the Roman Pliny the Elder (first century AD), to construct there was no European vernacular name upon which to draw.
universal naming and classification systems. The Linnaean As a consequence, the English common name corn was
binomial system, consisting of genus and species names, adopted into the American English vernacular to refer to the
resolved the problem of concisely and uniquely naming a only major New World cereal grain known in the eighteenth
plant. Continuing with the scientific heritage that preceded century.
him, Linnaeus used Latin as the language of taxonomic
names and descriptions. In his time, Latin was the universal
language by which European scholars could communicate. The Use of Genus and Species Names
Linnaeus took classical Latin and created a new botanical
form with its own terminology for plant structures and the Tables 1 and 2 list common and scientific names for grain
flexibility for adopting Latinized words taken from other lan- species. The convention is to write both genus and species
guages. Modern attempts to replace botanical Latin with names in italics, with the genus name having a capital as the
English for taxonomic descriptions have so far been unsuccess- first letter, followed by the species name (known as the specific
ful, leaving this particular Linnaean heritage intact. epithet) all in lower case. When a grain consists of a group of
species, the plural abbreviation spp. is used. In text, the genus
name component of the binomial is spelled out the first time
Origins of Taxonomic Names and abbreviated to its first letter, except when beginning a
sentence, for each following use as was done earlier in the
Many Linnaean scientific names for agricultural plants have text with the species examples of Amaranthus and Triticum. In
their roots in the vernacular of the ancient Greeks and Romans. cases where two genera with the same first letter are discussed
For example, the genus names for grains were derived from together, then the second letter of the species name is included
terms that often related to a particular function, activity, or trait in the abbreviation, for example, in discussions of the millets,
or to an association with a person. Thus, Triticum comes from Pa. for Panicum and Pe. for Pennisetum.
the Latin tero meaning I thresh; Cicer, which translates to Following the species and subspecies epithets is the name
chickpea, likely has its origin in the Greek kikus meaning (usually abbreviated and not in italics) of the botanist or
strength and power, a feature of chickpeas embodied in their researcher who named the species, for example, L. for Lin-
high nutritional value and a reputation as an aphrodisiac. For naeus in T. monococcum L. and Desf. for R. L. Desfontaines in
Cicer, the story is also tied up in family names. In time, the T. durum Desf. Known as the authority, this part of the scien-
early Greek name for chickpea evolved to kiros, which became tific name is important for synonymy, the component of tax-
cicer in Latin. Early Roman families would take surnames that onomy that deals with the historical naming of a species. When
related to their prosperity. Given the importance of legumes, it the name and classification of a grain species are in question
is not surprising that Cicero was the family name taken from due to competing treatments, synonymy provides a way to
cicer; likewise three other famous Roman families carried trace its taxonomic handling and to determine which is the
legume names: Fabius (faba: fava bean), Lentulus (lens: len- correct and legitimate name according to the ICBN. Authority
til), and Piso (pisum: pea). names are not included in Tables 1 and 2 as they are not
In the original Linnaean system, species names usually essential for the completeness of the binomial in text discus-
conveyed information about a trait. For example, cereale traces sions such as this. However, it is useful in research articles and
to Ceres the Roman goddess of agriculture and more specifi- literature reports to include the authority citation at the first
cally to Cerealis (of or relating to Ceres) from which the term mention of the species to clarify which taxonomic concept is
cereals was derived for collectively naming the edible grains of being followed.
grass species. The genus and species names together produce an
identifying designation for taxa as these examples illustrate:
Amaranthus caudatus translates as the amaranth ending with a Taxonomy: Rank and Hierarchy
taillike appendage, a reference to this species long, taillike
inflorescence (Figure 2); Cicer arietinum as chickpea shaped In classification, taxonomy provides a system for grouping
like a rams head, a reference to its seed shape; Fagopyrum species into ranked hierarchies. All grain species are flowering
esculentum as edible buckwheat (Figure 1); Triticum monococ- plants and thus are classified together as angiosperms. Mem-
cum as one-grained wheat (Figure 3); and Secale cereale as bers of this group reproduce sexually with the male and female
cereal rye. organs located in the flowers, which can be showy (pseudocer-
For species named by Linnaeus, the intertwining of scien- eals, pulses, and oilseeds) or inconspicuous (cereals). Their
tific names with the original vernacular helps to explain their seeds are protected by a fruit, which can be fleshy or dried. In
respective derivations. The origin of the term common wheat the case of cereals, the fruit is reduced to a thin outer tissue that
THE BASICS | Taxonomic Classification of Grain Species 35
Table 1 Classification of the grass family, Poaceae (Gramineae), all monocotyledonous plants: cereal grain species and their common namesa
is adherent to the seed. In botanical terminology, the cereal monocots (subclass Monocotyledoneae) and dicots (subclass
grain is called a caryopsis. Dicotyledoneae). From a more current phylogenetic perspective,
Traditional Linnaean classifications have followed the hier- angiosperms are classified as a clade with three subgroupings:
archical structure of KingdomClassOrderFamilyTribe dicots, monocots, and eudicots. Monocots and eudicots (true
GenusSpecies with subdivisions in each of these ranks, for dicots) are each designated as a clade (Table 3). Dicots are not
example, subclass. Largely morphological in concept, these treated as a clade because they are a nonmonophyletic group of
classifications do not always track well with current evidence primitive angiosperms with various members ancestral to both
on evolutionary relationships. On the other hand, phyloge- monocots and eudicots, respectively.
netic classifications are structured around monophyletic Grain species covered in this article fall within the monocot
groups that each consist of all descendants of a common and eudicot clades. Therefore, the biological features associated
ancestor, which share the groups identifying characteristics. with monocots and dicots are useful for distinguishing among
Given that ongoing molecular genetic research leads to chang- cereal (monocot) and noncereal (dicot) species. The terms
ing evolutionary concepts, phylogenetic classifications are in monocot and dicot refer to the presence of one or two embryonic
flux and have unresolved groupings. While some of the lower leaves (cotyledons) in the seed and young seedling. Other easily
Linnaean hierarchical rankings have been retained in these identifiable morphological traits differentiate between them.
classifications, groupings above Order, as well as unresolved Monocots have flower parts in threes, parallel-veined leaves,
groups at lower ranks, are designated as clades. As a general scattered vascular bundles, and fibrous adventitious root system.
term denoting a group consisting of an ancestor and all its Dicots have flower parts in fours or fives, net-veined leaves,
descendants, clade is used in this latter context to indicate a vascular bundles located in a ring, and a primary tap root.
group whose phylogenetic status is still open to question. Tables 13 present current taxonomic treatments for cereal
Differences distinguishing traditional versus phylogenetic and noncereal grain and oilseed species. For additional detail
classifications come into play for the monocot and dicot evolu- on traditional and phylogenetic classification approaches.
tionary lines of grain species. In older classifications, flowering Family is the most useful of the higher taxonomic ranks.
plants belong to the Class Angiospermae, which is divided into Family names end in -aceae except for several older names
36 THE BASICS | Taxonomic Classification of Grain Species
Table 2 Dicotyledonous grain and oilseed species and their common namesa
Amaranthaceae Amaranthoideae Amaranthus caudatus Amaranth, grain amaranth, Inca wheat, kiwicha
Amaranthus cruentas Purple amaranth, red amaranth
Amaranthus hypochondriacus Princes feather
Atripliceae Chenopodium quinoa Quinoa
Asteraceae (Compositae)b Cynareae Carthamus tinctorius Safflower
Heliantheae Guizotia abyssinica Niger, niger seed, ramtil
Helianthus annuus Sunflower
Brassicaceae (Cruciferae)b Brassiceae Brassica napus Canola, oilseed rape, rape
Brassica spp. Mustard
Crambe abyssinica Crambe
Fabaceae (Leguminosae)b Dalbergieae Arachis hypogaea Peanut, groundnut, goober
Cicereae Cicer arietinum Chickpea, garbanzo bean, gram
Genisteae Lupinus albus White lupin
Lupinus angustifolius Blue lupin
Phaseoleae Cajanus cajan Pigeon pea, Congo pea, red gram
Glycine max Soybean
Phaseolus lunatus Lima bean, butter bean
Phaseolus vulgaris Bean, kidney bean, pinto bean, navy bean, cannellini bean
Vigna angularis Adzuki bean, azuki bean, aduki bean
Vigna mungo Black gram, urad bean
Vigna radiata Mung bean, golden gram, green gram
Vigna unguiculata Black-eyed pea, cowpea
Fabeae Lathyrus sativus Chickling pea, chickling vetch
Lens esculentus Lentil
Pisum sativa Garden pea, field pea
Vicia faba Faba bean, broad bean
Lamiaceae (Labiatae)b Mentheae Salvia hispanica Chia, salba
Linaceae Linum usitatissimum Linseed, flax, Linola
Malvaceae Gossypieae Gossypium spp. Cottonseed
Pedaliaceae Sesamum indicum Sesame, sesame seed, benne, benne seed
Polygonaceae Fagopyrum esculentum Buckwheat, Japanese buckwheat, silverhull buckwheat
a
Following the classification system to family rank outlined in Heywood VH, Brummitt RK, Culham A, and Seberg O (2007) Flowering Plants of the World. Ontario, Canada: Firefly
Books and updated by the Angiosperm Phylogeny Group.
b
Original family name predating ICBN standardization.
that were in use before the ICBN standardized the suffix spell- particular species, then the ICBN requires that this subspecies
ing. Thus, in grain taxonomy, two family names are occasion- (or variety) repeats the species name for example, T. aestivum
ally encountered Gramineae (old) and Poaceae (new), ssp. aestivum. In general discussions, it is not necessary to
Leguminosae (old) and Fabaceae (new), Cruciferae (old) and include the repetitious infraspecific form. Thus, in this exam-
Brassicaceae (new), and Compositae (old) and Asteraceae ple, it is sufficient to cite only the species name T. aestivum
(new). The other useful higher taxonomic ranking is tribe, a when referring to bread wheat. Forma is the smallest category
subdivision of family, that is identified by an -eae ending. All and is infrequently encountered.
the cereal grains (e.g., maize, rice, sorghum, and wheat) are The infraspecific ranks are intended to describe variants that
monocots and belong to the grass family Poaceae. Dicot grains form distinct groups defined by genetics, morphology, ecology,
and oilseeds comprise a diverse mixture of families. The pulses and distribution. How a particular taxonomist chooses to clas-
or grain legumes (e.g., beans, peas, and soybeans), which are sify infraspecific taxa can appear arbitrary. For example, a current
the largest group of dicot grains, are classified in Fabaceae. taxonomic treatment of the wheats uses variety for the wild
Other families include one or just a few species, for example, wheat genus Aegilops (e.g., A. speltoides var. ligustica) but subspe-
Amaranthaceae (grain amaranths and quinoa), Polygonaceae cies for the domesticated genus Triticum (e.g., T. turgidum ssp.
(buckwheat), Asteraceae (niger, safflower, and sunflower), durum). Here, the infraspecific ranks should be viewed as
Malvaceae (cottonseed), and Brassicaceae (canola, mustard, approximately equivalent. This example also illustrates the dif-
and crambe). Subdivisions into subfamilies and/or tribes ficulty in reflecting evolutionary relationships within the frame-
reflect the size and diversity of the family. work of taxonomic classification. The infraspecific variation in
Grain species can be subdivided into infraspecific categories domesticated Triticum species is the product of human selection
known as subspecies (ssp.), variety (var.), and forma (f.). pressure, which is arguably on a different evolutionary scale
Because there is no clear agreement on use of subspecies versus than the variation in the wild species of Aegilops. Thus, in this
variety, the two are often used interchangeably, although it is example, the taxonomists choice of variety for Aegilops and
possible to treat variety as a subrank to subspecies. When the subspecies for Triticum should not be construed as a definitive
subspecies (or variety) is the taxonomic type that defines a reflection on evolutionary status.
THE BASICS | Taxonomic Classification of Grain Species 37
Table 3 Hierarchical classificationa for grain and oilseed speciesb with their use-category assignments
Monocots
Commelinids Poales Poaceae Ehrhartoideaed Oryzeae Oryza Cereal
Zizania Cereal
Chloridoideae Eragrostideae Eleusine Cereal
Eragrostis Cereal
Panicoideae Andropogoneae Coix Cereal
Sorghum Cereal
Zea Cereal
Paniceae Digitaria Cereal
Echinochloa Cereal
Panicum Cereal
Paspalum Cereal
Pennisetum Cereal
Setaria Cereal
Pooideae Poeaed Avena Cereal
Phalaris Cereal
Triticeae Hordeum Cereal
Secale Cereal
Triticosecale Cereal
Triticum Cereal
Eudicots
Core eudicots Caryophyllales Amaranthaceae Amaranthoideae Amaranthus Pseudocereal
Chenopodioideae Atripliceae Chenopodium Pseudocereal
Polygonaceae Fagopyrum Pseudocereal
Asterids Asterales Asteraceae Lactucoideae Cardueae Carthamus Oilseed
Asteroideae Heliantheae Guizotia Oilseed
Helianthus Oilseed
Lamiales Lamiaceae Nepetoideae Mentheae Salvia Oilseed
Pedaliaceae Sesamum Oilseed
Rosids Fabales Fabaceae Papilionoideae Aeschynomeneae Arachis Pulse, oilseed
Cicereae Cicer Pulse
Genisteae Lupinus Pulse
Phaseoleae Cajanus Pulse
Glycine Pulse, oilseed
Phaseolus Pulse
Vigna Pulse
Vicieae Lathyrus Pulse
Lens Pulse
Pisum Pulse
Vicia Pulse
Geraniales Brassicaceae Brassiceae Brassica Oilseed
Crambe Oilseed
Malpighiales Linaceae Linoideae Linum Oilseed
Malvales Malvaceae Malvoideae Gossypieae Gossypium Oilseed
a
Following the classification system to family rank outlined in Heywood VH, Brummitt RK, Culham A, and Seberg O (2007) Flowering Plants of the World. Ontario, Canada: Firefly
Books as updated by the Angiosperm Phylogeny Group.
b
For additional information on individual taxa.
c
To view the associated phylogenetic trees and clade subdivisions, see http://www.mobot.org/MOBOT/research/APweb/welcome.html.
d
Alternative subfamily and tribal classification assignment following the Grass Phylogeny Working Group (GPWG) revision.
Taxonomic Disagreements difficult to sort out. In the case of domesticated species, debates
usually deal with both the taxonomic status of taxa (usually
With taxonomic classification, there is not necessarily a right or species versus an infraspecific rank) and the relative impor-
a wrong treatment. Taxonomists and the researchers working tance of genetic evidence in defining species concepts and
with a particular group can often disagree with each other. As a evolutionary relationships. There is no clear separation of
consequence, different names for the same species can be these two points, partly due to the historical tradition on
found in use. For those who are not schooled in the ICBN or which taxonomy has been built and the inconsistencies in
unfamiliar with the underlying details of a taxonomic contro- how domesticated plants are viewed within an evolutionary
versy, conflicting classifications are confusing and can be context.
38 THE BASICS | Taxonomic Classification of Grain Species
The opposing species concepts at play in wheat taxonomy by agronomic traits such as disease resistance, maturity, yield,
offer an example of the different directions that a taxonomic and cold tolerance. The mixture also may include different
debate can take. Morphological diversity in the domesticated species. For example, wheat landraces collected during the late
tetraploid and hexaploid wheats illustrates the potential for 1940s in southeastern Turkey can be composed of ten or more
variation that developed with humans acting as evolutionary different forms of the two species T. aestivum (T. aestivum ssp.
selection agents. This variation originally correlated with eco- aestivum) and T. durum (T. turgidum ssp. durum). The advantage
geographic distributions and, if allowed to continue in isolation of landraces in subsistence farming systems is the guarantee of a
over geologic time, would have possibly led to new species. crop under all types of growing conditions. For example, if the
Wheat geneticists stress the need to portray accurate genetic farmer faces a particularly bad disease year, the resistant types of
relationships, circumscribing wheat species on the basis of the landrace will survive and yield a harvestable grain. Unfortu-
their close genetic similarities. Thus, in the AB- and ABD- nately, a clear-cut distinction between landrace and cultivar is
genome groups, geneticists accept only two species, each with not always maintained. It is not uncommon for a landrace to be
several infraspecific forms T. turgidum and T. aestivum. On the cataloged as a cultivar by one gene bank and as a landrace by
other side of the fence are those who stress the importance of another. While vernacular names are useful for identifying land-
recognizing the distinct variation among domesticated forms by races, they do not have taxonomic standing.
naming multiple tetraploid and hexaploid wheat species
(Figure 3 and Table 1). Provided that the rules of nomenclature
are followed, differing treatment concepts are acceptable. On Taxonomy: In Use
this basis, T. spelta and T. aestivum ssp. spelta are both correct as
are T. durum and T. turgidum ssp. durum, respectively, describing As a standardized system for classification of plants, taxonomy
the same wheats, albeit from different taxonomic perspectives. provides a useful way for tracing the origins of grain species. By
knowing the names of wild progenitors and related domesti-
cated species, researchers can make informed decisions when
Cultivated Plant Taxonomy selecting material for study or for breeding programs. Likewise,
knowing the taxonomic traits by which species are classified
Grain and oilseed species are classified as botanical entities enables researchers to verify the identity of research material.
under the ICBN. Their man-made forms derived through arti- This is particularly important given the possibility that gene
ficial selection, known by the general term cultigens, are clas- bank and research collections may hold and distribute misla-
sified under the ICNCP in two possible categories: cultivar beled or misidentified wild and domesticated germplasm.
(deliberately selected plants, either domesticated or wild) and The wheats offer a practical example of the importance of
group (an assemblage of cultivars). The term cultivar was taxonomy in the selection and use of research material.
developed from cultivated variety by combining the first five Because there are several conflicting taxonomic concepts for
and three letters of these two terms, respectively. According to the wild and domesticated wheats, a seed request may not be
the ICNCP, a cultivar is designated by one or more distinct, filled with the taxon actually requested. A general, unspecified
uniform, and stable characters that persist through propaga- request for accessions of the diploid A-genome progenitor of
tion. Although often used interchangeably, the terms cultivar tetraploid and hexaploid wheat species may be incorrectly
and variety are not equivalent, that is, variety is not a nomen- filled with either T. boeoticum or T. monococcum. While the
clatural term under the ICNCP. However, in the context of wild T. boeoticum and domesticated T. monococcum both have
cultivar registration, the term variety is often used to refer to a a form of the A-genome, neither was involved in the evolution
cultivar. Of note, variety is a legal term under the Interna- of T. durum (AB genomes) and T. aestivum (ABD genomes)
tional Union for the Protection of New Varieties of Plants (Figure 3). The correct species for this request would be
convention, which oversees the recognition of plant breeders T. urartu. Since T. urartu and T. boeoticum are morphologically
rights for a particular domesticated cultivar. Regarding termi- very similar to each other, a researcher, who is unfamiliar
nology differences between the ICBN and the ICNCP, the with the taxonomic characters that differentiate them, may
botanical term variety (also as varietas) applies only as a erroneously mistake T. boeoticum for T. urartu.
name for one of the infraspecific ranks under the ICBN.
A domesticated cultivar has a non-Latin name (cultigen
epithet) that is not italicized and enclosed by single quotation Taxonomy and Allergies
marks. Several examples should help to clarify this usage Zea
mays Bronze Beauty or maize cultivar Bronze Beauty, Lens Taxonomic relationships have a practical application for iden-
culinaris Laird, Avena sativa Wallaroo, Arachis hypogaea ssp. tifying potential allergic cross-reactions. For example, a person
fastigiata Georgia Red, Amaranthus cruentus K343, and Cicer with a proven allergic reaction to wheat grain would be more
arietinum Kranthi. Following a cultivars first mention, the likely to have similar problems with rye but less with the more
single quotation marks can be dropped. distantly related cereal grain species such as rice or maize
The term landrace is often encountered in discussions of (Table 1). This consideration is particularly relevant to celiac
crop germplasm. It refers to an indigenous grain species within disease, for which the prime cause is the gluten proteins found
a local farming system. Landraces predate modern agriculture in wheat. Celiacs are also warned to avoid eating the grain of
and often are ancient infraspecific forms that may have been in barley, rye, and triticale, closely related species in the tribe
cultivation for hundreds or thousands of years. Landraces are Triticeae. Although oats, which are classified in tribe Poeae,
usually composed of a mixed population whose members vary are considered to be less harmful for celiacs, some such
THE BASICS | Taxonomic Classification of Grain Species 39
individuals find that they must avoid them. On the other hand, Group and its followers offers a viable example of combining
celiacs may eat foods based on other cereal grains, which are the two systems: structuring higher-order relationships in an
classified in more distantly related tribes (e.g., maize, millet, ordinal phylogenetic system while drawing on the stability of
rice, and sorghum) and any of the dicot species. Within the the Linnaean system for treating the lower ranks (Family
context of taxonomy, the possible cross-reaction with oats but GenusSpecies) and revising their classifications as robust
not with maize, millet, rice, and sorghum is understandable. phylogenetic evidence becomes available.
The tribes Triticeae and Poeae are located in the same subfam-
ily, the Pooideae. On the other hand, maize, millet, and sor-
ghum are classified in the subfamily Panicoideae and rice in Exercises for Revision
the subfamily Ehrhartoideae.
Attempts at prediction of allergic cross-reaction may not be What is the function of scientific names for the grains and
valid if making comparisons with respect to different vegetative why are they preferred over common names in research,
or reproductive parts of the plant and different routes into the germplasm conservation, and communication among
body for example, the inhalation of pollen versus the inges- scientists?
tion of grain. It may thus be possible for a person, who is How do common names convey cultural and historical
allergic to wheat pollen, to suffer no adverse reactions to the information about the societies that cultivated domesti-
wheat grain. However, there is at least anecdotal evidence that cated grains? What role have common names played in
individuals, who experience mild ingestion intolerance to the taxonomic naming system?
cereal grains, may have these symptoms exacerbated in pollen What are the distinct differences among these taxonomic
season. Also, allergic cross-reactions are observed between pol- ranks: family, genus, species, subspecies, variety, and culti-
lens of different species, implying that sensitization by one var? In what case is variety and cultivar used interchange-
species can confer allergy to others. Thus, a positive test reac- ably and why is this practice not advisable?
tion to pollen from one species does not necessarily mean that Why is the ICNCP important as a separate set of rules for
it is the one causing clinical symptoms, nor need it have been naming and classifying domesticated grains? For grain spe-
the one responsible for the original sensitization. The real cies, what types of issues is the ICNCP able to better address
culprit may be another cross-reacting pollen. Here, taxonomy than possible with the ICBN?
offers a practical tool for elucidating such situations. For the How does taxonomic classification help in distinguishing
monocot grains, only the grass family Poaceae is necessary to between cereal and dicot grains and the species classified
consider. Within the very diverse dicot group of pulses, pseu- within each of these two major grain groups?
docereals, and oilseeds, the higher taxonomic rankings above With the much-enhanced ability to elucidate phylogenetic
Family are of practical use for evaluating allergic cross- relationships with data from molecular genetic studies,
reactions. In providing only a brief introduction into the tax- what types of challenges does the traditional Linnaean
onomy of grain species, Table 3 illustrates the hierarchical system face when incorporating this information into tax-
relationships that should be considered in dealing with poten- onomic classification?
tial cross-reactions. How do competing taxonomic classifications create diffi-
culties for grain researchers? What types of solutions can be
developed to address the controversies associated with tra-
Future Trends in Taxonomy ditional versus natural phylogenetic classifications?
How do evolutionary groupings of grain species in taxo-
As is evident in the previously mentioned discussion, taxon- nomic classifications aid in assessments of potential allergic
omy is subject to conflicting, and often changing, interpreta- cross-reactions?
tions. When taxonomic disagreements prevail, the focus shifts Since landraces are often mixtures of multiple species, what
away from taxonomys real value in aiding communication are the challenges faced in cataloging and classifying land-
among scientists. For example, a long-running debate among race diversity? Why is it important to differentiate among
wheat researchers dating from the 1950s over how to classify individual types within a single landrace collection?
wild and domesticated taxa by evolutionary relationships has
created a confusing situation with many different classifica-
tions and no consensus for names or species concepts. This Exercises for Readers to Explore the Topic Further
controversy illustrates the possible problems that lie ahead
should the current trend continue among molecular systema- Explore the advantages and conflicts posed by the applica-
tists and research geneticists to restructure traditional taxon- tion of molecular technologies to taxonomy.
omy of lower-ranking groups into a natural system that For example, examine and expand on the statement in
displays phylogenetic relationships even when they are not Future Trends: molecular systematists (may) suggest
fully resolved and/or open to widely varying interpretations. other naming systems and other ways of classifying, which
Because concepts of plant phylogeny are changing rapidly with could lead to a re-invention of taxonomy.
the sophisticated tools of molecular biology, an attempt to Refer to the article on variety identification, which describes
structure taxonomy on molecular evidence may lead to a situ- the use of protein composition and DNA analysis to differ-
ation of constant taxonomic flux. Traditionalists argue that entiate cultivars in agriculture. Consider how these
taxonomy should continue as it now stands guided by the methods might be expanded to compare a wider range of
ICBN. The current approach of the Angiosperm Phylogeny taxonomic relationships.
40 THE BASICS | Taxonomic Classification of Grain Species
Wiersema JH and Leon B (2013) World Economic Plants: A Standard Reference, 2nd
See also: Breeding of Grains: Barley: Genetics and Breeding;
edn. Boca Raton, FL: CRC Press.
Canola/Rapeseed: Genetics and Breeding; Food Grains: Williams JT (ed.) (1995) Cereals and Pseudocereals London: Chapman and Hall.
Intolerance, Allergy and Diseases: Celiac Disease; Genetics of
Grains: Maize: Genetics; Rice: Genetics; Wheat Genetics; The Cereal
Grains: Cereals: Domestication of the Cereal Grains; The Legumes Relevant Websites
and Pseudocereals: Amaranth: Overview; Buckwheat: Overview;
Pseudocereals: Overview; The Oilseeds: Oilseeds: Overview. http://www.ars-grin.gov The United States Department of Agriculture website for GRIN
(Germplasm Resources Information Network) is set up for taxonomic queries on
holdings of economic plants and with links to other relevant websites for the
taxonomy of economic plants. Economic plant listings include vernacular names
and economic uses. Information on this website is drawn from World Economic
Plants: A Standard Reference.
Further Reading http://www.eti.uva.nl The Expert Center for Taxonomic Identification (ETI) is an on-
line taxonomic database service supported by an international consortium. The site
Angiosperm Phylogeny Group (1998) An ordinal classification for the families of hosts a searchable taxonomic database for taxonomic hierarchies, species names,
flowering plants. Annals of Missouri Botanical Garden 85: 531553. synonyms, and descriptions.
Angiosperm Phylogeny Group II (2003) An update of the Angiosperm Phylogeny Group http://www.iapt-taxon.org/nomen/main.php The online version for the most
classification for the orders and families of flowering plants: APG II. Botanical recent edition of the ICBN (2012), which is known as the Melbourne Code, is in
Journal of the Linnean Society 141: 399436. English.
Angiosperm Phylogeny Group III (2009) An update of the Angiosperm Phylogeny Group http://www.ildis.org Location for the International Legume Database and Information
classification for the orders and families of flowering plants: APG III. Botanical Service (ILDIS), this website has a searchable database for species information
Journal of the Linnean Society 161: 105121. about the grain pulses.
Brickell CD, Alexander C, and David JC, et al. (eds.) (2009) International Code of http://www.ipni.org The International Plant Names Index (IPNI) is a comprehensive
Nomenclature for Cultivated Plants, 8th edn. In: Regnum Vegetabile, 8th edn., online database of plant names and bibliographic information.
151: Belgium: Drukkerji Geers Scripta Horticulturae 10. https://www.ksu.edu.wgrc.Taxonomy.taxintro.html This website has comprehensive
Hanelt P and Institute of Plant Genetics and Crop Plant Research (eds.) (2001) listings of historical and current taxonomic treatments of the wheats.
Mansfelds Encyclopedia of Agricultural and Horticultural Crops (Except http://www.mobot.org/MOBOT/research/APweb/welcome.html The Angiosperm
Ornamentals) Germany: Springer-Verlag, English edition. Phylogeny Group website has up-to-date revisions to the Angiosperm phylogenetic
Heywood VH, Brummitt RK, Culham A, and Seberg O (2007) Flowering Plants of the trees, extensive reviews of current evidence, and references.
World. Ontario, Canada: Firefly Books. http://www.plantnames.unimelb.edu.au The Multilingual Multiscript Plant Name
Judd WS, Campbell CS, Kellogg EA, Stevens PF, and Donoghue MJ (2007) Plant Database has a searchable database for locating scientific and vernacular names in
Systematics: A Phylogenetic Approach, 3rd edn. Sunderland, MA: Sinauer Associates. all applicable languages for a particular grain species.
Linnaeus C (1753) Species Plantarum. Sweden: Laurentius Salvius. http://tolweb.org.tree.phylogeny.html The Tree of Life Web Project is a searchable
Mangelsdorf PC (1953) Wheat. Scientific American 189: 5059. internet source for information on biodiversity, phylogeny, and characteristics of
McNeill J, Buck WR, and Burdet HM, et al. (eds.) (2012) International Code of Botanical biological groups.
Nomenclature (Melbourne Code). In: Regnum Vegetabile154: Konigstein, Germany: http://www.umsl.edu.services.kellogg.gpwg.default.htm As a collaborative group of
Koeltz Scientific Books. systematic botanists, the Grass Phylogeny Working Group (GPWG) produced a
Stuessy TF (2009) Plant Taxonomy: The Systematic Evaluation of Comparative Data, phylogeny of the grass family by combining existing data sets. Although GPWG is
2nd edn. New York: Columbia University Press. no longer active, this website provides access to its results.
Vietmeyer ND (ed.) (1989) Lost Crops of the Incas Washington, DC: National Academy http://upov.int/about/en/benefits_upov_system.html The International Union for the
Press. Protection of New Varieties of Plants is an intergovernmental organization that
Watson L and Wrigley CW (1984) Relationships between plants relevant to allergy. oversees the convention for protecting intellectual property rights to new plant
Medical Journal of Australia 141(5 Suppl.): S18. cultivars introduced into commerce.
Grain: Morphology of Internal Structure
H Corke, University of Hong Kong, Hong Kong, Pokfulam, China; Hubei University of Technology, Wuhan, China
2016 Elsevier Ltd. All rights reserved.
Topic Highlights the plant, the rachis (where the spikelet is attached to the axis)
breaks naturally and the seed (still surrounded by a tightly
Grain structure determines the necessity for dehulling and bound palea, lemma, and awn) falls from the plant. This is
the consequences of wet and dry milling. an essential survival feature for a plant whose seeds are dis-
The major cereal grains (wheat, rice, and maize), although persed in nature through the action of wind, insects, etc. In a
botanically similar, have significant differences in organi- farmers field, a brittle rachis would not be viable, as harvesting
zation of starch and protein reserves. would be extremely difficult. The genetic step from brittle to
Wet milling results in separation of major grain compo- nonbrittle rachis is the key to domestication of small-grain
nents; dry milling is generally a size reduction process. cereals (e.g., wheat, rice, and barley). However, wheat has a
further useful adaptation to domestication, which differs from
rice. In wheat, the palea and lemma naturally separate from the
Learning Objectives grain this is called a naked grain characteristic. In rice, the
grain is covered, that is, the hull is attached to the grain and
To be able to visualize and explain the major anatomical requires a separate processing stage (dehulling) to remove it.
features of cereal grains.
To understand the distribution of major nutritional com-
ponents in the grains. Bran
To understand how the specific structure of different cereal
grains drives the choice and consequence of wet and dry The wheat grain has been described as a single-seeded fruit
milling. called a caryopsis, in which the ripened ovary wall is fused to
the seed. The classical diagram for hexaploid common (bread)
wheat (Triticum aestivum) grain structure was produced by the
Introduction Wheat Flour Institute, Washington, DC (Figure 2). The objec-
tive of flour milling is to remove the embryo (germ) outer
This is a wide topic because virtually every aspect of grain layers (epidermis, hypodermis, cross cells, tube cells, seed
technology is related to grain structure. Handling, processing, coat, and nucellar tissue), which collectively are termed the
and utilization of grains depend on specific details of their bran. The aleurone layer, part of the endosperm, is also
structure. For example, in dry milling of rice and wheat, removed with the bran and from a millers point of view may
removal of the bran layers is necessary. The wheat grain is not be included as a bran layer. Much of the vitamin and mineral
covered with a tightly attached hull, so unlike in rice, initial content of the grain is contained in the bran. In many coun-
dehulling is not required. Bran removal in wheat is part of the tries, wheat flour is enriched or fortified to replace some of the
process of producing fine particulate flour by crushing the nutrients lost in milling. Removal of the bran (which, includ-
endosperm. Bran removal in rice should leave an unbroken ing the aleurone layer, comprises 15% of the grain by
polished rice grain. In both wheat and rice, endosperm should weight) permits production of white flour. Apart from appear-
be at a specific hardness to achieve the desired outcome. Wet ance, white flour has different technological properties to
milling is a fractionation process. In maize, starch, oil, and whole-wheat meal (that produced by grinding the whole
protein components are separated. Again, the physical struc- grain without bran separation). The presence of large amounts
ture of the grain is exploited in the design of the milling of bran dilutes the proportion of gluten in the product and
process. Issues such as these will be covered in detail in this results in a denser, more fiber-rich product. The presence of
article. More specifically, the following aspects will be covered: small amounts of bran (as bran specks) results in off-white
wheat grain morphology and its effect on dry and wet milling, color in flour and in many final products (e.g., steamed bread).
rice grain morphology and its effect on dry milling and In fresh white noodles, discoloration to gray is associated with
nutrition, maize grain morphology and its effect on handling polyphenol oxidase (PPO) content of flour, and most PPO is
and wet milling, barley structure and its effect on brewing, and present in bran. In Chinese steamed bread, formation of
a comparative summary of general effects of grain structure. brown spots on the surface may also be associated with PPO
effects from bran contamination (and the use of NaCl in the
formulation). Because of the high mineral content of bran
Wheat Grain Morphology and Its Effect on Dry and relative to endosperm, bran content in flour can be estimated
Wet Milling by ashing (furnace heating) a flour sample and weighing the
residue. For the same flour yield, a variety that gives lower ash
The relationship of the grain to overall spike structure is impor- content (less bran) is of course preferable.
tant in milling (Figure 1). Historically, the wheat ancestors In wheat flour milling, grain is tempered to a uniform
(e.g., the diploid Triticum monococcum) had a brittle rachis moisture content (usually 15.5%). The effect of this is to
trait. This means that after ripening of the grain and drying of make the bran layers flexible so that under the crushing action
Awn
Spikelet (not always
present)
Rachis
Palea Pales
Lemma Floret
Grain
Rachilla
Glume
(a) (b)
Figure 1 The structure of the wheat spike (ear). Cultivated wheat requires a nonbrittle rachis so that mechanical harvesting is possible and a
loosely attached palea and lemma so that a dehulling step is not required. Reproduced from Evers AD and Bechtel DB (1988) Microscopic structure of
the wheat grain. In: Pomeranz Y (ed.) Wheat: Chemistry and Technology, 3rd edn., vol. 1, pp. 4795. St. Paul, MN: American Association of
Cereal Chemists.
of roller milling, bran will break into large flakes, not be Endosperm
crushed into powder. This formation of flakes of bran results
The product of wheat dry milling is flour, the result of crushing
in pieces that are of different sizes, shapes, and densities to the
endosperm. The endosperm consists of cells surrounded by a
endosperm particles, which on size reduction become flour
pentosan-rich cell wall (mostly arabinoxylans) and contains
particles. Without this physical flexibility in bran, milling
starch granules embedded in a protein matrix. The size of the
would not be possible. However, with crushing, some endo-
cell and the thickness of the cell wall affect the proportion of cell
sperm is likely to be attached to some of the bran flakes.
wall material relative to starch and protein in the endosperm.
Continuing milling to remove the last particles of endosperm
The cell wall absorbs water during dough mixing, so it is signif-
(i.e., aiming for a high flour yield or percentage yield of flour
icant in determining optimum water absorption of flour. The
relative to starting weight of grain) will increase contamination
dimensions of the cell wall are genetically variable. The starch
of flour with bran. The flour yield also depends on several
granules consist of two types, larger A granules and smaller B
other properties of bran:
granules. Starch composition (proportion of amylose to amy-
1. The thickness of the bran layers, which is under genetic lopectin in starch) determines physical properties of any gela-
control tinized starch product resulting from cooking the flour. This is
2. The size of the grain (bran will be a relatively lower fraction most important in Japanese white-salted noodles (udon) where
of a larger grain; also, a plump well-filled grain will be a low-amylose (partial waxy), high-swelling starch gives favor-
preferable) able texture. In many other products, such as baked bread, the
3. The characteristics of the crease starch composition, within normal limits for wheat flour, does
not have much impact on product quality. Starch granules are
The crease in a wheat grain is the lateral fold (Figures 2 and 3)
subject to shear or breakage, termed starch damage in milling, if
ending with the pigment strand. A deep crease results in more
the endosperm is too hard (e.g., genetically hard texture or too
problems with bran removal in milling more bran particles
low moisture content in milling) or if milling equipment is not
attached to endosperm chunks usually result from a deep
adjusted correctly. A broken granule, exposing the interior, is
crease. Also, the crease can trap contaminants, for example,
much more readily subject to hydrolysis by amylases during the
dust and fungal spores, which can cause problems in storage
bread-making process (during fermentation and early stages of
and in cleaning the grain before milling.
baking before the enzyme is inactivated). Some starch damage
Grain color in wheat, usually due to red pigmentation in the
may be desirable to provide a substrate for yeast growth, but
bran (seed coat), is usually classified as red (the presence of
excessive damage adversely impacts on quality and may result in
pigmentation) or white (absence). White wheat flour, if contam-
a sticky texture in the final loaf. Endosperm hardness is largely
inated with bran, will show less discoloration than red wheat
due to the structure of two puroindolines (components of fria-
flour. This is why hard white-type wheat is becoming more
bilin), which are amyloplast membrane proteins present in the
widely accepted as suitable for Asian noodles. Other colors are
endosperm. There is a widespread confusion about grain hard-
possible but rare in commercial trade for example, blue, pur-
ness (protein-influenced) and noodle texture (starch-
ple, or even black wheat, with a dark purple pigment in the seed
influenced). A high-swelling starch, but not necessarily a soft
coat, which is used in some specialty products in China.
THE BASICS | Grain: Morphology of Internal Structure 43
Hairs of
brush
Endosperm
Cell filled with
starch granules
in protein
matrix
Cellulose walls
of cells
Aleurone cell
layer (part of
endosperm but
separated
with bran)
Nucellar tissue
Seedcoat
(testa)
Tube cells
Cross cells
Hypodermis
Epidermis
Scutellum
Sheath of shoot
Rudimentary
shoot
Rudimentary
primary root
Root sheath
Root cap
Crease
Endosperm
Pigment
strand
Bran
Germ
Cross-sectional view
Figure 2 The structure of the wheat grain longitudinal and cross-sectional views. Reproduced from Pomeranz Y (1987) Modern Cereal Science
and Technology, pp. 2539. Weinheim, Germany: VCH Publishers.
44 THE BASICS | Grain: Morphology of Internal Structure
Figure 3 Outer kernel layers, central starchy endosperm, and crease region of kernel cross sections of wheat, barley, oat, and rye stained with acid
fuchsin calcofluor. Reproduced from Dornez E, Holopainen U, Cuyvers S, et al. (2011) Study of grain cell wall structures by microscopic
analysis with four different staining techniques. Journal of Cereal Science 54: 363373.
endosperm, is required to make a soft-textured noodle (such as as in the endosperm. Thus, the wheat germ protein is of higher
Japanese udon). quality (more favorable amino acid balance, such as higher
lysine content) for human nutrition. The lysine level in protein
in the embryo is 8%; in endosperm protein 2%, and in
Embryo (Germ)
bran protein 4.5%. In addition, 20% of the lipids of the grain
The embryo consists of the rudimentary structures (shoot and are in the embryo (oil-containing spherosomes are present in
root) of the plant, which can grow from the grain, plus the root the aleurone layer and in scutellar cells), but there are also high
and shoot sheathes, the root cap, and the scutellar tissue. The levels of lipase and lipoxygenase enzymes, which hydrolyze
embryo, 3% of the grain weight, contains 8% of the grain and oxidize oil, causing rancidity. Wheat germ hence has uses
protein, but it is mostly in the form of enzymes related to as a food material or supplement for human nutrition and is
subsequent growth of the seedling, not storage protein (gluten) valuable as an additive to animal feed (particularly for
THE BASICS | Grain: Morphology of Internal Structure 45
monogastric animals such as pigs and poultry). On the other a genetically low proportion of B-type compared to A-type
hand, wheat germ, unless treated, has a short shelf life. Whole- granules would be a strong advantage for increased starch
wheat meal also has a shorter shelf life than white flour. yield in wet milling. Considerable effort is being expended to
Removal of germ to produce flour results in a product that is identify more suitable wheat varieties for this use.
stable for long periods of time. A significant recent development has been the application
of Raman microscopy (in combination with further develop-
ment of selective staining for optical microscopy) to image and
Other Features localize the presence of, for example, b-glucans, protein, starch,
and ferulic acid in wheat and barley grain. The advantage of
Dry milling of durum (Triticum durum) wheat (a tetraploid Raman microscopy, applied alone, is the absence of required
species) to make semolina (larger particle size than flour pretreatment or staining. An example of the capability of high-
from common wheat) has a few special features related to the quality selective staining can be seen in Figure 3; such infor-
durum endosperm texture. The physical hardness (vitreous- mation as the cell wall size and thickness, the size and packing
ness) of durum endosperm makes the grain liable to break of starch granules, the nature of the bran layers, and the geom-
during combine harvesting. These broken grains should be etry of the crease can be invaluable for genetic improvement of
separated and recovered during cleaning of grain before mill- wheat.
ing, or a high level of wastage may occur. Optimum tempering
of durum grain before milling is also essential and varies with
the vitreousness of the endosperm. Careful milling (e.g., more
break rolls than for bread wheat) is needed to achieve suitably Rice Grain Morphology and Its Effect on Dry Milling
fine semolina for high-quality pasta. and Nutrition
In wheat wet milling (to produce wheat starch, gluten, and
Grain Structure
wheat germ oil), separation is by soaking the grain, crushing,
and sedimenting off the starch component. B-type starch gran- The anatomical structure of rice (Figure 4) is of course very
ules are too small to easily sediment, and their complete similar to that of wheat. However, grain shape is of more
removal would require expensive centrifugation steps. Hence, significance. Short-, medium-, and long-grain rice are common
Awn
Lemma
Palea
Pericarp
Seedcoat
Nucellus
Hull Aleurone layer
Subaleurone layer
Endosperm
Starchy endosperm
Scutellum
Plumule
Embryo
Radicle
Epiblast
Sterile
lemmae
Rachilla
Figure 4 The structure of the rice grain. Reproduced from Juliano JB and Aldama MJ (1937) Morphology of Oryza sativa L. Philippines
Agriculturalist 26: 1134; Juliano BO (1993).
46 THE BASICS | Grain: Morphology of Internal Structure
classifications. In the United States, texture can be recognized steamed, driving nutrients from the hull and the bran into the
from grain length short grain is low amylose (sticky) and endosperm, inactivating enzymes that would cause rancidity,
long grain is intermediate amylose (2025%) (harder cooked and reducing microbial load. However, after steaming, the
texture such that grains are separate after boiling). This is a parboiled rice must be dried slowly before it is dehulled and
marketing phenomenon, not a genetic relationship, and in polished, or cracks will form with the same effect as before.
other countries, long grain may well be waxy or sticky types, Parboiled rice, despite the steaming treatment, does not
sometimes a source of confusion to traveling Americans. Grain require less time to cook by the consumer than ordinary
shape (expressed as ratio of length to width) affects milling. A white rice, because water penetration to the center of the
rounded grain with a low length/width ratio will be more grain is slow. Instant rice is prepared by steaming polished
resistant to breakage than a slender type of grain. Rice lacks white rice. It is dried rapidly, forming numerous internal
the crease found in wheat grains. This enables abrasive milling cracks. These cracks, not the starch gelatinization induced by
to remove the bran and leave only the endosperm, greatly steaming, allow the product to be cooked by the consumer in a
simplifying the milling process. few minutes because they aid water penetration. Instant rice is
The hull, which covers the rough (also known as paddy) more physically liable to breakage in transportation than raw
rice, comprises 1724% of the weight of the paddy rice. It polished white rice, and the softer texture resulting from the
consists of lemma and palea, awn, the rachilla, and lemmas. cracks is unattractive to traditional rice consumers in many
Varietal differences in percentage of hull affect yield of brown parts of the world.
rice, and this in turn affects milled rice recovery (yield of white
polished rice). The hull (husk) is silica-rich and indigestible for
human nutrition, even after heat treatment, even though it is Rice Bran Layers
rich in minerals.
The bran layers (pericarp, seed coat, nucellus, and aleurone
layer) contain most of the minerals (represented as ash), vita-
mins, fiber, fat, and much of the protein present in the rice
Cracking
grain (Figure 5). Considerable debate over the years has taken
In producing polished white rice, the control of cracking is place over the suitability of white polished rice, compared to
essential. Cracks caused by poor drying conditions after har- brown rice, in nutritionally deficient diets in some developing
vest, for example, will result in grain that breaks easily in nations. In fact, bran also contains phytate (phytic acid), which
milling (dehulling or polishing) producing a lower yield of binds to available minerals provided by brown rice, and may
head rice and consequently much lower value product. In the also bind and remove additional minerals from the gastroin-
production of parboiled rice, rice with the hull still attached is testinal tract. Brown rice consumption may therefore cause a
100
Starch
80
60
(%)
40
Fiber
Ash
20 Fat
Protein
20 40 60 80 100
Outer layer Center
Proportion of the kernel
Figure 5 The effect of progressive surface abrasion of brown rice on content of major constituents. Reproduced from Barber S (1972) Milled rice and
changes during aging. In: Houston DF (ed.) Rice: Chemistry and Technology, pp. 215263. St. Paul, MN: American Association of Cereal Chemists;
Juliano BO (1993).
THE BASICS | Grain: Morphology of Internal Structure 47
Large
spherical
Crystalline Protein
Compound bodies
starch
granule Small
spherical
1 m
Figure 6 The rice endosperm indicating structure of spherical and crystalline protein bodies. Reproduced from Coffman WR and Juliano BO (1987)
Rice. In: Olson RA (ed.) Nutritional Quality of Cereal Grains: Genetic and Agronomic Improvement. Madison, WI: Agronomy Society of America;
Juliano BO (1993).
net loss of minerals to the body and in addition is susceptible plant and consists of a central cob with 800 seeds attached in
to rancidity in storage (lipids and lipases still present) and rows by the pedicel and covered by a husk. The pedicel acts as a
more susceptible than polished rice to storage pest attack. conduit for photosynthetic products supplying the developing
grain. The male part of the flower the tassel produces pollen
and is at the top of the plant. The maize grain (Figure 7) is 10
Rice Endosperm times larger than small-grain cereals, an average dent maize
Rice starch granules are small (24 mm), polygonal, and com- grain weighing 300 mg. The flattened sides of the mature
pound, that is, many small granules are compacted together. grain are due to pressure from the tight packing in rows on
Compositionally, rice starch varies from 0% (waxy, sticky, and the cob. After removal of the husk, the maize grain is exposed
glutinous) amylose to over 35%. This range is much wider than (naked characteristic) and easily separated from the cob.
found naturally in hexaploid wheat starch. Expression of natu-
rally occurring waxy mutants is more likely in a diploid cereal
Handling
(e.g., rice, maize, and sorghum) than in a hexaploid where the
effect is buffered by the other two sets of chromosomes present. Shelled maize grain (grain separated from the cob and packed
Rice differs from wheat and maize in having crystalline protein in bulk) is more likely than other grains to enter export mar-
bodies rich in glutelin, in addition to the prolamin-rich protein kets. A high proportion of the US maize harvest is exported,
bodies common to all cereal endosperms (Figure 6). The aleu- often to highly quality-conscious markets such as Japan. A
rone layer (considered here as the botanical part of the endo- typical export grain may enter the transportation system by
sperm, although it is processed as part of the bran) differs in traveling by barge down the Mississippi, entering the Gulf of
thickness (usually one to seven cell layers) depending on vari- Mexico for the long voyage to Asian markets. This is physically
ety. The aleurone cells contain protein bodies and lipid bodies. stressful on the grain, and a poorly dried grain with internal
cracks is liable to breakage. This contributes to quality loss and
increased susceptibility to insect or fungal attack and moisture
Rice Embryo damage. The primary criteria for successful shipment of maize
The rice embryo is smaller relative to the whole grain than in are suitable endosperm texture (hard enough) and suitably
wheat. It contains several types of parenchymatous cells, con- controlled harvesting and drying to avoid cracking.
taining protein bodies and/or lipid bodies. The main parts of
the embryo are the cotyledon and the embryonic axis. The
plumule and radicle are similar to other cereal grains. The Wet Milling
high lipid content of the embryo contributes to susceptibility Most maize that is not fed to animals is processed by wet
to rancidity in storage of brown rice. milling to produce starch (with many diverse applications,
such as further processing high fructose corn syrup, bioetha-
nol, or modified starch for food and nonfood industries), oil
Maize Grain Morphology and Its Effect on Handling (a high-quality cooking oil), and protein or fiber-rich fractions
and Wet Milling that are used in livestock feed. The composition of these com-
ponents in the grain is of course of the utmost importance. As
Maize is monoecious, having separate male and female flower- the maize seed is much larger than wheat or rice, the relative
ing parts on the same plant. The female seed-bearing part of the proportion by weight taken up by the bran layers (including
plant (called the ear) develops off the lower-middle side of the aleurone layer) (5%) is less than in these other cereals.
48 THE BASICS | Grain: Morphology of Internal Structure
Hull
Epidermis
Mesocarp
Cross cells
Tube cells
Seedcoat (testa)
Aleurone layer
(part of endosperm
but separated with bran)
Horny endosperm
Floury endosperm
Walls of cells
Scutellum
Plumule or rudimentary
shoot and leaves
Longitudinal section
of a grain of corn Radicle or
(enlarged -30 times) primary root
Tip cap
Pericarp
Horny Floury
endosperm endosperm Cross-sectional view
Figure 7 The structure of the maize grain longitudinal and cross-sectional views. Reproduced from Pomeranz Y (1987) Modern Cereal Science
and Technology, pp. 2539. Weinheim, Germany: VCH Publishers.
The embryo of maize is a much higher proportion of the additional protein sources. In practice, spread of opaque-2 vari-
grain (11%, containing 33% of the oil) than in other eties has been limited because in most cases, production of
cereals; hence, the overall oil content of the grain is quite normal endosperm maize, and supplementation with soy pro-
high (typically 5%). Also, a higher proportion of the total tein, is easier for animal feed production. Similarly, a classic
protein is nonstorage in nature (e.g., enzymes of the embryo), long-term selection experiment at the University of Illinois
so overall, the protein of maize is higher quality for human showed that the oil content of maize could be increased to
nutrition than that of wheat flour. However, maize protein, extremes of over 40%. This was basically a result of increased
like other cereals, is deficient in lysine. High-lysine mutants embryo size relative to endosperm.
(opaque-2) give more favorable amino acid balance suitable The maize endosperm, the source of starch and most of
for feeding people or monogastric animals even without the protein, has several interesting features. Maize types are
THE BASICS | Grain: Morphology of Internal Structure 49
classified by endosperm hardness. Floury is a soft endosperm binding needs to be loosened by soaking in a solution,
that can be easily crushed to a friable powder. It is physically such as in alkaline wet milling, where the physical properties
susceptible to storage pest (mainly insect) and fungal attack. of the starch may be affected. Care should be taken that the
An example of floury maize is the opaque-2 mutant. Dent has a fractions separated still achieve proper functionality in the
high proportion of floury endosperm, but some horny or hard intended use. For a useful further overview of the develop-
endosperm (Figure 7) and is the most widely produced com- mental aspects of cereal grain structure, the reader may refer
mercial maize. The dent refers to the indentation at the top of to Evers and Millar.
the grain on drying when it contracts due to the floury endo-
sperm component. Flint maize (of which popcorn is one
special type) has only horny and no floury endosperm. The Exercises for Revision
top of the seed is rounded, not indented, and it is physically
very hard, even after cooking meal prepared from it. Draw a typical cereal grain representing the major anatom-
The outer layers of maize (the seed coat in particular) ical features common to all cereal species.
contribute the color, which is white or yellow in nearly all Explain how starch, protein, and oil are stored in different
commercial maize production. Mass production of maize for cereal grain species.
feed and industrial use, for example, in the United States, is
predominantly yellow dent. Some white dent is produced for
breakfast cereal (e.g., corn flake) production (which follows
Exercises for Readers to Explore the Topic Further
maize dry milling). The carotenoids (xanthophylls) of yellow
maize are nutritionally important and also, in poultry feed,
contribute yellow color to egg yolks and to skin and fat of meat Investigate the concept of domestication for a selected
cereal grain crop, showing the necessary and desirable
poultry. People in regions where maize is a daily food staple
changes from wild relatives to modern cultivated plants.
may have strong color preferences; for example, in southern
Africa, white (dent) maize is strongly preferred because yellow Design an ideal maize grain for different processing appli-
cations (e.g., bioethanol and high oil), and indicate what
maize is historically associated with flintiness (although there
genetic approaches have been and could be taken to pro-
is no genetic relationship).
duce it.
The bran is often discarded after processing. What unusual
Maize Dry Milling and interesting variants in bran compositional features exist
This also requires a physically sound clean sample with appro- (e.g., in pigmentation), and how could these be better
priate hardness. Dry milling in the United States usually exploited for value-added uses?
implies use of white maize (a minor proportion of total US
production) to make endosperm chunks, which may be
steamed and flaked to make corn flakes. Larger dry milling
industries exist to serve the needs of maize tortilla production See also: Barley, Rice and Maize Processing: Barley: Malting;
(Mexico) and maize porridge production (a staple diet in Maize: Wet Milling; Beverages from Grains: Beverages: Distilled;
much of sub-Saharan Africa) requiring maize meal. Breeding of Grains: Maize: Breeding; Food Grains and the
Consumer: Fortification of Grain-Based Foods; Genetics of Grains:
Maize: Genetics; The Basics: Grain and Plant Morphology of Cereals
Barley Grain Structure and Effect on Malting
and how characters can be used to identify varieties; The Cereal
Grains: Rice: Overview.
The barley grain is similar in structure to the wheat grain. For a
diagram and further information on this topic, see elsewhere in
this encyclopedia.
The same principles apply when considering the structure Barber S (1972) Milled rice and changes during aging. In: Houston DF (ed.) Rice:
Chemistry and Technology, pp. 215263. St. Paul, MN: American Association of
and utilization of other grains, such as sorghum, millets, Cereal Chemists.
oats, or even the pseudocereals buckwheat and Amaranthus. Coffman WR and Juliano BO (1987) Rice. In: Olson RA (ed.) Nutritional Quality of
First, consideration should be the required product, flour, Cereal Grains: Genetic and Agronomic Improvement. Madison, WI: Agronomy
separation of starch/protein/oil, or malt, which drives the Society of America.
Dornez E, Holopainen U, Cuyvers S, et al. (2011) Study of grain cell wall structures by
choice of process wet or dry milling or malting. The size
microscopic analysis with four different staining techniques. Journal of Cereal
and shape of the grain, particularly the dimensions of the Science 54: 363373.
outer layers will mostly affect dry milling. In some cases, the Evers AD and Bechtel DB (1988) Microscopic structure of the wheat grain.
seed is extremely small (e.g., with Amaranthus) and dry mill- In: Pomeranz Y (ed.) Wheat: Chemistry and Technology, vol. 1, pp. 4795. St. Paul,
ing flour yield will be low. For such crops, selection for larger MN: American Association of Cereal Chemists 3rd edn.
Evers T and Millar S (2002) Cereal grain structure and development: Some implications
grain size can greatly increase flour yield. For wet milling, the for quality. Journal of Cereal Science 36: 261284.
fraction of most value (usually starch) will drive the design Hoseney RC (1994) Cereal Science and Technology, pp. 128. St. Paul, MN: American
of the separation process. Usually, the starchprotein Association of Cereal Chemists 2nd edn.
50 THE BASICS | Grain: Morphology of Internal Structure
Topic Highlights characters of each species are expressed, they share a general
commonality in growth habit and flowering parts. By under-
An introduction summarizing the definition of plant mor- standing the morphology of each species, it is possible to use it
phology, how it has evolved, and how it can be used to to identify the varieties (cultivars) that exist in each classification.
identify cereal varieties Recognition of the importance attached to identifying varieties is
A table giving a list, in order of importance, of the species highlighted by the existence of UPOV (The International Union
covered in the manuscript for the Protection of New Plant Varieties). The test guidelines put
A general section outlining the characters that are shared by together by the 71 members of UPOV provide useful informa-
the different cereal species tion of the major characters used in establishing the morpholog-
Specific sections for each species on general growth habit ical identification of the different varieties of cereal species.
and inflorescence The following section begins with a description of the main
How specific parts of a mature grain can be used to identify morphological characters common to all the cereal species and
many characters is followed by further sections specific to each species. Illustra-
Drawings and photographs to illustrate the text tions and photographs are used to help in the identification of
Identifies some of the differences between the Triticum the characters described.
species
(n)
(h) (g)
(e) (f)
(e)
(e)
(d) (d i)
(d ii) (c)
(b)
(c)
(b)
(a)
(b) (a)
(a)
Figure 1 (a) Typical growth habit of maize. (b) Typical growth habit of a cereal.
(d)
(c)
(b)
(d)
(a)
(d)
(d)
hairs present
rice (numerous)
wheat (sparse)
triticale (occasional)
(a)
(c)
Weak Strong
(a)
Weak Strong
(b)
Photo 1 (a) Flag leaf glaucosity in winter wheat. (b) Ear glaucosity in winter wheat.
54 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties
present, they encircle the stem and are thought to provide a high level of gray coloration have an extra layer of a waxlike
barrier within the node to prevent moisture and fungal patho- material (epicuticular wax) on the surface, giving the plant an
gens from entering. Generally, the stems develop as a hollow additional defense mechanism against moisture loss.
stem and continue elongating until the uppermost leaf appears Color differences due to a difference in glaucosity can be
(flag leaf), from which the inflorescence emerges. observed in different species and in different parts of the plant,
The number of leaves on the stem is dependent on the ranging from green to blue gray in the leaves and stem
eventual height of the plant with the ultimate leaf (flag leaf) (Photo 1(a) and 1(b)). It is also possible to observe different
usually smaller than the other leaves. The exterior surface of a levels of glaucosity on the glumes and grain. Anthocyanin
leaf is covered by the cuticle, a structure that helps prevent coloration (pigment), if present, is found in different levels of
moisture loss and the penetration of the external surface by intensity and at different locations within the plant, depending
dirt and harmful microorganisms. Those plants that display a on species (Photo 2(a)2(c)).
(a)
(b)
Photo 2 (a) Pigment on grain of barley. (b) Pigment on anthers of maize. (c) Basal leaf sheath barley.
THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties 55
(h)
(g)
(f) (b)
(d)
(a)
(e)
(a) glumes
(b) lemma
(c) rachis
(d) ovary
(e) lodicule
(f) stigmas (c)
(g) filament
(h) anther
Each species shares common ground in having an inflo- The structures are known as awns (barley) or beards (wheat,
rescence made up of a series of florets attached to a central durum, rye, and triticale) if they are present on each spikelet.
stem, which contain (in all species) the male and female Where they appear in a reduced form on the apex or upper part
reproductive organs (Figure 4). Wheat, rye, triticale, durum of a wheat ear, they are referred to as scurs. Once fertilization
wheat, and barley have a central stem (rachis) with a series has taken place and providing that no outside influence affects
of spikelets attached to it. Each spikelet is made up of the structure, the grains will go on to produce viable seeds.
several florets, and once fertilization has taken place, these Each caryopsis (grain) is made up of (i) the embryo that
florets develop into individual grains arranged alternately contains all the structures essential for development and (ii)
on the stem (Figure 5). The first rachis segment attaches the the endosperm that provides nutrients essential for the germi-
ear to the stem. Rice, maize, and oats share a similar inflo- nating seed, mostly in the form of starch. The whole is held in a
rescence with a series of branches forming from the upper- covering (pericarp) and remains dormant until the right con-
most node (panicle). Panicles can be one-sided (unilateral), ditions for germination are present, normally moisture and the
spreading, or triangular (equilateral) in appearance optimum temperature.
(Figure 6) with a series of florets lying along the branches.
Maize has a separate male and female inflorescence, where
numerous male florets (tassel) are to be found at the apex Species Specific Plant and Grain Morphology
of the plant and the female florets (silk) located within the
Rice (Oryza sativa) (Table 1(a))
stem node(s) (Photo 3).
All cereal species are able to self-pollinate with male and In terms of human consumption, rice is the most important
female flower parts on the same plant (monoecious), although cereal grown. There are two subspecies: O. japonica that pro-
pollen is accepted from other sources if their own pollen duces a short-grained sticky rice and is more tolerant of tem-
supply is scarce. Once fertilization has taken place, the front perate climates and O. indica that produces a longer-grained,
(ventral view) of the developing caryopsis is protected by the nonsticky type of rice that has a high moisture and temperature
palea, the back (dorsal view) by the lemma, which overlaps requirement and is more suited to tropical conditions. There
the palea at the side. In wheat and oats, the whole structure is are two main types: rice that grows on flooded land (paddy)
protected further by the glumes. In order to assist pollination, and deepwater rice that requires more than 50 cm of rain for at
two swollen structures (lodicules), on either side of the ovary, least 1 month (Photo 4). Deepwater rice varieties are capable
prise open the lemma and palea. Once fertilization has of floating in depths from 80 to 100 cm and, by having longer
taken place, the lodicules shrivel but remain attached to the leaves than the traditional tall paddy varieties, ensure that a
lemma base. supply of oxygen is available to sustain the whole plant. As the
For species other than maize, the developing grain may stems elongate, they are capable of producing their own roots
have a bristle of varying lengths extending from the lemma. (nodal roots) and becoming independent from the primary
56 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties
(c)
(b)
(a)
Figure 5 (a) Rachis segments of barley. (b) Single line of rachis segments of maize cob, showing truncation of the rachis.
one-sided panicle
(unilateral)
ear
e.g., oat
e.g., wheat
barley many-branched spreading panicle
e.g., rice
stem. The stems are especially adapted to contain a large num- three leaves. The adventitious roots develop from underground
ber of air cells that allow oxygen to pass through to the root stem nodes and form the basis of the root system. Whether the
system. A typical rice plant has a hollow stem and will attain an root or the shoot emerges first is determined on whether germi-
average height of 11.8 m. Depending on environmental nation takes place in aerobic conditions (when the root will
conditions, the fertility of the soil, and the variety, it can appear first) or in flooded conditions (anaerobic) when the
grow to 5 m in length. shoot will appear first. As the main (primary) tiller develops
Rice is the only cereal species that is germinated before and if growing conditions are favorable, a secondary tiller will
planting and set out once the seedling has a minimum of grow from the primary tiller, followed by a third (tertiary) tiller.
The earlier a variety matures, the shorter the plant and the
distance between internodes. A typical flag leaf encloses the
emerging panicle. The flower (floret) structure differs from
other cereals in having six well-developed stamens with short
filaments but follows other cereals in having a pistil with a
feathery stigma and lodicules (Figure 7). Depending on the
variety, the color of the inflorescence and the subsequent grain
can be gold, brown, light to dark red, purple, or black. Virtually,
all varieties show basal leaf sheath coloration and colored
auricles, which can extend into the lemma tip and keel.
Table 1 List of the cereal species covered by this article in order of their importance worldwide. The common name is given first, followed by the
Latin declination and a further column, which highlights additional information on the usage of the species. It will be noted that in terms of human
consumption, rice is the most important cereal but that maize is the most important in terms of worldwide production
Species (common
name) Latin name Usage
(1(a)) Rice Oryza sativa (subsp. O. indica and Most important cereal in terms of human consumption
O. japonica)
(1(b)) Maize Zea mays Most important in terms of worldwide production but not all the grain produced goes for
human consumption
(1(c)) Wheat Triticum aestivum Second to rice in terms of human consumption. Both winter and spring types exist
(1(d)) Durum Triticum durum Second most important Triticum species after wheat
(1(e)) Triticale Triticale (X Triticosecale) Hybrid of wheat () and rye () has the yield of wheat with the additional advantage of
tolerance to diseases and environmental changes
(1(f)) Rye Secale X cereale Suited to much poorer soils than wheat and durum
(1(g)) Barley Hordeum vulgare Ranks fourth in importance and used primarily as a food source and as a raw material in
the brewing/distilling industry
(1(h)) Oats Avena sativa A higher tolerance to cool conditions and rainfall
58 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties
(f)
(g)
(d i)
(e)
(c)
(d)
(a) pedicel
(b) glumes
(c) palea Photo 5 Brace roots of maize.
(d) lemma (b)
(d i) lemma nerves
(e) feathery stigma
(f) six anthers
(a) style. Pollen is released from the male anthers in the tassel and
dispersed by the wind (anemophilous dispersal). As a result of
Figure 7 Typical floret of rice. the relative heaviness of the pollen, it drops the short distance
to the silk below or onto a neighboring plant. Once fertiliza-
tion has taken place, each individual maize grain develops on
Maize (Zea mays) (Table 1(b)) the shortened rachis, the whole structure forming the cob
(Photo 7). The color of the cob is related to the variety and
In terms of the volume grown, maize is the most important
can vary from white, to yellow, to orange, to red, to purple, to
cereal species, but much is used for purposes other than food,
brown, to black, to variations of the color range within the cob.
which places it second to rice in terms of worldwide staple food
consumption. It displays major differences when compared
with the other species covered in this article. The main stem
Wheat (Triticum aestivum) (Table 1(c))
grows to 1.23 m in height. In the early stages of development,
a mass of underground adventitious roots are produced just Wheat is grown in all the temperate climates of the world and
below the soil surface, and as the plant grows, a series of is second to rice in terms of human consumption. True winter
additional brace roots (Photo 5) emerge above the surface wheat varieties require a period of cold to initiate the repro-
from the two lowermost leaf nodes. Although the brace roots ductive stage of development (vernalization). Spring wheat
have the facility to grow quickly and provide stability for the varieties do not need a cold period for ear formation although
plant, maize is still very susceptible to wind damage. Depend- they can respond to cold by flowering earlier. Wheat has a
ing on the eventual plant height, an average of twenty alternate higher protein content than any other major cereal, and a
leaves will develop up the stem. further distinction is made in the type of flour produced.
Maize is classified as a hermaphrodite plant in having the Hard milling varieties produce flour suitable for bread produc-
male and female reproductive organs sited in different loca- tion. Soft milling varieties are used in the production of com-
tions on the same plant. The staminate (male) inflorescence modities such as animal feed or biscuits. Because winter wheat
(tassel) develops at the apex of the plant, normally after the has a longer growing season, they are usually denser in ear
emergence of the 14th or 15th leaf, as a panicle producing a formation than spring wheat (Photo 8). The average height of
series of spikelets in which fertile and sterile florets sit. The wheat is 75150 cm, and unlike other cereal species, the stems
pistillate (female) inflorescence (silk) lies normally in the leaf can be solid and hollow (Photo 9).
axils of the 11th13th leaves (Photo 6(a) and 6(b)) with pairs The wheat ear is made up of a series of spikelets arranged
of spikelets sitting on a much truncated rachis. From each silk, alternately on the central rachis and each one is made up of a
a series of tubelike structures emerge with small leaves, which cluster of six florets (Figure 8). Close study will identify the
will form the covering of the cob at maturity (husk leaves). At different structures that form the spikelet, the most important
one end is the undeveloped grain and at the other a feathery being the lower glume, which lies opposite the lemma with the
THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties 59
Tassle Silk
The wheat grain can be separated by shape into those that Durum Wheat T. turgidum (syn T. durum) (Table 1(d))
are round, elliptical, or ovate. At the opposite end to the
Commonly known as durum wheat and developed through
embryo, brush hair may be identified. If present, a distinction
artificial selection many thousands of years ago, it is the
can be made as to whether the hair is short or long in length
hardest milling variety of all the wheat species and used
(Figure 10).
extensively in the production of pasta. It grows well in coun-
tries bordering the Mediterranean where precipitation is low
and temperatures high. Similar in height (1.21.5 m) and
appearance to bearded wheat, but usually with a shorter
denser ear (Photo 10), durum wheat is characterized by
having well-developed, firm glumes and a pronounced keel
along the glume length (Figure 11(a)). The lemmas have a
rounded appearance due to the absence of a keel. The yel-
lowish, brown-colored caryopsis is often ridged. Brush hairs
are either absent or very short (Figure 11(b)).
(e)
Floret 6
Floret 5
Floret 4
(d ii)
(d i)
Floret 1
(c ii) Floret 2
(c i)
(f)
(a)
Extent of glume beak
Extent of shoulder
(b)
(d)
(e)
(f)
(c)
(b)
(b)
(a)
(a)
(a)
Caryopsis
Glume
elliptical, often
long and narrow ridged, brush hairs
firm, strongly keeled short or absent,
throughout yellow/brown in
color
(b)
Figure 11 (a) Triticum turgidum (syn T. durum) spikelet. (b) Triticum turgidum, glume and caryopsis.
64 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties
(a)
(b)
Glume Caryopsis
Note: strongy keel, may have Elliptical, often ridged, brush
weak hairs hairs present. (a) prominent
(b) embryo greyish brown in color
(a)
(a)
(b)
(a)
(a)
Caryopsis
(b) (a)
Figure 13 (a) Spikelet of rye (Secale X secale). (b) Rye (Secale X secale).
66 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties
(b)
(a)
(a) basal leaf sheath hair
(b) hairs on base of leaf
blade
Figure 14 Barley plant showing basal leaf sheath hairs, and base of leaf blade.
THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties 67
At the base of the grain and lying within the ventral furrow is observed on the dorsal side at the awn end, where it is
is a small rodlike structure covered in hairs (rachilla) and used possible to identify very small protrusions that create a rough
extensively in the identity of varieties. In some varieties, there surface on the areas between the lateral and median nerves
are an additional three distinct linked characters to observe in (sharkskin). It should be noted that this area may also be
barley grain. It is rare for these characters to be separated, so if associated with many rows of spicules.
one is present, then the other two will also be present. They are
fence hairs (ventral view) that lie in a vertical position at the
base of the grain and ventral furrow hairs that are extremely
small white hairs lying along the edge of the ventral furrow and
often extending beyond the rachilla. The third linked character
2 row 6 row
(e)
(a) lemma
(b) palea
(c) ventral furrow
(d) lemma base (attachment
point to rachis)
(e) broken off awn
(f) rachilla
(c)
(g) gape may not always be
present
(h) inter glume hair
(a)
(b)
(g)
(f)
(h)
(d)
Figure 15 Ventral view of grain of barley.
THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties 69
(a)
(b)
(g)
(c)
(d)
(d)
Conclusion
Photo 17 Barley: dorsal view showing spicules on inner lateral nerves. From the earliest times, man recognized the different cereals as
an important food source. The crops they produced were sub-
ject to continual change and selection over the decades result-
ing in the gradual improvement of yield and quality. At the
same time, others were recognizing similarities between plants
and evolving a system that placed them into taxonomic groups.
The more detailed and precise the study of plants and their
morphology became, the more demand it placed on using a
greater range of identifying morphological characters. It was
this assessment of morphological characters that gave the dis-
criminatory evidence essential for laying the foundation for
varietal identification, which could in turn be used for estab-
lishing variety ownership through the introduction of plant
variety rights. Modern techniques such as the use of DNA
fingerprinting are already being used to determine varietal
identity, but for the purposes of UPOV and plant breeders
rights, they can only be used as an addition to the use of
morphological characters.
Equilateral Unilateral
(spreading) (one sided)
Spikelet structure
(a) glumes
(b) primary grain
(a)
(c) secondary grain
(d) awn
(b)
(e) basal callus hair present, 1-2 per
side of in tufts
(f) lemma
(g) palea
(h i) rachilla (note length)
(h ii) rachilla medium long in length
(i) basal callus (d) (c)
(i)
(e) (i)
(h i)
(h
ii)
(g) (f) (f) (g)
Exercises for Readers to Explore the Topic Further See also: Grain Marketing and Grading: Identification of Varieties
of Food Grains; The Basics: Grain: Morphology of Internal Structure;
Currently, variety descriptions are produced using methods Taxonomic Classification of Grain Species; Wheat-Based Foods:
that have remained largely unchanged since their introduc- Cereals: Overview of Uses: Accent on Wheat Grain.
tion. Some biotechnological methods have now been
incorporated into existing systems. Using a species with
which you are familiar, identify any available biotechnol- Further Reading
ogy methods and undertake research to assess the advan-
Detailed descriptions of characters for varietal identification can be found in the UPOV
tages that could be gained by their inclusion.
Guidelines under the following headings: (upov.int/test guidelines/choice of
Could varietal identification be carried out partly or wholly language). The guidelines are available worldwide and provide a standardized
by biotechnology? format for the characters used in varietal identification.
72 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties
Barley (Hordeum vulgare) UPOV TG/19/10. NIAB: Botanical descriptions of varieties info@niab.com.
Durum wheat (Tritium turgidum syn T. durum) UPOV TG/120/3. Oat (Avena sativa) UPOV TG 20/10.
Hervey-Murray CG (1980) The Identification of Cereal Varieties. Cambridge: Cambridge Rice (Oryza sativa) UPOV TG16/8.
University Press. Rye (Secale x secale) UPOV TG58/6.
HGCA: Milling wheat: quality control criteria and tests. The biology of maize www.ogtr.gov.au.
Ikisan: morphology of rice. The biology of Triticum turgidum www.inspection.gc.ca/triticum turgidum.
Ikisan: morphology of maize. Triticale (triticale) X triticosecale UPOV TG/3/11.
Institute of rice research www.irri.org. Wheat (Triticum aestivum) UPOV TG/3/11.
Maize (Zea mays) UPOV TG2/7.
The Cereal Grains
Contents
An Overview of the Family of Cereal Grains Prominent in World Agriculture
Cereals: Domestication of the Cereal Grains
Maize: Overview
Wheat: An Overview of the Grain That Provides Our Daily Bread
Durum Wheat: Overview
Rice: Overview
Wildrice, Zizania: Overview
African Rice (Oryza glaberrima): A Brief History and Its Growing Importance in Current Rice Breeding Efforts
Barley: An Overview of a Versatile Cereal Grain with Many Food and Feed Uses
Sorghum Grain, Its Production and Uses: Overview
Rye Grain: Its Genetics, Production, and Utilization
Triticale: Overview
Oats: Overview
Coix: Overview
Millet Pearl: Overview
Millet Minor: Overview
Teff: Overview
Cereal grains are produced by flowering plants of the grass Colloquially, we often use the word cereal to mean a breakfast
family. cereal, such as cornflakes. The manufacture of cereal in this
Wheat, rice, and maize (corn) are the major cereals, each narrow sense is described in this encyclopedia, but more
produced in quantities of about 600 million tonnes annually. broadly, the term cereal applies to the many grain genera
Other cereals are barley, rye, triticale, oats, sorghum, and that are flowering plants of the grass family (Poaceae or Gra-
the millets. mineae), whose seeds are used as food. It may be difficult to
Cereal grains have a high proportion of starch, present as imagine wheat, for example, as a flowering plant, but wheat
distinct granules in the floury endosperm. flowers are illustrated in Figure 1. (And do not get confused
Cereals are a valuable source of protein; their low fat con- between this flower and the flour of similar-sounding name,
tent is mainly in the germ (embryo), from which the new meaning crushed grain.)
plant emerges during germination.
Processed foods from cereals include a wide range of bread
types, pasta, noodles, breakfast cereals, beer, and porridge. Seed or Grain?
Cereals are major sources of animal feed (grain and plant)
and fuel for transport. The name cereal derives from Ceres, the Roman goddess of
grain. The words seed, kernel, and caryopsis are also used to
denote cereal grain. In addition, the term corn is sometimes
used for the grains of cereal species in general, but more often
Learning Objective (especially in the United States), the word corn denotes the
cereal species maize, also known as Indian corn. Wheat and
To achieve understanding of the agriculturally significant barley seeds are shown in Figures 2 and 3, as they appear by
species and uses of cereal-grain family. scanning electron microscopy.
Figure 3 Scanning electron micrographs of (a) a barley grain, covered by the integrated glumes (lemma and palea); (b) the rachilla lying in the
groove of the crease; (c) the base of the rachilla with many of the long hairs broken off as a result of grain handling; (d) dust from the movement
of barley grain showing the spear-like rachilla hairs. Note the different degrees of magnifications: the black bar at the bottom right represents a length of
(a) 1 mm, (b) 400 mm, (c) 40 mm, and (d) 100 mm.
Historical Perspective A porridge left overnight would gain wild yeasts, leading to
the discovery of leavening as a means of making the baked
Cereal grains have the great distinction of being the catalyst product lighter. Further refinements would have been the siev-
that transformed humans, as the hunter-gatherers into agricul- ing step, to remove husks and bran particles after grinding, and
turalists no longer living as nomads. The origins of civilization the identification of wheat as the ideal grain for the baking of
involved the enormous step of primitive mankind discovering leavened bread due to its dough-forming properties.
that it was possible to remain in one place where cereal seed, There is archaeological evidence that primitive baked prod-
sown intentionally, would yield a reasonably reliable source of ucts were being made in the late stages of the Stone Age.
food, which could be stored indefinitely. As a result, people Purpose-built ovens were used in ancient Egypt as far back as
found that a fixed dwelling place could be established near 2700 BC, with reasonably sophisticated milling and baking a
cereal crops. Further consequences were the development of millennium later. As a result of this long history of cereal-grain
permanent forms of housing, the domestication of animals use, diverse uses have been developed worldwide for their
(fed with cereal grain), and the spare time to develop the processing, taking advantage of the specific characteristics of
many other characteristics of civilization. the individual cereal species (see Table 1 and Figures 5 and 7).
Early consumption of cereals involved merely chewing the
raw grain. Primitive forms of processing were developed, such as
grinding the grain between stones and soaking the grain in water World Production of Cereals
to make it easier to chew. Next, heat was discovered as a further
means of easing mastication and digestion, producing a porridge Cereal grains are grown extensively in all continents, with the
or gruel. Placing a stiff porridge in the hot stones or ashes of a fire obvious exception of Antarctica. They are the major sources of
would have made it more palatable, resulting in baked foods. food for mankind, especially in developing countries. In
76 THE CEREAL GRAINS | An Overview of the Family of Cereal Grains Prominent in World Agriculture
Brush
Aleurone
cells
Cuticle Storage
protein
Epidermis
Hypodermis Cell nucleus
Cell wall
Thin-walled
cells
Endosperm
Cross cells
Testa Starch
granules
Nucellar
epidermis Figure 5 Some of the many foods made from cereals (mainly wheat).
Figure 6 Grains of major cereal species: (a) bread wheat; (b) durum wheat; (c) triticale; (d) rye; (e) barley; (f) oats; (g) sorghum; (h) corn; (i) Japanese
millet. Adapted from Fitzsimmons RW and Wrigley CW (1984) Australian Barleys: Identification of Varieties, Grain Defects and Foreign Seeds, 2nd edn.,
pp. A11A13. Melbourne: CSIRO.
of milled rice, annual world production averages over 500 (production averaging about 150 million tonnes annually), sor-
million tonnes. International trade in rice is much less than ghum (50 million tonnes), and oats (30 million tonnes).
for wheat and maize. Rice, together with maize, provides the
major energy and protein source for many cultures.
Annual production of maize (950 million tonnes in 2013/ Morphology of Cereals: What They Look Like
14) is greater than that of wheat. Statistically, maize is included
in coarse grains, production of which totals 1.2 billion tonnes Cereal grasses have narrow leaves, hollow jointed stems, and
annually (Table 2). World trade in coarse grains is about 100 spikes or clusters of membranous flowers (Figure 1). Figure 9
million tonnes. In addition to maize, coarse grains include barley shows how the mature grain of wheat is held within glumes on
78 THE CEREAL GRAINS | An Overview of the Family of Cereal Grains Prominent in World Agriculture
Common
names Genus and species Uses
Pooids
Common Triticum aestivum Breads (leavened and unleavened), noodles, breakfast cereals (processed porridge), cakes, cookies,
wheat chapatis, snack foods, bulgur, ingredients for a wide range of foods, animal feed, industrial uses
Durum Triticum turgidum Pasta (e.g., spaghetti and macaroni)
wheat subsp. durum
Rye Secale cereale Bread (often together with wheat), crispbread, spirits
Triticale Triticosecale Bread (often together with wheat), animal feed
Barley Hordeum vulgare Malt, beer, spirits, porridge, cooked grains (e.g., soup ingredient)
Oats Avena sativa Porridge (as rolled oats, oatmeal), muesli, oatcakes, animal feed
Oryzoids
Rice Oryza sativa Cooked and fried grain, sake, extruded snack foods
Wild rice Zizania aquatica Cooked grain
Chloridoids
Teff, Indian Eragrostis tef Cooked grain, porridge
millet
Finger millet Eleusine coracana Cooked grain, porridge
Panicoids
Corn Zea mays Polenta, breakfast cereals, tortilla, cornbread, porridge, snack foods, a wide range of industrial uses
(maize) (e.g., corn syrup and food ingredients)
Sorghum Sorghum bicolor Animal feed, cooked grain, porridge, beer, flat cakes
Pearl millet Pennisetum glaucum Cooked grain, porridge
Common Panicum miliaceum Cooked grain, porridge
millet
Japanese Echinochloa species Cooked grain, porridge
millet
Foxtail millet Setaria italica Cooked grain, porridge
The flowers of oats and rice, on the other hand, are arranged
in an inflorescence called a panicle (Figure 11). The grains of
oats and rice do not thresh as the free seed. The term groat
applies to the oat seed with the glumes removed. The term
paddy rice, or just paddy, refers to the rice grain with its
husks in place, the form in which it is harvested. The familiar
cob (ear) of corn (maize) is distinct from the seed-bearing
structures of the other cereals described earlier (Figure 12).
Table 2 World production in the year 2013/14 of cereal grains by countries/regions, in millions of tonnes (metric tons)
Argentina 11 35 1 47
Australia 26 11 1 37
Brazil 5 73 8 86
Canada 33 26 59
China 121 218 142 480
EU 143 159 2 304
India 93 42 105 240
Indonesia 9 38 47
Mexico 3 29 33
Pakistan 24 4 6 34
Russia 52 36 1 88
South Africa 2 14 15
Thailand 5 21 26
Turkey 18 13 1 32
Ukraine 22 38 60
The United States 58 372 6 436
World 706 1252 473 2432
a
The category coarse grains includes maize, barley, rye, triticale, and oats.
b
Calculated as milled rice (not paddy).
Source: Data from United States Department of Agriculture.
2300 600
2100 400
Awn
Grain
1700 0
2003/04 2005/06 2007/08 2009/10 2011/12 2013/14
fcast Lemma
Figure 10 (a) Heads of wheat. (b) Heads of two-row barley. (c) Heads of six-row barley.
Spikelets
Branch
Rachis
Neck Like wheat, rice is another cereal with a special place in mans
diet, providing a dietary staple for over half of the worlds pop-
Flagleaf ulation, thus constituting their major source of energy and pro-
Top ligule tein. Unlike other cereals, rice is usually consumed as the whole
grain, after removal of the bran layers. Rice can be grown under a
wide range of climates, either in flooded irrigation or under
Flagleaf
sheath dryland conditions. Some 90% of the worlds rice production is
Uppermost
stem node in Asia, especially in developing countries. International trade in
rice accounts for only about 4% of world production.
Prices vary greatly depending on the type of rice. For exam-
ple, Basmati scented rice from Pakistan and northwest India
Figure 11 The head (panicle) of oats. Reproduced from Fitzsimmons RW, might command four times the domestic price of ordinary
Roberts GL, and Wrigley CW (1983) Australian Oat Varieties: Identification of rice. Quality grades of rice are partly based on dimensions
Plants, Panicles and Grains, p. 11. Melbourne: CSIRO, with permission. long-, medium-, and short-grain rice. Quality attributes relate
THE CEREAL GRAINS | An Overview of the Family of Cereal Grains Prominent in World Agriculture 81
Barley
Maize (Corn)
The main uses of barley (Figures 3 and 6) are for animal feed
Maize, indigenous to the Americas, is produced worldwide and for the production of pearled barley, malt, and beer
mainly in tropical and warm regions. It is an annual plant with (Table 1). These two uses (feed and food) form the basis of
82 THE CEREAL GRAINS | An Overview of the Family of Cereal Grains Prominent in World Agriculture
the two main grades for trade in barley (known as feed barley Rye is a popular human diet in Europe. The production of rye
vs. malting barley). Another major distinction is based on head in Germany exceeded that of wheat from about 1940 to 1960,
morphology, namely, two row or six row depending on but more recently, the production of wheat has increased
whether there are grains in the full set of florets in the head considerably, and a higher proportion of the rye crop has
(six row) or if there is only one set of grains on either side of the been used for animal feed.
head, with the other florets sterile (Figure 10). In some barley- Rye is particularly susceptible to contamination with ergot,
growing countries (Europe and Australia), six-row barley vari- a toxic fungus that replaces the grain in the rye head during
eties are often of feed quality, but six-row varieties are also of development, so that a long black ergot body protrudes from
malting quality in North America. the mature head. Ideally, the production of ergot in the crop is
Barley is the grain used for Scotch and Irish whiskies. In avoided, but there are techniques for the removal of intact
addition, a proportion of the barley crop is used directly for ergot bodies prior to milling.
human consumption; after pearling (abrasion) to remove the In addition to the manufacture of rye breads, rye-based
adhering lemma and palea, the barley grain is boiled in soup as crispbread is an important use of rye, generally with rye whole-
the whole pearled grain, or it is coarsely ground and cooked as meal or from flaked rye. Other baked products include Ryvita
gruel. Barley flour is also used for a range of baked products, and pumpernickel. In human nutrition, the rye grain offers an
but barley is not suitable for the production of leavened bread, unusually high level of pentosans. This source of soluble fiber
lacking the gluten proteins of dough-forming quality. Never- offers advantages of giving a feeling of satisfaction, thus help-
theless, barley appears to have been used for bread making by ing in slimming diets, as well as reducing the rate of rise in the
ancient civilizations. Barley, for example, appeared on Greek blood sugar level after ingestion.
coins several centuries BC, and it was the staple diet of Roman Triticale is a man-made crop that combines the genomes of
gladiators, who were thus known as hordearii (reflecting the wheat and rye (Table 1). Most cultivated triticales are hexa-
origins of the genus name, Hordeum). ploid types, containing the genomes of durum wheat and rye.
It thus tends to be intermediate between rye and wheat in its
grain composition and characteristics. The expectations for the
Oats
grain have been that it would contribute the disease resistance
The oat grain and plant and grain are illustrated in Figures 6 and growth hardiness of rye to wheat while also combining the
and 11. Oat plants thrive in temperate climates, with cooler baking quality of wheat with the unique flavor of rye bread.
moister conditions, compared to other cereals. Oats are often Triticale has thus been used for baking; nevertheless, its main
used as fodder for stock, grazing off the growing plants. In such use is for animal feed.
cases, it may also be possible for the farmer to harvest a crop of
grain, which in turn may also be stored on-farm for stock feed. Sorghum and the Millets
In addition, oats may be grown primarily as a grain crop,
The grains of sorghum and of various millet species (Table 1,
generally using varieties that have been bred specifically for
Figure 6) constitute a major source of protein and energy for
use as grain. A large proportion of harvested oats is used for
many people in Africa and Asia, where these grains are used as
animal feed, especially for horses.
porridge, boiled in water after grinding. In such situations,
Compared to the other cereal grains, the oat grain tends
these cereals offer the advantage for subsistence farming of
to have higher contents of protein, lipid, and soluble fiber
tolerance to drought and suitability for tropical growth envi-
(b-glucan). Oats are reported to have beneficial hypocholester-
ronments. Pearl millet is the most widely grown of the millet
olemic properties and are useful in the management of the
family, with an estimated area worldwide of about 30 Mha.
insulin response in diabetics. For human consumption, the
Second in importance are common millet and finger millet
adhering glumes are abraded from the oat grain. The resulting
(Table 1). Many millets, due to small grain size, are used as
groat is cut, rolled, or ground to yield products such as oatmeal,
feed for wild and domesticated birds.
rolled (flaked) oats, and oat flour, for processing into instant
In addition to the use of sorghum in cooking, it is also dry-
oats for porridge, oat cakes, breakfast cereal, and infant foods.
milled and fermented for various forms of beer. On the other
The groat has a higher content of lipid (fats) than wheat, plus
hand, the main use of sorghum in developed countries is for
significant lipase activity (fat-splitting enzyme), and these fac-
animal feed and, to a lesser extent, for industrial processing.
tors must be taken into account in oat processing. There is the
There are various mutant forms of sorghum with distinct char-
consequent risk that after milling, fat rancidity, bitterness, and a
acteristics that include high lysine content, waxy starch, high
soapy taste will be produced by the actions of various endoge-
sugar content (stalks), and even scented endosperm. The grain
nous enzymes, namely, lipase, lipoxygenase, and peroxidase. To
of many sorghum varieties is pigmented, largely due to tannins
prevent this, it is usual for milled oats to undergo some form of
located in the outer layers (testa) of the grain. The high-tannin
heat treatment to inactivate the lipase activity. This is often done
varieties are also somewhat less nutritious, because the tannins
by steam treatment, together with kiln drying.
interfere with protein digestibility. Some sorghum varieties
have bitter flavors, which confer resistance to attack by birds.
Rye and Triticale
Rye is the only cereal grain that approaches wheat with respect Composition
to bread-making properties, although the achievement of
bread quality in rye breads usually requires the incorporation Starch, a polymer of glucose, is the major component of the
of a considerable proportion of wheat flour (Table 1, Figure 6). endosperm of cereal grains (Table 3). It is the stored form of
THE CEREAL GRAINS | An Overview of the Family of Cereal Grains Prominent in World Agriculture 83
Table 3 Approximate composition of the major cereal grains, as percentages of dry weight of whole grain, except for rice
Figure 16 Pollen grains from the flowers of cereal species, revealed by scanning electron microscopy.
species distantly related to the species of interest. Novel genetic resistance, drought tolerance, and insect resistance. The various
technologies (genetic modification (GM)) facilitate these cross aspects of improved grain composition include increased levels
species transfers, thus augmenting conventional breeding, of specific vitamins (e.g., vitamin A) and minerals (e.g., iron)
leading to new varieties that are better at surviving stresses, and a reduced level of gluten-type protein in barley, permitting
plus providing novel grain qualities. the production of gluten-free beer.
GM technologies have been more successful in commercial The composition and functional properties of the edible
practice for noncereal species, especially soybean, cotton, and grains offer a wide diversity of possibilities for new and varied
canola. Nevertheless, GM corn is grown widely, especially in end uses. Given this genetic diversity, and the consequent
the United States, accounting for about one-third of the area access to the genes responsible, combined with the capabilities
worldwide for GM crops. of novel methods of genetic manipulation, we have many
For other cereal species, GM and conventional breeding possibilities for extending the compositional diversity of any
offer useful traits, such as abiotic stress tolerance, pathogen grain to provide it with new functional properties.
THE CEREAL GRAINS | An Overview of the Family of Cereal Grains Prominent in World Agriculture 85
Anther
Stigma
Stamen
Pistil
Style
Filament
Ovary
Lodicule
Rachilla
Palea
Lemma
Upper (second)
glume
Lower (first)
glume
Palea
Lemma
Rachilla
joint Callus
Figure 1 Stylized drawings of a grass flower (upper left), floret with adjacent rachilla internode (lower left), and spikelet with subtending rachis
internode (right). Reproduced with permission from Clark LG and Pohl RW (1996) Agnes Chases First Book of Grasses. Smithsonian Press.
The floret is attached to a secondary stem axis, the rachilla. Pollination, Fertilization, and Grain Development
Subtending the rachilla are two empty bracts, the glumes. One or
Cereal species can be either self- or cross-pollinating. In the
more florets, packaged within the glumes, form a spikelet
latter species, the lodicules swell at anthesis, forcing the florets
(Figure 1). Spikelets are alternately positioned at attachment
open to expose the stigmatic surfaces of the pistil to incoming
points (nodes) located along the main inflorescence axis (rachis)
pollen. While the florets are open, the stem-like filaments on
or on branches extending off the rachis. The interval between
the stamens elongate to push the anthers upward and out of
two nodes on the rachis is the internode. Within the spikelets,
the floret where they dehisce and shed pollen into the wind
florets are similarly arranged, alternately along the rachilla.
and air currents. The glumes can bend outward at their bases,
Although grass inflorescence morphology is complex, a sim-
enabling a wider opening of the florets. For self-pollinating
plistic classification designates three basic types (Figure 2),
cereal species whose florets do not stay tightly closed at anthe-
each differing in rachis branch development and spikelet
sis, cross-pollination can occur at low levels. This functional
attachment: (1) panicle with long, open branches or dense
morphology of the floret and spikelet at play during pollina-
branches, each bearing a pediceled spikelet; (2) raceme with
tion sets the stage for the hybrid speciation that characterizes
spikelets borne on short pedicels along the rachis; and (3)
cereal evolution.
spike with spikelets sessile along the rachis.
88 THE CEREAL GRAINS | Cereals: Domestication of the Cereal Grains
Diaspores generally move only a short distance from the mother Domestication Syndrome
plant unless aided by animals, humans, or water.
Under domestication, features of dispersal, reproduction, plant
Awns, which are extensions of the vascular nerves of
growth, and survival of wild grasses were transformed into
glumes and lemmas, can move the dispersal units in a variety
desirable agricultural traits designated as the domestication syn-
of ways. Natural humiditydrying cycles can cause awns to
drome. The two leading transitions were: (1) the retention of
spread open and close, thereby moving the diaspore by pas-
the mature grain on the mother plant due to the development
sive mechanical force with the seed end pointing forward for
of a nonshattering, tough rachis and (2) the easy release of the
short unidirectional distances or down into the soil (wild
grain from the bracts due to morphological changes reducing
wheat: Triticum urartu). Other hygroscopic awns will coil
or eliminating glume tenacity and adherence of the lemma and
and uncoil in response to moisture changes, thereby drilling
palea to the mature grain. Following these initial changes,
the diaspore into the ground (wild oat). Tiny barbs on awns
humans manipulated a suite of desirable physiological and
or bristles can ratchet the movement of diaspores across the
agronomic traits to further advance early forms into fully agri-
ground for short distances, lock the diaspore in place in the
cultural species:
soil, or catch the diaspore on the fur of animals or clothing of
humans. Awns can also vibrate in the wind to push the (1) Reduction of seed dormancy either by loss of physiological
diaspore downward into the soil or soil crack. Lightweight, dormancy or by loss or reduction of enclosing bracts that
feathery bristles of a fascicle can move a diaspore in the wind contained germination inhibitors
(wild pearl millet). (2) Increase in seed production by development of reproduc-
In contrast to their wild relatives, domesticated cereal species tively functional florets from sterile or vestigial ones
retain the mature caryopsis on the plant and have more easily (3) Increase in seed size
extractable grains due to reduced or absent lemmas, paleas, and (4) Synchronous development of tillers and ripening of the
glumes. Thus, humans must remove the mature grains from the grain
mother plant by hand or mechanical harvesting, thresh the (5) Changes in photoperiod sensitivity
harvested heads to free the grain from their enclosing bracts, (6) Greater agricultural fitness for soil and climatic conditions
protect the grain from predation and environmental threats, and pest resistance.
and plant it for a new crop. Some of the more primitive cereal
species still retain wild-type features. For example, semibrittle The speed and spread of the transition to agriculture is still
forms of emmer and spelt wheat (T. dicoccum and T. spelta) can debated with differing interpretations of domestication as
naturally disarticulate or break under relatively little mechanical rapid versus gradual and spread of initial domesticates as an
pressure into spikelet diaspores enclosed by tough glumes. outward diffusion from a primary place of origin versus a series
of relatively concurrent events in different locales.
Reduction or full loss of the rachis nodal abscission zones is
Evolution and Domestication responsible for the tough rachis trait. Semishattering forms
naturally break into diaspores in the upper portion of the
Domestication constitutes an evolutionary change and specia- inflorescence or break when pressure is applied in partially
tion mediated by humans acting as selection agents. For the formed abscission zones. Changes in bract morphology led to
cereal grains, early humans channeled the inherent genetic free-threshing cereals. The tough tenacious glumes softened or
variation of wild grasses in the direction of agricultural traits became more brittle, losing their clasping feature that enclosed
that eased work to cultivate, harvest, and extricate grain. Then, the spikelet at maturity. Lemmas and paleas lost their tendency
through their ancient migrations and trade routes, they spread to adhere to the mature grain. Size reduction, anatomical thin-
domesticated cereals and associated weed species beyond the ning, and increased brittleness of the bracts allowed them to
original sites of domestication. With the expansion of early easily fall away with mechanical threshing to release a naked
agriculture, opportunities opened for hybrid speciation events grain. The term hulled cereal refers to different protective features
between otherwise allopatric species or between weedy wild of the bracts that have not completely lost wild-type features:
forms and crop species. These expanded associations led to Hulled grains either are encased within the lemma and palea,
new domesticated forms (for example, T. aestivum) or intro- which either form a hard shell around the grain (pearl millet) or
gressed forms with more complex phylogenies (maize, rice, and adhere to the grain surface (barley); hulled spikelets have hard-
sorghum). Ancient farmers further exploited the domestication ened, tough glumes that tightly enclose one or more grains in a
process by developing unique land races adapted to the grow- spikelet package (spelt wheat, that is, T. spelta).
ing conditions and food needs of their locales. As a result of
these factors, discerning phylogenetic relationships can be chal-
Domestication Parallels: Crop and Weed Races
lenging. Thus, taxonomic classifications constructed to reflect
phylogenetic evidence can be dynamic, changing as the evi- Domestication followed different routes for each cereal species
dence changes. The practice of classifying the wild and domes- and did not necessarily involve a straightforward, one-time
ticated cereal forms as one biological species, usually named transition from wild to domesticated species. Introgressive
with the domesticated species name, can lead to confusion if hybridization has played a significant role and followed a
the wild versus domesticated forms at the infraspecific levels of variety of pathways. Domesticated races intercrossed with
subspecies or variety are not specified (for example, T. turgidum each other (rice), with weedy wild relatives (barley), or with
and Sorghum bicolor can refer broadly to both wild and domes- wild species related to the original wild progenitor (sorghum,
ticated taxa). pearl millet, and maize). Polyploid speciation in the wheats
90 THE CEREAL GRAINS | Cereals: Domestication of the Cereal Grains
and oats illustrates how new genome material introduced a staple, barleys greater adaptability to poor soils and tolerance
through interspecific hybridization formed new species. These of soil salinity increased its culture in areas and climatic condi-
types of events increased the narrowed diversity arising from tions unfavorable to wheat. The barley spike (or spike-like
the domestication bottleneck, thereby expanding the adaptabil- raceme) has three spikelets at each rachis node. Spikelets con-
ity of domesticated forms and their distribution ranges. tain one floret. Their row-like appearance along both sides of
The common element among wild cereal progenitors is the spike and floret fertility is the basis for dividing domesti-
their amenability to cultivation in human-modified habitats. cated barley into two groups: (1) two-rowed barley with a
Along with domesticated forms came weeds, the unwanted central fertile spikelet and two adjacent sterile spikelets, each
species that also thrive in these habitats. The development of containing a male floret (primitive form; subsp. distichum)
weed races parallels crop evolution from the standpoint of and (2) six-rowed barley with all three spikelets fertile
unintended human selection of weedy forms through agricul- (advanced form; subsp. hexastichum). Most domesticated bar-
tural practices and introgressive hybridization between the ley has a hulled grain (adherent lemma and palea), although
crop species and its weedy wild relatives. Weed races in crop there are naked six-rowed cultivars with the botanical ranking
weed complexes can be serious pests of agricultural fields as is of var. coeleste (var. nudum).
the case for sorghum and rice. Or, weeds can develop into crop The domestication of barley has been traditionally treated
species as happened to the weedy races of oats and rye that as a single event occurring in the Fertile Crescent. Although
infested early wheat fields. As wheat and barley agriculture current genetic evidence has supported one independent ori-
spread outward from Southwest Asia, wild oat and rye traveled gin, recent studies show a second primary center of origin
along as grain contaminants into regions where they developed located in the region east of the Zagros Mountains. Thus, the
into domesticated forms suited to growing conditions that barleys of Europe and North Africa developed from the Fertile
were less favorable to wheat or barley. Crescent line whereas those of Central and East Asia developed
from the Zagros Mountains line. The H. spontaneum wild weedy
complex spread from its primary habitats in the Fertile Cres-
cent to secondary habitats westward in the Mediterranean
Major Cereal Species
Aegean regions and eastward across Central Asia to Tibet.
Major cereal species (Table 1) are those that are in interna- The weedy forms (H. spontaneum) have played a role in
tional commerce and serve as dietary staples beyond a local- building diversity through introgressive hybridization with
domesticated barley. For example, the fragile-rachis, six-
ized region. Wheat and barley are temperate cereal crops of the
Fertile Crescent, and along with oats and rye, formed the major rowed domesticated barley discovered in Tibet, and originally
grain complex for Western Asia, North Africa, and Europe. proposed as a distinct wild progenitor species (H. agriocrithon),
is an introgressed form. Although barley has a lengthy agricul-
Maize and rice are the tropical cereals upon which civilizations
in the New World and Eastern Asia developed. Although sor- tural history and development of an extensive number of
ghum, pearl millet, and African rice, all tropical African species, cultivars and land races, its morphological diversity is relatively
limited.
have historically been given a lesser role, they are equally
important in the establishment of early human civilizations
in Africa and South Asia.
Maize
Among the domesticated cereals, maize (Zea mays subsp. mays)
Barley
has an unusual reproductive biology. It is a diploid, cross-
Domesticated barley (Hordeum vulgare) is an annual diploid pollinating monoecious species (staminate and pistillate inflo-
species derived from the wild progenitor species rescences on the same plant) whose pistillate inflorescence is a
H. spontaneum. In preagricultural archaeological finds, barley highly compressed raceme that develops into a cob bearing
is a universal companion of wheat. While considered inferior as naked grains. Maize originated from a single domestication
event in the lowland regions of southwestern Mexico where its annual, pearl millet was domesticated in the arid regions of
wild annual progenitor, Balsas teosinte (Z. mays subsp. parvi- West Africa approximately 5500 BP, moving to East Africa and
glumis) is still endemic. Interestingly, maize ancestry has then India 3000 BP and to Southern Africa 2000 BP. In West
proved to be more complex than a straight wild to crop species Africa, pearl millet still forms an integral component of decrue
development. Genetically, all maize cultivars in the Americas (flood stage to dryland) agriculture in the Central African river
can be traced to highland maize, which also is the maize most deltas where it is cultivated along with African rice (Oryza
closely related to the wild progenitor parviglumis. This ancestral glaberrima) and sorghum (Sorghum bicolor). Pearl millet is
relationship sets up a geographic contradiction due to a low- more drought-tolerant than sorghum and maize.
land origin but a highland ancestral line. With parviglumis Low genetic variation suggests a relatively limited region of
distribution restricted to mid- and lowland regions, a highland origin, although there are a diversity of morphological forms
ancestral form is difficult to explain. Recent genetic research, varying in seed shape, seed color ranging from red to gray to
which tracks with ecological and archaeological findings, has black, and a spike-like compressed panicle that can measure up
helped resolve the conflicting evidence by identifying a sec- to 2 m. Pearl millets are grouped into four ecogeographic races
ondary teosinte progenitor, Z. mays subsp. mexicana. Highland with characteristic morphological traits: typhoides (Africa and
teosinte grows sympatrically with highland maize and is inter- India), nigritarum (eastern Sahel), globosum (western Sahel),
fertile with maize and parviglumis. While mexicana is not the and leonis (West African coast). Active introgression makes
direct progenitor of maize, evidence now points to gene flow it difficult to draw clear phylogenetic lines between pearl
from this wild relative as the explanation for highland maize millet, its wild progenitor P. violaceum, and the weedy species
holding the ancestral genetic position. In this respect, maize P. sieberanum, which evolved from crosses between P. violaceum
offers an informative example of the complexities of discerning and domesticated millet. Crop mimicry is a problem in African
crop origins when geography and introgressive hybridization fields of pearl millet where weed races of P. sieberanum, known
between the domesticated form and secondary wild relatives as shibras, are almost indistinguishable from the domesticated
play a role. species.
Oats
Pearl Millet
Oats form a polyploid series (2, 4, and 6) of self-
Millet in a broad sense comprises a loose grouping of small- pollinating species following different evolutionary lines. Four
seeded, drought-tolerant species that can be cultivated on poor genomes characterize this group: A and C evolved independently
soils (Figure 4). Collectively, the millets belong to nine genera of one another; B and D are derivatives of A. The dominant
in the grass subfamilies Paniceae (eight genera) and Chloridoi- domesticated species is the common oat, Avena sativa (ACD),
deae (one genus) (Table 2). Due to their small size and a which developed from a Southwest Asian ecogeographic race of
hardened lemma and palea, millet grains are insect-resistant, the wild progenitor species A. sterilis. The less important but
a feature that adds to their value for the African and Asian closely related red oat, A. byzantina ( A. sativa subsp. byzantina)
cultures that cultivate them. Only pearl millet (Pennisetum was likely derived from a different disjunct regional A. sterilis
glaucum) is a major cereal grain on an international scale. It is race. The diploid sand oat, A. strigosa (As, a variant of A), evolved
a largely cross-pollinating plant due to the unsynchronized in Spain from the wild form of this species. The semi-
maturity of the respective female and male florets. A diploid domesticated, weedy tetraploid Ethiopian oat, A. abyssinica
(AB), evolved in Ethiopia from the wild species A. vaviloviana.
A. strigosa and A. abyssinica did not play a role in the evolution of
the hexaploids.
More water-tolerant than the other major temperate cereals,
common oat is suited to cultivation in temperate regions with
wet, cool summers. It has an open, nonshattering panicle with
hulled (adherent lemma and palea) seeds except for two naked
forms: the hexaploid A. sativa subsp. nuda with origins in
Mongolia and northwest China and the diploid A. strigosa
subsp. nudibrevis, which developed in England.
With its center of origin located in the western Mediterra-
nean, A. sterilis has a distribution extending from the Atlantic
coast to the Zagros Mountains. A species of primary habitats
often growing alongside wild wheat and barley, wild oat also
populates abandoned and actively cultivated agricultural land
and sites associated with human disturbance. Its initial domes-
tication began as a weed in the newly opened habitats of
cultivated wheat and barley fields. As A. sterilis spread eastward
Figure 4 Diversity and form in the millets. From left to right: panicles of and westward with wheat and barley agriculture, a complex of
pearl millet, broomcorn millet, foxtail millet, and Japanese millet. weed races evolved alongside the domesticated common oat,
Reproduced with permission from Hancock JF (2004) Plant Evolution which began to appear c. 40003000 BP in Europe. In addition
and the Origin of Crop Species 2nd edn., Portland, OR: Book News Inc. to these domesticated and weedy A. sativa forms, the A. sterilis
92 THE CEREAL GRAINS | Cereals: Domestication of the Cereal Grains
Table 2 Minor cereal grains either cultivated or harvested as cultivated or uncultivated wild species
line gave rise to other interfertile weedy wild hexaploid species: Neolithic archaeological sites suggest its status as a tolerated
A. ludoviciana, A. occidentalis, and A. fatua. Of these, A. fatua is weed of wheat fields. Domesticated rye began to appear circa
an aggressive agricultural weed whose extensive distribution is 95008500 BP and likely originated in eastern Turkey and
aided by its spring type habit, that is, it does not require western Armenia. From there, it spread across the Mediterra-
vernalization. Similarly, the wild tetraploid A. barbata (AB) is nean Basin, into Europe, and eastward into Central Asia. An
a noxious weed worldwide wherever it has been introduced interfertile crop complex of wild, weedy, and domesticated
into Mediterranean-like habitats. races, cereal rye is classified into four groups according to
spike fragility or shattering: (1) nonshattering domesticated
varieties (S. cereale subsp. cereale); (2) weedy forms (segetale)
Rice
that mimic wheat in spike and grain morphology and are
Although there are two species of rice, only one of these, Oryza harvested and threshed as part of the wheat crop; (3) semi-
sativa, which was domesticated in Asia, has become a major shattering weeds (afghanicum) whose upper spike shatters,
cereal staple in international agriculture. In contrast, spreading seed in the field while the lower, intact portion is
O. glaberrima is a minor cereal whose cultivation is confined harvested with the wheat; (4) shattering wild-types (ancestrale)
to a small area in the equatorial belt of its native West Africa. found in secondary habitats whose spikes fully disarticulate
Both domesticated species and their wild relatives are diploid, into spikelet dispersal units enclosed by brittle glumes.
semiaquatic plants whose premodern cultivation was tied to Included in the wild type group is the wild species S. vavilovii
seasonal flooding and rains. Of the two, O. sativa has a greater (S. cereale subsp. vavilovii) that grows in primary basaltic
climatic and latitudinal diversity with a range of forms found bedrock habitats in Turkey. Evidence points to S. vavilovii as
from tropical to temperate zones and cultivated in different ryes wild progenitor.
habitats defined by water availability: semiaquatic, either The wild annual relatives S. iranicum and S. sylvestre, both
rainfed or irrigated (lowland); deep water (floating); coastal self-pollinating species, are reproductively isolated from cereal
wetland; and dryland (upland). It is a highly diverse species rye. All the wild perennial rye forms belong to S. montanum
with two major races, sometimes treated as subspecies indica (S. strictum), a diverse species with intergrading and interfer-
and japonica. Based on current archeological and genetic evi- tile ecogeographic races, among them dalmaticum (Mediterra-
dence, the japonica form was first domesticated from its wild nean Basin), anatolicum and ciliatoglume (Anatolia), and
relative O. rufipogon in the middle area of the Pearl River in kupriyanowii (Iran and Transcaucasia). On the Anatolian
southern China c. 13 5008000 BP. Indica subsequently devel- plateau of Turkey where weedy cereal rye and S. montanum
oped from hybridization events between O. rufipogon and the come into contact, the two species hybridize to produce inter-
initial japonica cultivars as they spread into Southeast and mediate, introgressed forms.
South Asia. Diversity within the two races and distinctive dif-
ferences between them are attributable to introgression
Sorghum
between the wild and domesticated forms as well as introgres-
sion of indica into japonica forms. This variation is grouped into Sorghum (S. bicolor) is a drought-tolerant, self-pollinating C4
five subpopulations whose boundaries are not always well cereal grown throughout the arid and semiarid tropics and
defined: temperate japonica, tropical japonica (javanica), bas- particularly important in subsistence farming cultures. In cul-
mati-type (aromatic), ausboro, and indica. tivation, sorghum is treated as an annual although most forms
Domestication of the considerably less diverse O. glaberrima are perennial. Domestication took place in the northeastern
is dated to 3500 BP in the upper Niger River Delta. Its direct savanna/steppe region of Africa in an area stretching from
wild progenitor is the wild annual species, O. barthii, which present-day Sudan to Ethiopia around 8000 BP. From there,
populates savanna water holes and has weedy forms that infest sorghum spread to West Africa, down into southern Africa, and
rice fields located in the high rainfall belt of the African eastward to India, Pakistan, and China. The early extensive
savanna. Oryza sativa was introduced into West Africa from range of sorghum has led to a diverse mix of regional forms.
Europe in the fifteenth century CE and now predominates. The wild progenitor S. arundinaceum; (S. bicolor subsp. verti-
Hybrid cultivars known as NERICA (NEw RICe for Africa) cilliflorum), is still found in primary grassland habitats in East
developed from crosses combining the stress and disease tol- Africa. Secondary introgression from hybridization events
erance of O. glaberrima with the yield potential of O. sativa between domesticated sorghum and S. arundinaceum may be
show promise for improving food security in the rice-growing responsible for expanding sorghums ecological range from
areas of Africa. arid savanna habitats into areas with higher rainfall and
humidity where it can tolerate up to 5000 mm of rain during
the growing season. The cropweed complexes of sorghums
Rye
native Africa also characterize its cultivation in Asia, Australia,
Cereal rye (Secale cereale) belongs to a small Old World genus and the Americas due to the introduction of contaminated seed
of mostly annual diploid species whose phylogenetic relation- stocks. Thus, the bidirectional gene flow between the crop and
ships are complicated by nondomesticated taxa with an obli- weed essentially occurs worldwide.
gate cross-pollinating reproductive biology and perennial Through domestication, the shattering, open panicles of
habit. Its spikes are nonshattering, producing free-threshing S. arundinaceum evolved toward nonshattering, compact pani-
grains at maturity. Cultivation extends into areas requiring cles with larger grains. Traditionally, sorghum is classified into
winter hardiness, drought-resistance, and tolerance of sandy five races bicolor, guinea, caudatum, kafir, and durra and ten
and acid soils. Preagricultural forms of wild-type rye found in two-by-two race combinations, all 15 with a distinctive
94 THE CEREAL GRAINS | Cereals: Domestication of the Cereal Grains
S.R.
Bicolor Guinea
S.R.
ecogeographic range and place in indigenous African and Asian species S. propinquum, which is interfertile with domesticated
agriculture as well as morphological and use differences sorghum, both offer new, untapped genetic resources for
(Figure 5). Bicolor, the most primitive and similar in appear- breeding programs.
ance to wild sorghum with its loose, open panicle, was the first
domesticated race from which the other three races were
Wheat
derived. Caudatum, which is found from Lake Chad to Ethiopia
in bicolors primary domestication zone, is unique for its grain The wheats comprise a large allopolyploid complex of wild and
shape flat on one side, convex on the other. Kafir is associated domesticated species taxonomically classified in the genera
with the Bantu-speaking peoples of East and Southeastern Triticum (wild and domesticated) and Aegilops (wild). They
Africa who migrated there from West Africa. Durra, which has are annual, self-pollinating species. Together, hexaploid bread
dense, compact head-like panicles, was developed in India, wheat (T. aestivum) and durum wheat (T. durum; T. turgidum
where it is now largely grown. African durra likely arrived subsp. durum) are the two major wheats cultivated worldwide,
from India during Islamic migrations into Africa. Recent ranking third in cereal production after corn and rice. The
research confirms that the fifth race (guinea), now designated evolution and domestication of wheat traces an interesting
the guinea margaritiferum group, is the product of a second path in diploid genome development and polyploid specia-
domestication event in West Africa. Divergent from the four tion. Bread wheats three ancestral genomes A, B, and D are
other sorghum races, the guinea margaritiferum group is inter- closely related. A and B diverged approximately 7 million years
mediate between sorghum and its wild progenitor and suited to ago (mya), giving rise to the D genome 56 mya through
the poor soils and erratic rainfall conditions of West Africa. homoploid speciation. The wild diploid species that carry the
Guinea margaritiferum sorghum and the wild Southeast Asian respective genomes are T. urartu (A), Ae. speltoides, the putative
THE CEREAL GRAINS | Cereals: Domestication of the Cereal Grains 95
B donor species whose subgenome within the Aegilops genus is to continental temperate regions with colder winters and
traditionally distinguished as S: and Ae. tauschii (D). The evo- humid summers. As wheat cultivation of the respective tetra-
lutionary lines leading from ancient ancestral diploid wheats ploid and hexaploid forms diffused across Asia, Europe, and
to tetraploid and hexaploid wheats constitute a multilevel North Africa, humans developed more advanced, specialized
phylogenetic mosaic built on multiple rounds of homoploid naked grain forms (for example, tetraploid durum and rivet
and polyploid hybrid speciation events. wheats) (Figure 6; Table 2) as well as a diverse array of land-
The diploid progenitor T. urartu has no domesticated form. races unique for their regional distribution and morphological
T. dicoccoides (T. turgidum subsp. dicocccoides), the wild tetra- and agronomic features. Of note, the transition from wild-type
ploid wheat (BA), originated from an Ae. speltoides x T. urartu shattering spikes and hulled grain to tough rachis, free-
cross. Emmer wheat (T. dicoccum; T. turgidum subsp. dicoc- threshing domesticated forms follows two paths in the bread
cum) (Figure 6) was domesticated from T. dicoccoides circa wheat line. The spikelet dispersal units for T. urartu and
10 000 BP in the Fertile Crescent. A primitive form, emmer T. dicocccoides are wedge-shaped, with the rachis internode
has a semifragile spike and hulled grains (clasping glumes). piece subtending the spikelet. In the case of the semishattering
Its spread eastward brought it into contact with Ae. tauschii, T. spelta, its hulled spikelet units are cylindrical (barrel-
which populated primary habitats in the Caspian Sea Belt. The shaped), with the rachis internode adjacent to the spikelet
hexaploid hybrid speciation event that formed T. aestivum (Figure 3). Aegilops tauschii also exhibits the cylindrical mode
(BAD) (Figure 6) likely took place along the edges of cultivated of spikelet dispersal. All other Triticum (sensu stricto) wild
fields where emmer and weedy Ae. tauschii subsp. strangulata wheats shatter into the wedge units.
came into contact. Bread wheat has no known wild hexaploid Two other evolutionary lines characterize the wheats. A
progenitor. Evidence suggests that the preliminary hexaploid different hybrid speciation event between T. urartu and Ae.
form was a primitive hulled T. spelta type. Naked T. aestivum speltoides produced the wild tetraploid species T. araraticum
forms likely developed within a short period afterwards (GA). While T. araraticum is almost identical in morphology
c.80007000 BP. European forms of T. spelta may have arisen to T. dicoccoides and shares the same distribution ranges
independently from T. aestivum T. dicoccum crosses. in Turkey, Iraq, and Iran, the two species are reproductively
The diploid and tetraploid wheats of the bread wheat line isolated from each other. Thus, the B and G assignments for
are suited to the Mediterranean-type habitats of Southwest Asia the Ae. speltoides genome contribution recognize the respective
with mild winters and dry summers. The addition of the D origins for the two tetraploid species. Two domesticated forms
genome to hexaploid forms extended the ecogeographic range complete this evolutionary line: T. timopheevii (GA) and
T. zhukovskyi (GAAm). Triticum zhukovskyi has no known wild
hexaploid progenitor. The third evolutionary wheat line has
two diploid species: the wild T. boeoticum and its domesticated
offshoot, T. monococcum (Figure 6). Their A genome is different
from that found in the bread wheat line and variously noted at
Ab or Am. Triticum monococcum (einkorn) is an important player
in the early archaeological record but today is considered a relic
domesticated species.
Minor Cereals
differences between the human-mediated changes and Overview; Rye Grain: Its Genetics, Production, and Utilization; Triticale:
those that would have happened by natural biological Overview.
processes.
How did phenotypic diversity in pearl millet and sorghum
become extensive and what features of their domestication
pathways have encouraged these trends?
Further Reading
How did habitat and the geographic spread of early domes-
ticated forms affect rice diversity and its phylogenetic Clark LG and Pohl RW (1996) Agnes Chases First Book of Grasses. Washington, DC:
relationships? Smithsonian Press.
Although African rice is a minor grain, why is it important Elbaum R, Zaltzman L, Burgert I, and Fratzl P (2007) The role of wheat awns in the seed
dispersal unit. Science 316: 884886.
and what are the negative effects of its reduction in acreage Doebley JF, Gaut BS, and Smith BD (2006) The molecular genetics of crop
in Africa? domestication. Cell 127: 13091321.
Why are minor grains greater in number and what is their Abbo S, Gopher A, Peleg Z, et al. (2006) The ripples of The Big (agricultural) Bang:
importance to the regions where they are cultivated? The spread of early wheat cultivation. Genome 49: 861863.
Allaby RG, Fuller DQ, and Brown TA (2008) The genetic expectations of a protracted
How will genetic evidence of phylogenetic relationships
model for the origins of domesticated crops. PNAS 105: 1398213986.
advance development of cereal cultivars that will improve Diamond J (2002) Evolution, consequences and future of plant and animal
food security under changing climatic conditions and loss domestication. Nature 418: 700707.
of arable land? Tanno K-I and Willcox G (2006) How fast was wild wheat domesticated? Science
Why is preservation of germplasm diversity, particularly 311: 1886.
Harlan JR (1992) Crops and Man, 2nd edn. Madison, WI: American Society of
wild cereal relatives and land race cultivars, of critical impor- Agronomy.
tance for plant breeding and what role can molecular genet- Zohary D, Hopf M, and Weiss H (2014) Domestication of Plants in the Old World: The
ics play in building a more complete picture of cereal Origin and Spread of Cultivated Plants in South-West Asia, Europe and the
diversity? Mediterranean Basin, 4th edn. New York: Oxford University Press.
How will a functional grass model guide future cereal
Badr A, Muller K, Schafer-Pregl R, et al. (2000) On the origin and domestication history
of barley (Hordeum vulgare). Molecular Biology and Evolution 17: 499510.
research to improve agronomic and physiological traits Morrell PL and Clegg MT (2007) Genetic evidence for a second domestication of barley
and what benefits will be gained from moving a C4 photo- (Hordeum vulgare) east of the Fertile Crescent. PNAS 104: 32893294.
synthetic system into rice and wheat? Van Heerwaarden J, Doebley J, Briggs WH, et al. (2011) Genetic signals of origin,
spread, and introgression in a large sample of maize landraces. PNAS
108: 10881092.
Exercises for Readers to Explore the Topic Further Mathur PN (2012) Global Strategy for the Ex Situ Conservation of Pearl Millet and Its
Wild Relatives. Rome, Italy: Global Crop Diversity Trust.
Harlan JR (1996) The tropical African cereals. In: Harris DR and Hillman GG (eds.)
Examine the advantages of multidisciplinary arch- Foraging and Farming: The Evolution of Plant Exploitation. London: Unwin
aeologicalgenetic studies in discerning the timing of Hyman Ltd.
domestication and the location of primary and secondary Loskutov IG (2008) On evolutionary pathways of Avena species. Genetic Resources and
centers of cereal domestication and discuss how this work Crop Evolution 55: 211220.
Yuan X, Xu Q, Dong G, et al. (2012) A map of rice genome variation reveals the origin of
will inform germplasm conservation strategies and plant cultivated rice. Nature 490: 497501.
breeding programs. Sackville-Hamilton R (2010) Global Strategy for the Ex Situ Conservation of Rice
NERICA hybrids combine the yield benefits of Asian rice Genetic Resources. Manila, Philippines: International Rice Research Institute.
with the drought tolerance and weed and pest resistance of Linares OF (2002) African rice (Oryza glaberrima): History and future potential. PNAS
99: 1636016365.
African rice. Consider the advantages and disadvantages of
Shang H-Y, Wei Y-M, Wang X-R, and Zheng Y-L (2006) Genetic diversity and
NERICA introductions relative to a balance between diver- phylogenetic relationships in the rye genus Secale L. (rye) based on Secale cereale
sity conservation and food security in Africa. microsatellite markers. Genetics and Molecular Biology 29: 685691.
Explore the role of mapping initiatives and other compre- Mace ES, Tai S, Gilding EK, et al. (2013) Whole-genome sequencing reveals untapped
genetic potential in Africas indigenous cereal crop sorghum. Nature
hensive genetic studies of cereal species in advancing mod-
Communications 4: 2320. http://dx.doi.org/10.1038/ncomms3320.
ern agriculture. What is the importance of educating the Marcussen T, Sandve SR, Heier L, et al. (2014) Ancient hybridizations among the
public in their importance to global food security both ancestral genomes of bread wheat. Science 345: 287-12287-15. http://dx.doi.org/
from the perspective of public funding and collaborative 10.1126/science.1250092.
international projects? Kilian B, Ozkan H, Deusch O, et al. (2007) Independent wheat B and G genome origins
in outcrossing Aegilops progenitor haplotypes. Molecular Biology and Evolution
24: 217227.
Luven P (1988) Traditional Food Plants. FAO Food and Nutrition Paper 42. FAO.
Vietmeyer ND (1996) Lost Crops of Africa: Grains. vol. 1. Washington, DC: Board on
See also: Agronomy of Grain Growing: Sorghum: Production and
Science and Technology for International Development, National Research Council,
Improvement Practices; Breeding of Grains: Barley: Genetics and National Academy Press.
Breeding; Genetics of Grains: Rice: Genetics; Wheat Genetics and Wiersema JH and Leon B (2013) World Economic Plants: A Standard Reference, 2nd
Genomics; The Basics: Grain and Plant Morphology of Cereals and edn. New York: CRC Press, Taylor & Francis Group.
how characters can be used to identify varieties; Taxonomic Williams JT (ed.) (1995) Cereals and Pseudocereals London: Chapman and Hall.
Brkljacic J, Grotewold E, Scholl R, et al. (2011) Brachypodium as a model for the
Classification of Grain Species; The Cereal Grains: An Overview of grasses: Today and the future. Plant Physiology 157: 313.
the Family of Cereal Grains Prominent in World Agriculture; Maize: Brutnell TP, Wang L, Swartwood K, et al. (2010) Setaria viridis: A model for C4
Overview; Millet Minor: Overview; Millet Pearl: Overview; Oats: photosynthesis. Plant Cell 22: 25372544.
98 THE CEREAL GRAINS | Cereals: Domestication of the Cereal Grains
Relevant Websites on the taxonomy, common names, distribution, and other miscellaneous
information about crop plants.
http://www.ars-grin.gov A searchable United States Department of Agriculture crop http://www.public.iastate.edu/imagefpc/IBSC%20Webpage/IBSC%20Templatehome.
germplasm database providing information on scientific and common names, html International Barley Sequencing Consortium.
distribution, economic importance, and references. http://rgp.dna.affrc.go.jp/IRGSP/ International Rice Gene Sequencing Project (Oryza
http://www.ceg.icrisat.org/ipgsc.html International Pearl Millet Genome Sequencing sativa ssp. japonica cv. Nipponbare).
Consortium (global reference genotype Tift 23D2B1). http://www.theplantlist.org/ Searchable database offering current listings of
http://www.cgiar.org/cgiar-consortium/ Consultative Group on International accepted plant names and direct links to checklist data held by collaborating
Agricultural Research (CGIAR) with five independent international centers conducting institutions.
basic and applied research on barley, maize, millet, rice, sorghum, and wheat. http://wheat.pw.usda.gov The United States Department of Agriculture searchable
http://www.croptrust.org/ A CGIAR partner with crop diversity programs targeting database for a variety of topics dealing with the genomics, mapping, germplasm,
conservation of the major cereal grains. pathology, and taxonomy of barley, rye, triticale, wheat, and oats.
http://mansfeld.ipk-gatersleben.de An internet-searchable, version of Mansfelds http://www.wheatgenome.org/ International Wheat Genome Sequencing Consortium
World Database of Agricultural and Horticultural Crops, which is set up for queries (Triticum aestivum).
Maize: Overview
MP Scott and M Emery, Iowa State University, Ames, IA, USA
Published by Elsevier Ltd.
Maize is a member of the grass family Poaceae and the Maize is a monoecious plant, meaning it has both male and
closest known relative is the grassy weed teosinte. female reproductive organs on the same plant. Flowers mature
Maize is a major crop, accounting for 32% of the total cereal after 6070 days of vegetative plant growth. Male staminate
production worldwide in 2013. flowers develop into tassels and are found on the uppermost tip
Maize kernels provide nutrition in the form of starch, oil, of the main stem. Female pistillate flowers are found in one or
and protein. more ears located at nodes along of the stem. Typical maize
Culinary dishes incorporating maize as an ingredient are varieties are diploid, containing two sets of ten chromosomes.
consumed worldwide. Copious amounts of pollen (up to 1 billion grains per plant) are
Maize is processed via dry milling and wet milling and shed from anthers and dispersed by air currents. While the
nixtamalization. majority of the pollen falls close to the plant, a small portion
Maize is a model organism in research. of the pollen can be carried over great distances via the wind.
Maize cultivars include inbred lines, single-cross hybrids, Industry standards typically consider plants separated by a dis-
double-cross hybrids, and open-pollinated varieties. tance of 200 m to be reproductively isolated. Fertilization occurs
Biotechnology has allowed for reduced inputs and higher by the process of double fertilization common to angiosperm
outputs in maize production; however, some societies have species. A pollen grain carrying two nuclei lands on a silk and
concerns regarding the use of genetically modified maize. germinates to produce a pollen tube. The pollen tube grows
down the length of the silk until it reaches the embryo sac
where it ruptures releasing the two sperm nuclei. The first
sperm cell fuses with the egg cell to produce the embryo, the
Learning Objective organ that ultimately develops into the next-generation plant.
The second sperm cell fuses with the central cell of the embryo
To acquire a greater understanding of the role maize plays sac giving rise to the endosperm, the storage tissue that nour-
in the world and how technology seeks to enhance maize ishes the developing seedling until it is capable of living inde-
quality and profitability. pendently. Grain fill to maturity takes about 40 days.
Zea mays, more commonly referred to as maize, is a member of The mature maize kernel is referred to as a caryopsis and is not
the grass family Poaceae, or true grasses. Maize is thought to a true seed but rather a one-seeded fruit. Kernels are composed
have originated 5570 million years ago in what is now Cen- of four organs: the pericarp, embryo, endosperm, and pedicel.
tral or South America and has since diversified into nearly Physical properties, such as hardness, shape, size, color, and
10 000 nondomestic relatives. Figure 1 shows a phylogenetic composition, vary among maize varieties.
tree of grass species related to maize. There exists no direct The main organs of a maize kernel are shown in Figure 2. The
ancestor for maize; however, to date, the closest relative to outer layer of the kernel is the pericarp and encloses the kernel
maize are the teosintes. Prehistoric selection has resulted in for protection. The endosperm comprises the majority of the
ears lacking seed cases called glumes and seeds that adhere to kernels inner contents. The endosperm itself is composed of
the cob until manual removal. These alterations limit the four tissue types: the aleurone (outer) layer, the starchy endo-
ability of maize to survive without human intervention. sperm, the basal endosperm transfer layer, and the embryo-
Maize is an annual plant with C4 metabolism, making it very surrounding region. The endosperm provides nutrients in the
efficient at carbon fixation. It has the greatest global produc- form of sugars and amino acids to the growing embryo.
tion of any crop species around 800 million tonnes was pro- The embryo is composed of the following: the scutellum (the
duced worldwide in 2013, accounting for 32% of the total monocotyledon that absorbs nutrients during germination), the
cereal production. The top three producers include the United coleoptile (protective sheath of the emerging shoot), the plu-
States, China, and Brazil. Maize is grown on more areas of the mule (young plant), the radicle (primary root), and the coleo-
planet than any other crop and is grown on every continent rhizae (protective sheath of emerging root). The tip cap serves to
except Antarctica. The grain of maize is used for food, feed, and attach the kernel to the cob and protect the kernel.
industrial products including biodegradable foams, plastics, In terms of nutritive composition, the kernel can be further
and adhesives. Additionally, maize stover, the leaves and classified into five main components. The typical number 2
stalk of the maize plant, is used for forage, biofuel production, yellow dent maize kernel contains 72% starch, 9.5% protein,
and chemical production. 4.3% oil, 1.4% ash, and 2.6% sugar.
Starch is known for its slow digestion in vivo. The sugary-1 (su1) and
shrunken-2 (sh2) lead to kernel phenotypes that are sweeter
Starch is the most abundant component in maize kernels and
than field corn and are used to produce sweet corn varieties
serves as an efficient storage molecule for glucose. Starch accu-
for canning and fresh consumption.
mulates in the form of dense-insoluble granules. It is com-
posed of two main components: amylose and amylopectin.
Amylose is predominantly a linear polymer composed of 1,4- Oil
linked a-D-glucan chains. In contrast, amylopectin is highly
Oil from a kernel of typical Corn Belt maize, number 2 yellow
branched by a-1,6-glycosidic bonds. Starch biosynthesis
dent, contains 62% linoleic acid, 25% oleic acid, 10%
requires the coordinated activities of a myriad of enzymes,
palmitic acid, 2% stearic acid, and 1% linolenic acid; saturated
including starch synthases and starch branching enzymes and
fatty acids equate to 12% of the total lipid content. The oil
starch debranching enzymes. Enzymatic activity within the
within the maize kernel provides 9 cal g1. Linoleic, linolenic,
kernel alters starch content and properties. The degree of
eicosapentaenoic, and docosahexaenoic fatty acids are shown
branching and branch chain length are starch properties that
to have a positive correlation with cardiovascular health.
can vary considerably among maize varieties. Maturity also
A ratio of 6:1, linoleic to linolenic, is recommended.
affects starch quality. Traits sought in a commercial setting
Similar to starch content and quality, studies demonstrate
include gel strength, viscosity, and thermal properties such as
that exotic germplasm possesses extensive ranges of fatty acid
gelatinization. Maize starch provides 4 cal g1.
composition. Exotic lines are crossed to yield varieties with
Genetic mutations can confer altered starch phenotypes.
increased oil content. Oil content varies across inbred maize
Mutant alleles of waxy1 (wxy1) produce 100% amylopectin
lines and across varied environments. Total fatty acid compo-
starch, which is useful as a thickening agent in foods. Mutation
sition varies throughout kernel development and ultimately
of the amylose-extender (ae) gene leads to high-amylose starch
increases as the kernel matures. Oil content is believed to be
(HAS) with a range of amylose values from 25% to 80%. HAS
affected by a large number of loci and is a highly heritable trait.
Certain breeding schemes aim solely at increasing lipid content
A phylogeny of diploid grass species and/or quality. Duvick (2003) altered fatty acid content by
introducing Tripsacum genes, a wild relative of maize, into
Barley
various maize lines.
Goat Grass Fatty acid stability is directly correlated to saturation level.
Einkorn Wheat Linolenic is the least stable fatty acid, containing three points
Tall Fescue of unsaturation. Oleic fatty acids are much more stable and less
prone to oxidation. Oleic fatty acids are monounsaturated.
Asian Rice
Teosinte Once oxidation begins, it cannot be stopped or reversed and
ultimately leads to rancidity.
Maize
improved amino acid balance. Less abundant components of then used to coarsely grind the maize kernels. Several steps of
the maize kernel include fiber, minerals, vitamins, anthocya- size and weight separation, in addition to regrinding, yield
nins, and antinutrients. maize grits, flour, and fiber. The quality, content, and end use
of the maize must be considered before entering the wet- or
dry-milling process.
Maize kernels with a softer endosperm perform better in the
Maize in Food
wet-milling process. The wet-milling process entails steeping
maize in a dilute sulfur dioxide solution to soften the kernel
Maize is a food ingredient that brings commonality to culinary
and separate it into its smaller components. The germ can be
cultures across the world. Cultural traditions and corn varieties
first removed and later processed for oil. The remaining com-
dictate how maize is incorporated into a wide variety of foods.
ponents are ground and separated further into grits, flour, and
The main kernel components can be separated and processed
fiber. Further processing yields corn gluten meal, and starch.
into products such as corn starch for thickening and binding
Maize starch and high-fructose corn syrup are a main end
agents and corn oil for frying and baking; whole grain kernels
product of the wet-milling process in the United States.
are used in popped popcorn or ground into corn meal and
used in breads, biscuits, and cereals. From enchiladas, tamales,
totopos, tostaditas, and tortillas, virtually every Mexican dish
Maize Kernels
uses maize. Maize porridges are seen across the world: referred
to as puliszka and malderash in Hungary, posho in Africa,
polenta in Europe, grits in the United States, and kpekple in
Ghana. Maize meal can be ground and fermented into sora, a
Dry milling Wet milling Nixtamalization
maize beer in Peru, or used to make hard alcohols such as
whiskey and bourbon. Maize is truly a cross-cultural food.
Figure 3 The US maize grading scale. Reproduced from USDA (2013) Grain Inspection Handbook. Pamphlet. USDA Federal Gain Inspection Service,
Washington, DC.
102 THE CEREAL GRAINS | Maize: Overview
Nixtamalization, dating back from 1200 to 1500 BC, is an homogeneous. Due to inbreeding depression, inbred lines
ancient type of maize processing that includes rendering ker- have low yield and are not used for grain production. Their
nels into a paste to increase the bioavailable nutrients such as main purpose is in the production of hybrid seed. When two
calcium and digestible iron. Kernels are steeped in a water/lime inbreds are cross-pollinated, a single-cross hybrid results.
solution over heat and ground into masa, also known as maize Single-cross hybrids are genetically heterozygous and pheno-
dough, that is used to produce tortillas, corn chips, and other typically uniform. Because of the difficulties of producing seed
food products. on inbred lines, several types of hybrids have been developed.
An open-pollinated variety is a population of plants that is
genetically heterozygous and phenotypically nonuniform. As
Maize in Science the name implies, the seed of open-pollinated varieties is pro-
duced by allowing natural pollination to occur in the popula-
Maize is an important model organism in genetic research. It tion. Synthetic populations are derived from intermating
has several attributes that make it attractive for this purpose. It several varieties that are frequently used in breeding programs
is a large plant and phenotypic analyses are easily done. Each to produce inbred lines.
plant produces an ear typically containing 100400 kernels. It Mechanized agriculture has led to a preference for hybrids
is broadly adapted and has tremendous genetic diversity. Maize because of their uniformity and high yields. The process of
has a moderately sized genome of 2.5 gigabase pairs. A vast hybrid improvement and seed production has become highly
collection of mutant stocks have also been developed that industrialized. Industrial maize breeding has led to greatly
assist in research; this has allowed for many genes to first be increased yields. Open-pollinated varieties require much less
characterized molecularly in maize. Being a diploid species, infrastructure for seed production and genetic improvement
genetic manipulation and analysis is less complex than in and are often grown in developing countries.
species with a higher ploidy level. Additionally, the large phys-
ical size of the maize chromosomes is a great benefit to cyto-
genetic researchers. Hybrid Maize Breeding
Research on maize has led to several key discoveries. Per-
Hybrid maize breeding allows breeders to capture and fix
haps most notable is the discovery of transposons by Barbara
extremely productive genotypes by taking advantage of hybrid
McClintock, for which she was awarded the Nobel Prize in
vigor. Productivity and vigor in maize plants are generally
Physiology or Medicine in 1983. Cytogenetic studies in maize
proportional to the degree of heterozygosity. Thus, inbred
resulted in an understanding of genetic recombination and
lines, although uniform and reproducible, are usually poor
enabled genetic mapping. The role of telomeres was deter-
agronomic purposes. Heterozygosity and performance can be
mined in maize. Through collaborative efforts, it was one of
restored by crossing unrelated inbred lines to make hybrids.
the first crops to have its genome completely sequenced.
Inbred lines are classified into heterotic groups according to
their ability to form productive hybrids in combination with
other groups and their suitability as a male or female parent.
Maize Breeding, Genetics, and Biotechnology For example, nearly all inbreds used as females in North
Maize Cultivar Types American hybrids are in the Stiff Stalk heterotic group. Devel-
opment and maintenance of inbred lines and testing hybrid
A cultivar is a plant variety that has been developed for a combinations require a great deal of infrastructure and exper-
specific use. Several types of maize cultivars are grown includ- tise that are not available to most farmers or even small seed
ing inbred lines, single-cross hybrids, double-cross hybrids, companies and are therefore largely done by large seed
and open-pollinated varieties (Figure 5). Inbred lines are cre- companies.
ated by successive generations of self-pollination. The resulting Uniformity is essential in efficient and profitable produc-
plants are genetically homozygous and phenotypically tion of maize. Superior technology and machinery has assisted
with such uniformity. Improved accuracy in the evaluation of
cultivars has allowed for large genetic gains and the overall
Single cross Inbred line Inbred line creation and advancement of superior maize inbreds. Superior
A B
farm equipment equipped with GPS and computer monitoring
systems has led to optimal planting depth, density, and spacing
Modified single Inbred line Inbred line Inbred line
and precise measurements of grain yield during harvest. In the
cross A1 A2 B
future, precision agriculture will continue to increase produc-
tivity by optimizing inputs, such as maize variety and fertilizer
Three-way cross Inbred line Inbred line Inbred line amount, on a per land area basis.
A B C
Double cross
Maize Biotechnology
Inbred line Inbred line Inbred line Inbred line
Biotechnology gives the ability to introduce genes from any
A B C D
source into the maize genome. Two types of traits derived from
Figure 5 Types of hybrids grown commercially in North America. genetic engineering methods are currently in commercial pro-
Reproduced from Encyclopedia of Grain Science. duction: insect resistance and herbicide tolerance.
THE CEREAL GRAINS | Maize: Overview 103
Insect-resistant maize decreases the need of pesticide appli- recognized as safe under the Federal Food, Drug, and Cos-
cations directly to the plant. The use of pesticides in the United metic Act; therefore, foods made with approved GM varieties
States has been reduced to 6% since 1996, a total of 172 do not require premarket approval. Acceptance of GM maize
million kilograms per year. Fewer tonnes of chemical are by the consumer varies by country. The European Union (EU)
applied, benefiting the health of the environment and proving regulates GM crops based on the process in which they are
economically beneficial for the farmer. From 1996 to 2010, the produced. The EU tends to be cautious of GM crop consump-
income of US farmers increased to a total of $21.7 billion tion. The British Press often refers to such crops as Franken-
dollars, 23% of that profit was derived from 2010 alone. The stein Foods. Protester of third-world countries have been
percentage of genetically engineered (GE) maize has increased known to destroy entire fields of GM crop despite starvation
almost fourfold in 12 years (Figure 6.). The US Department of in the country. The major concerns over the production of GM
Agriculture (USDA) Economic Research Service reports that maize are pollination of weedy species or non-GM maize by
Bacillus thuringiensis (Bt)-resistant maize decreased the amount GM pollen resulting in undesirable transfer of the transgene
of insecticides per planted acres of Bt maize by 8% in the and the impact of transgenes on nontarget species, particularly
United States. Herbicide-tolerant maize is amenable to low to beneficial insects, and the development of insects resistant to
no-till agriculture. Minimal to no tillage results in decreased the mode of action of the insecticidal transgenes in use.
fuel usage and reduction of greenhouse gas emissions, as well Researchers and regulatory agencies continue to develop new
as less soil compaction and erosion. Additionally, crop residue deployment strategies in an effort to minimize these risks.
left on top of the soil increases the levels of organic carbon
sequestration. Soil and water quality are increased due to
decreasing soil erosion and nutrient loss (Committee on the Exercises for Revision
Impact of Biotechnology on Farm-Level Economic and
Sustainability and National Research Council, 2010; National How has human selection led to the evolution of maize?
Research Council, 2010). What countries are the top three producers of maize in the
Of the 159 million hectares of maize grown globally in world?
2012, 55.1 million hectares (35%) was genetically modified What kernel characteristics must be taken into consider-
(GM) maize. Legislation regulating such crops varies among ation when processing? Why?
countries. The United States regulates genetically modified What characteristics make maize a good model organism?
organisms based on the end product. Three groups with differ- Imagine a scenario where a GM maize line is created that
ing perspectives and expertise regulate GM crops in the United corrects a serious nutritional deficiency; however, the pol-
States: the US Environmental Protection Agency, the Food and len of the maize line is known to be compatible with weeds
Drug Administration (FDA), and USDA. GM crops must be grown in the area where the nutritional deficiency is found.
verified free from environmental and human toxins and for- Should this maize line be released in this area? Explain your
eign proteins deemed allergenic. The FDA policy established in reasoning.
1992 considers the currently approved GM crops to be sub- Does GM maize support or hinder progression toward a
stantially equivalent to non-GM crops and deemed generally sustainable agriculture?
GE Maize in the Unites States Exercises for Readers to Explore the Topic Further
100
Research continually strives to produce maize that requires
90 fewer inputs and produces higher outputs while maintaining
kernel quality, yield, and nutritive composition. The creation
80 of new maize lines with novel traits will continue to be seen.
What novel traits do you think should be introduced?
Percent GE Maize
70
Continued improvements in breeding and biotechnology
will allow the development of maize varieties with
60 improved yield and agronomic traits. What do you think
are the practical limits to yield improvement.
50
Educating consumers about the food they consume and
advocating individuals to take an active interest in the
40 production of their food will be helpful in ensuring sus-
tainable food production. Design educational material for
30 maize-based food in your community.
20
2000 2002 2004 2006 2008 2010 2012 See also: Barley, Rice and Maize Processing: Maize: Dry
Year Milling; Maize: Wet Milling; Oil from Rice and Maize; Carbohydrates:
Figure 6 Percentage of all maize grown in the United States that is Resistant Starch and Health; Starch: Starch Architecture and Structure;
genetically engineered (GE). Reproduced from USDA (2014) Adoption of Starch: Synthesis; Food Grains and the Consumer: Genetically
genetically engineered crops in the U.S. Modified Grains and the Consumer; Genetics of Grains: Maize:
104 THE CEREAL GRAINS | Maize: Overview
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Campbell MR, White PJ, and Pollak LM (1994) Dosage effect at the sugary-2 locus on Betran J, and Runge ECA (eds.) Corn: Origin, History, Technology, and Production,
maize starch structure and function. Cereal Chemistry 71: 464468. pp. 273303. Hoboken, NJ: Wiley.
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maxto9&minto1. Sema-Saldivar SO (2004) Foods from maize. In: Wrigley C (ed.) Encyclopedia of Food
Crawley MJ, Brown SL, Hails RS, Kohn DD, and Rees M (2001) Transgenic crops in Grains, 1st ed., pp. 242253. Oxford: Academic Press.
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pp. 225241. Oxford: Academic Press. Tucker J (2011) U.S. Regulation of Genetically Modified Crops. Case Studies in
Flint-Garcia SA, Bodnar AL, and Scott MP (2009) Wide variability in kernel Agricultural Biosecurity. 2014, from http://www.fas.org/biosecurity/education/
composition, seed characteristics, and zein profiles among diverse maize inbreds, dualuse-agriculture/2-agricultural-biotechnology/us-regulation-of-genetically-
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Wheat: An Overview of the Grain That Provides Our Daily Bread
CW Wrigley, University of Queensland, Brisbane, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights in the symbol of the FAO (Food and Agriculture Organization
of the United Nations); a wheat ear is the logo and the motto in
Wheat is unique in its suitability for bread production Latin Fiat panis, which means Let there be bread (Figure 1).
because of its gluten protein. Likewise, the word bread has entered our everyday languages
Bread has a major place in world language and culture as a symbol of food more generally.
because of its importance in our diet. Bread is international, but in its variations, there are
Wheat, a member of the cereal family, is a monocotyledon- national characteristics, as illustrated in Figure 2. In the lives
ous grass of the genus Triticum. There are two species of Polish and Russian people, bread (chleb) is a sacred item.
common (bread) wheat and durum (suited to pasta). In such cultures, an honored guest is welcomed by the presen-
Wheat is grown in a wide range of environments, including tation of a loaf of bread, covered with a lace cloth on a platter.
tropical and very cold regions. In Iran, sweet bread (Persian naan) may be taken as a gift
Worldwide wheat production is equivalent to about 300 g when visiting friends. In the Jewish Passover celebration, the
of wheat per day for each person on Earth. bread must be unleavened, as a reminder of the Israelites
Wheat-based breads include leavened loaves, Arabic and hurried departure from slavery in Egypt, when there was no
Middle Eastern flatbreads, and Chinese steamed breads. time to leave the bread to rise overnight. From Jewish and
Wheat has many other food uses, including noodles, pasta, Christian cultures, various bread-related expressions have
pastry, donuts, cakes, and cookies. entered everyday English: Cast your bread upon the waters,
Nonfood uses include animal and fish feeds, starchgluten Give us this day our daily bread, and Man shall not live by
manufacture, sweeteners, adhesives, and biofuels. bread alone. Similar expressions are found in other languages,
The main component of wheat flour is starch, followed by cultures, and religions.
protein, which is largely the gluten-forming gliadins and Wheat is even more than bread, because its dough-forming
glutenins. The fat content is low. Wheat is a major source of properties suit its use for the further range of foods in Figure 3
energy, fiber, vitamins, and minerals in our diet. crackers, cookies, pastries, and noodles of many types
Some individuals are intolerant to wheat in the diet. The (Figure 4). Pasta types (such as spaghetti, macaroni, and taglia-
best characterized intolerance is celiac disease. telle), distinct from noodles, are correctly made from durum
wheat (Figure 5). In addition, wheat is an important feed source
for animals, birds, and fish. Industrial uses include the separa-
Learning Objective tion of starch from gluten protein. The starch fraction is used for
paper, adhesives, and even to produce ethanol as a fuel. Gluten
To achieve understanding of wheat as a grain species, includ- is used to augment dough quality in baked goods and as an
ing its origins, genetics, production, utilization, and its value in additive to a wide range of foods.
our diet. The Greek goddess Demeter (depicted in Figure 6 with
awned wheat heads and stalks) was probably adopted and
renamed Ceres by the Romans, thus providing the origin of
Introduction
our word cereal, which now includes all the grasslike grains,
for example, barley, rye, rice, and maize.
Wheat is unique among the food grains. Only the wheat grain
contains storage proteins that form into a cohesive dough after
milling, adding water, and mixing. Only the dough from wheat
flour has the extensibility and elasticity to allow the growth of gas Wheat Species
bubbles (generally air and carbon dioxide) and to allow the
retention of the bubble structure throughout the bread baking The ancestors of modern wheats still grow in the regions where
process. Wheat proteins dough-forming ability also accounts for they had their origins in the Fertile Crescent (Egypt through
its value for making the wider range of foods, including noodles, Syria to the TigrisEuphrates valley) and further eastward. They
pasta, pastry and pizza crust, cakes, cookies, and even candies. include the diploid einkorn wheats (Triticum boeoticum and
These are the reasons for the success of wheat not only as one of T. monococcum) and the tetraploid emmer wheats (T. dicoccoides
the three most popular grains, together with rice and maize, but and T. dicoccum). Hexaploid bread wheat arose from natural
also as the grain with the most stringent quality specifications. hybridizations between these tetraploid wheats and the dip-
loid species T. tauschii.
There are two modern wheat species: bread (or common)
Wheat, Bread, and the Human Race wheat, T. aestivum, and durum wheat, T. durum (also desig-
nated T. turgidum ssp. durum), which is less common and is
Despite the wide range of uses of wheat, it is the word bread mainly used for pasta and whole-grain uses such as couscous.
that epitomizes the use of wheat. Bread and wheat are central Figure 7 shows stands of common wheat almost ready to
Figure 1 The logo of the Food and Agriculture Organization uses bread
(panis) as a symbol of food in general. Reproduced with FAO
permission.
Figure 5 Pasta types, such as spaghetti, are made from durum wheat, a
distinct species from bread wheat.
Figure 7 Wheat in the field: (a) unawned wheat, (b) awned wheat, and (c) wheat that has been swathed to hasten the drying of the grain ready for
harvesting.
statistic suggests that there should be no problem with hunger. ovum, resulting in a grain, which grows through the series of
However, about one-third of world wheat production is used anatomical stages shown in Figure 10. The diagram in
for a combination of seed use, animal feed (about 20%), and Figure 10(c) corresponds to the transmission electron micro-
industrial processing (about 10%). Furthermore, the places of graph in Figure 11.
excess production are far from the places of human need. The lowest third of Figure 11 is the outer part of the endo-
Many countries produce more wheat than their domestic sperm, showing cell walls that enclose starch granules (both
requirements, resulting in world trade of about 140 million large (A-type) and small (B-type)). These outer endosperm
tonnes annually. The main wheat-exporting nations are the cells have a larger proportion of storage protein (forming the
United States, Canada, Europe, Australia, and Argentina. Recip- matrix between the starch granules) than the inner endosperm.
ient countries that import grain include Brazil, Egypt, Japan, Enclosing the endosperm is a layer of aleurone cells; beyond
South Korea, Indonesia, Iran, and Iraq. Such international trade that are the bran layers, which are shriveled in the mature
and transport over thousands of kilometers contrast with the grain, having served their functions for the developing grain
subsistence farming of wheat in some regions, where farm- (as seen in Figure 10).
grown grain is milled locally for the farm family. It is the aim of the flour miller to remove these outer layers
and the germ, to produce white flour made up of the endo-
sperm, and then to reduce the endosperm to a fine powder.
The Grain That Becomes Our Food Moisture is used to render the bran layers leathery so that they
leave the grain during milling as large flakes that are readily
The wheat grain is the result of internal fertilization of the sieved from the endosperm particles. Flour milling involves
wheat flower. The glumes that enclosed the flower can be repeated passes between rolls, first fluted and later smooth,
seen in Figure 9 in which other florets have been removed with sieving steps between the rolling. From one stage to
from the rachis (the backbone of the wheat head) to show how another, the flour stocks are conveyed pneumatically through
the mature grain is held within the protecting outer glumes, the complex system of pipes shown in Figure 12.
lemma, and palea (the husks of the mature wheat head). At the Grain hardness is a significant factor to the miller. The
time of fertilization, anthers released pollen to fertilize the endosperm of a soft wheat has air pockets that render the intact
THE CEREAL GRAINS | Wheat: An Overview of the Grain That Provides Our Daily Bread 109
Country/region Wheat production (Mt) Wheat yields (tonnes ha 1) Production of all cereal grains (Mt)
Table 2 Production and trade involving wheat (and all cereals) over five recent years
endosperm opaque and whitish. On crushing, the endosperm Starch damage results; some granules may be split into pieces.
of a soft wheat readily disintegrates into a fine powder, with In any case, some of the protein matrix will remain on the surface
little damage to the starch granules. of the granules, as is shown by the rough surface of the large A
In contrast, the intact endosperm of a hard wheat or durum granule in Figure 13. A small B granule was caught on top of the A
grain is translucent and vitreous. The starch granules are held granule in the process of sample preparation. The scanning elec-
firmly in the matrix of storage protein. More energy must be tron micrograph thus shows the contrast in size of the two types of
put into milling a hard grain, with the initial mill pass produc- granule. A granule from a soft flour would have a smooth surface.
ing a coarse powder (semolina for durum), which must be Starch damage is desirable for most bread making as it
further reduced with repeated milling. facilitates access by starch-degrading enzymes (amylases) to
110 THE CEREAL GRAINS | Wheat: An Overview of the Grain That Provides Our Daily Bread
700
400
300
200
100
0
61 964 967 970 973 976 979 982 985 988 991 994 997 000
19 1 1 1 1 1 1 1 1 1 1 1 1 2
Figure 8 Worldwide wheat production compared to area sown. Reproduced with permission from Marshall DR, Langridge P, and Appels R (2001)
Wheat breeding in the new century Preface. Australian Journal of Agricultural Research 52: iiv.
PERICARP
OUTER INTEGUMENT
INNER INTEGUMENT
ANTIPODAL OUTER INTEGUMENT
EMBRYO SAC
THE CEREAL GRAINS | Wheat: An Overview of the Grain That Provides Our Daily Bread
ENDOSPERM INNER INTEGUMENT
NUCELLUS VASCULAR BUNDLE
CREASE
EMBRYO NUCELLAR EPIDERMIS
FUNICULUS
(a) 0.8 mm
EMBRYO SAC
0.5 mm
(b) DEGENERATING NUCELLUS
ALEURONE
CELLS
PERIPHERAL
PERICARP
ENDOSPERM
ALEURONE
CELLS PRISMATIC
ENDOSPERM
PERIPHERAL
ENDOSPERM
CENTRAL
PRISMATIC ENDOSPERM
ENDOSPERM
VASCULAR
BUNDLE
(c) CREASE
111
Figure 10 Diagrams of the wheat grain through stages of development. (a) The wheat ovule prior to fertilization. (b) The developing grain. (c) The mature grain.
112 THE CEREAL GRAINS | Wheat: An Overview of the Grain That Provides Our Daily Bread
400
Breakdown
Bandwidth
Resistance
at peak
200
Peak
resistance
Mixing
time
0
0 200 400 600
(c) Time (seconds)
Figure 15 (a) The mixograph (original mechanical design) used for measuring dough properties. (b) The mixing bowl is connected to a swinging arm,
which creates (c) the trace of dough properties.
Grain Composition
Wheat and Health Issues
Starch is the main component (up to 85%) of the mature
wheat grain. The white granules of starch are visible in the The many and various wheat-based foods provide a major
micrograph of endosperm (Figure 11) as larger A-type and source of nutrition for the world. They are an important source
THE CEREAL GRAINS | Wheat: An Overview of the Grain That Provides Our Daily Bread 115
In your local region, find out the origins of wheat used for
the many food and nonfood products available. Is there
wheat growing nearby? Are there local milling and
manufacturing facilities?
What are the meanings of bread in your home, district,
and country?
Compare the advantages and problems of using whole-
grain flour versus white flour for wheat-based foods.
Investigate the origins and progenitors of modern wheats.
Further Reading
Exercises for Revision Anderson RP and Wieser H (2006) Medical applications of gluten-composition
knowledge. In: Wrigley CW, Bekes F, and Bushuk W (eds.) Gliadin and Glutenin.
What features distinguish wheat from other cereal grains? The Unique Balance of Wheat Quality, pp. 387409. St Paul, MN: AACCI Press.
Review the range of growth conditions under which wheat Arendt EK and Zannini E (eds.) (2013) Cereal Grains for the Food and Beverage
Industries. Oxford: Woodhead Publishing Ltd.
is grown around the world. What problems and adapta- Baldo BA and Wrigley CW (1984) Allergies to cereals. Advances in Cereal Science and
tions are needed to accommodate these extremes? Technology 6: 289356.
How do wheat and bread impact our lives and culture? Carver B (ed.) (2009) Wheat: Science and Trade. Ames, IA: Wiley-Blackwell.
In your local region, what are the most popular uses of Cauvain SP (ed.) (2003) Bread Making: Improving Quality. Oxford: Woodhead
Publishing Ltd.
wheat? Consider both food and nonfood uses.
Cauvain SP (ed.) (2012) Bread Making: Improving Quality, 2nd edn. Oxford: Woodhead
What are the factors that affect the yield and quality of Publishing Ltd.
wheat grain? Day L, Batey IL, Wrigley CW, and Augustin MA (2006) Gluten uses and food industry
needs. Trends in Food Science and Technology 17: 8290.
Faridi H and Faubion J (1995) Wheat End Uses Around the World. St Paul, MN: AACCI.
Hammed AM and Simsek S (2014) Hulled Wheats: A Review of Nutritional Properties
Exercises for Readers to Explore the Topic Further and Processing Methods. Cereal Chemistry 91: 97104.
Henry RJ (2014) Wheat genomics for grain quality improvement. Cereal Foods World
The gluten protein of wheat can readily be separated from 59: 2224.
the starch of flour by making a dough and kneading it Khan K and Shewry PR (eds.) (2009) Wheat: Chemistry and Technology, 4th edn. St
Paul, MN: AACC International.
under a gentle stream of water. Attempt to perform this Wrigley CW (2012) Proteins The basis of life. Teaching Science 58: 5659.
separation to gain a better understanding of the nature of Wrigley CW and Batey IL (2010) Cereal Grains: Assessing and Managing Quality.
a bread-making dough. Cambridge: Woodhead Publishing.
Durum Wheat: Overview
GP Kadkol and M Sissons, NSW Department of Primary Industries, Calala, NSW, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights East, southern Europe, North Africa, the former Soviet Union,
North America (including Mexico), and India. Worldwide,
Durum wheat is a related but separate species of wheat. It is durum is grown approximately over 13 m ha with an approx-
a cereal and it belongs to the same genus as bread wheat. Its imate annual production of 30 mt (Figure 2). Although durum
origin is similar to bread wheat. constitutes only 58% of the world wheat production, it is an
Durum is a tetraploid with A and B genomes represented, economically important crop because of its unique character-
which are also found in bread wheat. istics and its use in making important food products such as
Durum is adapted to more diverse environments than pasta.
bread wheat, and it performs very well in semiarid regions.
World production of durum wheat is 58% of bread wheat
production. Origin and Spread of Durum Wheat
Durum grain is typically very hard with high protein con-
tent, amber-colored, and vitreous. It is thought that modern durum evolved from the wild tetra-
Durum has a special niche as a food crop being the main ploid, T. dicoccoides. This tetraploid, in turn, originated in the
grain used for making pasta, couscous, burghul, and several Near East through hybridization of diploid Triticum urartu (Au
other traditional foods. genome) and another diploid species, possibly Aegilops spel-
Durum breeding objectives include higher yield, disease toides. T. dicoccoides possesses a brittle rachis and is hulled
resistance, higher grain protein content, increased grain making it difficult to harvest and hard to thresh the grain out
weight, low screenings, higher grain vitreosity, higher sem- of the spikelets. Gradually, during domestication, these types
olina yield, semolina color, stronger dough characteristics, evolved into a tough rachis but hulled grain form, T. dicoccum,
and improved pasta-making characteristics. commonly known as cultivated emmer, which is considered to
Durum, similar to bread wheat, is self-pollinated, and be the first cultivated tetraploid. Emmer is still hard to thresh
therefore, pedigree breeding is a common method of breed- but can be gathered more easily compared with T. dicoccoides
ing used in durum. because the spikes stay intact. Emmer spread from the Near
The main diseases of durum are Fusarium head blight, East to various European, African, and Asian regions. Archaeo-
crown rot, stem, leaf, and stripe rusts. logical evidence has shown the arrival of emmer into regions
Management of nitrogen nutrition to the crop is an impor- far away from the Near East (its center of origin) into Britain, to
tant aspect of durum agronomy because grain protein con- the West, around 3500 BCE and in the Indian subcontinent, to
tent together with physical grain quality determines the the East, around 5000 BCE. Emmer is still cultivated on a small
delivery grade and price for the farmer. scale in Ethiopia, India, Iran, Italy, Spain, eastern Turkey, and
Future prospects for durum appear to be good as pasta the Balkans.
production has increased substantially over the previous The process of domestication of emmer influenced by its
decade. wide distribution and local cultural practices gradually resulted
in selection of mutations that resulted in the loss of the tight
covering of the kernel by the glumes giving rise to naked free
Learning Objectives threshing forms. This change is thought to have occurred
approximately around 70005000 BCE. Domestication also
To learn about durum wheat as a related but separate influenced other traits such as seed size. The first archaeologi-
species of wheat and as a world food crop cal occurrence of modern durum was recorded in Egypt during
To learn about the science behind the durum crop includ- the Greco-Roman period (332 BCE395 CE). Durum evolved
ing breeding, quality evaluation, and agronomy in the eastern Mediterranean region and it had established as a
major crop by 2300 BCE. Gradually, durum replaced emmer in
cultivation and spread to other parts of the world and became a
Introduction food crop of antiquity in Mediterranean Europe, several North
African and Middle Eastern countries, and the Indian subcon-
Durum wheat (Triticum durum) is a separate tetraploid species tinent and is an important ingredient of traditional foods in
of wheat. Durum has 28 chromosomes representing A and B these regions.
genomes compared with 42 in the hexaploid (bread) wheat
with A, B, and D genomes. The typical durum grain is very
hard, vitreous, and amber-colored with high protein content. Adaptation of Durum
Durum wheat is used to produce a range of food products, such
as semolina, pasta, burghul wheat, couscous, and desserts As a species, durum is adapted to a wide range of conditions
(Figure 1). The main durum-growing regions are the Middle from temperate colder climates to warm tropical and semiarid
Figure 1 Images from left to right: Durum wheat plants, field plots of durum wheat, durum grain, semolina (http://shop.icnussa.com.au/p/8670107/
semolinadurum-wheat-1kg-.html), pasta shapes (https://thewindroseblog.wordpress.com/2014/10/24/happy-world-pasta-day/), and couscous
(http://adashofflavour.blogspot.com.au/2010/10/moroccan-couscous-salad.html).
40000 4.00
35000 3.50
Area (1000ha) production (1000t)
30000 3.00
25000 2.50
Yield (t/ha)
20000 2.00
15000 1.50
10000 1.00
5000 0.50
0 0.00
3
1
00
00
00
00
00
00
00
01
01
/2
/2
/2
/2
/2
/2
/2
/2
/2
02
03
04
05
06
07
08
09
10
20
20
20
20
20
20
20
20
20
Figure 2 Worldwide area (brown), production (blue), and yield (blue line) statistics for durum wheat. USDA Foreign Agricultural Service http://www.
pecad.fas.usda.gov/highlights/2010/11/global%20durum/images/GlobalDurumChart.jpg.
conditions and is thus able to be grown in regions where bread harvested in summer. In North America, spring varieties are
wheat may not be profitable. The majority of the world pro- planted in spring and harvested in late summer. In the colder
duction of durum occurs under rainfed conditions in semiarid parts of Turkey and certain other Middle Eastern countries,
regions with relatively dry climate, warm days, and cool nights winter durum varieties are planted in autumn and harvested
during crop growth. Drought stress occurs frequently in these in summer. Winter durum is shown to be well adapted to some
regions. areas of North America and Germany, and there is interest to
Most durum varieties are spring types. In the warmer develop winter durum cropping in these regions.
regions of southern Europe, North Africa, and Australia, spring Generally, durum yields are competitive with bread wheat
durum is planted in late autumn or early winter and is under favorable soil conditions. However, under low pH and
THE CEREAL GRAINS | Durum Wheat: Overview 119
in the presence of mineral toxicities in the soil, durum yields In addition to the methods mentioned earlier, some pro-
are typically lower than bread wheat. grams also use doubled haploids (DHs) to accelerate progress
to homozygosity. DHs are completely homozygous lines gen-
erated by artificially doubling the haploid set of chromosomes
in pollen mother cells. DHs are more expensive to produce in
Durum Breeding
durum, compared with bread wheat, because of a lower success
rate. Single seed descent is a cheaper acceleration strategy and it
Improving productivity of durum by enhancing the inherent
could typically achieve progression from F1 seed to F4 seed in a
ability of durum varieties to produce higher yields combined
year compared with 4 years required in the field-based conven-
with high quality is essential for the continued viability of the
tional approach.
durum industry. Good varieties combining high yields and
All the previously mentioned strategies could be com-
high quality to meet the market demand have been the basis
bined with molecular marker-assisted selection to achieve
of strong durum industries. In durum breeding, as with many
more rapid breeding progress. Molecular markers are avail-
other cereals, the breeder must try to satisfy the requirements of
able in durum to screen for several quality traits (e.g., grain
the entire production chain consisting of farmers, grain han-
yellow pigment content, favorable lipoxygenase alleles, grain
dlers, millers, pasta makers, and consumers and is usually
cadmium content, and grain protein concentration), disease
assisted by experts from other fields like cereal chemistry,
resistance (mainly rust resistance genes), and genes involved
agronomy, pathology, molecular biology, and biometry.
in determining phenology such as vernalization and photo-
Durum breeding, historically, has been mainly conducted
period response.
by public institutions such as government departments of
Advanced evaluation of experimental lines is conducted in
agriculture and universities, while in the United States and
replicated yield trials in several key locations in the target
EU, private breeding companies are common. Durum breed-
regions. Detailed testing of grain, semolina, and pasta-making
ing commenced in EU in the early decades of the twentieth
quality is conducted at this stage of breeding. Testing for dis-
century. Italy was a pioneer in durum breeding with the first
ease resistance is normally conducted at all stages of breeding,
variety released in 1915. Breeding in the early years involved
but in advanced stages, the tests would include seedling resis-
the use of European, Near Eastern, and North African land-
tance evaluation. In some countries, varieties selected through
races. However, these early varieties were tall, responded
this process are progressed to independent variety tests where
poorly to nitrogen fertilizers, were prone to lodging, and were
they are evaluated against check varieties and the best new
relatively low yielding. In the 1960s, semidwarf durum germ-
varieties/breeding lines from other breeders.
plasm became available from International Maize and Wheat
Improvement Centre, Mexico (CIMMYT). Use of this germ-
plasm in durum breeding has delivered large improvements
in yield in most countries. The modern durum varieties are
Breeding for Yield and Agronomic Traits
shorter, more resistant to lodging, responsive to nitrogen
application, and better adapted to their target environments.
Substantial improvement in the yield potential of durum has
been achieved through reducing crop stature, improving lodg-
ing resistance, and manipulating phenology to suit local grow-
Durum Breeding Methods ing conditions. Reduction of plant height was achieved by
incorporating the reduced height gene, Rht1, from CIMMYT,
Durum wheat is a self-pollinated crop and therefore a pure-line which makes plants insensitive to the plant growth hormone,
breeding approach is used, that is, the final product is a uniform gibberellin. The height reduction from Rht1 also resulted in
true-breeding variety. This is achieved by several generations of resistance to lodging and in turn made it possible to apply
selection in the segregating material from crosses, ensuring self- higher rates of fertilizers, use irrigation (if available), and prac-
pollination at each stage. Crosses are made between carefully tice better agronomy to obtain higher yields.
chosen parent lines that have performed well in evaluation trials Reducing plant height also improved the harvest index with
or lines that carry particular traits. Many programs use not more of the plants resources going into grain formation. Fur-
only simple two-way crosses but also three-way crosses and ther improvements in productivity have come from other
backcrosses. Most durum breeding programs use variations of improvements in plant type such as erect leaves and fine tuning
the standard pedigree breeding method (which involves making of crop development and maturity. Some breeders have pre-
selections under self-pollination while maintaining lineage and ferred medium height varieties in an effort to obtain higher
identity information on the selections in each segregating gen- yields through the tall dwarf plant type, which produces a
eration until the breeding material reaches homozygosity), and higher biomass. Breeding has also produced changes in repro-
lines are generally bulked at F4 or F5 stage. Some programs ductive development phases of modern durums leading to
conduct early generation yield testing and then reselect from increases in the duration of the period between initiation of
high yielding bulks. The North American programs make use of the terminal spikelet and ear emergence resulting in improved
off-season nurseries in the southern hemisphere (New Zealand floret fertility and grains per spikelet.
or Chile) to speed up breeding. Quality testing in early stages is A disadvantage of Rht1 is the reduced coleoptile length in
limited to visual inspection of grain quality and evaluation of varieties containing this gene that makes the varieties unsuita-
protein and yellow pigment using near-infrared spectroscopy ble for deep sowing. It is considered that gibberellin-sensitive
(NIR). For further information on breeding methods, please reduced height genes, such as Rht18, would result in longer
refer to the chapter on wheat breeding. coleoptile length and provide better adaptation to deep
120 THE CEREAL GRAINS | Durum Wheat: Overview
sowing, which would allow seed placement in moist soil, breeding process that determines the amount of seed available
deeper down, under dry sowing conditions. for testing. Generally, the earlier the generation cycle, where
seed availability is limited, the more the requirement for tests
that use only small amounts of grain/semolina with preference
Breeding for Tolerance to Abiotic Stresses for nondestructive testing. It is at these stages where there may
be several thousand lines to evaluate that the test should allow
Tolerance to moisture stress is very important considering the effective screening and reduction of the population to more
majority of the durum crop is rainfed and the durum-growing manageable quantities for replicated field testing. At the later
regions of the world are semiarid. In general, breeding durum stages of breeding, more grain is available allowing more
wheat for tolerance to moisture stress is currently conducted by detailed testing.
evaluating yield potential of the experimental lines over several
years and locations. Such evaluation will expose the experi-
mental lines to moisture stress conditions and provide oppor- Grain Size and Test Weight
tunities to select for higher but stable yields combined with
maintenance of grain quality under drought conditions. Test weight (TW), which is affected by the size and shape of the
Tolerance to salinity has been recently researched in grain, is a widely used specification in wheat grading because it
Australia. Two genes, Nax-1 and Nax-2, have been incorpo- is internationally recognized as an index of wheat soundness
rated into durum wheat from conventional crosses with a salt- and wheat milling potential. TW is lowered by grain weath-
tolerant Triticum monococcum line. Nax-2 has been found to be ering, caused by rain at harvest, and grain shriveling due to
more useful for breeding because, unlike Nax-1, there was no terminal drought. Shriveled, high-protein grain produces a
evidence of yield loss associated with the presence of Nax-2. reduced semolina milling performance. This is because small
Molecular markers have been developed for these genes to kernels contain a lesser percentage of endosperm, the main
increase efficiency of selection. economically valuable part of the grain. Generally, a high
Lack of tolerance to acid soils in durum germplasm has thousand grain weight (TGW >40 g) and TW (>76 kg/hl) are
limited the growth of durum in low-pH regions. The impact desirable and further improvements of grain size are sought.
of acid soils is through increased concentration of aluminum
in the soil solution, which is toxic to durum and limits root
growth. Recent research in Australia has achieved transfer of a Percentage of Hard Vitreous Kernels
gene from bread wheat, TaALMT1, conferring tolerance to
aluminum toxicity to a popular durum cultivar, Jandaroi, Durum wheat kernels should have a translucent, vitreous
using conventional breeding methods. There is scope also for appearance to ensure good semolina yield and to avoid pasta
using variation within durum germplasm as there are reports of issues such as weak strands, poor cooking quality, and white
variation for Al tolerance comparable to bread wheat. specks, which occur when grain vitreousness is very low
(<50%). Nonvitreous or starchy or piebald kernels tend to
produce more flour than semolina upon milling because they
Breeding for Quality Traits are softer and the flour can give undesirable white specks in
pasta (Figure 3). Where the grain protein content is high, hard
Durum wheat is mostly used to prepare pasta, either dried or vitreous kernels (HVK) is not usually an issue and has become
fresh. Other products made from durum wheat include bread less important where millers use finer granulations. Genotypes
(flat and leavened), bulgur, freekeh, couscous, pastry, that produce low HVK grain compared with control varieties
pancakes, and some traditional Indian savory dishes and des- having similar protein achievement across multiple environ-
serts. The following discussion will focus on the selection ments should be selected against, but the influence of environ-
methods and objectives in breeding for technological quality ment on protein content makes selection for HVK difficult to
to meet customer requirements for pasta products. In addition, achieve in practice.
there is new interest in breeding for nutritional quality but this
will be covered elsewhere in this book.
% Hard vitreous kernels (HVK)
The choice of testing procedures to select for technological
quality used in breeding programs around the world will vary
depending on the program objectives, but many breeding Vitreous
targets are common across the world and will be described
here. Piebald
Screenings been obtained over the years leading to stronger gluten types.
Gluten strength can be measured by several methods (mixo-
These are undersized grains that fall through a 2 mm screen.
graph, extensograph, farinograph, glutograph, alveograph, and
Grain with high percentage of such grain is undesirable for
gluten index) that use from 10 to 300 g of semolina that can
milling because it reduces semolina yield and creates a less
rank cultivars with diverse strength equally well.
uniform particle size, and this causes white spots in pasta due
to uneven hydration. Therefore, lines with a tendency to high
screenings are avoided by breeders. Semolina Yield
A miller is mainly concerned with the milling yield or percent-
Grain Protein age of the grain converted to semolina of acceptable purity
with minimal flour production because semolina commands
A key quality parameter in durum is grain protein content that a higher price. TW appears to be a better predictor of durum
should be above 13% (11%moisture basis) to ensure about wheat milling performance than kernel weight. Purity is
12% protein in the semolina and pasta that is important to assessed as ash content (mineral content remaining when all
obtain a good texture of cooked pasta. Dry pasta made from the organic content has been removed by combustion at very
high-protein semolina (12%) is physically stronger and more high temperatures) and the number of brown specks per unit
elastic than pasta from lower protein semolina. Generally, as area in the semolina. High mill extraction rates produce sem-
protein content increases, cooked pasta becomes firmer and olina with higher ash (0.91.1% dry weight basis) due to
less sticky and resists overcooking better. Negative aspects of contamination by the bran and this reduces semolinas bright-
high protein content are lower TW and semolina yield, which ness and yellowness. Ash content is affected by the genotype
produces duller pasta. With very low levels of protein, fragile and environment. It is desirable to avoid genotypes with a
spaghetti is produced with low firmness. High-protein durum tendency to higher ash contents. There are currently no satis-
wheat allows spaghetti to swell when cooked, reduces cooking factory small-scale predictive measures of milling performance,
loss, and allows retention of firmness with overcooking, which and typically, a minimum of 100 g of grain has to be milled but
is also associated with less stickiness. Typically, farmers use larger quantities (3 kg) are desirable when using a Buhler mill,
nitrogen fertilizers to achieve these protein levels, but due to for example. The higher grain quantities required and the cost
the increasing cost and environmental concerns about the use of such milling have meant that these tests are conducted on
of too much fertilizer, breeders select for inherently high pro- small elite sets of lines toward the end of the selection process,
tein content. Significant advances have been made by breeders and this necessity has made improvements in this trait difficult.
with new varieties from Canada (Strongfield and CDC Verona) Nevertheless, a small improvement in semolina yield has been
and Australia (EGA Bellaroi and Jandaroi) that are inherently achieved in Australia.
high-protein achievers. Grain protein content is easily mea-
sured using NIR allowing large numbers of experimental lines
to be assessed in breeding programs. Pasta Quality
Evaluation of the quality of the final pasta product is the
Yellow Pigment ultimate test of acceptability of a new variety. In traditional
pasta-consuming countries, the consumer is concerned about
This is important in the market place as it contributes to the the aroma, color, appearance, texture, flavor, and nutritional
desirable yellow color typical of pasta products that consumers value of the pasta. The color of the pasta is very important
desire and provides valuable antioxidants. The yellow pigment and selection of lines giving high brightness and yellowness
content in the grain is strongly inherited, which has allowed in uncooked and cooked pasta is desirable. This can be
large improvements in this trait worldwide over the last decade achieved easily using a Minolta chroma meter or spectropho-
from breeding. Yellow pigment can be measured using NIR or tometer. A high level of yellow pigment in the semolina does
extraction of the yellow pigment from wholemeal using sol- not guarantee a high color in the pasta because it depends on
vents or measurement of the semolina color with a camera to the activity the enzyme lipoxygenase (LOX) in the semolina
measure the color space parameters, brightness (L*), redness during pasta processing. LOX levels in durum wheat are
(a*), and yellowness (b*). These methods allow measurement affected by genotype and environment, and assays for the
on small amounts of grain/semolina at very early stages in the enzyme or molecular markers exist to screen genotypes for
breeding program allowing rapid genetic gain. low LOX levels.
The pasta after cooking should maintain its texture and not
become a thick, sticky mass. Mechanical texture is typically
Gluten Quality
described by a range of terms (firmness, elasticity, stickiness,
Gluten strength is important in pasta manufacture and influ- chewiness, and bulkiness) and can be measured by a sensory
ences cooking quality (loss of solids into the cooking water) panel or by objective tests with the latter typically used. Also,
and texture (bite, stickiness, and adhesiveness) of the cooked cooking tests are employed to measure water absorption,
product. Gluten strength is a property of the dough that has degree of swelling, and solid loss to the cooking water to assess
become sought after in many markets where a higher price can pasta quality.
be commanded. This is because semolina from extra strong Selection for end use quality as discussed earlier has led to
durum varieties is thought to produce firmer pasta. The heri- release of several varieties in many durum-growing countries
tability of gluten strength is moderate and genetic gain has that combine high yield and high quality. Some examples are
122 THE CEREAL GRAINS | Durum Wheat: Overview
Svevo and Normanno in Italy; Strongfield, Eurostar, and most important diseases of durum as they can produce sub-
Brigade in Canada; and EGA Bellaroi, Caparoi, and Jandaroi stantial and widespread yield losses as they are airborne dis-
in Australia. eases. Ensuring an adequate level of genetic resistance to all
current rust races is a major breeding objective in most breed-
ing programs. There have been significant changes of race from
Durum Diseases and Breeding for Disease Resistance time to time, and it is a significant challenge to durum breeders
and pathologists to maintain resistance in the current varieties.
The most important disease of durum, worldwide, is Fusarium For example, a new stem rust race, UG99, also known as TTKS,
head blight (FHB) caused by several fungi including Fusarium has evolved in Africa overcoming two previously effective resis-
graminearum, F. culmorum, F. avenaceum, and F. sporotrichioides. tance genes, Sr24 and Sr31. Most Australian durum varieties
While the first two are more common causes of FHB, often, carry three stem rust resistance genes, Sr8b, Sr9e, and Sr13. This
there are more than one species present in infected crops. FHB combination is expected to be effective against UG99 due to
development is generally favored by wet conditions during Sr13, but it would be important to pyramid additional genes
flowering and grain filling. Durum is considered more suscep- into new varieties to improve the durability of resistance.
tible to FHB than bread wheat. Infected heads are often sterile Resistance to leaf and stripe rust is based on adult plant resis-
and any seed formed is shriveled and may have a pinkish color. tance (APR) genes. Leaf rust is normally not observed because
FHB produces mycotoxins that make the infected grain unsui- it develops later in the season when the weather is warm and
table for human and animal consumption. The most common APR genes are effective.
mycotoxin is deoxynivalenol for which there are legislated Kernel discoloration (black point and red smudge) is asso-
maximum acceptable levels for grain trade. While there are ciated with the same fungi that cause leaf spots but are not
sources of resistance available in bread wheat and closely associated with mycotoxins. Their economic impact on durum
related tetraploid species, resistance to FHB has been difficult is from the loss of clean, vitreous appearance of the grain that
to characterize and incorporate into durum due to the influ- reduces the value of the grain. Durums are more susceptible to
ence of the environment in expression of the disease. Control black point than bread wheat, but there is some genetic varia-
measures include adding nongrass crops in rotations and tion in susceptibility. The main control measures are to prac-
avoiding corn that is highly susceptible to Fusarium and fungi- tice good rotations to reduce fungal diseases and in irrigated
cide sprays. crops reduce watering after flowering.
Fusarium spp. are also involved in causing crown rot (CR) in
durum. F. pseudograminearum and F. culmorum are the main
species involved. CR-infected plants have a honey-brown Agronomy of Durum Production
discoloration on lower leaf sheaths and stem bases. There
could also be plant death at all growth stages. Affected heads For successful durum crop production, it is important to start
ripen prematurely and produce none or shriveled grain and are with selecting a suitable variety for the quality type targeted.
known as whiteheads. Moisture stress from the flowering Varieties differ in their adaptation, maturity, lodging and dis-
stage onward promotes the disease expression and can produce ease resistance, and their ability to achieve the required grain
heavy yield losses. CR expression is also higher under higher protein content and quality.
rates of nitrogen application that promotes vegetative growth It is also important to choose a suitable paddock that does
and leads to higher levels of moisture stress upon postanthesis not have a high disease inoculum load to reduce the incidence
drought. Durum varieties are considered more susceptible to of diseases such as crown rot. Rotation with broad-leaved
CR than bread wheat. There is some variation for susceptibility crops, such as grain legumes and canola, is very important to
to CR in durum germplasm, and some progress has been reduce buildup of disease inoculum, provide opportunities for
achieved in developing varieties tolerant to CR, but it is diffi- weed control, and, in the case of grain legumes, build up soil
cult to characterize the genetic effects due to large environmen- fertility. A good diversified crop rotation would be beneficial
tal effects on expression. Currently, rotation with broad-leaved economically and environmentally. It is also important to
crops is the best control measure for CR. avoid low-pH soils because of likely aluminum toxicity.
Yellow leaf spot (tan spot) caused by Pyrenophora tritici- Minimum tillage or no tillage systems are gaining popular-
repentis and Septoria leaf blotch (Mycosphaerella graminicola) ity in cropping as chemical weed control is widely available.
are important leaf diseases in several countries. These diseases, These systems conserve moisture, improve soil structure and
being stubble-borne, are increasing in their importance as soil organic matter, and reduce the cost of production. In many
a consequence of the increased popularity of minimum tillage. regions, no tillage produces higher yields than the conven-
A single recessive gene, tsn1, on chromosome 5BL controls tional tillage systems. A disadvantage of no tillage systems is
resistance to races 1 and 2 of tan spot. A marker is available to the likelihood of disease inoculum buildup leading to higher
screen for this gene. Resistance to races 3 and 5 is controlled by incidence of leaf spot diseases and crown rot. However, this
another recessive gene, tsn2, on 3BL. The Australian varieties, disadvantage can be overcome by adopting suitable rotations.
Caparoi, Jandaroi, and EGA Bellaroi, are resistant to tan spot. The timing of sowing for achieving optimum yield depends
While currently there are minimal yield losses in Australia upon location of the crop and should be determined to avoid
and North America from rust diseases due to good genetic periods of stress including frost and terminal drought. In most
resistance, in EU countries, resistant varieties are not widely durum areas, early sowing is desirable to avoid loss of yield
available, and as a consequence, there is substantial reliance on potential and drought during grain development. Early sowing
fungicides. All three rusts (stem, leaf, and stripe rust) are the allows the crop to establish and flower under favorable
THE CEREAL GRAINS | Durum Wheat: Overview 123
conditions, and this helps reduce the impact of crown rot. East/Africa regions with great potential for pasta consumption
However, these benefits should be balanced against the risk in India and China. Pasta is a low-glycemic-index food and it
of flowering during the frost-risk period. Sowing rates are lower contains high levels of lutein, which is an antioxidant and a
for rainfed crops to conserve moisture, but in high-rainfall and vitamin A precursor. Regular intake of lutein could help pre-
irrigated conditions, higher plant populations are targeted. vent macular degeneration.
Irrigation, whether protective (strategic application of water There are significant opportunities for continued improve-
during spells of drought or at key growth stages) or for inten- ments in productivity and quality of durum crops through
sive crop production, increases the yield potential of durum breeding and better agronomic management by developing
crops. The modern durum varieties with short, stiff straw with- specific management practices to suit particular varieties.
stand irrigation and higher rates of fertilizers and produce high World durum production has a potential to expand despite
yields compared with older varieties. It is important to avoid the challenges from the changing climate, loss of arable land to
irrigation after ear emergence to achieve high grain protein other uses, and competition from other crops.
content, HVK; to maintain good yellow pigment content; and There is considerable scope to improve the nutritional
to reduce the incidence of kernel discoloration. quality and functional food properties of durum wheat to
Higher rates of nitrogen nutrition are required, especially expand consumer demand. Some examples of functional
under irrigation, to achieve the required grain protein levels. It food properties in durum under development are high amy-
is possible to manage nitrogen requirements by conducting lose content (to increase resistant starch) and soft durum (pos-
soil tests and calculating the extra nitrogen required to achieve sible applications in noodles and biscuits). Development of
the grain protein content required for the expected level of such durum lines could provide scope for an expanded durum
yield. Split applications of nitrogen are more efficient at raising industry.
grain protein content than applying all fertilizer at sowing.
With the availability of yield mapping, it is possible to apply
fertilizers more judiciously according to the observed variation
in fertility in the paddocks. Exercises for Revision
Depending upon the nutrient status of the soil, it may be
necessary to apply other nutrients such as phosphorus, sulfur, How is durum wheat different from bread wheat?
and zinc. There are reports of both boron deficiency (Turkey) Why do we need ongoing breeding work?
and toxicity (Australia) in durum. Chloride deficiency and Why is grain quality important in durum wheat?
consequent need to use chloride containing fertilizers have Which is the most dangerous disease of durum from a
been reported in the United States and it is expressed occasion- human consumption point of view?
ally in Australian crops. What is the importance of plant height in durum breeding?
The future for durum as a world crop appears bright as pasta Check the various dry and fresh pasta products in the local
production in the EU has increased since 2010 (Figure 4) supermarket and note any differences in their appearance
although global growth is coming from Asia Pacific and Middle and origin of the product.
4.600
4.500
4.400
4.300
4.200
4.100
4.000
3.900
3.800
3.700
3.600
2003 2004 2005 2006 2007 2008 2009 2010 2011 2012
(000 tons)
Check for the less popular durum foods in the local super- Engel RE, Bruebaker L, and Emborg TJ (2001) A chloride deficient leaf spot of durum
market. Try finding semolina, couscous, burghul, freekeh, wheat. Soil Science Society of America Journal 65: 14481454.
Grant CA, Di Fonzo N, and Pisante M (2012) Agronomy of durum wheat production.
and durum bread. Note whether the semolina is white or
In: Sissons M, Abecassis J, Marchylo B, and Carcea M (eds.) Durum Wheat
yellow and work out the reason. Chemistry and Technology, 2nd edn., pp. 3755. Eagan, MN: American Association
Check whether durum wheat is part of any traditional of Cereal Chemists International.
preparations in your family. Isidro J, Alvaro F, Royo C, Villegas D, Miralles DJ, and Garca del Moral LF (2011)
Discuss whether you can cross durum wheat with bread Changes in duration of developmental phases of durum wheat caused by breeding
in Spain and Italy during the 20th century and its impact on yield. Annals of Botany
wheat. 107: 13551366.
Compare the glycemic index of pasta with other foods Joppa LR and Williams ND (1988) Genetics and breeding of durum wheat in the United
including bread. States. In: Fabriani G and Lintas C (eds.) Durum Wheat Chemistry and Technology,
Investigate the chemical nature of yellow pigment in durum 1st edn., pp. 4768. Eagan, MN: American Association of Cereal Chemists.
Kimber G and Sears ER (1987) Evolution in the genus Triticum and the origin of
grain and its importance for human nutrition.
cultivated wheat. In: Heyne EG (ed.) Wheat and Wheat Improvement, pp. 154164.
Madison, WI: American Society of Agronomy.
http://en.magazinebbm.com/?p235 The future of pasta is in less developed
Acknowledgments countries.
Martia J and Slafer GA (2014) Bread and durum wheat yields under a wide range of
The authors acknowledge assistance from various colleagues environmental conditions. Field Crops Research 156: 258271.
Matthews P, McCaffery D, and Jenkins L (2014) Winter crop variety sowing guide 2014.
who were very generous with information. NSW Department of Primary Industries. http://www.dpi.nsw.gov.au/agriculture/
broadacre/guides/winter-crop-variety-sowing-guide.
Morris CE and Sands DC (2006) The breeders dilemma Yield or nutrition? Nature
See also: Grain Harvest, Storage and Transport: Contaminants Biotechnology 24(9): 10781080.
of Grain; Wheat: Harvesting, Transport, and Storage; Grain DePauw R and OBrien L (2014) Wheat Breeding: Exploiting and Fixing Genetic
Variation by Selection and Evaluation. In: Wrigley C, Seetharaman K, Corke H, and
Marketing and Grading: Wheat: Marketing; The Basics: Grain and Faubion J (eds.) Encyclopedia of Food Grains, 2nd edn, vol. 4, pp. 279286.
Plant Morphology of Cereals and how characters can be used to Elsevier Ltd.
identify varieties; The Grain Crops: An Overview; The Cereal Grains: Park RF (2014) Wheat: Biotrophic Pathogen Resistance. In: Wrigley C, Seetharaman K,
Wheat: An Overview of the Grain That Provides Our Daily Bread; Corke H, and Faubion J (eds.) Encyclopedia of Food Grains, 2nd edn, vol. 4,
pp. 264272. Elsevier Ltd.
Wheat-Based Foods: Pasta.
Periyannan SK, Qamar ZU, Bansal UK, and Bariana HS (2014) Development and
validation of molecular markers linked with stem rust resistance gene Sr13 in durum
wheat. Crop & Pasture Science 65: 7479.
Royo C, Nachit M, Di Fonzo N, Araus JL, Pfeiffer M, and Slafer JA (2005) Durum Wheat
Further Reading Breeding Current Approaches and Future Strategies. volume 1, New York, NY:
Food Products Press.
Abecassis J, Cuq B, Boggini G, and Namoune H (2012) Other traditional durum derived Sissons MJ (2004) Pasta. In: Wrigley C, Corke H, and Walker C (eds.) Encyclopedia of
products. In: Sissons M, Abecassis J, Marchylo B, and Carcea M (eds.) Durum Grain Science, pp. 409418. Australia: Elsevier.
Wheat Chemistry and Technology, 2nd edn., pp. 177200. Eagan, MN: American Sissons MJ (2014) Pasta. In: Wrigley C, Seetharaman K, Corke H, and Faubion J (eds.)
Association of Cereal Chemists International. Encyclopedia of Food Grains, 2nd edn, vol. 3, pp. 7990. Elsevier Ltd.
Bariana HS, Brown GN, Bansal UK, Miah H, Standen GE, and Lu M (2007) Breeding Sissons M, Abecassis J, Marchylo B, and Cubadda R (2012) Methods used to assess
triple rust resistant wheat cultivars for Australia using conventional and and predict quality of durum wheat, semolina and pasta. In: Sissons M, Abecassis J,
marker-assisted selection technologies. Australian Journal of Agricultural Research Marchylo B, and Carcea M (eds.) Durum Wheat Chemistry and Technology,
2007(58): 576587. 2nd edition, pp. 213234. Eagan, MN: American Association of Cereal Chemists
Blanco A, De Pace C, Porceddu E, and Mugnozza GTS (1988) Genetics and breeding of International.
durum wheat in Europe. In: Fabriani G and Lintas C (eds.) Durum Wheat Chemistry Torrey G Lutein May Decrease Your Risk of Macular Degeneration. American Macular
and Technology, 1st edn., pp. 1746. American Association of Cereal Chemists: Degeneration Foundation, https://www.macular.org/lutein.
MN, USA. Troccoli A, Borrelli GM, De Vita P, Fares C, and Di Fonzo N (2000) Durum wheat
Bozzini A, David J, and Natoli V (2012) Origin and distribution of durum wheat genetic quality: A multidisciplinary concept. Journal of Cereal Science 32: 99113.
diversity in the world. In: Sissons M, Abecassis J, Marchylo B, and Carcea M (eds.) Wallwork H (2000) Cereal Leaf and Stem Diseases, GRDC and SARDI, ISBN 1 875477
Durum Wheat Chemistry and Technology, 2nd edn., pp. 114. Eagan, MN: 86 1.
American Association of Cereal Chemists International. Wallwork H (2000) Cereal Root and Crown Diseases, GRDC and SARDI, ISBN 1 875477
Clarke JM, DeAmbrogio E, Hare RA, and Roumet P (2012) Genetics and breeding of 74 8.
durum wheat. In: Sissons M, Abecassis J, Marchylo B, and Carcea M (eds.) Durum http://herbarium.usu.edu/webmanual/info2.asp?
Wheat Chemistry and Technology, 2nd edn., pp. 1536. Eagan, MN: American nameTriticum_durum&typetreatment Wheat taxonomy.
Association of Cereal Chemists International. Wrigley C (2016) Wheat - An Overview of the Grain that provides Our Daily Bread. In:
http://www.ipmcenters.org/cropprofiles/docs/NDwheat.pdf Crop profile for hard red Wrigley C, Corke H, Faubion J, and Seetharaman K (eds.) Encyclopedia of Food
spring and durum wheats in North Dakota. Grains, 2nd edn, vol. 1, pp. 105116. Oxford, UK: Elsevier.
Rice: Overview
BO Juliano, Philippine Rice Research Institute Los Banos, Laguna, Philippines
2016 Elsevier Ltd. All rights reserved.
Learning Objectives
Structure of the Rice Grain
To appreciate the wide adaptability of the rice plant and the
unique properties of the rice grain The rice grain (rough rice or paddy) consists of an outer pro-
To be able to relate structure and composition of rice to its tective covering, the hull (husk), and the edible rice caryopsis
nutritional properties or fruit (brown, cargo, dehulled, or dehusked rice) (Figure 1).
To understand the diversity in rice texture and the causes of Brown rice consists of the outer layers of pericarp, seed coat,
textural differences nucellus, and germ or embryo, which are the maternal tissues,
and the endosperm. The endosperm consists of the aleurone
layer and the starchy or inner endosperm. Pigment is confined
Commercial Importance, Global Distribution, to the pericarp, but there is a varietal difference in the extent of
and Varieties retention of pigment with the degree of milling. The aleurone
layer encloses the embryo.
Rice is the most important crop in the world in terms of total The inedible hull constitutes 1628% (mean 20%) of
developing world production (480 106 tonnes of rough rice rough rice weight. Brown rice consists of 12% pericarp,
in 2012) and the number of consumers (3.5 billion) depen- 46% aleurone plus nucellus and seed coat, 1% embryo, 2%
dent on it as their staple food. It is widely grown in over 100 scutellum, and 9091% endosperm. The aleurone and embryo
countries in every continent (except Antarctica), from 53 N to cells are rich in lipid bodies (spherosomes, 0.21.5 mm) and in
40 S and from sea level to an altitude of 3 km. The total protein bodies (aleurone grains containing inclusions of phytic
production in 2012 was 697 106 tonnes of rough rice. Asia acid bodies or globoids (13 mm)).
accounts for 149 106 of the 158 106 ha world area that is The endosperm cells are thin-walled and packed with amy-
used for rice cultivation. The mean rough rice yield was 4.40 - loplasts containing polyhedral compound starch granules about
tonnes ha1 in 2012. About 90% of the worlds rice is grown 39 mm in size. Protein occurs mainly in the form of large
and consumed in Asia, in 200 million rice farms most of which (12 mm) and small (0.50.8 mm) spherical protein bodies
are smaller than 1 ha. The major rice producers in 2012 were and crystalline protein bodies (24 mm). Spherical protein bod-
China, India, Indonesia, Bangladesh, Vietnam, and Thailand. ies (PB I) are rich in prolamin (alcohol-soluble protein), and
Rice can grow in both flooded (anaerobic soil) and upland crystalline protein bodies (PB II) are rich in glutelin (alkali-
(aerobic) conditions. The plant has air channels that connect soluble protein). High night ripening temperature increases PB
to its roots. In terms of water regime, 54% of the total rice area II/PB I ratio. Spherosomes are present in the subaleurone or the
was irrigated in 1991, 25% rainfed lowland, 13% upland, and two outermost cell layers of rice endosperm.
Awn
Lemma
Hull
Palea
Pericarp
Seedcoat
Inner Nucellus
endosperm Aleurone layer
Starchy
endosperm VENTRAL
Subaleurone
layer
Scutellum
DORSAL Epiblast
Plumule Embryo
Radicle
Rachilla
Sterile lemmae
Figure 1 Longitudinal section of the rice grain. Reproduced with permission from Juliano BO (1984) Rice starch: Production, properties and uses. In:
Whistler RL, BeMiller JN, and Paschall EF (eds.). Starch Chemistry and Technology, 2nd edn., p. 509. Orlando, FL: Academic Press.
Grading, Handling, and Storage aleurone layer, and the germ. In the Engelberg mills, hull and
bran are removed together in one step with high grain break-
There is no international standard for milled rice size and age. Milling is done in several steps in modern cone mills with
shape. The International Rice Research Institute (IRRI) uses tempering in between to minimize breakage. Many modern
the following scale for size: extra long, >7.50 mm; long, mills have shifted to milling at >14% moisture to minimize
6.617.50 mm; medium, 5.516.60 mm; and short, breakage and to moisture mist treatment (through hollow
<5.50 mm. For grain shape based on the length/width ratio, shaft) during milling to soften the bran and improve surface
the following scale is used: slender, > 3.0; medium, 2.13.0; gloss.
bold, 1.12.0; and round, 1.0. Grades are based on grain size Only about 45% of the worlds rice production enters the
and shape, degree of milling, percentage of head or whole- international trade. The major exporters in 2012 were Thai-
grain milled rice, immature grains, damages (discolored) and land, India, Vietnam, the United States, and Pakistan. The
heat-damaged grains (chalky grains, red grains, and red- major importers in 2012 were Nigeria, China, Iran, the Philip-
streaked grains), aroma, and organic extraneous matter and pines, Iraq, Saudi Arabia, Ivory Coast, and Malaysia.
inorganic extraneous matter.
An important property of harvested grain is its moisture
content: 14% on a wet-weight basis is considered a safe storage Chemical and Nutritional Composition
value; grains become susceptible to fissuring from moisture
adsorption stress below critical moisture contents of 1216%, Rice has one of the lowest protein contents (7%) among the
depending on the variety. Aging of sun-/oven-dried rough rice cereals. The bran layers and embryo are richer in nonstarch
for up to 2 months after harvest at ambient temperature constituents than the milled (white) rice (Table 1). The major
>15 C improves milling yields and makes milled rice expand nutritional advantage of brown rice over milled rice is its
more during cooking, with less dissolve solids, and become higher content of B vitamins. Although higher in minerals,
more flaky. Heating during grain drying accelerates aging. bran phytic acid forms complexes with minerals and proteins,
About 20% of rice is consumed as parboiled rice. Parboiling reducing their bioavailability. Thus, the available iron (mg per
consists of boiling or steaming steeped aged rough rice until meal) is similar in brown and milled rice meals, but probably
the hull starts to open and then cooling and drying the gelati- higher in brown rice for zinc (mg per meal) than for milled
nized grain. Diffusion of bran B vitamins into the endosperm rice. The energy content of brown rice and bran is higher than
occurs during parboiling but with some heat decomposition. that in milled rice, owing to the higher fat content. Rice has no
Mycotoxins are a potential problem in parboiled rice. vitamin A, C, or D.
Milling involves dehulling, followed by removal of the Although cereal proteins are deficient in lysine, rice protein
outer 710% of the brown rice, either by friction or by abra- has one of the highest lysine contents among them, corre-
sion, removing most of the pericarp, seed coat, nucellus, sponding to an amino acid score of 67% in milled rice based
THE CEREAL GRAINS | Rice: Overview 127
Table 1 Comparison of nutrient composition of brown rice, milled rice, and rice bran
Juliano BO (2007) Rice Chemistry and Quality. Munoz, Nueva Ecija: Philippines Rice Research Institute; Champagne ET, Wood DF, Juliano BO, and Bechtel DB (2004) The rice grain
and its gross composition. In: Champagne ET (ed.). Rice Chemistry and Technology, 3rd edn., pp 77108. St. Paul, MN: American Association of Cereal Chemists; United States
Department of Agriculture (2012) National Nutrient Database for Standard Reference, Release 25. http://www.ars.usda.gov/ba/bhnrc/ndl.
Table 2 Amino acid profile and energy and nitrogen balance in five growing rats of raw brown rice, milled rice, and rice bran
Means in the same line followed by a common letter are not significantly different at the 5% level by Duncans (1955) multiple range test. lys lysine is the first-limiting amino acid.
United States Department of Agriculture (2012) National Nutrient Database for Standard. Reference, Release 25. http://www.ars.usda.gov.ba/bhnrc/ndl; Eggum BO, Juliano BO, and
Maningat CC (1982) Protein and energy utilization of rice milling fractions by rats. Qualitas Plantarum-Plant Foods for Human Nutrition 31: 371376.
on the US Food and Nutrition Board amino acid requirement in milled rice. Bran proteins are 6698% albumins. Prolamin
pattern for the 13-year-old child of 5.1% lysine as 100% is poor in lysine but rich in sulfur amino acids. The high lysine
(Table 2). The solubility fractions of protein are about 15% content of rice protein is due to the low prolamin content.
of albuminglobulin (water- and salt-soluble), 20% prolamin The energy digestibility is higher in milled rice than in
(PB I, alcohol-soluble), and 65% glutelin (PB II, alkali-soluble) brown rice owing to the lower dietary fiber and phytic acid
128 THE CEREAL GRAINS | Rice: Overview
levels, as verified by the poor energy digestibility of rice bran Essential fatty acid content of rice oil is 33.4% linoleic acid
(Table 2). The true digestibility (TD) of milled rice protein is and 1.6% linolenic acid. Levels of oryzanol, tocopherols,
also higher than that of brown rice, but the biological value tocotrienols, and unsaponifiable matter differ among crude
(BV) is lower, resulting in a similar NPU. Bran protein has a oils, and oryzanol, tocopherols, and tocotrienols may be
lower TD but a higher BV than brown and milled rice proteins. reduced by up to 90% by conventional refining and deodoriz-
The amino acid score corrected for TD in rats proposed by the ing. Contents of these antioxidants are also lower in Philippine
Food and Agriculture Organization as a protein quality index rice than in US rice.
shows similar values to NPU for the rice proteins. Black or Breeding efforts to improve the nutritional value of rice
purple rice has a lower brown rice NPU (72%) and higher grain include a higher micronutrient density (higher iron,
phenolics level (0.6%, anthocyanin) than red rice (NPU; zinc, and vitamin A), low phytic acid (high inorganic phos-
83% and 0.2% phenolics, proanthocyanidins) and phate) content, low prolamin content, the absence of
nonpigmented rice (NPU 97% and 0.02% phenolics), but lipoxygenase-3 activity and allergenic globulin, and transgenic
their milled rice have identical NPUs. Rice complements rice endosperm with vitamin A, with the soybean glycinin gene
legumes (deficient in sulfur amino acids) in amino acid com- and with the soybean ferritin gene.
position for human diets.
Cooking and parboiling reduces the TD in growing rats by
515%, with a corresponding increase in BV but little change Processing and Food Uses
in NPU. However, lysine digestibility remains close to 100%,
and cystine digestibility drops to about 82%. The fraction that The mean worldwide per capita consumption of milled rice
remains in the feces as fecal protein particles represents the was 53 kg per year in 2009. It was 77 kg in Asia, 31 kg in South
lipid-rich core of large PB I, with < 1% lysine in the protein but America, 21 kg in Africa, 14 kg in Oceania, and <10 kg in
rich in cystine. PB II is readily digested. Europe and North America. Per capita, rice consumption was
Starch varies in apparent AC (as determined by iodine highest in Southeast Asia at 129 kg per year, followed by South
colorimetry in acetate buffer): waxy, 02%; very low, 210%; Asia at 71 kg per year. More than 3.5 billion people depended
low, 1020%, intermediate, 2025%; and high, >25%, all on a on rice for more than 20% of their daily energy in 2009. Rice
milled rice dry-weight basis since the 1950s. Actual AC values provides more than 50% of energy supply for about 520 mil-
by differential scanning calorimetry (by melting or crystalliza- lion people in Asia, most of them poor to very poor.
tion enthalpy of amyloselysolecithin complex) are slightly There are various ways of cooking milled rice. In tropical
lower and closer to AC values with iodine colorimetry in Asia, they are prewashed to remove dirt, which results in losses
ammonium buffer pH 9 of low, 1017%; intermediate, of B vitamins and fat. Presoaking for 30 min reduces cooking
1722%; and high, >22%. The glycemic index of cooked time (particularly of parboiled and brown rice). Grain cracking
brown rice tends to be lower than that of cooked milled rice during soaking is minimized by adjusting milled rice moisture
owing to the higher amounts of phytic acid and fiber in brown content to 15% by high-pressure hydration to minimize crack-
rice. However, the lowering in glycemic index from milled rice ing. Rice may be cooked in the amount of water that it will
to brown rice is highest in waxy and low-AC rice. Among absorb or boiled in excess water and the cooking water dis-
cooked milled rice, the glycemic index decreases with increas- carded. A steaming step is used in Indonesia and also for waxy
ing AC regardless of the cooking method. Processing, including rice. Oil or ghee may be added in the Middle East, and this
parboiling, tends to reduce the glycemic index. In vivo resistant reduces surface stickiness. Enriched rice premixes containing
starch in an ileostomate (large intestine surgically removed) is iron and B vitamins resistant to washing have been developed
5% and may be not as discriminating as the glycemic index for rice, but have not been popular because of additional
in humans. Short-term satiety may also be influenced by AC. expense; the enriched rice premix can be readily distinguished
The use of rice bran and brown rice in cereal products has from ordinary rice.
increased in recent years owing to the US Food and Drug Rice is hypoallergenic. Its AC correlates positively with
Administrations recent approval of brown rice as a whole water absorption and volume expansion during cooking and
grain and the hypocholesterolemic effect, in humans, of the with hardness of boiled rice. South Asian consumers prefer
factors in the high (up to 4.4%) unsaponifiable fraction of high-AC intermediate-high gelatinization temperature (GT
its oil, such as g-oryzanol, (cycloartenol, 24-methylene- >70 C) rice; Southeast Asians prefer intermediate-AC,
cycloartanol and campestrol esters of ferulic acid), tocopherols, intermediate-GT rice. Japonica (temperate) short-grain rice pre-
and tocotrienols, an analog of tocopherol (vitamin E). Defatted ferred in Japan, Korea, Taiwan, and northern China have
bran has no hypocholesterolemic activity, unlike in oats, mainly low AC and low GT (70 C), with some having an
where the active principle is soluble b-glucan. Inactivation of intermediate AC in Europe. Only in Laos and north and north-
antinutritional factors trypsin inhibitor, oryzacystatin, and east Thailand is steamed waxy (glutinous) rice consumed as a
hemagglutininlectin and lipase and lipoxygenase that are staple. The low GT and intermediatehigh GT result from the
concentrated in the rice bran by heat treatment and extrusion differences in proportion of debranched amylopectin with
cooking improves the shelf life of the bran and its nutritional chain lengths below 24 glucose units: Low GT has more of
value to poultry. By contrast, the antinutritional factor phytic the shorter chains. The higher longer chains increase not only
acid is heat-stable. Phytic acid in rice bran and brown rice has GT but also the staling of cooked starch. Among waxy and low-
been reported to have some medicinal activity in preventing AC rice, low-GT rice is a softer cooked rice than high-GT rice
some types of cancer. Phytic acid content of rice bran is highest (intermediate GT is rare at low AC), but among intermediate-
among cereal brans (38% phytic acid). and high-AC rice, intermediate-GT rice is a softer freshly
THE CEREAL GRAINS | Rice: Overview 129
cooked rice than low-GT rice. High-AC rice with high content How may mutants low in lipoxygenase and lipase improve
of long-chain amylopectin (blue staining with iodine) have shelf life of rice?
hard gel consistency, as long-chain amylopectin behaves like How does amylose differ from long-chain amylopectin in
amylose. Native long-chain amylopectin crystallizes from properties?
water-saturated 1-butanol only after debranching with Speculate on the binding of 2-acetyl-1-pyrroline with
isoamylase. starch.
Various rice products are prepared for which specific AC
types are preferred. Freshly and well-milled rice is preferred for
rice products to prolong shelf life by minimizing fat rancidity in
See also: Barley, Rice and Maize Processing: Oil from Rice and
the stored products. Parboiled rice are preferably aged high- and
Maize; Rice Processing: Beyond the Farm Gate; Beverages from
intermediate-AC, whereas extruded and flat noodles use mainly
Grains: Beverages: Asian Alcoholic Beverages; Breeding of Grains:
aged, high-AC, low-GT rice. Rice with a low GT of starch is
Rice: Breeding; Carbohydrates: Glycemic Index, Glycemic Load, and
preferred in rice puddings, breads and cakes, and beer adjuncts.
Their Health Benefits; Resistant Starch and Health; Starch: Analysis of
Waxy and low-AC rice are preferred for rice wines (for a higher
Quality; Starch: Starch Architecture and Structure; Food Grains:
ethanol yield) and in frozen sauces, desserts, snacks, and sweets,
Intolerance, Allergy and Diseases: Nutrition: Beriberi, A
because of their slow staling rate. Rice crackers are prepared
Deficiency Related to Grains; Food-quality Testing: Rice: Eating
from waxy and nonwaxy rice. Parboiled rice is preferred over
Quality; Genetics of Grains: Rice: Genetics; Non-wheat Foods:
raw rice for idli (pudding) and dosai (cake) with rice-black gram
Rice: Chinese Food Uses; Scientific Organizations Related to
usually fermented at a 3:1 weight ratio. The thermophysical
Grains: Research Organizations of the World: CGIAR; The Cereal
properties of starch such as glass transition temperature, GT,
Grains: African Rice (Oryza glaberrima): A Brief History and Its
enthalpy of raw starch, staled amylopectin melting
Growing Importance in Current Rice Breeding Efforts; An Overview of
(4560 C), amyloselipid complex I (<100 C) and II
the Family of Cereal Grains Prominent in World Agriculture.
(>100 C) melting, and staled amylose melting (>130 C) of
gelatinized starch affect the properties of rice products in addi-
tion to the amyloseamylopectin ratio and protein content.
Specialty rice include Italian Arborio rice for making risotto, Further Reading
waxy rice (sweet rice with an opaque grain), Thai Jasmine rice
Atkinson FS, Foster-Powell K, and Brand-Miller JC (2008) International tables of
(aromatic low-AC long-grain), and Punjab Basmati (aromatic glycemic index and glycemic load values. Diabetes Care 31: 22812283.
intermediate-AC long-grain that elongates when presoaked rice Biliaderis CG, Tonogai JR, Perez CM, and Juliano BO (1993) Thermophysical
is cooked). 2-Acetyl-1-pyrroline is the first major aroma prin- properties of milled rice starch as influenced by variety and parboiling method.
ciple identified in raw and cooked aromatic rice, and others are Cereal Chemistry 79: 512516.
Champagne ET, Wood DF, Juliano BO, and Bechtel DB (2004) The rice grain and its
being characterized. Endosperm 2-acetyl-1-pyrroline bound gross composition. In: Champagne ET (ed.) Rice: Chemistry and Technology,
to starch requires a higher extraction temperature than the 3rd edn., pp. 77108. St Paul, MN: American Association of Cereal Chemists.
free form. Eggum BO, Juliano BO, and Maningat CC (1982) Protein and energy utilization of rice
milling fractions by rats. Qualitas Plantarum-Plant Foods for Human Nutrition
31: 371376.
Food and Nutrition Board Institute of Medicine (2002) Dietary References Intakes for
Exercises for Revision Energy, Carbohydrates, Fiber, Fat, Fatty Acids, Cholesterol, Protein and Amino
Acids (Macronutrients). Washington, DC: National Academy Press.http://www.nap.
While total maize production exceeds that of rice, why is edu/opcabook/html/.
rice a more important food crop than maize? Hanashiro I, Matsugasako J, Egashira T, and Takeda Y (2005) Structural
characterization of long unit-chains of amylopectin. Journal of Applied Glycoscience
Does waxy or glutinous rice contain gluten?
52: 233237.
Why was the approval by the US Food and Drug Adminis- International Rice Research Institute, 2013. World Rice Statistics. irri.org/world-rice-
tration of brown rice as a whole grain delayed as compared statistics.
with the other cereal grains? Juliano BO (1985) Rice starch: Production, properties and uses. In: Whistler RL,
The N content of rice glutelin is 16.8%. What Kjeldahl N BeMiller JN, and Paschall EF (eds.) Starch Chemistry and Technology, 2nd edn.,
p. 509. Orlando, FL: Academic Press.
factor would one use for rice glutelin? Note: The factor 6.25 Juliano BO (2004) Rice/overview. In: Wrigley C, Corke H, and Walker CE (eds.).
is used for most proteins containing 16.0% N. Encyclopedia of Grain Science, vol. 3, pp. 4148. Oxford, UK: Academic Press/
Why is it harder to manipulate the composition of a whole Elsevier.
grain such as rice than that of a flour such as wheat? Juliano BO (2007) Rice Chemistry and Quality. Munoz, Nueva Ecija: Philippine Rice
Research Institute.
Juliano BO, Tuano APP, Monteroso DN, et al. (2012) Replacement of acetate by
ammonium buffer to determine apparent amylose content of milled rice. Cereal
Foods World 57: 1419.
Exercises for Readers to Explore the Topic Further
Tuano APP, Xu Z, Castillo MB, et al. (2011) Content of tocols, g-oryzanol and total
phenolics and grain quality of brown rice and milled rice applied with pesticides and
What factors would influence satiety index of cooked organic and inorganic nitrogen fertilizer. Philippine Agricultural Scientist
milled and brown rice? 94: 211216.
How would a low-phytic acid mutant affect iron absorption United States Department of Agriculture, 2012. National Nutrient Database for Standard
Reference, Release 25.http://www.ars.usda.gov/ba/bhnrc/ndl.
from brown rice? Yoshihashi T, Huong NTT, Surojanametakul V, Tungtrakul P, and Varanyanond W
Speculate on how global warming would affect rice grain (2005) Effect of storage conditions on 2-acetyl-1-pyrroline content in aromatic rice
properties. variety, KhaoDawk Mali 105. Journal of Food Science 70: S34S37.
Wildrice, Zizania: Overview
EA Oelke, University of Minnesota, St. Paul, MN, USA
RA Porter, University of Minnesota-NCROC, Grand Rapids, MN, USA
2016 Elsevier Ltd. All rights reserved.
Topic Highlights and some breeding efforts were initiated in the early 1960s
with a full-scale breeding program started in 1972 by the
Wildrice (wild rice), Zizania, belongs to the grass family, is a University of Minnesota. Today, wildrice is cultivated mainly
cereal grain, and has four species, three naturally occurring in Minnesota and California in the United States and outside of
in North America and one in Asia. the United States on a few hectares in Hungary. The Canadians
The grain from wildrice growing in lakes and rivers in North have expanded lake production by seeding new lakes in several
America has been hand harvested, using a canoe, and pro- provinces. In Canada, the natural stands as well as newly seeded
cessed by Native Americans for thousands of years as a lakes are harvested by airboats with catchers attached to the
carbohydrate food. front. There are three species of wildrice that grow naturally in
Most cereal grains have been under cultivation and domes- the North American continent: Zizania aquatica L., Zizania palus-
tication for centuries, but for wildrice, this first began in the tris L., and Zizania texana A.S. Hitchc. Z. aquatica grows along the
early 1950s in Minnesota. eastern seacoast and has thin kernels. Z. palustris grows around
Improved cultivars have been developed for better yields the Great Lakes region and has large seeds. It was gathered by
than natural plants and adapted for harvesting with large Native Americans for thousands of years as a food source. Both
combines. of these species are annual aquatic grasses. Z. texana is a peren-
Wildrice cultivation is similar to rice, Oryza, in diked fields nial with small seeds and grows only in a very small area in
(paddies) to allow flooding and draining. Texas. Z. latifolia Turcz. grows in Asia, and the fungal infected
Today, most of the cultivation is done in Minnesota and base of the stem is harvested as a vegetable, Makomo-taki.
California, the United States. Wildrice is a recently cultivated and partially domesticated
Yields per hectare of cultivated wildrice are about ten times cereal grain; thus, the purpose of this article is to acquaint the
that from hand harvested natural stands, but still well reader with this new crop. Wildrice is cultivated using similar
below that of rice. practices as is used for lowland rice, in that it only grows in
Processing of the cultivated grain follows similar steps used flooded soils. Harvesting the newly developed cultivars is done
by Native Americans of parching (roasting and drying) and with combines similar to ones used for rice in the United
hulling, however using modern equipment in inspected States. Processing is different, however, giving wildrice its
processing facilities. unique roasted flavor. The nutritional value of wildrice is con-
Processed wildrice is a unique, specialty cereal grain with a siderably better than rice having nearly twice as much protein
distinct flavor and has a good amount of protein, similar to and an amino acid complex similar to oat groats (hulls
wheat, and twice the dietary fiber of brown rice. removed). Wildrice is often mixed with rice and served as a
The processed grain (kernel) without hulls is very versatile blend. The pure product serves as a side dish similar to pota-
and is often featured in soups, salads, breads, side dishes, toes or rice. Continued improvement in yield/ha should make
entrees, and even desserts. wildrice more available in stores and restaurants not only in
In Minnesota, any packaged processed wildrice sold in the the United States but also in other countries.
state must be labeled as to where and how it is grown.
Zizania texana
Figure 1 The natural distribution in shallow lakes and rivers in North America of the three species of Zizania. Reproduced from Oelke EA, Porter RA,
Grombacher AW, and Addis PB (1997) Wild rice New interest in an old crop. Cereal Foods World 42: 234247. American Association of Cereal
Chemists, Minneapolis, MN, with permission.
was classed into three different species by Linnaeus and others. hundreds of years. After harvest, it was processed and stored
They are two annual species Z. aquatica L. and Z. palustris L. and and used as a carbohydrate source for the long, cold winters.
one perennial species, Z. texana Hitchc. The distribution of the Many battles were fought for control of the prized wildrice
three species in North America is shown in Figure 1. stands. In most cases, the Chippewa ended up controlling the
There are two varieties of annual Z. aquatica that grow along wildrice stands. The Ojibwa name for wildrice is Manoomin
the eastern Atlantic seacoast. Their seed is long and thin and meaning good berry or good seed, but the exact meaning is not
generally not harvested for food. Z. aquatica can also be found agreed upon. The early explorers, both English and French,
in the Z. palustris area but to a limited extent. The two varieties who often traded with the Ojibwa, gave various names to this
of annual Z. palustris have relatively large grains and have been plant; some were Indian rice, wild rice, Canadian rice, squaw
and are harvested as food by Native Americans. Today, the rice, water oats, march oats, and water rice. The French
most extensive natural stands occur in the states of Minnesota explorers called the plant folle avoine, fools oats, or wild
and Wisconsin in the United States and in the provinces of oats. The name that was most commonly used was wild rice
Manitoba and Ontario, Canada. Z. texana is a perennial and and is used in the trade today. To avoid confusion with the
only grows in a small stream in Texas; it has small grains and is wild species of Oryza genus, writers often use the hyphenated,
not harvested. Z. latifolia Turcz. is also a perennial and can be wild-rice, or the single word, wildrice. In this article, we are
found in Asia. There are accounts that the grain has been used using the single word wildrice. Today, stands of wildrice on
by the elite as a special food. However, today in Asia, the base reservations in Minnesota and Wisconsin are controlled by
of the plant, which is infected with a fungus, is used as a Native Americans, but in lakes and rivers outside of the
vegetable, Makomo-taki. A brief description of each species reservations, the stands are controlled by the Departments of
is given in Table 1. Natural Resources of each state. In Minnesota and Wisconsin, a
license is required by everyone to harvest from lakes and rivers
that are not on Native American reservations. Today, natural
Historical Use of Species
stands in Minnesota are harvested, by law, in the traditional
Zizania palustris L. method using a flat bottom boat or canoe. The canoe is pushed
The large seed from this species, ranging from 8 to 16 mm through the stand by one individual using a long pole, while
long and 1.22.5 mm in diameter, has been harvested from the other individual uses two flails (knockers) to knock the
lakes and rivers in the Great Lakes region by the Chippewa grain into the canoe. Since the grain ripens at different times on
(Ojibwa) and Menominee Native American tribes for the same panicle and easily falls (shatters) from the plant when
132 THE CEREAL GRAINS | Wildrice, Zizania: Overview
Zizania aquatica var. aquatica L. southern Annual, tall, large panicles, many branched florets. Grows in muddy shores of streams in southern
wildrice Ontario and Quebec, southward to Florida and Louisiana. Average seed size: 14.3 mm long and 0.9 mm
diameter
Zizania aquatica var. brevis, Fassett, Annual, short, small panicle with few branched florets. Grows in tidal areas of St. Lawrence River estuary.
estuarine wildrice Average seed size: 6.7 mm long and 1.0 mm diameter
Zizania palustris var. palustris L. northern Annual, height ranges from short to tall, slender few-flowered panicle. Grows in water up to 120 cm deep
wildrice in southern Canada from New Brunswick to Manitoba and northern US states. Average seed size:
16.7 mm long and 1.4 mm diameter
Zizania palustris var. interior (Fassett), Annual, medium tall, medium to large panicle with numerous florets. Grows in water 3060 cm deep
Dore, interior wild rice along rivers in southeastern Manitoba and adjoining Ontario and in the North Central States. The
germplasm of this species and variety has been used to develop cultivars for cultivation. Average seed
size: 10.7 mm long and 1.8 mm diameter
Zizania texana A.S. Hitchc., Texas wildrice Perennial, decumbent with many long stems, panicles short, short seeds. Grows only in a localized area
in San Marcos River in Texas. Average seed size: 6 mm long and 1.2 mm diameter
Zizania latifolia Turcz. Manchurian water Perennial, spreads by subterranean runners, tall, medium panicles, seeds short to medium in length.
rice Native grass of Manchuria, Korea, Japan, Burma, and northeastern India. Base of plant often becomes
infected with a fungus that is used as a vegetable delicacy. This species is commercially grown in
Japan, Korea, and China for its vegetable. Average seed size of collections from Japan: 7 mm long and
1.3 mm diameter
Figure 2 The traditional way (canoe and flail) of harvesting natural stands of wildrice. One individual (standing) propels the canoe through the stand,
while the other uses two flails (knockers) to remove the grain from the stalks. Photographs by Oelke EA, left photograph appeared in Oelke EA
(2007) Saga of the Grain A Tribute to Minnesota Cultivated Wild Rice Growers. Lakeville, MN: Hobar Publications.
mature, stands are harvested by this method every other day for processed in this manner. Most of it is processed with mechan-
a period of 2 weeks. Enough grain falls back into the water ical equipment as described later.
before and during harvest to reseed the lake or river for the next
year. Figure 2 depicts the traditional harvest method. In Can- Zizania aquatica L. and Zizania texana Hitchc.
ada, natural stands are generally harvested by airboats fitted The seed from these species is either too thin or small to make
with a grain catcher onto the front. it worthwhile to harvest as a food source. Z. texana is on the
The traditional method of processing the harvested wildrice endangered species list.
was to lay it on the ground for drying since the grain has 40%
moisture when harvested. Later, the Native Americans used an Zizania latifolia Turcz.
iron kettle over a fire to parch (roast and dry) the grain This species is a perennial and is a native grass of Manchuria,
(Figures 3 and 4). The hulls (lemma and palea) were removed Korea, Japan, Burma, and northeastern India. The plant gener-
from the warm grain by placing it into a round-bottom con- ally is not used for grain production but grown commercially
tainer or leather-lined hole and then walking in place (jigging) for its vegetable delicacy, especially in Japan, Korea, and China.
in the container. The chaff was removed by tossing the grain The base of the plant becomes infected with a fungus. The
into the air (winnowing) and allowing the wind to blow it infected base of the plant is harvested and eaten as a vegetable.
away. The heavier grain was caught in the container. Today, In Japan, they call the vegetable Makomo-taki. The infected
very little of the grain harvested from natural stands is part is also dried and allowed to develop spores. The spores are
THE CEREAL GRAINS | Wildrice, Zizania: Overview 133
Figure 3 The traditional method of drying wildrice grain in a kettle Figure 4 The traditional way (jigging) to remove the outer hull of the
above a fire. The grain is stirred while drying. Photograph by Oelke EA, wildrice grains. The individual walks in place to rub the grain against each
appeared in Oelke EA, Grava J, Noetzel D, et al. (1982) Wild Rice other and also the kettle. Photograph by Oelke EA, appeared in Oelke EA,
Production in Minnesota. St. Paul, MN: University of Minnesota. Grava J, Noetzel D, et al. (1982) Wild Rice Production in Minnesota, St. Paul:
Agricultural Extension Service, Bulletin 464. University of Minnesota. Agricultural Extension Service, Bulletin 464.
fourth year. They continued their pioneering efforts and by were planted with seed collected from lakes. These fields were
1958 had a 50 ha diked field for growing wildrice. During the harvested with picker/harvesters on tracks. The fields had to be
mid-1950s and early 1960s, others started growing wildrice. In harvested several times during a 2-week period, due to seed
1965, Uncle Bens, Inc. started contracting for production, shattering, without cutting off the plants. Yields with this
which was a big impetus to produce wildrice. Today, there method were 6890 kg ha1 of processed grain. Fields
are 5000 ha in Minnesota and in California. A few ha are planted to the newer nonshattering cultivars could be har-
now also being grown in Hungary, Europe. vested once with a regular grain combine and yielded over
700 kg ha1 (Figure 7). The development of the nonshattering
cultivars resulted in a tremendous expansion of cultivated
Genetics and Breeding hectares in Minnesota and later in California. Figure 8 shows
Domestication of a wild species requires that plants are the cultivated wildrice production after processing for Minne-
selected from wild populations that maintain their grains on sota and California since 1968.
the plant even after maturity (nonshattering). The grain also Individual wildrice growers began their own selection pro-
needs to mature uniformly and has more grain production in grams to select better plant types in the 1960s, and some are
relation to vegetative production. One of the important factors continuing today. The wildrice breeding program at the Univer-
in the domestication process was the 1963 discovery, in a sity of Minnesota began in 1972. It continues till today and is
growers field, of a nonshattering plant by the grower and
two University of Minnesota scientists (Figure 6). All wildrice
species are mostly cross-pollinated with the male and female Female flowers (grain)
flowers separated on the same stem. Male flower retention and
Male flowers
seed shattering are almost always associated, facilitating selec-
tion for nonshattering. Subsequently improved cultivars were Panicle
developed from this germplasm by individuals, companies,
and the University of Minnesota. Up until that time, fields
Stem
Leaves
Tillers
Internode
section
Parchment-like
cross-partitions
Adventitious
root
Figure 6 The panicle on the left is a nonshattering type, while the one
on the right is a shattering type. The nonshattering type can be
recognized by loss of male flowers soon after pollen shed on the Figure 7 A drawing of a Zizania palustris plant. This type is a typical
branched (lower) portion of the panicle. Photograph by Oelke EA, nonshattering plant cultivated in fields. Reproduced from Oelke, EA,
appeared in Oelke EA, Grava J, Noetzel D, et al. (1982) Wild Rice Elliott WA, Kernkamp MF, and Noetzel DM (1973) Commercial
Production in Minnesota. St. Paul, MN: University of Minnesota. Production of Wild Rice. St. Paul, MN: University of Minnesota.
Agricultural Extension Service, Bulletin 464. Agricultural Extension Service, Folder 284.
THE CEREAL GRAINS | Wildrice, Zizania: Overview 135
14000
MN+CA
12000
CA
MN
10000
8000
6000
4000
2000
0
68
70
72
74
76
78
80
82
84
86
88
90
92
94
96
98
00
02
04
06
08
10
12
19
19
19
19
19
19
19
19
19
19
19
19
19
19
19
19
20
20
20
20
20
20
20
Figure 8 The amount of processed wildrice in 1000 kg from 1968 to 2013 from lakes and rivers in Minnesota (30%) and Canada (bottom red line),
Minnesota cultivated fields (yellow line), California cultivated fields (purple line), and the total from cultivated fields in Minnesota plus California
(top blue line). Graph by Porter RA (2014) Data provided by the Minnesota Cultivated Wild Rice Council, California Wild Rice Board, various wildrice
industry sources and Minnesota Department of Natural Resources.
the only public wildrice breeding program in the United States. up development resulting in a short, unproductive plant. The
To date, nine cultivars have been released all with some desir- high humidity also results in severe leaf disease problems.
able characteristics needed to further the domestication of wild- Cultivars have been developed that grow well in the warm day
rice. Improved shattering resistance is the primary goal, but but cooler night temperatures of Northern California. The cul-
disease resistance is a close secondary goal. Another goal is tivation practices for wildrice are similar to rice. Fields need to
shorter plants with higher yields. Yields have nearly doubled be kept flooded until several weeks before harvest. Wildrice will
from the original Johnson nonshattering (some resistance to grow well on organic or inorganic soils if nutrients are supplied.
shattering) cultivar developed from the nonshattering plants Wildrice requires less nitrogen than rice since it can get too tall
found in 1963. A private breeding program exists today in and lodge (fall down) if overfertilized. It will grow in deeper
California, which has developed cultivars for their warmer and and cooler water than rice; thus, water depth can be used as a
longer growing season. The genetic material for that program means of weed control in wildrice. Handling wildrice seed is
basically came from Minnesota. Thus far, all of the breeding is critical in establishing fields. Wildrice seed must be stored wet
done by conventional selection and not genetic engineering. and in cold (3 C) water for 3 months to release seed dormancy.
A program at Lakehead University, Thunder Bay, Canada, In Minnesota, new fields are seeded in the fall and the fields
selected types of plants that did well in specific lakes, but that flooded either in the fall or in the spring to a depth of 30 cm.
program has been reduced. Cultivated wildrice (Zizania palustris In Minnesota, fields can be kept in production for 23 years and
var. interior and palustris L.) of North America is an annual then rotated with other crops. After the first year, the fields seed
diploid with a chromosome number of 2n 2x 30; thus, it themselves from shattered seed even when initially seeded to
has 15 chromosome pairs. Compared with rice (Oryza sativa), the newer more shatter, resistant cultivars. In the Sacramento
wildrice has three more pairs of chromosomes and twice as much Valley of California, the fields are seeded in the spring of each
DNA content as rice. This research also indicated that when total year due to loss of viability of the shattered seed in the dry soil
wildrice DNA was used as a probe in southern hybridization to between seasons. Seed has to be stored in cool wet conditions
different Poaceae genera such as rice, oat, barley, wheat, and over the winter and then seeded the next spring. In the northern
maize, there was a strong hybridization signal with rice but higher elevations in the valleys between Mount Lassen and
relatively little with oat, barley, wheat, or maize. The close rela- Mount Shasta, the production practices are similar to Minne-
tionship of wildrice to rice will benefit the mapping of genes in sota (Figures 9 and 10).
wildrice. Similar probes used for mapping genes in rice have Fungal brown spot (Bipolaris sp.) is a severe disease in
been used in wildrice and some of the same genetic markers Minnesota, but not a problem in California. The wildrice
have been found to be located in the same area in wildrice as in worm (Apamea apamiformis Guenee), which is the larval stage
rice. Locating markers for important genes such as seed shatter- of noctuid moth, is the most serious insect pest in the Upper
ing has facilitated the wildrice breeding efforts for domestication. Midwest but not a problem in California. In Minnesota
and California, fields are drained 3 weeks before harvest.
Cultivation Practices Grain combines are adapted, with larger reels and often
full tracks, for harvesting wildrice. The grain has to be har-
Wildrice is well adapted to northern latitudes with its cooler vested at 2035% moisture since seed shattering will occur
climate. In southern latitudes of the United States, such as if harvested at lower moisture content. The grain is immedi-
Arkansas, shorter day lengths and higher temperatures speed ately transported to processing plants (Figure 11).
136 THE CEREAL GRAINS | Wildrice, Zizania: Overview
Processing consists of curing, parching (roasting and drying), The wildrice kernel with the hulls removed is similar to other
hulling, cleaning, scarification, and grading. When the grain cereals and consists of a pericarp, aleurone layer, endosperm,
arrives at the processing plant, it is put into piles or containers and embryo. The pericarp and embryo each represent 5% of
and kept for 1 or 2 days to allow for more uniform color. Then, the kernel weight. The rest, 90%, is largely endosperm and a
the grain is put into rotating, heated drums (parchers) that will smaller amount of the aleurone layer. The nutritional quality is
hold 225370 kg and heated for 1 h (Figure 12). The grain very good for a cereal grain. It is equal to or better than other
is dried to 7% moisture and to 135 C. After parching, the cereals such as wheat, barley, or rice. The protein content is
hot grain is passed through a huller consisting of two rubber about that of hard red spring wheat 1315%, while brown
rollers going at different speeds. The light chaff and other rice has 8%. The sum of lysine, threonine, and methionine
foreign material are evacuated from the kernels, and the hulled content is similar to hulled oats, which is considered one of the
kernels are graded with graders and gravity tables for size better cereals for humans. The mineral content of the wildrice
(Figure 13). Sometimes before grading, the kernels are scari- kernel varies but can be similar to oats. Processed wildrice
fied by passing through a tube with rubber paddles. This contains no vitamin A but is an excellent source of B vitamins.
removes some of the darker, impermeable outer layer of the Wildrice is nutritious, since it is not refined but consumed as a
kernel to allow a faster cooking time (Figure 14). The parched, whole grain (Tables 24).
graded kernels can then be stored for future packaging Wildrice has good antioxidant properties. There is consid-
(Figure 13). erable interest in antioxidants, which can slow diseases such as
coronary heart disease and certain types of cancer. Hydrated
wildrice mixed with ground beef has significant antioxidant
activity.
Figure 10 In Minnesota, wildrice fields in production for two or more years, plant populations are reduced by airboats. V-shaped blades mounted
on the rear are dropped into the water to the soil line to remove excess plants, which leaves rows of plants for production. Excessive amounts of seeds,
even from the more shatter resistant cultivars, fall onto the soil before and during harvest often producing too many plants the following year.
Photographs by Oelke EA.
THE CEREAL GRAINS | Wildrice, Zizania: Overview 137
Figure 13 Processing is done with modern grain handling equipment in clean processing plants. Storage is done in clean warehouses before sale or
packaging. Photographs by Oelke EA.
138 THE CEREAL GRAINS | Wildrice, Zizania: Overview
Figure 14 Processed kernels from cultivated fields that were slightly scarified for faster cooking (left). Kernels without and with hulls on the right. Hulls
are removed during processing. Hand-processed kernels, from lakes and rivers, most often are lighter in color than the ones in the left photograph.
Photographs Oelke EA.
Table 2 Composition of wildrice, medium grain brown rice, hard red spring wheat, and oats in 100 g
Nutrient Unit Wildrice Brown rice, medium grain Wheat, hard red spring Oats
Data from US Department of Agriculture, Agricultural Research Service (2013) USDA National Nutrient Database for Standard Reference, Release 26.
To facilitate the selection of better cultivars, better and Processing of wildrice is still a batch system that slows
new genetic markers are needed. See Further Reading processing; thus, there is a need to develop storage
references: Hass BL, Pires JC, Porter R, Phillips RL (2003) techniques for the high moisture grain before processing.
Comparative genetics at the gene and chromosome levels See Further Reading references: Marcum DB (2007) Wild
between Oryza sativa and wildrice Zizania palustris. Rice Production in California. Davis, CA: University of Cal-
Theoretical and Applied Genetics 107: 773782; Kennard ifornia. Extension Publication, No.21622; Oelke EA,
WC, Phillips RL, Porter RL (2002). Genetic dissection of Boedicker JJ (2000) Wild rice: Processing and utilization.
shattering, agronomic, and color traits in American wildrice In: Kulp K, Ponte JG (eds.) Handbook of Cereal Science and
(Zizania palustris var. interior L.) with a comparative map. Technology, pp. 275306. New York: Marcel Dekker; Oelke
Theoretical and Applied Genetics 105: 10751086; Kennard EA (2007) Saga of the Grain A Tribute to Minnesota
WC, Phillips RL, Porter RA, Grombacher AW (2000) A Cultivated Wild Rice Growers. Lakeville, MN: Hobar
comparative map of wild rice (Zizania palustris L. Publications.
2n 2X 30). Theoretical and Applied Genetics 101: Additional research needs to be conducted to verify the low
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and hard red spring wheat in 100 g Harmony with Nature. Saskatoon: Saskatchewan Agriculture and Food.
Cardwell VB, Oelke EA, and Elliott WA (1978) Seed dormancy mechanisms in wild rice
Amino acid Unit Wildrice Oats Brown rice Wheat (Zizania aquatica). Agronomy Journal 70: 481.
Chang TT (1989) Domestication and spread of the cultivated rices. In: Hillman GC
Tryptophan g 0.179 0.234 0.096 0.195 and Harris DR (eds.) Foraging and Farming: The Evolution of Plant Exploitation,
Threonine g 0.469 0.575 0.275 0.433 pp. 408417. UK: Routledge.
Fassett NC (1924) A study of the genus Zizania. Rhodora 26: 153.
Isoleucine g 0.618 0.694 0.318 0.541
Hass BL, Pires JC, Porter R, and Phillips RL (2003) Comparative genetics at the gene
Leucine g 1.018 1.284 0.620 1.038 and chromosome levels between Oryza sativa and wildrice Zizania palustris.
Lysine g 0.629 0.701 0.286 0.404 Theoretical and Applied Genetics 107: 773782.
Methionine g 0.438 0.312 0.169 0.230 Jenks AE (1900) The wild rice gatherers of the Upper Lakes: A study in American
Cystine g 0.174 0.408 0.091 0.404 primitive economics. In: 19th Annual Report of the Bureau of American Ethnology,
Phenylalanine g 0.721 0.895 0.387 0.724 189798, Part 2, pp. 10131137. Washington, DC: US Department of Interior.
Tyrosine g 0.622 0.573 0.281 0.441 Kennard WC, Phillips RL, Porter RA, and Grombacher AW (2000) A comparative map of
Valine g 0.858 0.937 0.440 0.679 wild rice (Zizania palustris L. 2n 2X 30). Theoretical and Applied Genetics
Arginine g 1.136 1.192 0.569 0.702 101: 677684.
Kennard WC, Phillips RL, and Porter RL (2002) Genetic dissection of shattering,
Histidine g 0.384 0.405 0.190 0.330
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Alanine g 0.825 0.881 0.437 0.555 with a comparative map. Theoretical and Applied Genetics 105: 10751086.
Aspartic acid g 1.419 1.448 0.702 0.808 Kosbau H (2005) The Taming of Wild Rice. Grand Rapids, MN: Treasure Bay Printing.
Glutamic acid g 2.565 3.712 1.528 4.946 Marcum DB (2007) Wild Rice Production in California. Extension Publication, Davis,
Glycine g 0.672 0.841 0.369 0.621 CA: University of California, No.21622.
Proline g 0.519 0.934 0.352 1.680 Nelson RN and Dahl RP (1985) Wild Rice Market Shows Vigorous Growth. St. Paul,
Serine g 0.778 0.750 0.388 0.663 MN: University of Minnesota, Minnesota Agriculture Economics, No.649.
Oelke EA (2007) Saga of the Grain A Tribute to Minnesota Cultivated Wild Rice
Data from US Department of Agriculture, Agricultural Research Service (2013) Growers. Lakeville, MN: Hobar Publications.
National Nutrient Database for Standard Reference, Release 26. Oelke EA and Boedicker JJ (2000) Wild rice: Processing and utilization. In: Kulp K and
Ponte JG (eds.) Handbook of Cereal Science and Technology, pp. 275306.
New York: Marcel Dekker.
Table 4 The amounts of nutrients based on 1/2 cup serving of Oelke EA, Elliott WA, Kernkamp MF, and Noetzel DM (1973) Commercial Production of
cooked product Wild Rice. St. Paul, MN: University of Minnesota, Agricultural Extension, Service
Extension Folder 284.
Nutrients (based on 1/2 Enriched white Brown rice Oelke EA, Grava J, Noetzel D, et al. (1982) Wild Rice Production in Minnesota.
cup serving (cooked)) Wildrice rice (long grain) (long grain) Agricultural Extension Service, St. Paul, MN: University of Minnesota, Extension
Bulletin 464.
Calories 83 103 108 Oelke EA, Porter RA, Grombacher AW, and Addis PB (1997) Wild rice New interest in
Protein (g) 3.27 2.13 2.52 an old crop. Cereal Foods World 42: 234247, American Association of Cereal
Chemists, Minneapolis, MN.
Fat (g) 0.28 0.22 0.88
Piperno DR (2012) Archaeobotanical information on early cultivation and plant
Dietary fiber (g) 1.5 0.3 1.8
domestication involving microplant (phytolyth and starch grain) rements.
Total carbohydrates (g) 17.5 22.25 22.39 In: Gepts P, Famula TR, Bettinger RI, Brush SB, Damania AB, McGuire PE, and
Minerals Qualset CO (eds.) Biodiversity in Agriculture: Domestication, Evolution, and
Phosphorus (mg) 67 34 81 Sustainability, p. 203. Cambridge: Cambridge University Press.
Iron (mg) 0.49 0.95 0.41 Steeves TA (1952) Wild rice Indian food and a modern delicacy. Economic Botany
Potassium (mg) 83 28 42 26: 107142.
Zinc (mg) 1.1 0.39 0.61 Vennum T (1988) Wild Rice and the Ojibway People. St. Paul, MN: Minnesota Historical
Vitamins Society.
Wu K, Zhang W, Addis PB, Epley RJ, Salih A, and Lehrfeld J (1994) Antioxidant
Riboflavin (mg) 0.07 0.01 0.02
properties of wild rice. Journal of Agricultural and Food Chemistry 42: 34.
Niacin (mg) 1.05 1.16 1.49
Vitamin E (mg) 0.2 0.03 0.03
Reproduced with permission from the Minnesota Cultivated Wild Rice Council, St. Paul. Relevant Websites
Data from the U. S. Department of Agriculture, Agricultural Research Service (2013)
USDA National Database for Standard References, Release 26. http://www.ars.usda.gov/ba/bhnrc/ndl This website is an excellent source for nutrient
content of many grains and foods including wildrice.
http://www.cawildrice.com/ This is the website of the California Wild Research and
See also: Breeding of Grains: Rice: Breeding; Genetics of Advisory Board of the California cultivated wild rice growers. Information on
Grains: Rice: Genetics; Non-wheat Foods: Rice: Chinese Food California wildrice cultivation, food preparation and recipes can be found on this
Uses; The Basics: Taxonomic Classification of Grain Species; The website.
http://www.mnwildrice.org This is the website of the Minnesota Cultivated Wild
Cereal Grains: An Overview of the Family of Cereal Grains Prominent
Rice Council whose members consist of Minnesota cultivated wildrice
in World Agriculture; Oats: Overview. growers. Information on wildrice history, Minnesota cultivation, research,
processing, nutrition, gluten content, cooking, and recipes can be found on this
website.
Further Reading http://www.wildrice.org This website is the one for the International Wild Rice
Association (IWRA). IWRA consists of wildrice growers, processors and marketers.
Aiken SG, Lee PF, Punter D, and Stewart JM (1988) Wild Rice in Canada. Winnipeg: Members include individuals from the US and Canada. Information on wildrice
Agriculture Canada. history and links to other wildrice websites can be found on this website.
African Rice (Oryza glaberrima): A Brief History and Its Growing Importance
in Current Rice Breeding Efforts
JT Manful and S Graham-Acquaah, Africa Rice Center, Cotonou, Republic of Benin
2016 Elsevier Ltd. All rights reserved.
Topic Highlights species, thus creating crosses that are better adapted, and
higher yielding under adverse conditions, than either parent.
The two cultivated species of rice are Oryza sativa (Asian This article discusses the history of glaberrima, its genetic and
rice) and Oryza glaberrima (African rice). morphoagronomic diversity, its use in conventional breeding,
O. glaberrima have tolerance to several biotic and abiotic and some of its physicochemical characteristics.
stresses and are particularly suited for low-input subsistence
farming as practiced in many African communities.
African rice has yield potential comparable to O. sativa as Origin and History of O. glaberrima
evidenced by yield-related morphoagronomic traits. Their
tendency to easily shatter is a responsible factor for the O. glaberrima is unique to Africa. It is believed to have been
lower yields observed. domesticated from the wild ancestor O. barthii over 3000 years
NERICA (New Rice for Africa) a product of interspecific ago. The many indigenous words for rice in several West Afri-
breeding between O. glaberrima and O. sativa developed by can language groups predate the Portuguese-derived words
the Africa Rice Center combines the stress tolerance of associated with Asian rice and offer linguistic evidence to the
African rice with the high yield potential of Asian rice. fact that rice was cultivated prior to the arrival of Europeans in
The recent release of O. glaberrima genome and the devel- Africa. Two distinct theories have been advanced as to the exact
opment of African rice-specific single-nucleotide polymor- origin and domestication of the African species. The first theory
phisms will enhance the identification and exploitation of propounded by Porteres suggests that O. glaberrima was first
adaptive traits in African rice in rice improvement domesticated in the Inland Delta of the Niger River in present-
programs. day Mali and then subsequently spread to other centers of
Very little work has been done on the physicochemical diversification, some being the coast of Gambia, the Casa-
properties of O. glaberrima. Ongoing studies by the Africa mance region of Senegal, and Guinea Bissau along the Niger
Rice Center show that a high level of genetic diversity exists River and then in the Guinea forest between Sierra Leone and
in the accessions of O. glaberrima from the West Africa sub- the western Ivory Coast. Others claim that ancient hunting and
region as evidenced by variations in both apparent amylose gathering human populations living in different localities
content and pasting properties. within the vast forest and savanna areas where O. barthii grew
may have selected for O. glaberrima. Recent studies into the
population structure and genomics of O. glaberrima accessions
Learning Objectives tend to support a single domestication origin of African rice in
areas of the upper Niger and Senegal Rivers.
At the end of this article, it is expected that readers would African rice, as has been asserted, was introduced into the
new world via the trans-Atlantic slave trade. Botanical evidence
have knowledge of the origin, domestication history, and and historical records suggest that African rice was grown in El
diversity in African rice; Salvador, French Guiana, Brazil, Jamaica, South Carolina, and
understand the traits that set O. glaberrima apart from O. Suriname and is to this day grown, eaten, and exported by the
sativa; Maroons. Although it has diminished in terms of cultivated
know how O. glaberrima is used in classical breeding; area as a result of the introduction of O. sativa, O. glaberrima
have an appreciation of ongoing research work on the remains an important crop for certain communities in West
physicochemical properties of African rice. Africa who still grow them for use in rituals and as food due to
its unique taste and texture. Their hardiness in the field also
makes them ideal for subsistence farming.
Introduction
Rice is a major staple in the world and belongs to the genus What Makes O. glaberrima Different?
Oryza. Out of over 20 species in the genus Oryza, only two
species are cultivated: Oryza glaberrima (African rice) and Oryza Differentiating between the two cultivated species of rice in the
sativa (Asian rice). O. glaberrima is native to sub-Saharan Africa field can be cumbersome for the untrained as they differ mar-
(SSA) thought to have been domesticated from the wild ances- ginally in their morphology. In general, O. glaberrima has short
tor Oryza barthii (formerly known as Oryza breviligulata). round ligules, while the panicles are erect and simply branched
O. glaberrima is presently being replaced everywhere in with small spikelets. The grains are small and pear-shaped with
Africa by O. sativa and is rapidly diminishing in importance. a reddish-brown to purple bran color and olive to black seed
Recent advances now allow for gene transfer between the two coats. As a result of this bran color, O. glaberrima are often
Plate 1 Panicles of O. sativa (a) and (b) and O. glaberrima (c) and (d).
referred to as red rice. Compared with Asian varieties, African silent sites). African rice is reported to have more than 70% less
rice usually gives lower grain yields, readily shatters, and is diversity compared to its wild progenitor resulting from a
difficult to mill because of the brittle nature of the grains severe genetic bottleneck during its domestication from the
(Plate 1). initial population of O. barthii.
These notwithstanding, O. glaberrima possess distinct char- An earlier characterization of 198 accessions using SSR
acteristics, perhaps due to the fact that they have survived with (simple sequence repeat) markers showed a range of 227
little interference from humans, which make them ideal for the alleles per locus with an average of 9.4 alleles per locus.
low-input subsistence farming still practiced in many African Based on this characterization, O. glaberrima were grouped
communities. The plants have wider leaves that shade out into five genetic groups with two of these segregating with O.
weeds and are more tolerant than O. sativa to biotic and abiotic sativa indicating intermediaries and thus gene flow between the
stresses such as diseases, pests, fluctuations in water depth, iron two species.
toxicity, infertile soils, severe climatic conditions, and neglect. A morphoagronomic and molecular characterization of
African rice varieties collected from the inland delta of the
Niger River (Mopti and Tombouctou), Sikasso, and Gao
Diversity in African Rice Region in Mali believed to be centers of domestication for O.
glaberrima revealed a higher average heterozygosity (0.18) in O.
O. glaberrima is largely being replaced by O. sativa because of glaberrima than previously reported. The population collected
the higher yields obtained from the latter. In spite of this, it is from Mopti and Gao Region had higher diversity than other
still being grown in many areas because of their preference for populations. Although there was no clear clustering of individ-
the preparation of certain traditional foods and also for their uals based on location or growing condition because of the
tolerance to biotic and abiotic stresses. This stress tolerance probable higher gene flow that occurs between different grow-
prompted efforts at its conservation and use as a gene pool ing conditions within a location, a higher differentiation was
for improving Asian rice varieties. The Africa Rice Center observed between populations from different locations than
(www.africarice.org) is leading efforts at conservation and from different ecologies, while within locations, similarities
exploitation of O. glaberrima in breeding. The gene bank of were observed between populations from similar growing con-
the center contains about 3000 accessions of O. glaberrima that ditions. The accessions RAM116, RAM100, and RAM122 out-
are being systematically screened. performed the best check varieties under drought conditions.
O. glaberrima has long been perceived to possess less genetic Recently, single-nucleotide polymorphisms (SNPs) specific
diversity in comparison with O. sativa. In fact, African rice is to African rice have been developed and used to genotype some
reported to have only one-sixth of the nucleotide diversity 279 O. glaberrima accessions, selected based on their impact in
found in Asian rice. Indica rice (O. sativa ssp. indica) alone current breeding programs and their geographic distribution,
has higher diversity (ysil 0.00280.0029) than either African and 101 O. barthii accessions, selected based on their
rice (ysil 0.0007) or its wild ancestor O. barthii (ysil 0.0024) geographic origins. A structuring of accessions into three popu-
(Wattersons estimator of y per base pair calculated based on lations of O. barthii associated with geography and two
142 THE CEREAL GRAINS | African Rice (Oryza glaberrima): History and Its Growing Importance in Rice Breeding Efforts
populations of O. glaberrima that are yet to be linked with any early-maturing varieties ensure that they escape drought in
phenotyped traits was reported. Also, an apparent misclassifi- drier areas and late-season fungal disease in wet ecologies.
cation in both O. glaberrima and O. barthii was identified in Rice plant height and stem strength are related to lodging.
addition to a new form of O. sativa within the African rice Generally taller varieties with weaker stems are more prone to
varieties. The complete genomic sequence of O. glaberrima lodging especially in high soil fertility conditions. Most O.
has been deciphered through immense global research effort. glaberrima varieties are deemed intermediate in stature
The release of O. glaberrima genome and the development of (80100 cm) but possess fragile culms that render them sus-
African rice-specific SNPs bring in their wake hitherto non- ceptible to lodging during their vegetative stage. A number of
existing opportunity for the identification and exploitation of O. glaberrima have, however, been observed to have thick leaf
adaptive traits in African rice in rice improvement programs. sheaths making them resistant to lodging. Such varieties may
Little variation has been reported in the morphoagronomic be useful in breeding.
traits of O. glaberrima compared with O. sativa. With respect to The greatest limitation to the cultivation of O. glaberrima is
their vegetative growth, more than 90% of O. glaberrima their lower yields. Yield-related characteristics have been
screened by the Africa Rice Center exhibited high seedling studied, and results point to the fact that O. glaberrima have
vigor. A very rapid vegetative growth culminating in large the same yield potential as O. sativa. Yield-related morpho-
number of tillers and droopy lower leaves was observed. agronomic traits such as average panicle number in O. glaber-
These vegetative growth characteristics (vigorous initial rima are comparable to O. sativa. Shattering could also be a
growth) typified by their high tillering ability, large leaf area, contributory factor to the inferior yields as spikelet number in
and droopy leaves provided rapid ground cover, thus suppres- O. glaberrima prior to shattering is no less than in O. sativa. O.
sing weeds and making them weed-competitive (Plate 2). glaberrima possess higher dry matter production and greater
O. glaberrima also vary greatly in their growth durations leaf area than O. sativa at least until heading. They have lower
with as much as 89 days between some varieties. Whereas photosynthetic rate per leaf area but a higher rate against the
traditional O. sativa and improved varieties have maturity same leaf nitrogen content in a low content range. Their
days ranging between 120140 days and 80120 days, respec- response to dry matter, leaf area, and leaf photosynthesis to
tively, some O. glaberrima can mature within 75 days. Such increases in nitrogen inputs is higher than in O. sativa.
AG: Vigor of selected O. glaberrima and O. sativa after 14 days of seeding dry season
AG: Vigor of selected O. glaberrima and O. sativa after 21 days of seeding wet season
Plate 2 Seedling vigor in O. sativa and O. glaberrima. Photo courtesy of Venuprasad Ramaiah; Africa Rice Center.
THE CEREAL GRAINS | African Rice (Oryza glaberrima): History and Its Growing Importance in Rice Breeding Efforts 143
Additionally, water use efficiency on dry matter accumulation determine the market value and reputation of rice varieties.
and gas exchange bases is lower compared with that of O. Hitherto, studies on the physicochemical properties of O. gla-
sativa, while both leaf senescence during maturity and grain berrima have not received as much attention as work on their
filling are faster in O. glaberrima than O. sativa. agronomic traits. The few studies conducted suggest that gen-
The greatest attribute of O. glaberrima is their resilience. erally, O. glaberrima tend to break easily during milling; thus,
Accessions of O. glaberrima have been found to be highly resis- head rice recovery is lower compared with O. sativa. The red
tant to pests and diseases such as African rice gall midge, the rice pericarps of African rice make them less attractive when
yellow mottle virus, and blast. O. glaberrima can limit water loss polished. O. glaberrima is preferred by certain groups of con-
through stomata closure and leaf curling. This trait in addition sumers, particularly in regions where they are still cultivated,
to the well-developed root system makes them well adapted to because of their distinct taste and flavor. Some aroma com-
drought conditions. The ability for strong leaf elongation under- pounds such as 4-ethylphenol (musty) and (E, E)-2,4-
water that leads to enhanced expansion of leaf area and photo- heptadienal (fruity) have been found to be unique to
synthesis, requirements for escaping complete submergence O. glaberrima. On cooking, O. glaberrima have firmer texture,
underwater over a prolonged period (up to 37 days), has also perhaps due to their higher apparent amylose content (AAC)
been observed in O. glaberrima, making them a promising res- and unique pasting properties (low peak, low breakdown, and
ervoir of genes for selecting and breeding rice genotypes for higher setback viscosities) compared with O. sativa. Asian rice
different flood-prone environments in Africa. Tolerance to iron however has greater diversity in AAC than African rice. While
toxicity and saline environments has also been documented in waxy varieties exist among Asian rice, waxy varieties are yet to
O. glaberrima. Although some O. sativa varieties are better at be encountered among O. glaberrima.
tolerating some specific stresses, a unique trait in O. glaberrima With the broad objective of contributing to better under-
is that some can possess multiple tolerances to various stresses. standing the physicochemical properties of O. glaberrima,
CG 14 is one such O. glaberrima variety that has weed AfricaRice and its partners characterized over 1000 O. glaber-
competiveness and iron toxicity and drought tolerance. These rima accessions from its gene bank based on data of their Rapid
make them ideally suited for cultivation in SSA where rice Visco Analyser (RVA) pasting characteristics and AAC. Based
farming is predominantly carried out by resource-poor farmers. on the AAC and RVA pasting parameters of the rice flours, the
accessions were hierarchically clustered into five groups C I,
C II, C III, C IV, and C V (Figure 1). The distribution pattern
O. glaberrima in Classical Plant Breeding indicated that the maximum number of accessions (282) was
included in C I, followed by C IV (242), C V (231), C III (161),
At the Africa Rice Center (AfricaRice), attempts have been made and C II (104), with all five clusters showing a large diversity of
to develop new interspecific rice varieties from crosses between AAC and pasting properties.
O. sativa and O. glaberrima. Initially, this proved difficult as the Cluster I had 282 accessions and constituted 27.6% of the
progenies were sterile. This problem was, however, successfully entire population. These accessions were further grouped into
overcome, and the first large-scale hybridization of the two five subclusters SC IA, SC IB, SC IC, SC ID, and SC IE with a
cultivated species of rice occurred. The resulting varieties were population of 44 (15.6%), 84 (29.8%), 107 (37.9), 26 (9.2%),
collectively named NERICA (New Rice for Africa). These varie- and 21 (7.4%), respectively. All the subclusters under cluster I
ties, developed for both upland and lowland ecologies, combine had similar ranges of AAC all of them fell under high-amylose
the high yield of O. sativa with the stress tolerance and adapt- rice category. However, their pasting characteristics differed
ability to local growing conditions of O. glaberrima. The NERICA greatly as shown in Table 1. Wide variations were observed for
development process began with an initial screening of 1721 peak and trough viscosities in all the five subclusters. With
accessions comprising 1130 O. glaberrima and 591 O. sativa respect to breakdown viscosity, while SC IB, SC IC, SC ID, and
based on their morphoagronomic performance. O. glaberrima SC IE were close, that of SC IA was relatively very distinct. SC IA,
(CG 14) and three O. sativa ssp. japonica (WAB56-104, WAB56- SC IC, and SC IE were likewise similar with respect to final and
50, and WAB181-18) are the parental lines for the first set of setback viscosities, but SC IB and SC ID showed diversity.
NERICAs (NERICA 118). These varieties were crossed to obtain Cluster II with 104 accessions composed the lowest propor-
interspecific lines that were backcrossed and the progenies sub- tion (10.2%) of the entire population. The broadest ranges of
jected to pedigree selection to obtain the first NERICAs (NERICA peak, trough, setback, and final viscosities were observed in this
118) for upland ecology. Further research and improved meth- cluster. However, it had the second broadest AAC range and the
odology led to the release of 60 varieties (NERICA L1L60) for narrowest pasting temperature range. The range of AAC for
lowlands. The NERICA-S lines have also been bred for the accessions in C II conferred them to intermediate and high-
Sahelian zones of West Africa. These NERICA varieties devel- amylose types of rice. Three subclusters SC IIA, SC IIB, and SC
oped and disseminated have affected farmers livelihoods and IIC were formed under cluster II with populations of 26 (25%),
contributed to poverty reduction in SSA. 54 (52%), and 24 (23%) accessions, respectively. SC IIA and SC
IIB had similar ranges of AAC, with SC IIC having the broadest
range (Table 1). SC IIC also had the broadest range of peak,
Physicochemical Properties of O. glaberrima final, setback, and trough viscosities, while SC IIB had the
narrowest range as regards peak, final, and trough viscosities.
The physicochemical properties of rice such as milling recov- With 161 accessions, cluster III constituted 15.8% of the
eries, physical appearance (grain dimensions and chalkiness), entire population of the O. glaberrima analyzed (Table 1) and
aroma, cooking and pasting properties are important as they had five subclusters, namely, SC IIIA, SC IIIB, SC IIIC, SC IIID,
144 THE CEREAL GRAINS | African Rice (Oryza glaberrima): History and Its Growing Importance in Rice Breeding Efforts
O. glaberrima
accessions
C I C II C III C VI CV
(N = 282) (N = 104) (N = 161) (N = 242) (N = 231
Figure 1 Graphical representation of distribution of 1020 African rice accessions into 5 clusters and 23 subclusters.
and SC IIIE, making up 23%, 24.2%, 26.7%, 11.2%, and second broadest range of setback and breakdown viscosities,
14.9% of the accessions, respectively. Cluster III had the nar- and the third broadest AAC range.
rowest ranges of peak, trough, setback, and final viscosities.
The accessions in this cluster belonged to the category of
intermediate- to high-amylose rice based on their AAC range. Conclusions
SC IIIA had the broadest apparent amylose range, SC IIID had
the narrowest range, and that of SC IIIB and SC IIIE was The tremendous amount of work on O. glaberrima at AfricaRice
similar. SC IIIA also had the broadest ranges with respect to notwithstanding the full potential of the species is yet to be
peak, trough, breakdown, and final viscosities (Table 1). exploited. There remains the need to consider intraspecific
Despite having the narrowest final viscosity range, SC IIIC breeding within O. glaberrima and ways in which the multiple
had the second broadest ranges of apparent amylose, peak resistance traits of O. glaberrima could be transferred into new
viscosity, and breakdown viscosity (Figure 2). varieties.
Accessions in cluster IV had the broadest range of apparent The studies of the physicochemical properties supported
amylose, the second broadest peak viscosity range, and the the idea that there is a high level of genetic diversity in the O.
fourth broadest ranges of both final and setback viscosities glaberrima accessions as evidenced by the variation in both
but the narrowest range of breakdown viscosity. Low-, AAC and pasting properties in and within the clusters and
intermediate-, and high-amylose rice types were all included subclusters. This suggests that there may be a valuable resource
in the accessions within this cluster. Cluster IV formed 23.7% of potentially very important gene pool that may be of benefit
of the entire population analyzed (242 accessions) and con- to interested rice breeding programs.
tained five subclusters, namely, SC IVA, SC IVB, SC IVC, SC Additionally, a detailed molecular analysis of a representa-
IVD, and SC IVE, which constituted 33.1%, 38.8%, 17.4%, tive sample of these accessions from the clusters such as those
5.4%, and 5.4% of the accessions, respectively (Table 1). with a high tendency to retrograde (high setback and poten-
Whereas SC IVA, SC IVB, and SC IVC shared some similarities tially slower rate of digestion due to retrogradation) or form
in AAC and peak time, very diverse peak, trough, breakdown, viscous paste/gel after cooking and cooling (high final viscos-
final, and setback viscosities were observed in all the five sub- ities) will ensure the use of the diversity in O. glaberrima in the
clusters except in SC IVE, which, though distinct with respect to future development of acceptable commercial cultivated rice
breakdown viscosity, showed very close similarity to SC IVB varieties that have the desired amylose content and pasting
with regard to peak, breakdown, final, and setback viscosities. characteristics to meet consumer preferences in Africa and
The 231 accessions in cluster were intermediate to high- elsewhere.
amylose rice types and formed 22.6% of the total number of
accessions analyzed. This cluster also contained five sub-
clusters, namely, SC VA, SC VB, SC VC, SC VD, and SC VE, Exercises for Revision
which constituted 25.5%, 15.2%, 32.9%, 9.1%, and 17.3% of
accessions, respectively (Table 1). Very similar AAC ranges Outline the theories of domestication of African rice.
were observed in the subclusters except for SC VE. The acces- How do the two cultivated species of rice differ from each
sions in cluster V also had the broadest range of final viscosity, other?
Table 1 Distribution of 1020 O. glaberrima rice accessions obtained through cluster analysis using SPSS
Range
I C I (282) 24.729.4 67.3123.8 66.7124.2 0.4to 6.6 149.6296.8 71.7173.0 5.97.0 89.694.5
SC IA (44) 25.329.4 76.8115.6 73.3113.4 1.06.6 160.1225.4 71.7124.5 5.96.5 90.793.6
SC IB (84) 24.827.9 67.387.4 66.787.3 0.4 to 2.2 149.6188.3 75.0104.3 6.17.0 91.994.0
SC IC (107) 24.728.5 76.3101.6 76.6101.8 0.4 to 0.8 170.4234.9 75.8139.5 6.37.0 91.994.5
SC ID (26) 26.029.0 92.9123.8 93.0124.2 0.3 to 2.2 217.7296.8 118.0173.0 6.37.0 91.593.6
SC IE (21) 25.427.7 89.4107.9 89.7105.8 0.2 to 2.2 176.3217.1 86.8116.8 6.47.0 89.692.8
II C II (104) 20.629.6 152.2230.7 113.0183.5 6.049.5 246.5421.5 94.9224.8 5.75.9 86.491.9
SC IIA (26) 23.029.2 137.8182.7 125.3160.9 6.032.4 308.1394.8 54.8224.8 5.76.2 88.391.9
SC IIB (54) 23.929.6 135.5165.8 115.9143.0 6.934.2 246.5327.3 95.3170.7 5.75.9 87.291.0
SC IIC (24) 20.628.5 152.2230.7 113.0183.5 21.949.5 247.0421.5 94.9190.8 5.75.9 86.489.2
III C III (161) 21.726.8 80.9125.5 80.4122.2 0.2 to 159.9258.8 68.3149.8 5.97.0 89.694.0
12.8
SC IIIA (37) 21.726.8 89.2125.5 88.2122.3 0.2 to 228.3258.8 120.8149.8 5.97.0 90.793.6
12.8
SC IIIB (39) 23.226.6 89.8110.8 89.8103.1 0.2 to 9.2 212.1232.8 107.8132.2 5.97.0 90.793.6
SC IIIC (43) 22.626.5 80.9110.0 80.4104.1 0.2 to 194.3211.5 92.1119.1 5.97.0 89.994.0
11.8
SC IIID (18) 24.626.4 108.2121.3 100.7115.5 2.410.1 213.6236.1 103.3117.2 5.96.2 89.692.0
SC IIIE (24) 23.426.4 82.9107.9 81.0105.0 0.16.5 159.9196.6 68.3101.2 6.07.0 90.392.4
IV C IV (242) 15.128.6 36.493.1 36.593.0 0.5 to 1.2 70.8205.8 33.0122.0 6.37.0 91.595.6
SC IVA (80) 22.427.8 53.673.4 54.073.8 0.5 to 0.0 119.8158.4 57.989.8 7.07.0 92.895.2
SC IVB (94) 15.128.6 63.683.9 63.784.0 0.4 to 0.2 135.8180.4 62.3105.0 6.87.0 91.594.7
SC IVC (42) 23.027.6 36.463.3 36.563.4 0.4 to 0.1 70.8118.7 33.067.8 7.07.0 92.495.6
SC IVD (13) 21.327.4 74.593.1 74.393.0 0.2 to 0.4 171.0205.8 81.2122.0 6.67.0 92.094.4
SC IVE (13) 21.725.6 50.883.3 50.482.9 0.4 to 1.2 123.4176.0 60.498.0 6.36.7 91.994.0
V C V (231) 21.028.8 96.3152.3 87.1141.4 0.822.2 194.2343.5 84.8197.7 5.76.4 87.192.4
SC VA (59) 25.328.8 117.3152.3 104.9139.3 6.322.2 212.4275.3 84.8137.7 5.76.0 87.191.1
SC VB (35) 25.728.8 114.3151.1 108.5141.4 1.614.8 255.7343.5 135.2197.7 5.86.3 89.992.4
SC VC (76) 25.528.5 100.8131.6 198.3124.8 0.810.1 209.7268.1 91.3153.2 5.96.4 89.192.4
SC VD (21) 25.628.1 96.3117.2 87.1104.2 3.122.0 194.2229.4 90.5117.3 5.76.0 89.391.5
SC VE (40) 21.027.5 108.3141.5 95.9127.5 5.717.0 237.5306.7 111.4167.7 5.76.2 88.792.0
C, cluster; SC, subcluster; PV, peak viscosity; TV, trough viscosity; BDV, breakdown viscosity; FV, final viscosity; SBV, setback viscosity; PT, peak time; PTemp, peak temperature;
AAC, apparent amylose content.
75
Temperature (C)
2500
Viscosity (cP)
65
2000
55
1500
1000 45
500 35
0 25
0
36
68
100
132
164
196
228
260
292
324
356
388
420
452
484
516
548
580
612
644
676
708
740
Time (s)
Figure 2 RVA profile of selected O. glaberrima and O. sativa varieties.
146 THE CEREAL GRAINS | African Rice (Oryza glaberrima): History and Its Growing Importance in Rice Breeding Efforts
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Provide reasons to support the assertion that O. glaberrima Steud. Plant Production Science 15(3): 151163.
Gayin J, Chandi GK, Manful J, and Seetharaman K (2015) Classification of rice based
could have the same yield potential as Asian rice. on statistical analysis of pasting properties and apparent amylose content: The case
Why is O. glaberrima an important food security crop in of Oryza glaberrima accessions from Africa. Cereal Chemistry 92(1): 2228.
Africa? Jones MP, Mande S, and Aluko K (1997) Diversity and potential of Oryza glaberrima
Briefly describe the variations in the physicochemical prop- Steud in upland breeding. Breeding Science 47: 395398.
Li ZM, Zheng XM, and Ge S (2011) Genetic diversity and domestication history of
erties of O. glaberrima. African rice (Oryza glaberrima) as inferred from multiple gene sequences.
Theoretical and Applied Genetics 123(1): 2131.
Linares OF (2002) African rice (Oryza glaberrima): History and future potential.
Exercises for Readers to Explore the Topic Further Proceedings of the National Academy of Sciences of the United States of America
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Does the nutritional profile of O. glaberrima differ from O. Biotechnology 9(44): 74097417.
Orjuela J, Sabot F, Cheron S, et al. (2014) An extensive analysis of the African rice
sativa and can we derive functional foods from African rice? genetic diversity through a global genotyping. Theoretical and Applied Genetics
127: 22112223. http://dx.doi.org/10.1007/s00122-014-2374-z.
Sahrawat KL and Sika M (2002) Comparative tolerance of Oryza sativa and O.
See also: Barley, Rice and Maize Processing: Rice Processing: glaberrima rice cultivars for iron toxicity in West Africa. International Rice Research
Beyond the Farm Gate; Breeding of Grains: Rice: Breeding; Notes 27(2): 3031.
Sakagami JI, Joho Y, and Ito O (2009) Contrasting physiological responses by cultivars
Carbohydrates: Glycemic Index, Glycemic Load, and Their Health of Oryza sativa and O. glaberrima to prolonged submergence. Annals of Botany
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Overview; Genetics of Grains: Rice: Genetics; Processing of Semon M, Nielsen R, Jones MP, and McCouch SR (2005) The population structure of
Grains: Grain Quality Attributes for Cereals Other than Wheat; The African cultivated rice Oryza glaberrima (Steud.) evidence for elevated levels of
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glaberrima Steud.) for breeding in Sub-Saharan Africa. Genes, Genomes and
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Further Reading Africans discovered in Suriname. Economic Botany 64(1): 110.
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Agnoun YB, Samadori SH, Sie M, Vodouhe RS, and Ahanchede A (2012) The African glaberrima) and evidence for independent domestication. Nature Genetics
rice Oryza glaberrima Steud: Knowledge distribution and prospects. International 46: 982988. http://dx.doi.org/10.1038/ng.3044.
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economic potentials of Oryza glaberrima and wild species in the genus Oryza. Paper Steud.) and Asian rice (Oryza sativa L.). Plant Production Science 9(3): 287293.
presented at the Meeting on African Rice Species. Watanabe H, Futakuchi K, Teslim I, and Sobambo BA (2002) Milling characteristics and
Futakuchi K, Fofana M, and Sie M (2008) Varietal differences in lodging resistance of grain quality traits of interspecific progenies of Asian rice (Oryza sativa) with African
African rice (Oryza glaberrima Steud.). Asian Journal of Plant Sciences 7(6): rice (Oryza glaberrima). Japanese Journal of Tropical Agriculture 46: 4755.
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Barley: An Overview of a Versatile Cereal Grain with Many Food and Feed Uses
AM Stanca, University of Modena and Reggio Emilia, Reggio Emilia, Italy
A Gianinetti, F Rizza, and V Terzi, Consiglio per la ricerca in agricoltura e lanalisi delleconomia agraria, CRA-GPG, Fiorenzuola
dArda, Italy
2016 Elsevier Ltd. All rights reserved.
Topic Highlights The Mediterranean lands featured two barley types with
very precise traits. The first was the six-row type because it
Barley was domesticated from its wild relative Hordeum grew rapidly in the mild Mediterranean climate, characterized
vulgare subsp. spontaneum in the Fertile Crescent about by low and erratic rainfall. The second, called two-row barley,
10 000 years ago. features high-quality grain and was sown in spring for direct
Barley is a model organism for genetic and genomic studies human consumption. Gladiators, whose diet consisted almost
in small grain cereals because it is a true diploid entirely of food made from barley flour, were even called
(2n 2x 14). hordearii, meaning barley men. Genotypes that had a naked
Barley is the fourth most important cereal crop in the world. kernel were often resorted to as foodstuffs because of their
Today, barley is used as grain for livestock feed (8090%) superior nutritional value. It was during the Roman era too
and as malt for beer and other alcoholic beverages (10%), (Figure 2) that all the criteria for a rational use of grain and
and a small fraction is used as human food. straw were defined and refined to perfection, Columella pro-
Specific characteristics of barley grain composition allow viding a detailed description in his Res rustica. The yield per
barley flour to be used for the production of functional hectare in the fertile lands was about tenfold the number of
food due to the high content of beta-glucan (48%). seeds employed. Barley further expanded its horizons in con-
Barley, in comparison with the other cereals, shows high junction with Columbuss second voyage, when it was intro-
genetic variability for positive adaptation to environmental duced to the New World. Later, Dutch, English, French, and
stresses and growth in extreme climatic conditions. Danish colonists promoted its spread throughout North Amer-
ica, as did their Spanish and Portuguese counterparts in Cen-
tral and South America. On the Altiplano of the Andean
Learning Objective nations of Peru and Bolivia, barley is grown for grain at higher
elevations over 4550 m than oat, wheat, and maize.
To achieve an overview of the botany, physiology, genetics,
and agronomy of barley and of the main uses of this small
grain cereal. Morphology and Botany
Since the turn of the twentieth century, our insights into plant
Barley Domestication and Spread growth, reproduction, and trait inheritance have been of indis-
pensable support to barley field management and the improve-
Barley, one of the earliest domesticated plants, has been crucial ment of cultivars. Several morphological forms have evolved,
to the development of mankind, providing both a regular including winter, spring, two-row, six-row, awned, awnless,
staple and a source of compounds essential for life carbohy- hooded, naked and covered, malting, feed (grain and forage),
drates, proteins, fat, and secondary metabolites. Its history as a and food type. The high diversity of barley phenotypes, true
crop begins in the Fertile Crescent, a cradle area that today diploid nature (2n 2x 14), and ease of hybridization and
includes part of Iran, Turkey, Syria, and Jordan. The domesti- cultivation have made barley a model genetic organism since
cation of its wild ancestor, with a two-row brittle rachis allow- the rediscovery of Mendels laws. Barley belongs to Triticeae
ing shattering of the seeds, Hordeum spontaneum (Figure 1), and has seven pairs of distinct chromosomes, five of them
which is still to be found in the Middle East, dates to the being without satellites (1H, 2H, 3H, 4H, and 7H) and two
Neolithic (10 000 years BC). with satellites (5H and 7H). Cultivated barley is an inbreeder
It is from here that the species set out first to colonize the and it has more or less cleistogamous flowers, but under stress
Mediterranean Basin and then, following the great river and sea conditions, it can be more open flowering. The normal out-
migration routes of the time, fanned out to Central and Northern crossing rate has been determined to a maximum of 1.71.8%.
Europe, the Himalayas, and the Far East. Human intervention Despite a very high inbreeding rate, a high degree of diversity is
has been a decisive factor in the breeding of this species down maintained, which is a sufficient basis for rapid adaptation to
through the millennia, the cultivars currently available manifest- varying environmental conditions.
ing completely different traits from those found in either its
original parent or those populations that have evolved in the
wild. During the process of domestication, barley has gradually Barley as Model for Triticeae Genetics and Genomics
accumulated traits that facilitated agricultural production. The
perfecting of crop management techniques during the period of Barley is a model organism for genetic and genomic studies in small
Roman dominion contributed to the spread of barley in the grain cereals because of several reasons, ranging from its high
fertile as well as in the poorer soils of the empire. degree of natural variation and wide adaptability to the diploid
regulatory networks models and reverse genetics. Genome-wide feeding: improving grain protein quality, reducing grain phytic
reverse genetics have been developed in barley to understand the acid content, reducing beta-glucan content in feed for mono-
genefunction relationships, including both transgenic strategies, gastric animals, reducing rumen grain fermentation rate, and
like insertional mutagenesis or activation tagging, and non- increasing grain starch content. Barleys economic importance
transgenic ones, like TILLING (targeting induced local lesions in also extends to the whole plant, which can be used for both
genomes). Even metabolomics that is, the characterization of livestock feed and industrial purposes. The use of the whole
metabolites present in a tissue in a specific environment plant harvested at the end of the milk-ripe stage for silage is a
condition can be very informative. Integrative genomics deals practice that is becoming increasingly common in a number of
with combining the different data from these multiple-omics barley-producing areas. This approach makes it possible to
technologies to study biological systems multilaterally. In partic- develop the double-cropping system in irrigated fields by grow-
ular, the combined analysis of a relevant amount of genotyping ing in the same soil and same year barley silage followed by a
and phenotyping data can give information useful for genetic crop of summer maize, sorghum, or soybean. Silage quality
mapping and advanced breeding programs of barley. Barley can be improved only when varieties exhibit an overall grain/
genetic resources remain the basis for this flowchart. For biomass ratio (harvest index) greater than 50%.
this reason, barley collections representative of existing natural As food, the human health benefit of consuming barley
genetic diversity are maintained worldwide in situ and ex situ, grain in food formulations is well known. The kernel of barley
in the gene banks. New barley genetic resources are now devel- has a level of beta-glucans that is much higher than that found
oped for more efficient phenotypinggenotyping, like nested in other grains. Beta-glucans are a kind of soluble dietary fiber
association mapping (NAM) populations obtained by crossing having hypoglycemic and hypocholesterolemic effects, as it
diverse genotypes to a reference parent or multiparent advanced swells in the presence of water to form highly viscous solutions
generation inter-cross (MAGIC) based on the cross of several that can trap glucose and cholesterol and slow down their
parentals. absorption. The reduction of the rate at which glucose is
Talking about genetics, it is required to talk even of epige- released to the intestine and to the blood is particularly inter-
netics: epigenetic effects have been shown to be strongly esting when barley flour is included in the preparation of
involved in developmental and physiological processes in bread, a high glycemic food. Thus, incorporation of beta-
model plants. Although barley has been poorly investigated glucans derived from barley into this widely consumed food
from this point of view, some evidence exists on the impact of can reduce the glycemic response to its consumption. The use
epigenetic factors on seed development and response to of beta-glucan-enriched barley flour is recommended to pro-
hormones. duce bread dough made of wheat/barley flour blends with a
Finally, from a practical point of view, what we need is not not-too-high proportion of barley flour. The majority of the
new: efficient and inexpensive tools to select for agronomic Waxy barley genotypes (which have a very low to null amylose
and quality traits. Molecular markers developed in the last content) also have high beta-glucan content and can therefore
decades have been demonstrated to be of great help, in partic- be used to improve the hypocholesterolemic effects of foods.
ular for pyramiding resistance genes in a marker-assisted selec- Barley flour can be easily incorporated into many wheat-
tion (MAS) scheme of breeding. What is around the corner is based baked or not baked products such as cookies, bread,
genomic assisted selection (GAS) that means to build breeding flatbread, tortillas, puffed sacks, and pasta for producing the
schemes based on the selection of large portions of the so-called functional foods (Figure 5). In addition, tocols and
genome. This strategy has been applied successfully in barley phenolic compounds of barley are also considered bioactive
prebreeding/breeding for selecting barley genotypes improved components.
for disease resistance and other agronomic traits.
Barley Use
Malting Quality during mashing. During the malting process, high production
and/or activation of hydrolytic enzymes is required for cell-
Barley is, and has been since the dawn of civilization, the cereal
wall degradation and protein solubilization, but starch break-
par excellence in the production of malt for beer, whisky, and
down should be minimal, to avoid unwanted consumption of
other alcoholic beverages. Indeed, ever since the development
utilizable dry matter by the embryo. To this aim, malting
of the most essential techniques by the ancient Egyptians and
should both accelerate modification and retard embryo
Babylonians, the art of brewing has continued to be perfected
growth, which are partially conflicting purposes.
down through the ages and its secrets jealously guarded by the
It is well known that the higher the level of protein in the
master brewers, despite the fact that fermentation technology
barley grain, the lower the percentage of carbohydrate and,
had remained entirely shrouded in the dark until the nine-
then, the extract yield of the malt. In addition to this negative
teenth century.
relationship with starch level, storage proteins have been sug-
The transformation of barley into malt and of the latter into
gested to play a significant part in determining the rate at
beer is a biotechnological process that is markedly dependent
which the whole endosperm is broken down during malting.
on the initial raw material, that is, on the barley grain that gives
More specifically, malting barley must have soft and mealy
rise to the malt. To this aim, barley has first to be converted
rather than hard and flinty endosperm and include such other
into malt, to increase its content of enzymes. Among them,
characteristics as high germinative energy, a certain degree of
amylases are needed to convert polymeric starch into soluble
dormancy to prevent pregermination in case of rain at harvest,
sugars, which can be used by yeasts for alcoholic fermentation.
adequate enzyme activity (amylase, beta-glucanase, and prote-
Malting is a controlled process of germination. As the first
ase), low beta-glucan content, low protein content, high starch
step of malting, barley is steeped to imbibe the grains and start
content, reduced glumeal hulls, and high extract percentage.
germination. After 13 days of germination, depending on the
The percentage of extract is directly linked to starch content
malting conditions, the aleurone layer responds to gibberellin
and seed size and inversely correlated to protein content. A
hormone from the embryo and releases hydrolytic enzymes
high nitrogen content adversely affects malt production by
that disaggregate and solubilize the endosperm. They can break
reducing the amount of extract and lowering quality. This is
down starch and storage proteins into simple molecules that
because a longer period is required for the modification pro-
nourish the growing embryo. Since starch and storage proteins
cess, which results in a greater development of rootlets and
are enclosed in the walls of the dead endosperm cells, these cell
coleoptiles and consequent respiratory losses, and because of
walls have to be dismantled to allow access of the hydrolytic
an abundance of soluble proteins that enter the extract and
enzymes to the internal reserves. Breakdown of endosperm cell
form suspensions in the beer. It is, however, important that the
walls mainly involves degradation of beta-glucans, which are
extract contain a permanent fraction of soluble nitrogen
the major constituent of the walls (>70%). This initial step is
(0.5%) to feed the yeasts during fermentation.
performed by endo-(1!3),(1!4)-beta-D-glucanase (shortly,
Malting begins with the soaking of the kernel in water to
beta-glucanase), which is the chief enzyme responsible for
raise its moisture content to 4245%, followed by the period
beta-glucan degradation and is therefore the enzyme leading
(4 days) of modification. The changeover from barley to malt
endosperm modification. Modification is a term used to
commences when the embryo secretes gibberellic acid, which
mean all the chemical and physical changes that occur when
immediately stimulates the production of enzymes (glucanase
barley is converted into malt, particularly the weakening in the
and pentosanase) that alter the endosperms structure by dis-
consistence of the endosperm consequent to the degradation
solving the walls of its cells and freeing starch grains. Other
of the cell walls. Prior to milling, malt appearance resembles
enzymes as phosphatase, phytase, glucanases, pentosanases,
the original barley grain but is friable when crushed, revealing
and protease also become active in the early stages of modifi-
the physicalchemical changes that have occurred during malt-
cation, while the amylases are triggered later. All of this activity,
ing. Secretion of beta-glucanase and other hydrolytic enzymes
which serves as a vehicle to move substances from the endo-
starts from the embryos scutellum and proceeds in the aleu-
sperm to the embryo for the formation of new tissues and
rone layer from near the embryo to the distal end of the grain.
organs (rootlet and coleoptile), is accompanied by a marked
Much research has focused on the role of endosperm compo-
increase in respiration: the resulting losses can range from 5%
nents in determining these aspects, and beta-glucans are con-
to 9% unless restricted by a rapid and uniform germination.
sidered to play a negative role. On the other hand, beta-glucans
From what we have mentioned thus far, it is not difficult to
have also received much interest recently with regard to poten-
see that malting quality depends on a host of organic factors.
tial health benefits. Thus, whereas high beta-glucans are
The analytic data of a single character are insufficient to judge
required for food functionality, low beta-glucans are usually
either the grain quality initially or the malt quality thereafter.
desirable for the best malting quality.
Genetically speaking, breeding for malt varieties is a com-
The endosperm of unmalted grains is compact and inacces-
plex issue because many traits at grain, malt, and wort level
sible to hydrolytic enzymes because large proteins cannot per-
have to be taken into account.
meate cell walls. Thus, in the germinating barley grain,
thorough breakdown of endosperm cell walls and partial
digestion of the protein matrix inside the endosperm cells are Biotic and Abiotic Stress Resistance
necessary steps to render starch granules accessible to amylo-
lytic enzymes. These two processes are the chief causes of the Barley, despite its proven adaptability to a range of ecological
physicalchemical modification of the endosperm that occurs situations throughout its evolution, is nonetheless susceptible
during malting and make the starch susceptible to degradation to extreme environmental situations and a number of
THE CEREAL GRAINS | Barley: An Overview of a Versatile Cereal Grain with Many Food and Feed Uses 151
Table 1 Major diseases and pests of barley Are the barley mutants useful to increase secondary metab-
olites for food?
Disease common name Causal agent
Which strategy is still active for disease resistance in barley:
Fungal disease monogenic or quantitative resistance?
Barley stripe Pyrenophora graminea
Powdery mildew Blumeria graminis
Net blotch Pyrenophora teres See also: Agronomy of Grain Growing: Barley: Agronomy;
Fusarium Head Blight Fusaria
Barley, Rice and Maize Processing: Barley: Malting; Barley:
Snow rot Typhula incarnata
Spot blotch Cochliobolus sativum
Milling and Processing; Beverages from Grains: Beverages:
Leaf blotch or scald Rynchosporium secalis Distilled; Fermentation: Foods and Nonalcoholic Beverages;
Loose smut Ustilago nuda Fermentation: Origins and Applications; Bioactives and Toxins:
Ramularia leaf spot Ramularia collo-cygni Bioactives: Antioxidants; Mycotoxins; Breeding of Grains: Barley:
Yellow rust Puccinia striiformis Genetics and Breeding; Carbohydrates: Health Effects of b-Glucans
Brown rust Puccinia hordei Found in Cereals; Cereals: Chemistry and Physicochemistry of Non-
Black stem rust (including Puccinia graminis starchy Polysaccharides; Glycemic Index, Glycemic Load, and Their
Ug99) Health Benefits; Resistant Starch and Health; Food Grains and Well-
Septoria leaf blotch Septoria passerini being: Functional Foods: Overview; Genetics of Grains: Genomics
Rhizoctonia root rot Pythium
of Food Grains; Grain Harvest, Storage and Transport: Barley:
Root rot Rhizoctonia solani
Ergot Claviceps purpurea
Harvesting, Storage, and Transport; Grain Marketing and Grading:
Covered smut Ustilago hordei Barley: Grading and Marketing; Non-food Products from Grains:
Loose smut Ustilago nuda Cereal Grains as Animal Feed; Processing of Grains: b-Glucans:
Take-all Gaeumannomyces graminis Measurement and Processing; The Basics: The Grain Crops: An
Eye-spot Pseudocercosporella herpotrichoides Overview; The Cereal Grains: An Overview of the Family of Cereal
Leaf non parasitic spots Unknown Grains Prominent in World Agriculture; Cereals: Domestication of the
Bacterial disease Cereal Grains.
Black chaff Xanthomonas translucens
Bacterial blight Xanthomonas campestris
Viral disease
Barley yellow mosaic virus- Potyviridae (soil borne disease
BAYMV, BAMMV transmitted by Polymixa graminis) Further Reading
Barley yellow dwarf virus Luteoviridae (transmitted by aphids)
BYDV Bulgarelli D, Biselli C, Collins NC, et al. (2010) The CC-NB-LRR-type Rdg2a resistance
Pests and nematodes gene confers immunity to the seed-borne barley leaf stripe pathogen in the absence
Cereal cyst nematode Heterodera avenae of hypersensitive cell death. PLoS One 5(9): e12599.
Demetriou K, Kapazoglou A, Tondelli A, et al. (2009) Epigenetic chromatin modifiers in
Root lesion nematode Pratylenchus spp.
barley: I. Cloning, mapping and expression analysis of the plant specific HD2 family
Green bug Elymus canadensis of histone deacetylases from barley, during seed development and after hormonal
Aphids treatment. Physiologia Plantarum 136: 358368.
Hessian fly Mayetiola destructor Fincher GB (2011) Biochemistry, physiology, and genetics of endosperm mobilization
Cereal leaf beetle Oulema melanopus in germinated barley. In: Ullrich S (ed.) Barley: Production, Improvement and Uses,
Cereal ground beetle Zabrus tenebroides pp. 449477. Chichester, UK: Wiley-Blackwell.
Frit fly Oscinella frit Finocchiaro F, Ferrari B, Gianinetti A, et al. (2012) Effects of barley beta-glucan-
Cutworms Agrotis enriched flour fractions on the glycaemic index of bread. International Journal of
Food Sciences and Nutrition 63(1): 2329.
Francia E, Tondelli A, Rizza F, et al. (2011) Determinants of barley grain yield in a wide
range of Mediterranean environments. Field Crops Research 120: 169178.
Is the whole barley plant used for forage? Graner A, Kilian A, and Kleinhofs A (2011) Barley genome organization, mapping and
Explain the difference between winter, alternative, and synteny. In: Ullrich S (ed.) Barley: Production, Improvement and Uses, pp. 6384.
Chichester, UK: Wiley-Blackwell.
spring barley varieties. Mathre DE (1997) Compendium of Barley Diseases. St. Paul, MN: American
Are there in barley grain secondary metabolites for human Phytopathological Society.
health other than beta-glucans? Paulitz TC and Steffenson BJ (2011) Biotic stress in barley: Disease problems and
Why is the barley plant considered more economically and solutions. In: Ullrich S (ed.) Barley: Production, Improvement and Uses,
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What is the barley genome size? representative barley genotypes cultivated in Europe. Crop Science 51: 27592779.
Stanca AM, Tumino G, Pagani D, et al. (2013) The Italian barley genetics mutant
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Exercises for Readers to Explore the Topic Further Thomas WTB, Hayes P, and Dahleen L (2011) Application of molecular genetics and
transformation to barley improvement. In: Ullrich S (ed.) Barley: Production,
WUE (water use efficiency) and NUE (nitrogen use Improvement and Uses, pp. 122143. Chichester, UK: Wiley-Blackwell.
efficiency) are considered two strategic traits to develop Ullrich S (2011) Significance, adaptation, production and trade of barley. In: Ullrich S
plants to cope with environments. Is barley a more efficient (ed.) Barley: Production, Improvement and Uses, pp. 313. Chichester,
UK: Wiley-Blackwell.
species within cereals? von Bothmer R and Komatsuda T (2011) Barley origin and related species. In: Ullrich S
How do the agronomic practices improve yield and quality (ed.) Barley: Production, Improvement and Uses, pp. 1462. Chichester,
of the barley grain? UK: Wiley-Blackwell.
Sorghum Grain, Its Production and Uses: Overview
A Cruickshank, DAFF Queensland, Warwick, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
The sorghum grain is a high-energy cereal grain that is a Botanically sorghum belongs to the grass family and the tribe
staple to many poorer farmers in Africa and parts of Asia. Andropogoneae. This is a tropical and subtropical group of
Sorghum is a genetically diverse and robust plant, posses- grasses, which also includes sugarcane (Saccharum spp.) and
sing resistances to many biotic and abiotic stresses. maize (Zea mays). Like all the grasses in this tribe, sorghum has
Sorghum is a (sub)tropical grass (like sugarcane and maize) a C4 photosynthetic pathway. The genus Sorghum has at least
with a C4 photosynthetic pathway. 20 species extending from Africa through Asia and including
Sorghum bicolor is classified into five races: bicolor, kafir, 12 species endemic to Oceania. Sorghum bicolor, a native of
caudatum, durra, and guinea. Africa that has spread throughout the world, has three subspe-
Major sorghum producers are Nigeria, the United States, cies: S. bicolor bicolor, S. bicolor verticilliflorum, and S. bicolor
India, Argentina, Ethiopia, and Australia. drummondii, with distinct morphology for the respective
Sorghum can grow in temperatures from 15 to over 40 C heads (Figure 1).
Flowering and seed set require temperatures between 20
and 39 C. S. bicolor subsp. verticilliflorum is the wild progenitor spe-
cies. These sorghums have small seed, < 1 mm thick. These
Significant diseases are downy mildew, ergot, and grain
seeds are shed soon after they ripen. This process is called
mold.
shattering, and selection of nonshattering off-types is one
Significant insect pests include sorghum midge and
of the first steps in domesticating a wild plant to develop
greenbugs (aphids).
a crop.
Sorghum grain can be white, yellow, red, or purple.
S. bicolor subsp. bicolor incorporates all the cultivated types.
Sorghum is consumed as a thin porridge or gruel, as a thick
The cultivated types have been further classified on the
porridge or dough, as various flatbreads, as popped or
basis of the shape of heads and grain into five races: bicolor,
puffed sorghum grain, or as alcoholic beverages.
kafir, caudatum, durra, and guinea. The range of head types
is illustrated in Figure 2.
Learning Objective S. bicolor nothosubsp. drummondii incorporates all material
derived from natural crossing between S. bicolor subsp.
bicolor and S. bicolor subsp. verticilliflorum types. In some
To achieve an overall understanding of sorghum as a grain
production systems, weedy S. bicolor nothosubsp. drum-
worldwide: its origins, breeding, agronomy, production,
mondii types (known as shattercanes) are problem weeds.
processing, and utilization
Analysis of DNA sequence data has led scientists to believe
there have been at least two, possibly more, domestications
Introduction of sorghum. There is little doubt that the original domestica-
tion occurred in East Africa somewhere in Ethiopia or possi-
All grain sorghum is the product of cultivated types of Sorghum bly Sudan. Evidence from DNA sequence analysis suggests that
bicolor (L.) Moench. Sorghum bicolor is an African species and there has been at least one other more recent domestication in
the crop was first domesticated in East Africa. In traditional West Africa, giving rise to the group that has been distinguished
agriculture, tall varieties of sorghum are grown for multiple by the name margaritiferum. These plants have a lot of wild
purposes including various uses of grain, fodder, and extract of plant features and are grown separately from other sorghums,
sweet syrup (by crushing stems in the same manner as with named as a different crop and prepared in different foods.
sugar cane) and use of the panicles for broom heads. Today,
sorghum is a staple for approximately half a billion people.
In developed agriculture, particularly in the Americas and Sorghum Production
Australia, sorghum has been selected and bred for specific
end uses grain sorghums as short plants suited to mechani- Sorghum is produced in the tropics and in subtropical or
zed agriculture and taller types for forage, fodder, and silage temperate regions throughout Africa, Asia, and the Americas
production. (only a small amount of grain sorghum is produced in
Grain sorghum is a crop with great genetic diversity and Europe). Prior to the colonial era, maize was the dominant
adaptation to drier and hotter environments. It has tradition- grain crop in the Americas and sorghum the most important
ally been used in many different ways in Africa and Asia. It is summer cereal in Africa. Since both crops have been available
now an important world crop and the most important crop worldwide, there has been a general trend for maize to be more
locally in some hot and dry regions. popular in higher rainfall areas and sorghum to be cultivated
Western Central
South-Eastern Asia Oceania
Asia Asia
Middle Africa Eastern Africa
Table 1 Production of sorghum by major countries, averaged for the the soil, but fertilization with phosphorus, potassium, and
decade 200413 other minor nutrients will often still be necessary to remove
limitations to yield.
Country Tonnes % The agronomy of sorghum production in mechanized agri-
culture is quite different to that in traditional production sys-
World 59 850 648 100
The United States 9 327 805 15.6 tems. In developed agriculture, sorghum is grown as a row crop
Nigeria 7 891 520 13.2 at plant densities suited to the available moisture. Nitrogen
India 6 883 880 11.5 fertilizer may be applied entirely preplanting or split between a
Mexico 6 361 639 10.6 starter application and a later targeted top-up when the likely
Sudan (former) 3 873 500 6.5 yield is more predictable. Herbicide application may be used
Argentina 3 056 573 5.1 to desiccate the sorghum plant and grain in preparation for
Ethiopia 2 943 216 4.9 harvest.
Australia 2 171 743 3.6
China, mainland 2 141 650 3.6
Brazil 1 813 631 3.0
Burkina Faso 1 673 123 2.8
Niger 1 018 760 1.7
Sorghum Breeding
Mali 993 038 1.7
Cameroon 961 818 1.6 Traditional farmers have been responsible for the selection and
maintenance of the great diversity that exists in the landraces of
Source: FAOSTAT. sorghum. When some of these landraces were introduced to
the Americas, any off-types that had dwarfing genes or different
maturities were selected and (just as important) were studied.
latitudes or even similar latitudes with different temperature An understanding of the differences in height and maturity
regimes. developed that allowed selection of types that were more suited
Sorghum is adapted to a wide range of soil conditions. to mechanized agriculture. Subsequently in the United States,
Some of the genetic characteristics that confer this adaptation there was a great plant breeding initiative called the sorghum
are being discovered: a general tolerance of salinity, greater conversion program in which landraces were converted by
tolerance to acidic aluminum ions, more efficient uptake of crossing them to short, photoperiod-insensitive types and the
essential phosphorus, and different angles of nodal root progeny back-crossed to the landrace while selecting for short
growth. As with other cereal grains, in most circumstances, photoperiod-insensitive plant type. This provided a resource to
nitrogen will be the limiting nutrient in grain production. sorghum breeders from which many improved traits could be
Sorghum will form mycorrhizal associations with fungi that bred into other genetic backgrounds including high yield,
improve the nutrient efficiencies of elements that are present in drought resistance traits, and midge resistance.
156 THE CEREAL GRAINS | Sorghum Grain, Its Production and Uses: Overview
Sorghum now has a published genome sequence that more than discoloration of the grain, but other species in some
allows breeders to develop understanding of the genetics of circumstances produce toxins that make the grain unsuitable
traits of which their predecessors could only dream. Given the for consumption. Many traditional varieties, particularly in
great diversity available within the species, this understanding West Africa, flower in response to changes in day length so
should allow development of grain sorghum with robust adap- that most if not all of their grain-filling period occurs after the
tation to extreme environments and grain qualities that pro- end of the rainy season.
vide a range of products fit for many purposes. The sorghum midge (Stenodiplosis sorghicola) is an insect
that is totally dependent on sorghum plants for its survival
and reproduction. The female midge seeks out flowering sor-
Sorghum Diseases and Pests ghum to lay its eggs at the base of florets. The larvae of the
midge then feed on the ovary and prevent the development of
Like all crops, sorghum productivity is affected by a range of grain. In susceptible sorghum types, if the environment is
pests and diseases. Some pests or diseases have a specific inter- warm and humid, losses can be substantial. The level of dam-
action with sorghum; others are more widely adapted and are a age is also dependent on the number of flying female midge
threat to other crops also. Some directly affect yield of usable present when a crop is flowering. Most commonly, these
grain through killing plants or reducing the amount of grain females come from wild or volunteer sorghum (particularly
produced per plant. Diseases also indirectly influence produc- Sorghum halepense) or an earlier crop of cultivated sorghum.
tivity through damaging and devaluing grain or by causing Pupae can overwinter in plant trash in a diapause state. There
farmers to sacrifice potential yield with management decisions are sorghum lines that resist the oviposition or the larval
that are part of a strategy to avoid disease. feeding. With resistant cultivars and cultural practices, such as
To show the diversity of pest and disease biology, three early sowing, good control of midge damage can be achieved
diseases, downy mildew, ergot, and grain mold, and two impor- without the use of insecticide. In Australia, the widespread use
tant insect pests, sorghum midge and greenbugs (aphids), will of midge-resistant sorghum has allowed the survival of many
be considered. more beneficial arthropods (predatory insects, spiders, etc.),
Downy mildew is caused by the pseudofungus Peronoscler- and this has aided integrated pest management in other sum-
ospora sorghi. This pathogen can infect other species including mer crops also.
maize. It can survive for years in the soil as a resistant oospore A range of aphid species will feed on sorghum but the most
and can do most damage when oospores infect a young crop in common pest is the greenbug (Schizaphis spp.). They are grass
moist soils. This is called systemic infection and the pathogen feeders and are also pests of other cereals in farming systems. In
can then grow throughout the plant. Symptoms include ini- sorghum, they cause damage by degrading tissue with their
tially leaf yellowing but eventually leaves are completely dead saliva, by depletion of nutrients and energy, by contaminating
and visibly shredded. When warm humid conditions favor harvested grain with honeydew, and by being a vector for plant
growth of the pathogen, it can produce short-lived spores pathogenic viruses notably maize dwarf mosaic.
called conidia that can infect the aerial plant parts. Conidial Greenbugs are capable of both sexual reproduction and
infection may cause minimal damage in mature plant or lead parthenogenesis. They also have a short generation time and
to systemic infection in a very young plant. Control is achieved a capacity for rapid population increase. This has led to rapid
by host plant resistance, seed treatment with fungicides, and emergence of new biotypes with different preferences in feed-
cultural measures such as crop rotation (to reduce oospore ing or resistances to pesticide, when control measures provided
infection) and early sowing (to reduce conidial infection selection pressure on the greenbug population. Resistant vari-
from plants infected earlier). eties and natural enemies, together with careful use of
Sorghum ergot is caused by the fungus Claviceps africana (in pesticide, can provide adequate control.
the Indian subcontinent by C. sorghi). It is host-specific to
plants of the genus Sorghum and has one very specialized
mode of infection: conidia of sorghum ergot can only infect Sorghum Grain Composition and Utilization
nonfertilized stigmas of flowering sorghum heads. Genetic and
environmental factors can contribute to nonfertilized stigmas The sorghum grain is a typical grass seed but it has been
being exposed to infection. Male-sterile lines used in the pro- selected over time for larger seed size and other particular
duction of hybrid seed are particularly vulnerable; and seed characteristics. Each grain is a caryopsis the botanical name
production is done in lower risk environments and protected for a fruit containing a single seed with the fruit wall (pericarp)
with fungicides if needed. The environmental factors that lead and seed coat (testa) fused. The color of the grain is mostly
to ergot disease are factors that prevent successful pollination: determined by the pericarp that can be white, yellow, red, or
rainfall and high humidity at flowering and cool temperatures purple (almost black), but the presence of condensed tannins
prior to and at flowering. The sensitivity of the flowering in the testa causes a strong brown subcoat. Tannin-rich sor-
period influences farmer decisions on when to sow sorghum, ghums offer the advantage of being bird-proof to a degree, but
often choosing not to sow sorghum after a certain date to avoid they also offer greater difficulty in most forms of processing.
a high risk of ergot infection. Variations in head shape and grain color are illustrated in
Sorghum grains are exposed to the weather, and moisture Figure 4.
on the grain can lead to infection by a number of species of The embryo of a cultivated sorghum grain is larger than the
fungi particularly Fusarium and Curvularia. Collectively, these embryo of wild sorghum, but the increase is not proportional
infections are referred to as grain mold. Some cause nothing to the increase in grain size with domestication. Most of the
THE CEREAL GRAINS | Sorghum Grain, Its Production and Uses: Overview 157
Figure 4 A range of sorghum grain colors, including some white grains Figure 5 Injera, a fermented flatbread popular in Ethiopia, made from
stained by grain mold. teff or sorghum. Source: Illustration from Wikipedia.
volume of the grain is made up of endosperm, but different produced by the absence or the presence (and nature) of
sorghum seed lines have very different endosperm structures fermentation. When sorghum is prepared as a heavy porridge,
and textures, which determine their value for different uses of it is usually eaten by hand with a savory sauce.
the grain. In Ethiopia, sorghum is made into injera (Figure 5), a
When the grain is mature, its endosperm is composed of fermented flatbread that looks to western eyes like a pancake.
starch granules, spherical bodies composed of kafirin proteins, The preparation of injera includes a lactic acid fermentation
and a protein matrix of glutelin. These three components can leading to a marked sour taste. The most valued grain for injera
be loosely packed in what is called floury endosperm or tightly is the Ethiopian grain teff, but sorghum is produced in much
packed in what is called corneous endosperm. Kafirins can greater quantities than teff and is preferred over wheat or barley
form disulfide bonds, with one another and with glutelin, for making injera. Consequently, Ethiopia is one of the few
which are very resistant to digestion. The resistance of these places in the world where sorghum attracts a higher price than
bonds to hydrolysis is believed to reduce the digestibility of wheat. Injera is usually made with little or no bran. In Sudan, a
both protein and starch, particularly in tightly packed corneous similar bread called kisra is made with whole grain meal.
endosperm. In India, sorghum is used in roti-style flatbreads or as a
There are other factors present in whole grain that can cause boiled grain substituting for rice. Rotis are often made with a
resistance to digestion condensed tannins in the testa and mixture with wheat flour. As in many other uses, the elastic
phytate in both the embryo and the pericarp. The tannins, nature of the gluten in wheat flour helps maintain the structure
when present, can interfere with protein digestion. Phytate of a roti for longer than if made with sorghum flour only.
can interfere with the digestibility of protein, phosphorus, In East Asia, particularly China, sorghum grain is used
and iron. (either on its own or in combination with other grains) in
The end uses of sorghum can be divided for convenience the production of highly alcoholic distilled spirits. These spirits
into traditional uses associated with cultivation of landraces in are produced for both high-value exclusive markets and more
traditional African and Asian societies and uses in the devel- traditional local consumption. Because the sorghum produc-
oped economies of the Americas, Australia, and Europe. tion in China has declined and there is a demand for sorghum-
Throughout Africa, sorghum grain is prepared as a wide derived spirits in the growing middle class, China may be
range of foods. Traditionally, the milling of sorghum grain developing as a market for sorghum-exporting countries like
has been a laborious manual task with tools that are variations the United States and Australia.
on the theme of a mortar and pestle. The milled grain can then Sorghum grain produced in the Americas, Australia, and
be used in a variety of ways; broadly, most foods can be classed Europe is used primarily as animal feed. It is most commonly
as a thin porridge or gruel, as a thick porridge or dough, as used in intensive production of poultry, pigs, and feedlot
various flatbreads, as popped or puffed sorghum grain, or as cattle. (The use of sorghum grain as supplementary feed for
alcoholic beverages. In West Africa, the margaritifera types are dairy cattle is comparatively rare.) Smaller volumes are used in
cultivated separately and often used in a manner similar to rice. pet food. The grain is prepared in various ways for use in
The variety in the foods produced is influenced by the choice of different animal production systems. The simplest preparation
sorghum, that is, use of the whole grain meal or some fraction is to break the grain in a hammer mill or similar device and
or use of a white-, red-, or brown-seeded sorghum containing offer as meal to animals. Due to the potential antinutritive
condensed tannins. Variation within these foods is also factors present in sorghum, this is not the most efficient use
158 THE CEREAL GRAINS | Sorghum Grain, Its Production and Uses: Overview
of the grain. More commonly sorghum is offered to poultry What climate conditions encourage or deter the diseases
and pigs in prepared complete-ration pellets and to cattle as and pests of sorghum?
steam-flaked grain. What agronomic management can be used to reduce the
When sorghum is used in pelleted rations, it is frequently as risks of damage by diseases and pests?
a cheaper substitute or complement for wheat. Due to different Describe the range of foods for which sorghum is used.
gelatinization temperatures, a greater degree of heat energy How can sorghums resistance to digestion be mitigated for
may be required to successfully bind a pellet-containing sor- humans and animals?
ghum. Many feed producers will add artificially produced
digestive enzymes to improve the digestibility of both the
protein and starch of the sorghum. Enzymes used in this way Exercises for Readers to Explore the Topic Further
include phytase, xylanase, and proteases thought to help break
down kafirins more efficiently. The protein complex of the sorghum grain is made up of
many proteins. What are their names and possible
functions?
Future Prospects for Sorghum Sorghum varieties differ in grain color. Find out the chem-
ical reasons and the possible advantages and disadvantages
Over decades, there has been the trend in many sorghum- of these differences.
producing countries for eating habits to move away from Sorghum, like sugarcane and maize, has a C4 photosyn-
sorghum-based foods as living standards have improved. On thetic pathway. It is thus distinct from C3 cereals such as
the other hand, these changes may be progressively comple- wheat and barley. What is the significance of this distinc-
mented by increased use of sorghum for livestock feed and tion for sorghum production now and into the future?
alcoholic beverages (mainly sorghum beers in the same
regions) and also the likely use of sorghum as a basis for
fuel ethanol production. In addition, the specific properties See also: Agronomy of Grain Growing: Sorghum: Production and
of sorghum starch may increase its use as an industrial source Improvement Practices; Grain Harvest, Storage and Transport:
of starch. Sorghum: Harvest, Storage, and Transport; Non-wheat Foods:
The environment favorable for sorghum production can be Sorghum: Utilization; Proteins: The Protein Chemistry of Cereal
expected to become greater with trends of climate change Grains; The Basics: Taxonomic Classification of Grain Species; The
toward warmer growth temperatures and elevated levels of Cereal Grains: An Overview of the Family of Cereal Grains Prominent
carbon dioxide in the atmosphere. Like plant breeders in gen- in World Agriculture; Cereals: Domestication of the Cereal Grains.
eral, sorghum breeders are planning to change breeding strat-
egies. However, the expected increases in grain yield for cereals,
such as wheat (a C3 plant), are not expected to be so great for Further Reading
C4 plants such as sorghum. Regarding climate change, the
Anonymous (2012) The U.S. Sorghum Industry. Washington, DC: US Department of
greatest concern of sorghum breeders is the probable more
Agriculture.
frequent occurrence of extreme heat events that have the capac- Dendy DAD (ed.) (1994) Sorghum and Millets: Chemistry and Technology St Paul,
ity to reduce seed set dramatically. MN: AACC International.
Doggett H (1988) Sorghum, 2nd edn. London: Longman Scientific & Technical.
Pereira TD (2012) Sorghum: Cultivation, Varieties & Uses. New York: Nova Science
Publishers Inc.
Exercises for Revision Ramirez JA and Manuel Vazquez M (2012) Sorghum: Food & Energy Source. New York:
Nova Science Publishers Inc.
List the major regions of sorghum production worldwide. Smith CW and Frederiksen RA (2000) Sorghum: Origin, History, Technology, and
Production. New York: Wiley.
How do they differ in their utilization?
What climatic and soil conditions suit these places to sor-
ghum production? Discuss rainfall, temperatures, day
Relevant Websites
length, and latitude with respect to stages of plant growth.
How have plant breeding and selection changed the origi- faostat.fao.org FAOSTAT.
nal growth conditions for sorghum? www.icrisat.org ICRISAT.
Rye Grain: Its Genetics, Production, and Utilization
HD Sapirstein and W Bushuk, University of Manitoba, Winnipeg, MB, Canada
2016 Elsevier Ltd. All rights reserved.
This article is a revision of the previous edition article by W. Bushuk, volume 3, pp. 8591, 2004, Elsevier Ltd.
The origins and history of rye production and uses are The primary center of origin of the rye plant is considered to be
discussed. in the Anatolian Plateau of the Middle East (Asia Minor),
Rye production worldwide is falling. encompassing the eastern region of Turkey and neighboring
International trade in rye is about 13% of world portions of what is now Georgia, Armenia, Azerbaijan, Iran,
production. Iraq, and Syria. This northern region of the Fertile Crescent is
Starch (stored in discreet granules) is the major component also the center of origin of wheat, barley, oats, and lentils.
of rye endosperm. There is no evidence of rye cultivation in ancient Egypt. Cereal
Secalin is the major protein of the rye grain, but rye is not as rye is believed to have diverged genetically from related Secale
suitable for bread making as wheat. species about 10 000 years ago. Rye grain is mentioned in early
The rye grain contains more fiber than wheat and other writings of northern Europe suggesting this region as an impor-
cereal grains. tant area of historic cultivation. Cultivation of rye migrated
Rye contains some compounds with antinutritional prop- from its center of origin in the Middle East to northern Europe
erties for monogastric animals. in the first millennium BC. One possible route of migration
Rye endosperm has a soft texture, releasing flour readily in was from Asia Minor to western Russia and, from there, west-
milling. ward to Poland and Germany where it is highly adaptable and
Breads may be made from 100% rye flour but up to 90% popular with farmers. Another possible route of rye plant
wheat flour may be added. migration was from Turkey by way of the Balkan Peninsula to
Baked goods include rye bread, rye rolls, crisp bread, and north-central Europe. Rye was brought to North America and
pumpernickel. South America by European settlers during the sixteenth and
seventeenth centuries. At about the same time, its cultivation
spread eastward in Europe to Siberia. During the nineteenth
Learning Objectives and twentieth centuries, its cultivation began in Argentina,
southern Brazil, Uruguay, Australia, and South Africa. Today,
To recognize the place of rye within the range of cereal rye cultivation like that of wheat is worldwide, but the largest
grains and to understand ryes genetics, morphology, production is in central and eastern Europe (Table 1).
composition, and utilization.
Botanical Classification
Introduction
Rye belongs to the grass family Poaceae (also called Grami-
Cereal rye (Secale cereale L.) is an annual grass plant that is neae) and the genus Secale. The most common cultivated
typically fall-sown and cultivated worldwide. Historically, rye species is S. cereale, which is presumed to have evolved from
was a major food crop in eastern European countries in the the wild perennial grass of the species S. montanum. Rye, along
Middle Ages due to its ample winter hardiness and lower soil with evolutionarily related cereal grains barley and wheat, is
moisture requirements compared to wheat. Rye grain is used botanically classified in the same tribe of Poaceae, that is, the
mainly for feed and food and also for ethanol production. Triticeae. Rye and wheat are placed in a separate botanical
Cereal rye is also used as a winter cover crop because it grows subtribe, called Triticinae, on the basis of the morphology of
quickly in the fall to produce ground cover that protects soil their inflorescences or spikes in which the grain forms after
from erosion. Cultivated area and production worldwide have flowering. DNA studies have indicated that barley originated
declined considerably in the past half century and currently from a common ancestor before the divergence of wheat and
stand at about 6 Mha and 15 Mt, respectively (200812 aver- rye. Cereal rye should not be confused with ryegrass that
age). Rye is second after wheat for production of bread, and the belongs to a different genus (Lolium) of the Poaceae family
grain contains the highest levels of fiber among all cereal crops. and is typically used for forage and production of turf.
Rye is also used as a grazing or forage crop and for silage. The
grain is also important in the production of mixed feeds for
livestock and as feedstock for the distillation of rye whisky. Worldwide Production and Trade
Unprocessed straw is used in livestock feeds as low-digestibility
roughage and as bedding in animal husbandry. Rye straw can Worldwide production of rye grain has declined significantly
also be used as a building material, for example, roof thatch over the past decade (Table 2), a trend that has continued for
and particle board, and for other applications. the past half century reflecting a declining area of harvested
Country 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012
Germany 2277 3830 2794 2644 2698 3744 4329 2903 2521 3893
Poland 3172 4281 3404 2622 3126 3449 3713 2852 2601 2875
Russian Federation 4152 2872 3628 2959 3909 4505 4333 1636 2971 2132
Belarus 1151 1397 1155 1072 1305 1492 1227 735 801 1082
Ukraine 624 1593 1054 584 563 1051 954 465 579 677
China 709 600 554 350 700 645 630 570 680 659
Denmark 169 146 132 130 135 152 238 255 294 384
Turkey 240 270 270 271 241 247 343 366 366 370
Canada 327 398 330 383 252 316 281 232 195 340
World total 14 610 17 695 15 165 12 650 15 174 18 127 18 294 11 963 13 029 14 562
FAOSTAT data, 2014. Food and Agricultural Organization of the United Nations. Accessed via the web site www.fao.org.
Grain 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012
Maize 615.5 604.9 645.2 729.0 713.7 706.8 789.9 830.3 820.0 851.2 888.0 872.1
Rice 597.5 569.1 585.1 605.8 632.2 639.1 654.8 686.2 687.0 703.2 725.0 719.7
Wheat 589.8 574.7 560.1 632.1 626.7 602.3 612.9 683.0 687.5 649.5 699.5 670.9
Soybeans 178.2 181.7 190.7 205.5 214.6 222.0 219.7 231.3 223.4 265.2 262.4 241.8
Barley 144.0 136.7 142.6 153.9 138.7 139.5 134.2 154.8 151.8 123.7 133.0 132.9
Canola 35.9 34.4 36.8 46.5 50.0 48.0 51.4 57.9 62.6 60.1 62.7 65.1
Sorghum 59.8 53.5 59.0 58.1 59.6 57.7 62.7 66.5 56.8 60.0 58.4 57.0
Millet 29.0 23.9 34.8 29.6 31.0 31.9 33.7 34.4 26.1 32.6 27.2 29.9
Rye 23.3 20.9 14.6 17.7 15.2 12.7 15.2 18.1 18.3 12.0 13.0 14.6
FAOSTAT data, 2014. Food and Agricultural Organization of the United Nations. Accessed via the web site www.fao.org.
grain over this period. Rye production is significant in abso- Plant and Grain Morphology
lute terms but ranks last of eight grain species, including
canola, grown for human food and/or animal feed (Table 2), The mature rye plant has a slender, tough, fibrous stem (straw),
the position it has occupied over the past five decades. The and elongated leaves. Plant heights vary widely from about
decline in harvested area and production of rye is mainly due 30 cm to over 2 m. The inflorescence (spike) is long and slen-
to a decrease in demand for both food and feed and, der with stiff long awns (beards). The appearance of rye spikes
undoubtedly, economic returns to farmers compared with at different stages of maturity is shown in Figure 1. The pro-
those for other grains. Rye is grown in over 30 countries truding anthers during flowering as shown in the figure tend to
worldwide, and 9 of these (Germany, Poland, Russian Feder- be a feature of cereal grain plants, like rye, which typically
ation, Belarus, Ukraine, China, Denmark, Turkey, and propagate by outcrossing.
Canada) are the main producers that together represent over Rye grain (caryopsis) is arranged in pairs alternately on a
85% of world rye grain production (Table 1). The four lead- zigzag-shaped rachis (Figure 2). The grain is covered with a
ing countries, Germany, Poland, Russian Federation and Bela- lemma, a palea, and a glume (chaff), which is normally awned.
rus, combined to produce close to 70% of world rye At maturity, individual grains can be easily seen by bending the
production in 2012 (Table 1). inflorescence (Figure 3). Like wheat, rye grain is free-threshing;
Most of the rye grain produced is used domestically. This is the mature grain separates easily from the glume during thresh-
reflected in the low proportion of worldwide production that is ing. Rye grains or kernels tend to be more slender and longer
traded internationally; that figure is approximately 13%. The than wheat (Figure 4) ranging in length from 4.5 to 10 mm and
majority of rye commerce (85%), considering both exports in width from 1.5 to 3.5 mm. The grains are normally of grayish-
and imports, is handled by 12 countries (Table 3). Based on yellow color but the color can vary widely (Figure 4) depending
most recent international statistics, between 2009 and 2011, on rye cultivar, region of cultivation, and harvesting conditions.
Poland, Germany, and Canada were the leading rye exporters As in wheat, a crease extends the full length of the ventral side of
that, when combined, supply 66% of the world market on the grain. The surface of the grain is usually shriveled and has a
average. In the same years, the major importers of rye were rough texture. A single grain weighs about 20 mg.
Germany, Netherlands, and the United States, which together Rye grains comprise three distinct morphological parts
accounted for 52% of total imports. (Figure 5). They are the starchy endosperm, bran (pericarp
THE CEREAL GRAINS | Rye Grain: Its Genetics, Production, and Utilization 161
Table 3 Rye grain exports and imports by leading countries (in thousands of tonnes)
Exports 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011
Germany 1001 1003 954 1344 1196 1392 584 367 465 355 225
Canada 85 58 50 213 108 158 246 97 100 160 200
Denmark 136 140 44 4 4 2 3 7 16 48 105
Poland 0 2 10 74 347 64 59 79 592 403 103
Russian Federation 0 49 400 1 0 3 129 23 8 9 85
Sweden 83 44 16 2 7 5 26 13 16 54 42
Imports 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011
Germany 14 17 79 20 145 101 167 198 354 322 268
The United States 132 129 94 162 124 142 139 144 120 120 171
The Netherlands 107 173 147 246 249 121 104 153 380 211 109
Japan 339 355 406 288 282 272 215 59 65 121 88
Poland 332 13 84 16 3 18 167 54 6 5 45
Lithuania 3 45 47 13 23 18 18 3 5 24 41
Finland 37 57 21 53 80 89 50 69 60 44 38
FAOSTAT data, 2014. Food and Agricultural Organization of the United Nations. Accessed via the web site www.fao.org.
Crease
Brush
Vascular
bundle
Central
endosperm
Prismatic
endosperm
Peripheral
endosperm
Aleurone cells
0.5 mm
Scutellum
Coleoptile
Cuticle
Plumule
Epidermis
Hypodermis
Thin walled
Rcot
cells
Sheath and cap Cross cells
Cell nucleus Testa
Nucellar
Starch epidermis
0.5 mm granules Aleurone cells
Storage Tube cells
protein Peripheral
endosperm
Prismatic
Cell wall endosperm
100 m Pericarp
(a) (b)
Figure 5 Diagrammatic view of rye grain in (a) longitudinal section and (b) transverse midsection. Reproduced with permission from Simmonds, D.H.,
Campbell, W.P., 1976. Morphology and chemistry of the rye grain. In: Bushuk, W. (Ed.), Rye: Production, Chemistry, and Technology, first ed.
American Association of Cereal Chemists, St. Paul, MN, pp. 63110.
sclerotia in commercial handling and grading of rye, removal a diploid and has seven pairs of chromosomes in its genome.
of ergot bodies along with other foreign material by modern Wheat is more complex as it has a polyploidy makeup com-
cleaning equipment prior to milling of rye grain, and the prising either two or three basic sets of seven pairs of
substantial inactivation of alkaloids that occurs during baking. chromosomes, which provide a much larger genetic framework
for plant breeding aimed at improving adaptation and agro-
nomic performance, protein content, and other grain compo-
Genetics, Plant Breeding, and Agronomy sition components and end-use properties. With this goal in
mind, there have been attempts to develop tetraploid rye by
Essentially, all of the commercial cereal rye grown is the dip- doubling the number of chromosomes with the chemical
loid S. cereale type. Cultivated rye contains seven pairs of colchicine, which inhibits cell division. While tetraploidy has
chromosomes belonging to a single genome designated by been achieved in rye, results have been mixed in terms of
the letter R. Accordingly, diploid rye has the genome makeup practical outcomes, and little progress has been made beyond
RR. This basic genomic composition of rye is similar to that the experimental stage.
that exists in the genetically related cereal grains, wheat, and Rye, unlike most other cereal grains, is considered to be an
barley, which are all botanically classified in the same tribe of allogamous breeding plant; that is, it cross pollinates naturally
the Poaceae family, that is, the Triticeae. Like rye, barley is also and extensively with other rye plants. Therefore, it is difficult to
164 THE CEREAL GRAINS | Rye Grain: Its Genetics, Production, and Utilization
Table 4 Chemical and nutrient composition of rye compared with other cereal grainsa,b
Rye Wheat Pearled or hulless barley Oat groats or oatmeal Brown rice Corn
maintain genetic purity in conventional rye populations. of straw and lodging resistance, and disease resistance (snow
Owing to the close botanical and, in essence, genetic relation- mold, mildew, leaf rust, Fusarium head blight, and ergot).
ship between wheat and rye, their cross fertilization can be Improving end-use quality of rye grain for baking is another
achieved relatively easily. This is the basis for the man-made goal of breeding efforts. The main quality characteristic that
development of the ryewheat hybrid grain called triticale. Like continues to be a detriment in rye processing is its tendency to
rye, triticale sees modest production levels worldwide but is a sprout in the head under wet harvest conditions. In contrast to
significant grain for food and feed uses. wheat, rye grain developing to maturity on the plant has rela-
Most of cultivated rye is a fall-sown annual crop called tively low dormancy, which leads to preharvest sprouting. The
winter rye although spring rye is grown in some countries, latter results in substantially increasing levels of the starch-
such as Canada. Rye has excellent winter hardiness and there- hydrolyzing enzyme, a-amylase, which transfers into flour
fore can be grown in areas where the climate is too severe for causing very sticky doughs that are difficult to handle in com-
winter wheat. Another distinct feature of rye that contributes to mercial bread making.
its robust agronomic performance is its drought tolerance, The most significant aspect of rye breeding to date has been
which is considered to be the best among all cereal grains the development of commercial hybrid rye varieties with con-
and is explained by its ability to produce an extensive root siderably enhanced grain yields, up to 1520% higher on
system. Rye has been used extensively as a source of disease average, compared with the best conventional open-pollinated
resistance in wheat breeding programs. The disease-resistant counterparts. Indeed, impressive grain yields of hybrid rye of
genes are incorporated through a translocation of 1R chromo- over 11 tonnes per hectare have been reported. In Germany,
some into 1A or 1B chromosome of wheat. Many commercial where hybrid rye research originated in the 1970s, hybrid rye
wheat varieties contain one of these two translocations. varieties today comprise 70% or more of total acreage. Hybrid
Rye breeding efforts, whether for conventional or hybrid breeding that is much more complex than conventional pop-
rye, are typically focused on improving grain yield as the most ulation breeding exploits the heterosis effect, that is, enhanced
important objective. Other yield-related objectives include vigor, when two carefully selected genetically uniform but
improving resistance to drought and nutrient stress, shortness divergent inbred parents are crossed by controlled pollination
THE CEREAL GRAINS | Rye Grain: Its Genetics, Production, and Utilization 165
to obtain fertile hybrid seed. Apart from the grain yield advan- grain is softer. For the same reason, the tempering time for rye
tage to producers growing hybrid rye, the breeder benefits by is shorter, typically 615 h compared with 1624 h for hard
being the exclusive commercial source of that seed and spring wheat. Because of its softer texture, rye endosperm
through royalties. releases a high proportion of its weight as flour during the
break operations. Reduction is achieved by corrugated rolls
instead of the smooth rolls used in wheat milling, as smooth
reduction rolls tend to flake rye flour, which minimizes proper
Primary Processing of Rye Grain flour generation. Rye milling requires a larger sifting area than
that for hard wheat types because rye flour particles tend to
Agricultural products, including grains, vary in composition clump together because of its finer particle size distribution
and processing quality caused by fluctuations in abiotic (e.g., arising from the softer endosperm texture.
weather) and biotic (pests) factors during the growing season. Rye mills produce a number of different grades of flour by
To facilitate marketing and processing, the grain is graded, systematic blending of product streams to produce flours con-
separated, or combined into parcels of relatively uniform prop- taining different proportions of endosperm and bran. In North
erties. In countries where rye grades exist, they are based on America, usually three flour grades are distinguished based on
physical characteristics of the grain and on the presence of degree of refinement (Table 5), which can be evaluated based
foreign contaminants. Test weight, a measure of bulk density on ash or mineral content, fiber content, or flour color. A
related to milling quality, is the only grading factor that is relatively refined flour such as white or light rye will contain
measured objectively. All other factors are assessed subjectively mostly endosperm, and as the ash (and fiber along with pro-
by visual inspection. Normal or straight grades usually have a tein) content of endosperm is lower than that for bran, this
prescribed maximum moisture content, for example, 14% in explains why light rye flour has lower ash (and protein) com-
Canada and the United States. The number of grades of milling pared with medium and dark rye flour (Table 5). There is no
rye varies among countries. Germany has one grade, whereas standard of identity of North American flours compared with
Canada has three, and the United States has four grades. the situation that exists in Europe and Germany in particular
In transforming rye grain into food for human consump- where rye milling and baking considerations are more precise.
tion, the primary processing step is milling the grain into flour, A modern German rye mill will produce as many as seven
which basically involves two multistep processing operations, different flours designated by ash content and several types of
that is, a break and reduction systems. The break system breaks meals processed by crushing and cutting over a range of parti-
open the grain using corrugated rolls to separate bran from cle size (Table 6). Regardless of where rye is milled, the main
endosperm and involves some additional grinding of endo- quality parameter for flour is ash content, which has a close
sperm into flour, while the reduction system basically involves relationship to flour extraction, that is, the proportion of the
reducing to flour, the so-called middling particles of more or whole grain that is blended into flour. Ash content in turn has
less pure endosperm. Despite some significant differences a close correspondence with flour color; that is, the higher the
between the physical structure of rye and that of wheat, the ash content, the darker the flour (Table 5).
milling processes for the two grains are similar. The first step is
the removal of all undesirable contaminants. This is achieved
by a series of special cleaning machines functioning consecu-
tively. Differences in shape, size, and density are all used to Rye for Bread Making
improve the efficiency of the cleaning. The final step of the
cleaning section is tempering that involves addition of water to Bread and other baked products are the main foods produced
toughen the bran and mellow the endosperm to facilitate their from rye. Many types of baked products are made including
separation. Moisture levels for tempering rye are usually in the bread from 100% rye flour or whole meal; ryewheat bread
range 14.515.5% depending on kernel texture, and this range containing at least 50% rye; wheatrye bread containing at
is about 1% lower than that used for hard spring wheat, as rye least 50% wheat and at least 10% rye, rye bread, rye rolls,
This article is reproduced from the previous edition, volume 3, pp 298303, 2004, Elsevier Ltd.
Topic Highlights extracted from the autumn crocus plant to create chromosome
doubling, overcame this sterility problem. In 1938, Arne
Triticale is a man-made crop derived from hybridization of Muntzing from Sweden applied colchicine to wheat/rye
wheat and rye. hybrids, obtaining fertile plants. Once a fertile hybrid was
Triticale has desirable agronomic traits including high established, it became possible to utilize modern plant-
yield, and resistance to biotic and abiotic stresses. breeding methodologies. Early varieties were primitive and
Most triticale is used for animal, use of triticale flour has had numerous agronomic disadvantages such as low grain
been limited by quality issues, although there are opportu- yield, poor seed set, shriveled grain, excessive height, low ger-
nities in new product development. mination, and late maturity. Triticale improvement com-
menced in the 1960s to create new and better combinations
between wheat and rye, triticale and triticale, and triticale and
wheat. Most notable were breeding programs at the Interna-
Learning Objectives
tional Center for Maize and Wheat Improvement (CIMMYT) in
Mexico and the University of California at Davis (UCD) for
To understand the origin and development of triticale
spring triticales and programs in Poland and the University of
and the underlying genetics behind this new crop devel-
Manitoba, Canada for winter varieties.
opment.
There are two main types of cultivated triticales: octoploid
To understand where and why triticale is produced, and its
types produced from the hybridization of bread wheat, Triti-
agronomic advantages and limitations.
cum aestivum L., with rye, Secale cereale L., and hexaploid types
To understand the basics of triticale processing and utiliza-
using durum wheat, T. turgidum L., followed by chromosome
tion, especially in relation to protein quality.
doubling of the hybrid plant (Figure 1). Octoploid triticales
(2n 56) contain the A, B, and D genomes of bread wheat and
the R genome of rye, while the hexaploid triticales (2n 42)
Introduction contain the A and B genomes of durum wheat and the R
genome of rye. However, most triticale cultivars are hexaploids.
Triticale is the first man-made cereal grain crop species result- There is a third type of triticale (2n 28) produced from the
ing from the hybridization of wheat (Triticum) with rye hybridization of diploid wheat, T. monococcum (2n 14) with
(Secale), the name of which combines the scientific names of rye, but is not considered to be important economically.
the two genera involved. This synthetic amphiploid is obtained Although triticale is a cross between wheat and rye, it is self-
by chromosomal doubling after artificial crossing to produce pollinating (similar to wheat) rather than cross-pollinating
fertile hybrids. It is a small-seeded cereal grain that is used (like rye). Most agronomically desirable triticales that breed
for both human consumption and livestock feed. As a hybrid true have resulted from several cycles of improvement, and are
species, it combines many of the better qualities of both of primarily from the durumrye crosses with some common
its parents. Triticale possesses wheats properties for food wheat parentage occasionally involved.
production and ryes adaptive properties, and under certain In general, triticales can be divided into three groups
conditions can out-yield both parents. This promising crop (Table 1):
species is grown on more than 3 million hectares (Mha) world-
1. Primary triticales are the initial product of the
wide. Furthermore, triticale is an important germplasm source
wheat rye hybridization followed by doubling of chro-
for wheat improvement, providing a vehicle to transfer desir-
mosome numbers to produce the hexaploid or octoploid
able rye characteristics to wheat.
types.
2. Secondary triticales are produced by intercrossing pri-
mary triticales or by crossing a primary triticale with wheat.
Origin and Types 3. Substituted secondary triticales are hexaploid with A, B,
or D genomes of wheat substituted for one or more R
The Scottish scientist Alexander Stephen Wilson produced the
genome chromosomes of rye.
first triticale in 1876. Triticale was initially developed to com-
bine the positive traits of both parent types: the vigor and Primary triticales are often found to be fragile, poor producers,
winter hardiness as well as the higher protein content of rye and genetically unstable. They are used as breeding stock to
combined with the higher-quality gluten and baking properties produce the more stable and agronomically favorable second-
of wheat. However, initial progress was limited by the fact that ary and substituted secondary triticales. Secondary triticales
resulting hybrid progeny was sterile. In the 1930s, the discov- can be either hexaploid or octoploid and often contain com-
ery and use of the chemical colchicine, a natural chemical plete genomes of wheat and rye, whereas substituted triticales
Table 2 Desirable characteristics of wheat, rye, and triticalea Table 3 World triticale production, 2002
High-yield potential Many grains per ear High yield China SW 500 000 16.1
Large, filled grain High biomass High-quality straw Poland W 920 523 29.7
High harvest index Low-temperature growth High feed value Germany W 560 466 18.1
Tillering efficiency Winter hardiness Disease resistant France SW 269 000 8.7
Short straw Drought tolerance Stress tolerant Australia S 264 000 8.5
Sprouting resistance Disease resistance Winter hardiness Hungary W 132 000 4.3
High-energy grain Grain high in lysine High lysine content Belarus W 94 200 3.0
Czech Republic W 53 093 1.7
a
Modified from Semundo Limited (1994) Triticale The Hybrid Evolution. Cambridge: Canada SW 47 282 1.5
Semundo Limited. Denmark W 37 657 1.2
Austria W 37 621 1.2
Sweden W 30 740 1.0
Spain S 29 900 1.0
does well in regions where wheat performs poorly, such as cold Portugal S 25 000 0.8
and infertile soils, extremely sandy soils, soils with high levels Lithuania W 20 000 0.6
of boron, salty soils, acidic soils, manganese-deficient soils, Slovakia W 18 372 0.6
and dry soils. One particular concern, however, is the presence Latvia W 15 500 0.5
of ergot infection (caused by the fungus Claviceps purpurea) in United Kingdom W 14 000 0.5
some areas. Switzerland W 13 500 0.4
The first commercial triticale cultivars were released in Belgium W 12 000 0.4
United States SW 8979 0.3
1969. Today triticale is becoming a crop in its own right and
Estonia W 6847 0.2
is grown on over 3 Mha worldwide and in at least 27 countries
Netherlands W 4618 0.1
(Table 3). This crop contributes more than 10 Mt year1 to Luxembourg W 4000 0.1
global cereal production. Since its introduction, the area har- Norway W 1000 0.03
vested has increased over 7 times and amount harvested has Tunisia S 1000 0.03
increased over 18 times (Figure 3). Mexico S 850 0.03
Although it is grown throughout the globe, the countries Algeria S 3 0.0001
that produce the most triticale are China, Poland, and World 3 122 151 100
Germany. There is also significant production in Canada and a
S: spring type; W: winter type.
United States.
Sources: United Nations FAOSTATS, Statistics Canada, and United States Census of
Agriculture.
Usage
Care should also be taken to insure that the crop is not infected
Use of triticale for human consumption has not yet become with ergot.
widespread. Although triticale flour and products are available Alternative uses of triticale include use as a cover crop to
commercially (namely in specialty markets such as health food prevent soil erosion and land reclamation. Triticale has also
stores), this availability is limited. It comes in several forms been used in limited amounts as raw material in bioethanol
including whole berry, flakes, and flour. Whole triticale can be production. Ethanol plants pay a premium for triticale over
cooked and used in a variety of dishes. Quality evaluations barley, since it has more starch and no hull, making alcohol
have shown triticale grain inferior to wheat for milling and production more efficient.
baking, making large-scale commercial baking not feasible.
Triticale flour is low in gluten, and bread made from it alone
is heavy. For that reason, it is usually combined half-and-half
with wheat flour. If mixed with wheat or rye flour, triticale Genetic Resources
flour can be used to make a number of breads and pastries. In
developing countries, triticale flour is often mixed with wheat As a synthesized species, triticale has no wild ancestors and
flour during wheat shortages. It is of course important that the there are no existing landrace varieties. In addition, the actual
crop is not infected with ergot, as this is highly toxic to wheat and rye parents used in triticale synthesis are often either
humans. unknown or no longer available. It is therefore often not
Most triticale production is used for animal feed. It offers possible to resynthesize unique triticale genotypes through
better amino acid balance, lysine content, and higher protein, hybridization. Genetic resources for the development and
particularly important for swine and poultry. However, triticale enhancement of triticale include existing triticales, wheat and
has lower energy content than other grains, and feeding of rye, and the ancestral species of both wheat and rye. Itself,
triticale must be supplemented with other grains. It can also triticale exists as a genetic resource for the improvement of
be used as forage, silage, or hay for ruminants, offering high wheat, providing a vehicle to transfer desirable characteristics
digestibility and out-yielding traditional crops in dry soils. from rye.
THE CEREAL GRAINS | Triticale: Overview 171
20 000 000
production (Mt)
area (ha)
10 000 000
8 000 000
6 000 000
4 000 000
2 000 000
0
76
78
80
82
84
86
88
90
92
94
96
98
00
02
72
74
19
19
19
19
19
19
19
19
19
19
19
20
20
19
19
19
Figure 3 World production (Mt) and area (ha) trends since the 1970s. (Source: United Nations FAOSTAT.)
International Board for Plant Genetic Resources (1985) Descriptors for rye and triticale. Qualset CO (2002) Triticale, McGraw Hill Encyclopedia of Science and Technology, 9th
Rome: IBPGR Secretariat. edn. The McGraw-Hill Companies, Inc.
Limited Semundo (1994) Triticale The Hybrid Evolution. Cambridge: Semundo Limited. Qualset CO, Rupert EA, and Prato JD (1973) Triticale in California: Review of current
Lorenz K (1974) The history, development, and utilization of Triticale. Critical Reviews research and appraisal as a new cereal crop. In: Yang SP (ed.) Proceedings of the
in Food Technology 5(2): 175280. International Triticale Symposium Lubbock, Texas: International Center for Arid and
MacIntyre R and Campbell M (eds.) (1973) Triticale Proceedings of an International Semi-Arid Land Studies.
Symposium Ottawa, Canada: International Development Research Center. Tsen CC (ed.) (1974) Triticale: First Man-Made Cereal. St. Paul, MN: American
Muntzing A (1979) Triticale Results and Problems. Berlin, Germany: Verlag Parl Parey. Association of Cereal Chemists.
National Research Council (1989) Triticale: A Promising Addition to the Worlds Cereal Yang SP (ed.) (1973) Proceedings of the International Triticale Symposium Lubbock,
Grains. Washington, DC: National Academy Press. TX: International Center for Arid and Semi-Arid Land Studies.
Oats: Overview
PK Zwer, South Australian Research and Development Institute (SARDI), Urrbrae, SA, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights Oat is used for animal feed, human consumption, and
nonfood uses. Figure 2 shows the uses of oat relative to the
The history of oat including speciation, domestication, and proportions utilized.
early cultivation Oat was a traditional feed on farms for centuries and pow-
Oat plant growth and morphology ered workhorses until the introduction of machinery powered
Growing oat including agronomy, soil factors, and climate by fossil fuels. Between 50% and 90% of the world oat pro-
Growing oat diseases and pests duction is used as animal feed for horses, cattle, and sheep.
Processing oat Naked oat is particularly suited for poultry, pigs, racehorses,
Using oat for feed, human consumption, industrial, and birds. Oat as an animal fodder is also important for
nonfood, and fodder end uses domestic and export markets. Oat can be grazed as a green
Breeding improved oat varieties feed, made into silage, utilized as straw, or baled for hay.
Oat for human consumption is used to produce traditional,
functional, and medicinal products. Oat is differentiated from
other cereal grains by using the entire kernel after the hull is
Learning Objectives removed for many food products. Porridge or oatmeal, hot
cereals, bread, biscuits, infant food, and muesli or granola
To become familiar with the development of oat as a culti- bars are a few examples of food products produced from oat.
vated crop Nondairy food uses have been developed resulting in oat milk,
To gain general knowledge of the morphological parts of an yogurt, and ice cream. Oat has been shown to have health
oat plant and developmental stages benefits for lowering blood cholesterol, normalizing blood
To learn production constraints such as agronomy, dis- glucose levels, and reducing the risk of colorectal cancer. Phar-
eases, and pests macological properties are reported in the literature and prod-
To understand basic processing of oat for human ucts are being developed for commercialization.
consumption Nonfood and industrial oat products vary from cosmetics,
To become familiar with the many uses of oat grain and to the production of cardboard products, to the manufacture of
fodder furfural and furan compounds used for solvents, adhesives,
To know the basic knowledge of oat variety improvement, filtering aids, and the construction of board material and cel-
breeding systems, and new breeding technology lulose pulp.
Although oat production is declining globally, specializa-
tion for animal feed, hay, food, industrial, and pharmaceutical
products is growing and provides opportunities for adding
Introduction value to the oat crop.
This article will briefly outline the significance of oat as a
Oat has played a significant role in farming systems from cereal crop including origin, domestication, plant morphol-
domestication to the present due to the versatile uses of the ogy, production, end uses, and variety improvement.
grain and plant (Figure 1). Oat currently ranks seventh in
world production of cereals after maize, rice, wheat, barley,
sorghum, and millet. World oat production exceeded rye and
triticale. Classification
Oat is primarily grown in cool temperate climates with
67% of world production occurring in the northern hemi- The genus of oat is Avena L. (Poaceae) and belongs to the tribe
sphere. The Russian Federation, Canada, Poland, Finland, Aveneae of the family Gramineae. The primary species culti-
Australia, and Finland were ranked as the top six countries vated is Avena sativa. However, Avena byzantina and Avena
for world oat production. Oat is also grown in the southern strigosa are also grown in some regions for animal feed and
hemisphere with Australia ranked first in production while fodder.
Argentina, Chile, and Brazil are also significant producers. The species described in Avena form a polyploid series vary-
There are two types of oats husked oat, with hulls sur- ing from 1 to 3 chromosome sets with the basic chromosome
rounding the kernel or groat after harvest, and naked oat, number (n) 7. Each chromosome set has genome designations
where the hull is removed when the crop is harvested. Naked A, B, C, and D donated by a different oat species in the evolution
oat has the free threshing character similar to wheat. Husked of the genus. The different species with varying ploidy levels
oats represent the majority of global oat production except in evolved over time and will be discussed in the next section.
China where naked oat is the most common type. Naked oat is The diploid species have one set of chromosomes and are des-
gaining prominence for specialist markets as improved varie- ignated as 2n 2x 14. This designation indicates the diploid is
ties are being developed. composed of one genome with 14 chromosomes. The tetraploid
Agronomy
Spring varieties are primarily sown in the southern hemisphere
during the winter months. Both spring and winter varieties are
grown in the northern hemisphere and the time of sowing will
vary with latitude. Sowing rates are dependent on a combina-
tion of climate, soil conditions, and use of the crop. Sowing rates
are higher for fodder production when compared to grain pro-
duction. Seed size also varies extensively in oat. Hence, sowing
rates need to be calculated as the number of seeds m2 rather
than the weight of seeds m2 to achieve optimum plant density.
Adequate nutrition of oat is essential to achieve maximum
yields. Inputs such as nitrogen, phosphorus, potassium, and
sulfur are applied based on soil tests. Micronutrient deficien-
cies for copper, zinc, manganese, molybdenum, and iron occur
in some soils. Micronutrient toxicity for boron, aluminum, and
manganese also occurs in some soils.
Oat generally tends to be more sensitive to herbicides than Figure 6 Oat stem rust.
wheat or barley. Sensitivity is also variety-dependent.
A harvester is used to cut standing plants and threshes the
grain from the plant. Wind rowing is another option for har-
vest and requires that plants are cut and put into rows for grain
threshing at a later date.
Nematodes
rust, the disease causes significant yield losses when it occurs. Cereal cyst nematode (CCN), Heterodera avenae Wollenweber,
The disease is reported in eastern Canada, the United States, causes a serious economic threat to oat production worldwide
Australia, Europe, Great Britain, and Israel. Septoria is a major (Figure 15). In regions where CCN is abundant, genetic resis-
production constraint in Western Australia (Figures 8 and 9). tance reduces CCN population sizes and oat varieties with
Pyrenophora leaf blotch caused by Pyrenophora avenae Ito & genetic tolerance ensure maximum productivity. Genetic resis-
Kuribayashi apud Ito occurs worldwide with varying economic tance and tolerance are independently inherited.
significance (Figure 10). Pyrenophora was the third most Stem nematode, Ditylenchus dipsaci (Kuhn) Filipjev, limits
important disease in Germany, significant in Brazil, and com- oat production in cool moist climatic conditions (Figure 16).
mon in Scandinavia. Red leather leaf caused by Spermospora Winter oat sown where mild winters occur and spring oat sown
avenae Sprague & A.G. Johnson is a minor disease identified in in the winter (Mediterranean climates) are affected by the
northwestern United States, Turkey, and Australia (Figure 11). nematode. Resistance and tolerance appear to be more closely
associated for stem nematode than CCN.
Root lesion nematode, Pratylenchus neglectus, affects oat
Diseases Caused by Bacteria crops in Europe, Iran, the United States, and Australia. Yield
losses up to 37% were recently demonstrated in South Austra-
Halo (Pseudomonas syringae pv. coronafaciens) and stripe blight lian trials where the nematode population was high. Resistance
(P. syringae pv. striafaciens) are collectively called bacterial and tolerance mechanisms are inherited similar to CCN.
blight (Figure 12). The disease is common while cool, moist
conditions persist. Severe foliar symptoms develop on suscep-
tible varieties. Dry conditions limit the spread of bacterial
Insect Pests
blight.
Russian wheat aphid (Diuraphis noxia), greenbug (Schizaphis gra-
minum), bird cherry-oat aphid (Rhopalosiphum padi), grain aphid
(Macrosiphum avenae), and rose grain aphid (Metopolophium dirho-
Diseases Caused by Viruses
dum) can attack oat with varying effects on productivity.
Barley yellow dwarf virus is the most yield-limiting viral disease Army worm (Phalaenidae spp.), fruit fly (Oscinella frit (L.)),
of oat and is an economic threat worldwide (Figures 13 and and wireworms (Agriotes spp.) are insect pests that affect oat
14). The virus is transmitted by a number of aphid species. production in varying capacities worldwide.
Figure 12 Bacterial blight in oat.
Figure 13 Barley yellow dwarf virus causing blasted florets and leaf
Figure 11 Red leather leaf in oat. discoloration.
180 THE CEREAL GRAINS | Oats: Overview
Oats corn, wheat, and barley. Lignin is the primary fiber fraction of
the hull and reduces grain digestibility in animals. Lignin
Stone milling content also varies in different varieties. Although hulls with
Impact milling Cleaning Impact milling high lignin reduce digestibility, varieties with lower-hull lignin
content can have a beneficial effect for horses, cattle, and
Kilning sheep. The hull reduces digestive problems in these animals.
The hull is a major constraint as a feed grain for poultry and
Grading Grading pigs. It reduces digestibility resulting in low protein and poor
energy. Because naked oat does not have a hull, the grain
Hulling Hulling provides a good source of energy for grower and weaner pigs,
Hull and fines Hull and fines broilers, and laying hens.
seperation seperation Overall, oat is a favorable feed for ruminants such as cattle
and sheep. Oat is also the preferred feed for horses due to the
Groat Groat
separation separation palatability, digestibility, and nutritive value of the grain.
Naked oat is also used for racehorses, due to the limited
Kilning requirement of grain intake and the need for a good source of
energy.
Broken or Large
small groats groats Oat Grain for Human Consumption
Cut Cutting Flaking Grinding Products produced from oat are generally made with the whole
groats grain after the hull is removed, whereas other cereal products
are produced from grain with the germ and bran removed.
Milling Flaking Premium Oat Whole oat has the highest protein and a favorable ratio of
oat flakes flour unsaturated to saturated fatty acids compared with other
Oat bran
Oat flour cereals. Whole oat also provides vitamins, minerals, and anti-
oxidants. The heat treatment used to stop rancidity also
Fast cooking enhances the unique sensory characters of oat.
oats
The unique composition of whole oat results in healthy
Figure 17 The milling process of an oat kernel. products. b-Glucan content in the groat varies between 1.8%
and 7.5% on a dry-matter basis. The high level of water-soluble
dietary fiber of which b-glucans are the main component helps
the proportion used. Animal feed is the largest use of oat; cereal
to reduce high blood cholesterol and normalize blood glucose
products and other human food the second most common
levels. The fiber also reduces the risk of colorectal cancer. The
use. Although pharmaceuticals represent the smallest group
groat also contains vitamins and antioxidants. Lactose-
of products, they have the greatest potential for future growth
intolerant individuals can now purchase nondairy oat
and value.
products.
The main product produced from oat is porridge or oat-
meal. Other food products processed from oat are cold cereals,
Oat for Feed Grain infant foods, muesli or granola bars, breads, biscuits or cook-
ies, thickeners, and specialty flour. New processes were devel-
Oat has been a traditional feed grain for centuries. Recent
oped recently to manufacture nondairy products such as milk,
advances in corn, wheat, barley, soybean, and canola as feed
ice cream, and yogurt using oat. Antioxidants in oat flour can
grains have resulted in a worldwide decline of oat production
be used to stabilize some milk and meat products sensitive to
for feed. Despite the decline, the primary use of oats remains as
fat oxidation during storage. Oat gum primarily composed of
a feed grain. Prior to mechanization, oat was the primary feed
b-glucans is used to stabilize ice cream. Oat proteins have been
for horses that powered farm equipment. Oat is a suitable feed
used in many food products including heat-resistant chocolate,
for dairy and beef cattle, sheep, and horses. Although not as
because of viscosity and emulsification properties.
prevalent, oat can also be used for poultry, pigs, cats, dogs,
birds, rabbits, bison, deer, and fish. In recent years, naked oat is
being developed as a feed grain with improved nutritional
Oat Grain for Industrial and Nonfood Uses
value for markets such as weaner and grower pigs, poultry,
racehorses, and birds. The following discussion centers on the There are many potential uses of oat grain fractions and hulls
traditional oat grain possessing a hull. in products such as adhesives, cosmetic products, pharmaceu-
The nutrient value for animal feed is based on the propor- ticals, and oil-spill cleanup. Some of the uses are proposed,
tion of groat to hull. The ratio varies with variety and environ- whereas the technology for other uses has been developed and
ment. Oat groats have a higher oil or lipid content than other implemented.
cereals, varying between 3% and 11%. The oil is composed The unique chemical composition of the oat groat is
primarily of unsaturated fatty acids, which can alter the fatty responsible for oat starch gels having more elastic, adhesive,
acid composition of the animal fat. Protein content in oat and translucent properties than wheat or maize starches. Oat
groats varies from 9% to 15% with higher lysine content than starch could replace products produced from wheat or maize
182 THE CEREAL GRAINS | Oats: Overview
with the waxy starches the exception. Products currently pro- Breeding Priorities
duced from starch include brown paper and cardboard
Breeding priorities determine selection criteria that eventually
products and coating agents for tablets in the pharmaceutical
result with the release of oat varieties with improved traits.
industry.
Understanding production and processing constraints of oat
Oat has been used in cosmetic products for some time. The
is the initial step in identifying breeding priorities for improv-
starch characteristics make oat a useful ingredient in bath
ing varieties. These priorities will differ or change in emphasis
products to relieve itching and nappy rash. Other products
depending on the growing environment and end use of the oat
produced from oat include cleansers to replace soap in facial
plant or grain. Breeding priorities are based on gross margin for
masks, soap for cleansing dry and oily skin, shampoo, lotions,
growers and processors, resulting in a superior product for the
bath additives, and skin care.
consumer and processor. Breeding priorities set to achieve
Pharmaceutical products have great potential, but are not
long-term goals are based on developing varieties that offer
currently tapped. The health benefits of oat fractions were
new products or increased value for both growers and industry.
discussed in the preceding section.
The long-term breeding priorities are particularly important for
The hull can also be used to produce industrial products.
oat due to the global decline of production.
Oat hulls are used to produce furfural and furan compounds.
Although there are many different growing environments
These compounds are used in the manufacturing process of
and end uses globally, the primary breeding priorities deal with
crude petroleum; nylons; formaldehyde furfural resins; sol-
productivity, disease resistance, agronomy, and quality.
vents for dyes, resins, lacquers, paints, and varnish; elastomers
Increased grain and hay yields are the primary aims of
and thermoplastics; phenolic resin glues and plywood adhe-
increased productivity. Improved disease resistance increases
sives; construction board material; and cellulose pulp. A very
yield and enhances quality.
different aspect of hull utilization is the cariostatic properties of
The significance of oat diseases varies with region and con-
tooth protection.
tinent and is prioritized by breeding programs. Diseases such
as stem and leaf or crown rust are two global diseases that cause
significant losses to the oat industry. Barley yellow dwarf virus,
Oat for Fodder
smut, and septoria also reduce yield and quality in certain
Fodder production is also an important component of animal production areas of the world. Cereal cyst, stem, and root
feed shown in Figure 2. Several types of oat fodder, such as lesion nematodes when present in the production environ-
hay, silage, grazing, and straw, are produced as an essential ment are equally as devastating, but more regionally localized
component of animal production systems worldwide. in the world.
Oat hay is produced for dairy and beef cattle, horses, and Important agronomic characters are standing ability, matu-
sheep. Hay not only is an important commodity domestically rity of the plant, and the ability of the plant to resist shattering
but also is traded internationally. A significant market is the or loss of seed at maturity. Winter hardiness is also an impor-
Asian dairy and beef cattle industry. The oat hay crop is cut at tant agronomic trait of the varieties that require a cold period
the watery-ripe growth stage (Zadoks 71) for optimum hay to produce seed.
quality. Hay quality characters that may be associated with Grain quality characters that are essential for the industry
palatability are digestibility, water-soluble carbohydrates, acid are high groat to hull ratio and groat color. Weather and
detergent fiber, neutral detergent fiber, and stem diameter. disease can cause groat discoloration. Other grain quality char-
Sensory quality characters are also important and include hay acters of importance for variety development are protein, oil,
color and aroma. b-glucan content, hectoliter weight, and screening percentage.
Cattle and sheep graze oat during the vegetative stage when The most important aspect of fodder quality is palatability. The
the plant height is from 20 to 40 cm. Animals are removed traits that comprise palatability are being researched but are
from the oat crop before the growing point of the plants is most likely a combination of digestibility, water-soluble car-
damaged, so the crop produces grain. Grazing is important in bohydrates, neutral detergent, and acid detergent fibers, color,
countries worldwide including New Zealand, Australia, South and smell.
America, northern Africa, Nepal, and the United States.
Silage production using oat is used in the dairy and beef
cattle industry worldwide. Oat is cut in the early heading stage, Breeding Methods
chopped, and ensiled.
Breeding systems should be flexible and easily modified for
changes in breeding priorities. A combination of breeding
methods is combined in a program to produce improved oat
Breeding-Improved Oat Varieties varieties. Classical breeding methods used for decades are ped-
igree, bulk, recurrent selection, backcross, and single-seed
History
descent. More detailed information about the breeding
Early oat improvement occurred when farmers selected the methods is available in Plant Breeding Methodologies by Neal
best-performing plants in their paddocks. The first attempt to Jensen.
introduce genetic variation by crossing different oat genotypes Doubled haploids (DH) are true breeding homozygous lines
was reported in England and accomplished by Shirreff in the produced in one generation. An economically viable method for
1860s. The first variety developed from hybridizing oat, Abun- DH production in oat is currently not available, but new devel-
dance, was released from Gartons Ltd., Warrington, England. opments using isolated microspore culture hold promise for
THE CEREAL GRAINS | Oats: Overview 183
Vegetative Organization
Learning Objectives
The coix plant is 70400 cm high. Its stems are erect or strag-
gling with branches and prop roots from the lower nodes. The
To understand the origin and taxonomy of coix and its
internodes of the culms are solid. Leaves are not basally aggre-
relationship to maize.
gated and nonauriculate, the leaf blades are lanceolate and
To understand the uses of coix, both as food and in tradi-
broad and 3070 mm wide, the shape is cordate or not
tional medicine.
cordate but not sagittate, they are flat and persistent without
To understand the basics of the chemical composition of
cross venation, they are rolled in bud with a ligule present, and
coix that contribute to its medicinal properties.
there is an unfringed to a fringed membrane.
Introduction
Reproductive Organization
Coix lacryma-jobi L., commonly called Adlay, Jobs tears, adley, The coix plant is monoecious with all the fertile spikelets being
adley millet, or coix millet, is an annual crop. Botanically, coix unisexual but without hermaphrodite florets (Figure 1). The
belongs to the tribe Maydeae (Tripsaceae) of the family Poa- spikelets of sexually distinct forms (female-only or male-only)
ceae, and it is classified as a close relative to maize. This annual are on the same plant, the glumes of the fertile spikelet are
grass is native to India, Burma, China, and Malaysia and has several-nerved, and all are enclosed finally in a bony beadlike
been grown extensively in Southeast Asia for several thousand involucre containing a minute fertile female flower and two
years. The plant is typically 1 m high with knobbly bamboo- sterile flowers. Pollen-bearing male flowers are produced on a
like stems, and the new tillers arise from the base of stems. The slender stalk that extends out of the bead through a tiny pore.
glossy deep green leaves are up to 5 cm wide with slightly wavy Two feathery stigmata from the fertile female flower also pro-
edges. The flowering and fruiting spikelets are insignificant; the trude from the pore ready to receive pollen from the male
grain is pear-shaped and 5 mm in diameter and has a hard, flowers:
shiny dark brown to grayblack hull. The grain is used in soups
1. Female-sterile spikelets: Male spikelets grow in pairs or tri-
and beverages and is popular in traditional Chinese medicine.
ads, several per disarticulating raceme. Rachilla of male
It is well known to have pharmacological properties. The
spikelets is terminated by a male floret. The male spikelets
ancient Chinese medical book Pen-tsao kang mu described it
have glumes and proximal incomplete florets or not (the
as an effective remedy for a number of maladies and particularly
lower floret sterile or male).
beneficial to the digestive system. Coix has been considered as a
2. Female-fertile spikelets: Spikelets fall with the glumes.
good potential germplasm resource for maize improvement
Rachilla is terminated by a female-fertile floret. Usually, it
because it is tolerant to lateritic soils and low pH, less prone
is pollinated by the wind.
to attacks by viruses, and less susceptible to waterlogging and
has a low variation in photoperiod. Accurate world production
figures are impossible to ascertain, but coix has a value beyond Seed
that of a typical commodity grain for food use.
The shiny gray to black beads are dispersed and planted like
seeds, but they are like little shells containing flowers and
Botany grains. The seeds are usually oval-shaped or egg-shaped, with
a milky white outer surface and a slightly sweet taste (Figure 2).
Habit
Fruit is medium sized with globular form and 612 mm long.
Coix is a light-sensitive and short-day-length plant and prefers Close-up view of flowering coix shows the hollow, beadlike
wet soils and cool weather. Although it is an annual crop, the involucres, which have a natural hole in them.
Thrumb
Female spikelet
Flower stalk Figure 3 The karyotype of Coix. Reproduced from Zhang F and Yi Z
(1994) Natural Science Journal of Changsha Normal University of Water
Male spikelet Resources and Electric Power 9(2): 187191.
Biochemistry
Like other members of the grass family, coix has seeds contain-
ing prolamin as the major storage protein. The coix prolamin is
known as coixin and represents more than 70% of the endo-
sperm protein. Based on differential solubility, coixins can be
separated into two factions: a-coixins and g-coixins. a-Coixins
Figure 1 Inflorescence of Coix. Adapted from the images from http://
are constituted by four size classes, while g-coixins comprise
www.desert-tropicals.com/plants/Poaceae/coix_lacryma-jobi.html. only one molecular weight class. a-Coixins are synthesized at
earlier developmental stages than g-coixins. The deduced
amino acid sequence of a full-length cDNA clone encoding a
sulfur-rich coix prolamin predicted a polypeptide of 194 resi-
dues, which shared 64% homology with the 17 kDa b-zein.
The mature protein contains the familiar composition of pro-
lamins and an unusually high content of the sulfur-containing
amino acids methionine (11.6%) and cysteine (5.2%).
Hydropathy analysis showed that a-coixin is slightly more
hydrophobic than b-zein.
The anticomplementary polysaccharides CA-1 and CA-2
purified from the seed of Coix lacryma-jobi L. var. ma-yuen
both consist of rhamnose, arabinose, xylose, galactose, galac-
turonic acid, and glucuronic acid but in different ratios. CA-2
contains more mannose and glucose. CA-2 showed more
potent anticomplementary activity than CA-1 in low dose.
The seeds of C. lacryma-jobi L. var. ma-yuen Stapf are used as a
traditional Chinese medicine reputably possessing antitumor
activity, attributed to an acidic fraction. Infrared spectroscopy
and gasliquid chromatography showed that this acidic frac-
tion was composed of four free fatty acids: palmitic, stearic,
oleic, and linoleic acids.
A protein inhibitor of locust gut a-amylase that was purified
from coix seeds consists of two major isomers, each dimer of
two closely similar or identical subunits of Mr 26 400 Da and
Figure 2 Photograph of flower and seed of Coix. http://www.biherb. associated by interchain disulfide bonds. These isomers
com/CoixSeedExtract.html.
also have closely similar amino acid compositions. The major
isomer shows no inhibitory activity against amylases from
Cytology other sources (human saliva, porcine pancreas, Bacillus subtilis,
Aspergillus oryzae, and barley malt). This highly specific func-
The standard karyotype formula is 2n 20 20m (Coix puel- tion may be relevant to protection of the grain from insect
larum). But there are other karyotype formulas outside of the feeding and fungal infection. The complete amino acid
standard types, such as 2n 20 18m 2sm (C. stenocarpa); sequence of the polypeptide of the major trypsin inhibitor
2n 20 18m 2sm and 2n 20 16m 4sm (Coix lacryma- from coix seeds showed 64 amino acids with a high content
jobi var. lacryma-jobi); 2n 20 12m 2m (sat) 6sm (Coix of cysteine. The sequence exhibits strong homology with a
186 THE CEREAL GRAINS | Coix: Overview
number of BowmanBirk inhibitors from legume seeds and Steady-state levels of DHPS mRNA are slightly reduced in the
similar proteins recently isolated from wheat and rice. endosperms and embryos of the maize lysine-rich O-2 mutants
The amylose contents of normal coix ranged from 15.9% to when compared with those in normal kernels. Also, the DHPS
25.8%, and those of waxy coix were 0.71.1%. Swelling power gene is not under the control of O-2.
of waxy coix starches varied between 28.6 and 41.0 g per g, Using random amplified polymorphic DNA (RAPD) markers,
generally higher than waxy maize. Normal coix starches 21 coix accessions were characterized for genetic variations and
had significantly higher gelatinization peak temperature relationships. The results indicated considerable variation at the
(71.171.4 C) than normal maize (71.975.5 C), similar to DNA level among the coix germplasm, and the classification by
waxy maize. Rapid Visco Analyser pasting profiles of normal RAPD data reflected the differences in geographic origins and
coix show some variation but closely match the normal maize evolution of coix. Recently, a coix genetic linkage map with
starch profile. Pasting profiles of waxy coix have more variation restriction fragment length polymorphism (RFLP) and amplified
and lower peak viscosities than waxy maize starch. Waxy coix fragment length polymorphism (AFLP) markers was constructed.
starches form very weak gels, while the gel hardness of normal The map consisted of ten linkage groups, consistent with the
coix starches was 11.431.1 g. The gelatinization temperature chromosome numbers observed cytogenetically.
of waxy coix starches varies between 63.4 and 76.4 C, and
water-binding capacity ranges from 103% to 108%. The starch
yield from coix grains averages 45%, the birefringence loss of Utilization
starch granules begins at 65 C and finishes at 75 C, and
breakdown during pasting in a Brabender Visco-Amylo-Graph Coix may be a source of ornamental beads, a staple sustenance,
is substantial. The properties of waxy coix starch are similar to and a productive fodder grass increasingly viewed as a poten-
those of waxy maize starch. Amylose content is the main factor tial energy source. It has long been consumed both as a nour-
controlling differences in starch properties of coix starches. ishing food and as an herbal medicine.
Oil yields ranged from 3.03 to 4.83 wt%, ferulate phytos-
terol esters in oils ranged from 0.109 to 0.119 wt%, free phy-
tosterol varied from 0.54 to 0.61 wt%, phytosterol fatty acyl Food
esters in the oil ranged from 0.79 to 0.98 wt%, total phytos-
terols were from 1.43 to 1.69 wt%, and g-tocopherol varied Before maize became popular in South Asia, coix was rather
between 0.030 and 0.045 wt%. Whole kernel oil yields of coix widely cultivated as a cereal in India and still occurs as a minor
are generally higher than Zea mays ssp. mays and teosinte cereal. Coix was used as food, particularly by peasants in the
accessions. Whole kernel oil yields and levels of four phytonu- Far East, very early in agricultural history. In wild varieties, the
trients (free phytosterols, fatty acyl phytosterol esters, ferulate fruit has a hard, shiny coat. After domestication, this coat
phytosterol esters, and g-tocopherol) in oils were measured became less hard and easier to cook, for example, into por-
from three coix accessions (C. lacryma-jobi L.). ridge. Coix ranks along with wheat and barley in the Near East;
beans, maize, squash, and pepper in the Americas; and rice in
Asia as one of the earliest domesticated plants. As with other
Molecular Biology cereals, there are many coix genotypes, including soft-shelled,
easily threshed types with a sweet kernel.
The seed storage proteins of coix, sorghum, and maize are Though the hard seed coat, removed by decortication,
coded by homologous genes, which are coordinately expressed makes extraction of the flour rather difficult, this is a poten-
in the endosperm in a temporal-specific fashion. Opaque-2 tially very useful grain because it has a higher protein to car-
(O-2), a bZIP protein originally isolated from maize, has bohydrate ratio than any other cereal (Table 1). Reported data
been described as a transcription activator of a-prolamin and differ somewhat. According to List and Horhammer, 100 g of
b-prolamin genes. The coding region of the coix O-2 gene is seed contains 358 cal, 11.2 g H2O, 15.4 g protein, 6.2 g fat,
interrupted by five introns and codes a polypeptide of 408
amino acids. Comparison of the deduced amino acid sequence Table 1 Typical composition of Coix seed compared with other
with two different sequences of maize O-2 protein showed that grains (on a 14% moisture basis)
the coix O-2 protein is similar to the maize O-2 isolated from a
Coix Rice Wheat Maize
maize inbred line, W22. The coix O-2 protein has the same
binding specificity and expression pattern of the maize O-2. Energy (cal) 1500 1460 1480 1510
Dihydrodipicolinate synthase (DHPS) is the main enzyme Protein (%) 14.2 7.8 9.9 8.4
of a specific branch of the aspartate pathway leading to lysine Fat 3.6 1.3 1.8 4.3
biosynthesis in higher plants. The open reading frame of the Starch (%) 67.2 76.6 74.6 70.2
DHPS-encoding DapA gene from C. lacryma-jobi is interrupted P (mg) 299 203 268
by two introns and encodes the 326-amino acid-long coix Mg (mg) 126
DHPS protein, which is 95% identical to the maize DHPS Ca (mg) 49.0 9 38 34
protein. DHPS transcripts are present in coleoptiles, embryos, Fe (mg) 2.90 4.2
Zn (mg) 1.89
endosperms, and roots but are almost undetectable in blades
Mn (mg) 1.89
of young leaves of both coix and maize. The 50 -flanking region
of DapA gene contains a TGACTC GCN4-like element located Per 100 g for fat and mineral contents.
372 bp upstream from the putative translation start codon. Source: www.fao.org, www.chinafeedbank.com.cn, and www.cmymy.51.net.
THE CEREAL GRAINS | Coix: Overview 187
aa Arg Cys Gly Hys Ile Leu Lys Met Phe Thr Try Tyr Val
4.4 1.7 2.8 2.3 4.0 14.4 1.9 3.0 4.8 3.1 nd 4.2 5.6
65.3 g total carbohydrate, 0.8 g fiber, 1.9 g ash, 25 mg Ca, warts. It is also used in the treatment of lung abscess, lobar
435 mg P, 5.0 mg Fe, 0 mg b-carotene equivalent, 0.28 mg thia- pneumonia, appendicitis, rheumatoid arthritis, beriberi, diar-
mine, 0.19 mg riboflavin, 4.3 mg niacin, and 0 mg ascorbic rhea, edema, and difficult urination. The action of coix seeds
acid. There is 5060% starch, 18.7% protein (mainly gluta- against many kinds of disease can be attributed to various
mine, leucine, tyrosine, arginine, histidine, and lysine), and components with pharmacologically different activities. Coix
510% oil with glycerides of myristic and palmitic acids. The roots can be also used in the treatment of menstrual disorders.
partial amino acid composition of crude protein in coix seeds A major development in the utilization of coix has been the
is shown in Table 2. It should be noted that despite the level of advance of Kanglaite, which is coix seed oil injected intrave-
lysine noted, it is still deficient relative to human dietary needs. nously as a treatment for many forms of cancer. Coixenolide
Coix seeds can be used for baking, mixed with wheat flour, and coixol (regarded as anticancer and anti-inflammatory,
and for making porridge and beer. They are usually pounded, respectively) are two major active fractions of coix oil (see
threshed, and winnowed. The pounded flour is sometimes Figure 3). It has become a widely used cancer therapy, and a
mixed with water, similar to use of barley for the beverage much-hyped treatment in the West. Clinical evidence is posi-
barley water. The pounded kernel is also made into a sweet tive to some degree, and many studies have recently been
dish by frying and coating with sugar. The grains are also conducted, for example, on gastric cancer, on pancreatic cancer
utilized in soups, broths, porridge, drinks, and pastries. In xenografts, and on stimulation of anticancer-related immune
India, the Nagas, a group of tribes near the Myanmar border, responses. In fact, numerous recent studies indicate possible
use the grain for brewing a beer called zhu or dzu. A Japanese anticancer and other medicinal roles of polysaccharide, protein
variety called Ma-Yuen is brewed into a tea and an alcoholic (antidiabetic), polyphenols, bran, and oil of coix. Some of
beverage, and roasted seeds are made into a coffee-like drink. these uses may come to lack support from future rigorous
In some cases, the hulled grain is adapted for parching or clinical trials. Some uses may depend on complex interactions
boiling like rice; it is also dehusked and eaten like a peanut. of coix components with other aspects of a long-term healthful
Beers and wines are made from the fermented grain. Continu- diet. But certainly, there is abundant evidence that coix may
ing recent interest in new product development from coix is have vast future potential as a nutraceutical source.
apparent, and coix bran may be particularly notable as a high-
potential food ingredient.
Fodder
The leaves and stems provide a useful fodder for cattle and
Medicine and the Rise of KanglaiteW
buffalo. The grass is very suitable for growing in waterlogged
areas. Its average yield is 29 tonnes ha1. The grass can also be
According to traditional Chinese medicine, coix seeds serve
turned into silage.
several functions. They stimulate function of the spleen and
lung, remove heat, help in the drainage of pus, and induce
diuresis. Coix seeds have been widely employed as a diuretic,
stomachic, analgesic, and antispasmodic agent from ancient Genetic Resources
times. They are also used to treat the symptoms of diarrhea and
arthritis. The fruits are anodyne, anti-inflammatory, antipy- Coix is reported to tolerate lateritic soils, low pH, slope,
retic, antiseptic, antispasmodic, hypoglycemic, hypotensive, viruses, and waterlogging and to be photoperiod-insensitive.
sedative, and vermifugal. According to Hartwell, the fruits are It is a potential gene resource for improvement of other cereal
used in folk remedies for abdominal tumors; esophageal, gas- crops. It is a perennial plant, which forms several stalks, and its
trointestinal, and lung cancers; various tumors; and excres- pollen can be collected throughout the year when the plant is
cences, warts, and whitlows. This folk reputation is all the maintained in a controlled environment. It can be used as the
more interesting when reading that coixenolide (Figure 4) pollen parent for wheat crosses for haploid production without
has antitumor activity. It is also a folk remedy for abscess, requiring synchronization of flowering dates.
helminthiasis, anthrax, appendicitis, arthritis, beriberi, bron-
chitis, cancer, catarrh, diabetes, dysentery, dysuria, edema,
fever, headache, phthisis, pleurisy, pneumonia, rheumatism, Cultivation
small pox, splenitis, etc., is an anodyne and tonic, and can be
used during puerperium. Coix seeds are reportedly effective in Generally, coix is propagated by seeds. The plant prefers light
treating verrucas caused by the human papillomavirus. A tea (sandy), medium (loamy), and heavy (clay) soils. It prefers
from the boiled seeds is drunk as part of a treatment to cure neutral and basic soils but can grow in very acidic soil, and it
188 THE CEREAL GRAINS | Coix: Overview
Name Coixol
Synonyms 6-Methoxy-3H-benzoxazol-2-one
H
N
Molecular Structure O
O O
Name Coixenolide
Synonyms [(2S,3R )-3-[(Z)-hexadec-9-enoyl]oxybutan-2-yl] (E)-octadec-11-enoate
O
H3C
O
Molecular O CH3
Structure H3C
CH3 O
C17566
Molecular
C38H70O4
Formula
Molecular
590.96
Weight
Figure 4 Molecular data and structure of coixol (from www.Chemblink.com) and coixenolide (from Kegg Brite via www.genome.jp).
requires moist soil. It is sown at rate of 610 kg ha1. Seed is Exercises for Readers to Explore the Topic Further
planted 2.5 cm deep, at a spacing of 60 60 cm. One intercul-
tivation, before the plants tiller and cast shade on the ground, Write a proposal for the development of a line of nutraceu-
may be necessary. The largest plants result when the plant is tical ingredients based on coix.
given partial shade although it will tolerate full sun. It adapts to Suggest how traditional food uses of coix could be mod-
wet soils and so may be used near water. Sufficient rains in the ernized and marketed as new products.
early stage of growth and a dry period when grain is setting are Investigate the potential for transfer of medicinally impor-
necessary for good yields. Plants respond well to liberal appli- tant components of coix grain to the high-yielding and
cations of organic manure. The grain is usually harvested in the related crop species, maize.
fall when the plant matures and is dried in the sun. Plants are
cut off at base and grain is separated by threshing. Coix yields
depend on varieties cultivated in different countries. Usually,
See also: Breeding of Grains: Maize: Breeding; Food Grains and
unhusked grain yields vary from 2.1 to 3.5 tonnes ha1.
Well-being: Functional Foods: Dietary Fibers, Prebiotics, Probiotics,
Leaf blight, caused by Bipolaris coicis, is one of the most
and Synbiotics; Functional Foods: Overview; Genetics of Grains:
destructive and major yield-limiting diseases in coix. Plant
Maize: Genetics; The Basics: Taxonomic Classification of Grain
response to leaf blight is quantitative rather than qualitative,
Species; The Cereal Grains: An Overview of the Family of Cereal
varying greatly among cultivars or lines tested. Yield reductions
Grains Prominent in World Agriculture; Maize: Overview; The
are primarily associated with decrease in kernel numbers per
Legumes and Pseudocereals: Amaranth: Overview; Buckwheat:
plant, thousand kernel weight, and percent kernel ripeness.
Overview; Pseudocereals: Overview; Quinoa: Overview.
Other diseases common in coix are rusts and smuts. Rusts are
caused by Puccinia, while smuts are caused by Tilletia and
Ustilago. Leaf-gall virus and the nematode Meloidogyne incognita
acrita also attack this plant.
Further Reading
Ary MB, Richardson M, and Shewry PR (1989) Purification and characterization of an
insect alpha-amylase inhibitor/endochitinase from seeds of Jobs tears (Coix
Exercises for Revision lachryma-jobi). Biochimica et Biophysica Acta 999: 260266.
Ary MB, Shewry PR, and Richardson M (1988) The amino acid sequence of a cereal
Compare and contrast the composition and reported Bowman-Birk type trypsin inhibitor from seeds of Jobs tears (Coix lachryma-jobi
L.). FEBS Letters 229: 111118.
medicinal claims for coix and other minor grain crops,
Dante RA, Neto GC, Leite A, Yunes JA, and Arruda P (1999) The DapA gene encoding
such as Amaranthus, buckwheat, and quinoa. the lysine biosynthetic enzyme dihydrodipicolinate synthase from Coix lacryma-
Prepare a map illustrating global production and uses jobi: Cloning, characterization, and expression analysis. Plant Molecular Biology
of coix. 41: 551561.
THE CEREAL GRAINS | Coix: Overview 189
Duke JA and Ayensu ES (1985) Medicinal Plants of China. Algonac, MI: Reference Vettore AL, Yunes JA, Neto GC, da Silva MJ, Arruda P, and Leite A (1998) The
Publications. molecular and functional characterization of an Opaque2 homologue gene from
Hartwell JL (1971) Plants used against cancer, a survey. Lloydia 34: 386425. Coix and a new classification of plant bZIP proteins. Plant Molecular Biology
Huang X, Qin J, and Lu S (2014) Kanglaite stimulates anticancer immune responses 36: 249263.
and inhibits HepG2 cell transplantation-induced tumor growth. Molecular Medicine Yamada H, Yanahira S, Kiyohara H, Cyong J-C, and Otsuka Y (1987) Characterization of
Reports 10: 21532159. anti complementary acidic heteroglycans from the seed of Coix lacryma jobi var ma
Jain SK and Banerjee DK (1974) Preliminary observations on the ethnobotany of the yuen. Phytochemistry 26: 32693276.
genus coix. Economic Botany 28: 3842. Yeung HC (1985) Handbook of Chinese Herbs and Formulas. Los Angeles: Institute of
Leite A, Yunes JA, Turcinelli SR, and Arruda P (1992) Cloning and characterization of a Chinese Medicine.
cDNA encoding a sulfur-rich coixin. Plant Molecular Biology 18: 171174. Zhan YP, Huang XE, Cao J, et al. (2012) Clinical safety and efficacy of KanglaiteW
Li SC (1596) Pen-tsao kang mu (Systematic Pharmacopoeia), China. (Coix Seed Oil) injection combined with chemotherapy in treating
Li J and Corke H (1999) Physicochemical properties of normal and waxy Jobs tears patients with gastric cancer. Asian Pacific Journal of Cancer Prevention
(Coix lachryma-jobi L.) starch. Cereal Chemistry 76: 413416. 13: 53195321.
Li XH, Huang YQ, Li JS, and Corke H (2001) Characterization of genetic variation and Zhang F and Yi Z (1994) Cytological studies of the genus Coix. Natural Science Journal
relationships among Coix germplasm accessions using RAPD markers. Genetic of Changsha Normal University of Water Resources and Electric Power 9(2):
Resources and Crop Evolution 48: 189194. 187191.
List PH and Horhammer L (1979) Hagers Handbuch der Pharmazeutischen Praxis.
Berlin: Springer, vols. 26.
Liu Y, Zhang W, Wang X-J, and Liu S (2014) Antitumor effect of KanglaiteW injection in
human pancreatic cancer xenografts. BMC Complementary and Alternative Relevant Websites
Medicine 14: 6, Article Number 228.
Moreau RA, Singh V, and Hicks KB (2001) Comparison of oil and phytosterol levels in http://www.fao.org The website of Food and Agriculture Organization of the United
germplasm accessions of corn, teosinte, and Jobs tears. Journal of Agricultural and Nations. Search for coix information within the site, such as: http://www.fao.org/ag/
Food Chemistry 49: 37933795. agp/AGPC/doc/Gbase/data/pf000205.htm.
Numata M, Yamamoto A, Moribayashi A, and Yamada H (1994) Antitumor components http://www.hort.purdue.edu The website of the center for new crop and plant products
isolated from the Chinese herbal medicine Coix lachryma-jobi. Planta Medica at Purdue University. Search for coix information within the site, such as: https://
60: 356359. www.hort.purdue.edu/newcrop/duke_energy/Coix_lacryma-jobi.html.
Schaaffhausen RV (1952) Adlay or Jobs tears: A cereal of potentially greater economic http://www.itmonline.org/articles/coix/coix.htm A useful article by Dr. Subhuti
importance. Economic Botany 6: 216227. Dharmananda on Coix: Food and Medicine.
Millet Pearl: Overview
JRN Taylor, University of Pretoria, Hatfield, South Africa
2016 Elsevier Ltd. All rights reserved.
Pearl millet is quantitatively the most important millet Pearl millets great advantage over other cereals is the fact that it
species. can be and is cultivated in areas of very low rainfall
Pearl millet is uniquely hardy and can be cultivated in hot (300500 mm per year). However, it requires evenly distributed
and dry conditions. rainfall during the growing season, and too much rain at flower-
Pearl millet grain has better nutritional value than most ing can cause a crop failure. Pearl millet can be cultivated in
other cereals. areas with very high temperatures (daytime temperatures above
Many traditional foods and beverages and specialty prod- 30 C), and it germinates well at soil temperatures of 2330 C.
ucts are produced from pearl millet. Its importance in food security in dry, marginal agricultural
Pearl millet is becoming widely used in commercial small- areas is immense. It is estimated that more than 90 million
scale food manufacture. people in the semiarid tropics depend on pearl millet food
Iron- and zinc-biofortified pearl millet has been developed and income. The major areas of pearl millet cultivation are
for improved nutrition. India and Western and Central Africa. It is also cultivated in
Eastern and Southern Africa. In Western and Central Africa,
there is a broad band of cultivation from Senegal in the west,
across the Sahel region. It is also cultivated in Sudan, South
Sudan, Eastern Africa, and Southern Africa where there is a
Learning Objectives
band of cultivation from Angola and northern Namibia, to
Zimbabwe and northern Southern Africa, to Mozambique.
To provide an introduction to the importance of pearl
There is also some production in Brazil and the United States.
millet as a staple food crop.
Figures for pearl millet production are only approximate as
To provide brief and accurate information about pearl grain
indicated by the wide reported ranges. FAOSTAT data do not
structure, nutritional composition, and traditional food
distinguish between the different types of millets. Pearl millet
and beverage uses.
is generally a subsistence grain crop and hence not commer-
To provide brief and interesting information on the tech-
cially traded in quantity. Thus, precise figures are not always
nologies used to produce traditional foods and beverages
available. Pearl millet accounts for about half of the world
from pearl millet.
millet crop, which is at least 29 Mt per year. The major pearl
To sensitize the reader to the potential of pearl millet as a
millet grain-producing countries are India, 8.3 Mt per year;
global staple food crop in the age of climate change.
Western and Central Africa, 14.1 Mt, with Niger being the
major producer, followed by Mali, Burkina Faso, Nigeria, and
Senegal; and Eastern and Southern Africa, 1.8 Mt. It is also
increasingly grown as a forage crop and as an ornamental grass.
Introduction Relative to other major cereal grains, yields are in general very
low, on average <1 t ha1. This is because pearl millet is grown in
Pearl millet, commonly known as bulrush millet (Pennisetum areas of low rainfall and high temperature, often on light, well-
glaucum (L.) R. Br.), also classified as P. typhoides, P. ameri- drained soils. Generally, traditional farming practices are used
canum, or P. spicatum, is a cultivated, small-grain, C4 tropical with very low inputs, that is, no biocides or inorganic fertilizers
cereal grass. Vernacular names include bajra (India), gero and little application of organic fertilizer, and use of traditional
(Nigeria, Hausa language), hegni (Niger, Djerma language), landraces. Pearl millet has, however, much greater potential. In
sanyo (Mali), dukhon (Sudan, Arabic), and mahangu areas, especially India, where new varieties and hybrids are being
(Namibia). It is diploid (2n 2x 14) with a large genome widely cultivated, with some irrigation and using higher-input
(2450 Mbp). Pearl millet is quantitatively the most important agriculture, yields can be well in excess of 2 t ha1.
millet, with a world annual production of some 1525 Postharvest practices also remain backward in many areas, for
million tonnes (Mt). It is cultivated mainly in the semiarid example, manual threshing and winnowing of the grain
tropics, almost exclusively by subsistence and small-scale (Figure 1). Pearl millet can also suffer severe storage losses,
commercial farmers. particularly as a result of insects such as the rice moth (Corcyra
This article will examine pearl millet production and agro- cephalonica). The larvae contaminate the grain by producing
nomic issues, grain structure, chemistry and nutritional webbing (silk) and consume the protein- and fat-rich grain
aspects, primary processing technologies, important foods germs. Control is traditionally affected by putting wood ash in
and beverages produced from pearl millet, pearl millets future the storage bin. Today, application of the fumigant phostoxin
prospects, and research and development challenges. (active ingredient phosphine gas) is becoming more widespread.
Style
Pericarp
Seed coat
Aleurone layer
Corneous
endosperm
Peripheral
endosperm
Floury
endosperm
Scutellar
Epithelium
Scutellum
Embryonic
axis
Hilum
Figure 3 Diagrammatic longitudinal section through a pearl millet
grain. Courtesy of Mr S C Barrion.
Proximate composition (db) (g 100 g1)a Minerals (db) (mg 100 g1)a Essential amino acids (g 100 g protein1)b
This is due to its high fat content, a consequence of the rela- contents of glutamine (2324 g 100 g protein1), alanine (89 g
tively large germ. 100 g protein1), and leucine ( 1012 g 100 g protein1), but
relatively low in proline ( 8 g 100 g1 protein1) and cystine
Carbohydrates and methionine ( 1 g 100 g protein1 each). Sodium dodecyl
As with all other cereals, the predominant carbohydrate in sulfate polyacrylamide gel electrophoresis under reducing condi-
pearl millet is starch. The starch content, typically 7172 g tions shows a major band of Mr 22 kDa and minor bands
100 g1 (db), is perhaps slightly lower than most other cereals, of Mr 20 and 10 kDa. Research indicates that, unlike the situa-
due to the fact that the endosperm is proportionally smaller tion with sorghum, wet cooking, as occurs in porridge making,
than other cereals. The percentage amylose in pearl millet does not substantially reduce the digestibility of pearl millet
starch is in the range 17.021.5%, which is relatively low protein. This is probably a consequence of the somewhat lower
for normal (nonwaxy) starches. Waxy (essentially 100% amy- percentage of prolamin proteins and the low content of cystine.
lopectin) pearl millet has not been reported. The starch gelati- However, like the other tropical cereals, the prolamins of pearl
nization temperature is in the range 6169 C, typical of millet differ so much from those of wheat that it is considered as
tropical cereal starches. Soluble sugars in pearl millet grain, as being gluten-free.
in all sound cereal grains, are low, in the range 1.42.8 The amino acid composition of pearl millet protein
100 g1, with the major sugars being sucrose and raffinose. (Table 1) is characterized by a relatively high content (com-
The dietary fiber content of pearl millet grain is 89 100 g1 pared with most cereal grains) of the essential amino acid
(db). The dietary fiber of pearl millet, as with sorghum, is lysine, in the range 2.83.2 g 100 g protein1. This is a conse-
mainly of the insoluble (water-inextractable) type. quence of the relatively high proportion of lysine-rich albumin
and globulins, due to pearl millets large germ. However, the
lysine content of pearl millet does not meet the requirements
Proteins and amino acids
of infants, but it is close to that of young school age children.
Pearl millet grain protein content, typically 14.5 g 100 g1
This emphasizes the importance of supplementation of cereals
(db), is high for cereal grains. This is because the grain has a
in the diet with meat, dairy, or legumes, as sources of lysine.
large germ. This, in turn, also affects the balance of Osborne
Pearl millet protein is, as shown, a satisfactory source of the
protein solubility groups and the overall amino acid composi-
other indispensible amino acids.
tion of the pearl millet protein. Considering the protein frac-
tions, the prolamin (aqueous alcohol-soluble) proteins,
3141% of total protein, are, as in most other cereals, the Fats
major fraction in pearl millet. However, this level is lower The fat content of pearl millet grain, typically 5.1 g 100 g1
than most other cereals as prolamins are endosperm-specific (db), is high compared with all other cereal grains, except
proteins. In contrast, the amount of albumin and globulin maize. In the case of both pearl millet and maize, the high fat
(saline-soluble) proteins, 2526% of total proteins, is proba- content is due to the large germ. As with all other cereals, the
bly somewhat higher than in most other cereals. major fatty acids are linoleic acid (C18:2), typically 4345% of
The prolamin proteins of pearl millet are called pennisetins the total; oleic acid (C18:1), 2627% of the total; and palmitic
and are similar to the prolamins of maize and sorghum, with high acid (C16:0), 2021% of the total. The high content of fat and
THE CEREAL GRAINS | Millet Pearl: Overview 193
in particular of polyunsaturated fatty acids means that whole- Pearl millet, like all grains, contains the phosphorus-
grain pearl millet, when milled, is highly subject to deteriora- containing compound phytate, myo-inositol hexaphosphate.
tion through oxidative rancidity. Phytate acts as the main phosphorus store in seeds. In grain
foods, it has the undesirable property of binding multivalent
Vitamins metal cations such as iron, zinc, and calcium and rendering
Pearl millet, like other cereal grains, is an important source of B them biologically unavailable. The level of phytate in pearl
vitamins (Table 1). These are concentrated in the aleurone millet grain, in the range 350800 mg 100 g1, is typical of
layer and germ. Hence, their content can be adversely affected cereal grains (Table 1). Phytate in pearl millet, as in other
by milling. Niacin occurs partially bound to carbohydrate. To cereal grains, is located in the aleurone layer and germ.
make it fully available, the flour has to be treated with alkali. Hence, dehulling the grain during milling substantially
This is done by treatment with wood ash in the preparation of reduces the level of phytate (but also the minerals) in flour.
the Western African porridge called to. Because of its high fat Malting and fermentation specifically reduce the level of phy-
content, pearl millet is a good source of tocopherols (vitamin E), tate by enzymatically degrading it, freeing the minerals.
located mainly in the germ. It should be noted that contrary to what is written in
some texts, pearl millet, unlike some varieties of sorghum
and finger millet, does not contain condensed tannins
Minerals
(proanthocyanidins).
Pearl millet grain, like other grains, is a good source of most
dietary minerals (Table 1), with the exception of calcium.
Quantitatively, the highest contents are of phosphorus and
potassium. However, the availability of the former and of the Primary Processing
multivalent metal ions is negatively affected by the presence of Milling
phytate (see the succeeding text). The minerals are concen-
trated in the pericarp, aleurone layer, and germ. Hence, their Pearl millet is milled into flour for porridge and bread making
content can be adversely affected by milling. and into semolina (coarse grits) for couscous making. The
milling process used in northern Namibia is shown in Figure 4
and will be described in detail as it has some unique features.
Phytochemicals
With regard to phenolic phytochemicals, when compared with
Dehulling
other millets, pearl millet seems to be very low in soluble and
To improve the palatability and storage quality of the flour, the
insoluble forms of hydroxybenzoic acids and flavonoids but
bran (pericarp and germ) is generally removed from the grain
contains reasonable levels of both soluble and bound hydro-
first. This process of bran removal is referred to as dehulling.
xycinnamic acids. However, the available data are very limited,
and it is probable that levels vary greatly with genotype and
cultivation environment. Cleaned pearl millet
grain
Antinutrients Decorticate
(PRL dehuller)
There is evidence that pearl millet contains goitrogens. It
appears that the goitrogens are primarily phenolic flavonoid-
type compounds, the C-glycosyl flavones: vitexin, glucosyl Lactic acid ferment
vitexin, and glucosyl orientin. Other phenolic compounds, (soak overnight)
such as phloroglucinol, resorcinol, and p-hydroxybenzoic
acid, could also be involved. These compounds apparently
inhibit the deiodination of the hormone thyroxine to its Partially dry
(solar drying)
more active form triiodothyronine. They are concentrated in
the outer layers of the grain and are considerably reduced when
the grain is dehulled during milling. The nutritional signifi- Mill
cance of the goitrogens in pearl millet should not be over- (hammer mill)
stated. Although some rural people, for example, in Sudan/
South Sudan and areas of India, who consume pearl millet as a
Fine flour
staple have been found to suffer from goiter, it is probable that
this is because their diet is very restricted and hence deficient in
iodine. Fully dry
The C-glycosyl flavones contribute to the brown/gray color (solar dry)
of pearl millet and also appear to be responsible for the char-
acteristic musty flavor of damp pearl millet flour, which is Pack flour
probably of greater significance. Some people describe the (paper or woven
flavor as being mousy or mouse-dropping like. In pearl sacks)
millet foods, such as porridges, it can be rather disagreeable
to those not familiar with the food. Figure 4 Pearl millet milling process used in northern Namibia.
194 THE CEREAL GRAINS | Millet Pearl: Overview
Figure 5 Small-scale commercial pearl millet mill in northern Namibia. Solar drying
Foreground hammer mill; background PRL dehuller. The dehulled, steeped grain is partially dried before milling,
and after milling, the flour must be fully dried to a shelf-stable
moisture content of 10%. Solar drying is used for both steps.
Strictly speaking, decortication is the correct term since the The grain or flour is dried on polythene sheets, either directly
pearl millet grain does not have a hull. on a concrete floor or on long tables made of wooden slats.
Figure 5 shows the equipment used in a small-scale com-
mercial pearl millet mill in Namibia. In the background is a Milling
Prairie Research Laboratory (PRL) abrasive-type dehuller. The partially dried, dehulled steeped grain is milled into a fine
Today, abrasive-type hullers are manufactured in a number of flour using a powerful hammer mill. The high moisture con-
countries in Southern and Western Africa. The PRL dehuller tent (3040%) of the grain necessitates relatively powerful
comprises an axle with 12 Carborundum disks mounted on milling. Thus, the process is actually semiwet milling. This
it, within a cylindrical box. The axle revolves at high speed and apparently rather inefficient process seems to be necessary in
the disks abrade off the bran from the grain. A suction fan order to mill the grain into fine flour. In Western Africa, disk
removes the bran. Alternatively, pearl millet dehullers in West- mills are generally used to produce semolina for couscous
ern Africa incorporate revolving brushes and screens to remove making.
and separate the loose bran from the dehulled grain.
The amount of dehulling is determined by the duration of
Malting
the dehulling process. Anything between 10% and 30% of the
grain is removed, depending on the desired final color and fat In sub-Saharan Africa, pearl millet is malted to produce malt
content of the flour. Dehulling has the effect of reducing the for brewing traditional opaque beer and nonalcoholic bever-
fiber, fat, and protein contents of the grain due to the removal ages. Pearl millet malting is almost exclusively done in the
of pericarp and germ material. The quality of the protein will home, although one large-scale commercial malting operation
also be adversely affected as the quantity of the lysine-rich germ in Zimbabwe malts pearl millet together with sorghum.
proteins is reduced. The starch content is increased slightly. Figure 6 shows pearl millet malt prior to drying. It can be
seen that there is extensive root and shoot growth. The opti-
mum temperature conditions of pearl millet malting are in the
Steeping range 2428 C, identical to that for sorghum. The malting
In the Namibian process, the dehulled grain is steeped in process involves three steps: steeping, germination, and drying.
ambient temperature water overnight. This steeping process is The steeping time for pearl millet is short, up to 8 h, as the
actually a lactic acid fermentation. A lactic acid bacterial cul- grain germinates rapidly. Germination is allowed to progress
ture, maintained by the process of back slopping, is used to for 45 days. Like sorghum, pearl millet must be watered
inoculate the steep water. The culture comprises a portion of during germination. Drying is at ambient temperature or up
the previous steep, with a high load of suitable lactic acid to 50 C in order to conserve malt enzyme activity.
bacteria. The purpose of the lactic acid steep seems to be Pearl millet malt quality seems to be somewhat higher than
severalfold. It lightens and brightens the flour. The low pH of that of sorghum malt. Diastatic power (total amylase activity)
the steep water is responsible for lightening the color and is similar, 3050 sorghum diastatic units g1. However, the
partially leaching out the brown/gray polyphenolic pigments activity of the b-amylase component is higher. b-Amylase is
of the grain. Goitrogens may also be leached out of the grain. the enzyme that produces the fermentable sugar maltose. The
However, it is probable that water-soluble proteins, vitamins, higher b-amylase activity indicates that pearl millet malt could
and minerals are lost in the steeping process. The steep softens have potential as partial substitute for barley malt for brewing
the grain, facilitating its reduction into a fine flour. The result- lager beer. Free amino nitrogen and extract are also higher in
ing flour has an acidic flavor, which consumers prefer. pearl millet malt than sorghum malt. Of particular significance
THE CEREAL GRAINS | Millet Pearl: Overview 195
is the fact that malting almost completely eliminates the mousy Western African countries such as Mali and Burkina Faso is
flavor associated with pearl millet flour. Some nutritional often produced from highly refined flour giving it a gel-like
improvements also occur, in particular improvement in carbo- consistency. It may be cooked with tamarind or lemon or with
hydrate and protein availability. However, against this, there is wood ash or potash, giving it acidic or alkaline taste, respec-
a substantial overall loss in mass of grain, 1015%, due to tively. These porridge products are traditionally eaten cool or
respiration during germination. cold with the hand and served with a vegetable relish.
Doughs and gruels made from pearl millet are invariably
lactic acid bacteria-fermented. The fermentation used in fura
dough making has been reported to produce bacteriocins with
Food and Beverage Production
inhibitory activity against a wide range of pathogens.
Worldwide, pearl millet is processed into many different tradi-
tional foods and beverages. Foods include rice and flatbreads Couscous
(roti) in India, couscous in Mali and Senegal, doughs such as In the Sahel region of Western Africa, couscous made from
fura in Ghana, thick porridges such as to in Western Africa pearl millet and sorghum is a very popular food (Figure 7).
and bogobe in Southern Africa, thin fermented porridges such Couscous making involves processing coarse endosperm par-
as uji in Eastern Africa, and gruels like ogi in Nigeria. Bever- ticles (semolina) in repeated steps of agglomerating, steaming,
ages can be nonalcoholic like oshikundu in Namibia or alco- and sifting to ultimately produce cooked granular particles of
holic like the opaque beer of Southern Africa. uniform size (Figure 8). The steaming gelatinizes the starch on
the surface of the particles so that they stick together. The
couscous is often served with yogurt, the product being called
Foods thiakri.
Porridges, doughs, and gruels
In sub-Saharan Africa, pearl millet is most frequently con- Flatbreads
sumed as a porridge. A traditional Pedi (Limpopo Province, In India, pearl millet is generally consumed as a crisp flatbread,
South Africa) recipe to make bogobe bia bupi bja leotsa called roti or chapatti. Figure 9 shows a flowchart for making
(literally porridge of pearl millet) involves cooking 1 kg of roti. A problem in making breads from non-gluten-containing
whole-grain pearl millet meal in 1.9 l water. The porridge has grains such as pearl millet is that the product tends to break
greenish-brown color and a thick, smooth viscous consistency. into pieces. It is notable that in the milling process, the flour is
Because of the high ratio of starch to water, the porridge gels on ground to fine particle size, 43% having a particle size
cooling. The flavor is musty, sweet, and bitter. The meal ration <75 mm. This increases the surface area of the flour particles
size is 1 kg, which provides some 2205 kJ, 2025% of an and damages the starch granules, increasing their water absorp-
adults recommended dietary allowance. To porridge from tion. Both factors increase the cohesiveness of the dough.
Mill
(disk or plate mill)
Sieve
(850 mm screen)
Flour (50 g)
+
Warm water (45 ml)
Roti
Variations on the roti-making process, which also alleviate the Figure 9 Roti-making process.
problem, can involve cooking part of the flour in water and
mixing it together with the uncooked flour. Cooking gelati-
nizes the starch making it into a binder, as with couscous, Nonalcoholic
which improves the cohesiveness and elasticity of the dough. A very popular nonalcoholic fermented beverage in Namibia is
Alternatively, pearl millet flour can be composited with flours oshikundu made from pearl millet and sorghum malt flour.
such as gram (mung bean) or other legume flours that give Some 200 g of pearl millet flour are thoroughly mixed into
better cohesiveness or wheat flour that gives elasticity. 500 ml of boiling water. Then, 100 g of sorghum malt flour
Because of the absence of gluten in pearl millet, breads is added. Sorghum malt provides amylase enzymes and lactic
made from pearl millet flour are unleavened. However, in the acid bacteria. a-Amylase thins the porridge into gruel by hydro-
roti-making process, as shown in Figure 9, the flatbread is lyzing the starch into dextrins, and b-amylase further hydro-
exposed on one side to high heat, which puffs it giving the lyzes the dextrins into maltose. The thinning and hydrolysis
bread some leavened texture. Puffing takes place as result of actions of the amylases improve food palatability and carbo-
the fact that the moistening water added during baking is very hydrate availability. The lactic acid bacteria use the fermentable
rapidly turned into steam. The resulting increase in volume sugars for respiration, producing lactic acid and other flavor
creates an air sac as the steam escapes. Roti is served with a compounds. After the mixture has cooled to room tempera-
small amount of hot pickle, dhal, or vegetable sauce. ture, 11.5 l of cold water is mixed in. The container is closed
and the mixture is allowed to ferment overnight before con-
sumption. Oshikundu is greenish brown in color and has the
consistency of drinking yogurt. It is slightly effervescent with a
Beverages
buttery sour taste. A lactic acid content of 0.6% has been
Throughout Africa, sorghum, maize, and millets are used to measured.
produce both nonalcoholic and alcoholic beverages. Invari-
ably, these beverages have undergone a lactic acid fermenta-
tion, giving them a refreshing sour taste. The low pH of the Opaque beer
beverages renders them free from foodborne pathogenic bac- In several southern African countries, pearl millet malt is used
teria and helps protect them against microbial spoilage. to brew traditional, opaque-type beer. Figure 10 shows a
THE CEREAL GRAINS | Millet Pearl: Overview 197
flowchart for a Pedi recipe for making bjalwa bja leotsa (lit-
Exercises for Revision
erally beer of pearl millet). The process has some interesting
features. It is noteworthy that only malt is used in the recipe. In Draw the structure of the pearl millet grain, naming the
grain tissues.
contrast to brewing traditional opaque sorghum beer, no
unmalted grain adjunct is used. In fact, part of the malt is Explain the major nutritional advantages of pearl millet
compared with other cereals.
used as adjunct by cooking it after the first fermentation pro-
cess. The first fermentation process is probably primarily a Name the antinutrients present in pearl millet and explain
their nutritional significance.
lactic acid fermentation, but some fermentable sugar produc-
tion will also take place. As there is no unmalted adjunct in the Draw a flow diagram showing the steps in producing a
traditional pearl millet food of your own choice.
recipe, it means that there is a proportionally higher ratio of
amylase enzymes to starch, producing more fermentable Draw a flow diagram showing the steps in producing a
traditional pearl millet beverage of your own choice.
sugars. Coupled with the fact that pearl millet malt intrinsically
contains a higher ratio of b-amylase, this probably accounts for Explain the science of how textured dry food products can
be produced from pearl millet, despite the fact that it does
the fact the pearl millet beer is claimed to be more intoxicating
not contain gluten.
than sorghum beer. Typical of traditional recipes, more than
one process takes place during a particular step. This is unlike
modern food and beverage manufacturing processes where
there is a system of unit operations. In the mashing and fer- Exercises for Readers to Explore the Topic Further
mentation stage, gelatinized starch is hydrolyzed by the malt
amylase enzymes into fermentable sugars, which are fermented There is value in biofortification of pearl millet with other
by both yeast and lactic acid bacteria to produce ethanol, essential micronutrients than iron and zinc, for example,
carbon dioxide, and lactic acid as the main products. The provitamin A.
beer is described as turbid, greenish brown in color, with a How can the yields of pearl millet in Africa be substantially
milk like consistency. The flavor is musty and bitter sour, with improved to help eliminate food insecurity?
a lactic acid content of 1.8%. Pearl millet beer seems to be The phenolic phytochemicals in a pearl millet need to be far
much thinner and more acidic than sorghum beer. better characterized across a wide range of genotypes and
198 THE CEREAL GRAINS | Millet Pearl: Overview
individual functional and antinutrient properties Germplasm Resources Information Network (GRIN). GRIN Taxonomy for Plants. http://
determined. www.ars-grin.gov/ (assessed 14.02.01).
A more efficient way to mill the pearl millet grain is ICRISAT EXPLOREit. Pearl millet. http://exploreit.icrisat.org/ (accessed 14.01.08).
ICRISAT and FAO (1996) The World Sorghum and Millet Economies. Patancheru, India/
required in order to reduce losses and eliminate the Rome: International Crops Research Institute for the Semi-Arid Tropics/Food and
mousy aroma in pearl millet foods. Agriculture Organization of the United Nations.
The real extent of the goitrogen problem in pearl millet Klopfenstein CF and Hoseney RC (1995) Nutritional properties of sorghum and the
must be established. millets. In: Dendy DAV (ed.) Sorghum and Millets: Chemistry and Technology,
pp. 124168. St Paul, MN: American Association of Cereal Chemists.
Murty DS and Kumar KA (1995) Traditional uses of sorghum and millets.
In: Dendy DAV (ed.) Sorghum and Millets: Chemistry and Technology,
See also: Beverages from Grains: Fermentation: Foods and pp. 185222. St Paul, MN: American Association of Cereal Chemists.
Nonalcoholic Beverages; The Basics: Grain and Plant Morphology of National Research Council (1996) Lost Crops of Africa, vol. 1. Washington, DC:
Cereals and how characters can be used to identify varieties; Grain: National Academies Press.
Obilana AB and Manyasa E (2002) Millets. In: Belton PS and Taylor JRN (eds.)
Morphology of Internal Structure; Taxonomic Classification of Grain Pseudocereals and Less Common Cereals, pp. 177217. Berlin: Springer.
Species; The Cereal Grains: Millet Minor: Overview; Teff: Overview. Quin PJ (1959) Food and Feeding Habits of the Pedi. Johannesburg: Witwatersrand
University Press.
Rai KN, Govindaraj M, and Rao AS (2012) Genetic enhancement of grain iron and zinc
content in pearl millet. Quality Assurance & Safety of Crops and Food 4: 119125.
Serna-Saldivar S and Rooney LW (1995) Structure and chemistry of sorghum and
Further Reading millets. In: Dendy DAV (ed.) Sorghum and Millets: Chemistry and Technology,
pp. 69124. St Paul, MN: American Association of Cereal Chemists.
Abdalla AA, El Tinay AH, Mohamed BE, and Abdalla AH (1998) Proximate composition, Taylor JRN and Emmambux MN (2008a) Millets. In: Arendt E and Dal Bello F (eds.)
starch, phytate and mineral contents of 10 pearl millet genotypes. Food Chemistry Gluten Free Cereal Products and Beverages, pp. 119148. London: Elsevier.
63: 243246. Taylor JRN and Emmambux MN (2008b) Products containing other speciality grains:
Chandkara A and Shahidi F (2011) Determination of antioxidant activity in free and Sorghum, the millets and pseudocereals. In: Hamaker BR (ed.) Technology of
hydrolysed fractions of millet grains and characterization of their phenolic profiles Functional Cereal Products, pp. 281335. Abington, UK: Woodhead Publishing.
by HPLC-ESI-MSn. Journal of Functional Foods 3: 144158. Taylor JRN, Barrion SC, and Rooney LW (2010) Pearl millet New developments in an
Emmambux MN and Taylor JRN (2013) Morphology, physical, chemical and functional ancient food grain. Cereal Foods World 55: 1619.
properties of starches from cereals, legumes and tubers cultivated in Africa: A World Health Organization (2002) Joint FAO/WHO/UNU expert consultation on protein
review. Starch/Starke 65: 715729. and amino acid requirements in human nutrition, WHO Technical Report Series; No.
FAOSTAT. Production: Crops. http://faostat.fao.org/ (access 14.02.01). 935. Geneva: WHO.
Millet Minor: Overview
GK Chandi, University of Guelph, Guelph, ON, Canada
GA Annor, University of Ghana, Accra, Ghana
2016 Elsevier Ltd. All rights reserved.
Topic Highlights 3. Foxtail or Italian millet (Setaria italica): This millet species is
a native of China and ranks first in its global production. Its
Minor millets are domesticated and used as food across cultivation in Europe dates to about 2000 BP. Although
different parts of the world. found in the tropics, it is adapted to temperate regions. It
Millets are generally used for pancakes, flatbreads, porridge, has a broad range of days of maturity (70120 days). Green
fermented beverages, boiled grains, gruel, etc. foxtail millet (Setaria viridis) is the wild antecedent of foxtail
Minor millets are decorticated before grinding them into millet and has very limited food usage.
coarse or fine flours or grits of various particle sizes. 4. Kodo millet (Paspalum scrobiculatum): This millet species is
Hypoglycemic properties of millets may be attributed to commonly harvested as a wild cereal in tropical Africa and
various factors such as phenolic compounds, soluble fibers, the Deccan region of India. It was introduced in India almost
and starchlipidprotein interactions. 3000 years ago. It is self-pollinating, and its florets generally
Owing to their potential therapeutic properties and resistance remain closed during the flowering period. It matures late
to severe weather conditions, minor millets have a great compared to other cereals. Sometimes, due to fungal infec-
potential to address the health and food security issues. tions after the rain, kodo millet grains become poisonous.
5. Little millet (Panicum sumatrense): This millet species grows
Learning Objectives up to an altitude of 2100 m in India. It is commonly grown
in India and cooked like rice. It is primarily a self-
pollinating crop with about 3.5% cross-pollination. It
Understand the processing and utilization of minor millets.
matures between 2.5 and 5 months.
Identify the key attributes responsible for nutritional bene-
fits of minor millets. Besides the aforementioned millets, there are other minor
millets that are cultivated in restricted geographic regions and
have no importance in world agriculture trade. Some of these
Introduction millets include the following:
Figure 1 Minor millet crops and grain. (a) Foxtail millet, (b) little millet, (c) kodo millet, (d) common millet, (e) barnyard millet, and (f) finger millet.
Reproduced from Balasubramanian S (2013) Processing of millets. In: National Seminar on Recent Advances in Processing, Utilization and
Nutritional Impact of Small Millets. Madurai Symposium, Thamukkam Grounds, Madurai, 13 September 2013.
Finger millet (Eleusine coracana) Uganda or neighboring Eastern Africa and in Asia (India and Nepal)
region
Proso (common) millet (Panicum Central and Eastern Asia Ukraine, Kazakhstan, the United States, Argentina, and Australia
miliaceum)
Foxtail millet (Setaria italica) Eastern Asia (China) China, India, Indonesia, the Korean Peninsula, and some parts of southern
Europe
Barnyard millet (Echinochloa utilis) Japan Tropics and subtropics of India
Little millet (Panicum sumatrense) Southeast Asia India, Nepal, Pakistan, Sri Lanka, eastern Indonesia, and western Myanmar
Kodo millet (Paspalum scrobiculatum) India Wild cereal in Western Africa and India
Source: FAO.org.
suited for intercropping cultivation practices with other grain consumption of wheat-based products. With total carbohy-
crops in dry lands under unreliable rainfall. drate and protein contents of about 70% and 12%, respec-
tively, there is no doubt that minor millets contribute
significantly to the energy and protein needs of populations
in regions where they are principal source of food.
Nutritional Importance of Millets
Figure 2 Minor millet grains. Reproduced from Balasubramanian S (2013) Processing of millets. In: National Seminar on Recent Advances in
Processing, Utilization and Nutritional Impact of Small Millets. Madurai Symposium, Thamukkam Grounds, Madurai, 13 September 2013.
Plant height
Millet (cm) Seed color Seed shape Seed size (mm) Thousand kernel weight (g, 12% mb)
Finger millet 40100 Yellow, white, red, brown, violet Globose 12 2.6
Proso millet 30100 White cream, yellow, orange, red, black, Oval 32 4.77.2
to brown
Foxtail millet 0.5200 Yellow, pale yellow Elliptical 23 1.86
Barnyard 50100 Yellow, red, brown Globose 1.3 4.17
millet
Little millet 3090 White, yellow, red, brown Elliptical 2.3 1.6
Kodo millet 55 Light red to dark gray
Source: Upadhyaya HD, Dwivedi SL, Singh SK, Singh S, Vetriventhan M, and Sharma S (2014) Forming core collections in Barnyard, Kodo, and Little Millets using morph agronomic
descriptors. Crop Science 54: 26732682 and FAO.org.
202 THE CEREAL GRAINS | Millet Minor: Overview
Parameter Finger millet Proso millet Foxtail millet Little millet Kodo millet Barnyard millet
Source: Shobana S, Krishnaswamy K, Sudha V, et al. (2012) Finger millet (ragi, Eleusine coracana L.): A review of its nutritional properties, processing, and plausible health benefits.
Advances in Food and Nutrition Research 69: 139; Gopalan C, Rama Sastri BV, and Balasubramanian SC (2009) Nutritive Value of Indian Foods. Hyderabad, India: National Institute of
Nutrition, Indian Council of Medical Research. Geervani P and Eggum BO (1989) Nutrient composition and protein quality of minor millets. Plant Foods for Human Nutrition 39(2): 201208.
calcium levels of about 344 mg/100 g, finger millet arguably choline. Finger and foxtail millets have been reported to con-
has the highest calcium content when compared with other tain 42 mg/100 g and 32 mg/100 g carotenoid, respectively
cereals. Using proton-induced x-ray emission spectrometry, (Table 3).
Kruger et al. confirmed the high calcium content of finger
millet and also showed that calcium was distributed through-
Fat
out the endosperm where phytate levels were low. Phosphorus,
iron, magnesium, potassium, sulfur, and zinc, for example, are Fat in minor millets, as in other cereals, is mainly composed of
all found in considerable amounts in minor millets. In terms of unsaturated fatty acids. Oleic and linoleic acids are the main
vitamins, some minor millets have been found to have low unsaturated fatty acids in minor millets, with palmitic acid as
amounts of carotenoids thiamine, riboflavin, niacin, and the main saturated fatty acid. Depending on the species of
THE CEREAL GRAINS | Millet Minor: Overview 203
millet palmitic, oleic and linoleic acids can range from 6% to increase in the soluble fiber content has special nutritional
23%, 20% to 66%, and 22% to 65%, respectively, of the total significance due to its physiological advantages in terms of
lipid content of the millet flours. Though these unsaturated hypoglycemic and hypocholesterolemic characteristics.
fatty acids may have health benefits, they are also responsible
for the rancidity that is normally observed in stored millet
StarchProtein Interactions
flours with relatively high fat contents.
Some researchers have attributed the hypoglycemic property of
finger millet to their rigid starch granule architecture. This
Fiber
rigidity slows down the rate at which the millet starch granules
The dietary fiber contents of minor millets also confer some are hydrolyzed by starch digestive enzymes. The effects of
health benefits when they are consumed. Total dietary fiber protein-encapsulated granule matrix have been suggested as
values of about 27%, 20%, and 30% of grain weight have been part of the reason for the hypoglycemic property of kodo
reported for barnyard, kodo, and little millets, respectively, out millet. Some protein fractions glue protein bodies into a matrix
of which 0.59%, 1.94%, and 1.33% are soluble fiber, surrounding starch granules, which then acts as a barrier to
respectively. amylases. The removal of protein from kodo millet flour
clearly results in a significant increase in its starch hydrolysis.
Amino Acids
StarchLipid Interactions
Just like most of the cereals, the minor millets are deficient in
lysine, which is usually complemented when mixed with The presence of starchlipid complexes, particularly lipid com-
legumes. plexes with millet amylose, also influences the susceptibility of
millet starch to enzymatic degradation. Annor et al. reported
significant effects of starchlipid complexes on the starch
Phytochemicals
hydrolysis rates of millets. Foxtail, little, barnyard, kodo, and
The pericarp of minor millets is rich in several phenolic proso millets still contain a considerable amount of lipids even
phytochemicals, which are associated with various health ben- after decortication. The residual lipids in the endosperm are
efits. The inhibitory effects of finger millet phenolic com- still sufficient to result in significant reductions in starch hydro-
pounds on a-glucosidase and pancreatic a-amylase have been lysis rates of the decorticated millet flour.
mentioned. It is thought that these millet phenolic compounds
noncompetitively inhibit carbohydrate-hydrolyzing enzymes.
Fatty Acid Profile
Millet phenolic compounds have also been suggested to delay
or interfere with the absorption of dietary carbohydrates in the In addition to starchlipid complexes influencing the glycemic
small intestine resulting in the suppression of postprandial properties of millets, the type of lipid and their amounts pre-
glucose spikes. A comprehensive review of phytochemicals in sent in millets also significantly contribute to their
minor millets and their health benefits has been reported by hypoglycemic property. The main fatty acids present in millets
Taylor et al. and Taylor and Duodu. in no particular order are palmitic, oleic, and linoleic acids.
These three fatty acids form more than 80% of the total fatty
acids present in millets. These fatty acids form complexes with
Hypoglycemic Properties millet starches and thereby reduce the rate at which the result-
ing starchlipid complexes are hydrolyzed by digestive
The hypoglycemic property of millets and its importance in the enzymes. The unsaturated fatty acids form more complexes
management of type II diabetes were reported as far back as than the saturated fatty acids. With regard to their effects on
1957. Since then, several other scientists have also reported on starch hydrolysis rates, all the fatty acids significantly reduce
the lowering effects the regular consumption of minor millets the millet starch hydrolysis rates and their expected glycemic
has on postprandial blood glucose levels. It will also be of index. Starcholeic acid complex are the most stable against
interest to understand the factors that contribute to the hypo- amylolysis. The amounts of these fatty acids present in the
glycemic property of millets to help maintain this property respective millet flour samples also significantly influence the
during processing and also to develop hypoglycemic food expected glycemic index of millet flours. The greater the quan-
products. Some attempts have been made in this regard. tity of a particular fatty acid that is present in the millet flour,
the more it reduces the starch hydrolysis rates and their result-
ing expected glycemic index. It is therefore important to retain
Role of Fiber
the fatty acids present in millet flours when they are being
Dietary fiber in millets has been mentioned as one of the processed to utilize their beneficial hypoglycemic property.
reasons for its hypoglycemic property. With millets containing Apart from hypoglycemic benefits, the regular consump-
about 2% soluble and 30% insoluble fiber contents, their role tion of minor millets has been associated to a myriad of health
in hypoglycemic property of millets cannot be overlooked. benefits. Germinated finger millet flour has been reported to
Fiber reduces the glycemic response of foods by increasing improve hemoglobin status of school children in Tanzania and
their viscosity, especially the soluble fiber. A considerable also have antimicrobial, antifungal, antioxidant free radical-
increase in soluble fiber content of finger millet is observed scavenging, antiulcerative, cholesterol-lowering, and wound-
after decortication, though total dietary fiber decreases. This healing properties.
204 THE CEREAL GRAINS | Millet Minor: Overview
Storage of Grains faster cooking of the grain. For food purposes, postharvest
processing of millets depends on the cultural preferences and
Minor millets are important food crops of a large group of traditions.
people living in rural, tribal, and hilly areas across the globe.
About 90% of millets produced are used as food, and the rest of
it is used as fodder or for medicinal purposes. The millet grains Traditional Processing
are stored either as unthreshed along with their ear heads or as
threshed and clean grains. The unthreshed grains are normally For the small-headed grains, husk (hull) is usually removed by
heaped on the ground and covered with straw. The threshed hand pounding and less often by mechanical dehusking or
and clean grains are packed in jute sacks and stacked in piles on polishing units. Grain is sometimes moistened before pound-
the floor or wooden platforms. Some farmers also utilize silos ing. This is done by adding about 10% water to loosen the
or bins to store grains when needed. The larger storage struc- fibrous bran and also to separate the germ and the endosperm,
tures are built with metal, wood, stone, or bamboo. In the if desired. The traditional processing of minor millets involves,
absence of silos, the grains are stored in pits lined with straw. in no particular order, dehulling, winnowing, washing, drying,
Once the pit is full, the grains are covered with straw and soil or milling, and sieving. Local small-scale mills are usually based
dung. Clay pots are also sometimes used to store the millet on crude stone mortar and pestle or one horizontal hand
grains. Occasionally, grains are exposed to sunshine to dry and millstone rotating on another. Both types of mills deliver
also to prevent them from insect infestation. whole meal. A wooden pestle and mortar (Figure. 4) is also
commonly used in which by dampening, pounding,
winnowing, and pounding again, millet is separated into fine
and coarse semolina, flour, and bran.
Postharvest Processing
HARVESTING
Stems/Panicles/long straws/weeds/leaves
STORAGE
Spikelets/same sized weeds
SOURING
POPPING MALTING
MILLING AND POUNDING (Lactic acid
fermentation)
Grain/chaff/small weeds Husk/chaff/weeds
Tempering Steep
Conditioning Cooking
Figure 4 Women husking fonio millet. Reproduced from Gari JA (2001) Figure 5 Tangential abrasive dehulling device. Reproduced from
Review of African millet diversity. IPGRI/IFAD, Bamako, Mali, 1922 Balasubramanian S (2013) Processing of millets. In: National Seminar on
November 2001, www.FAO.org. Recent Advances in Processing, Utilization and Nutritional impact of
small millets. Madurai Symposium, Thamukkam Grounds, Madurai, 13
September 2013.
process of partial cooking the grain along with its husk or bran.
corners or surface irregularities being knocked off. These
The resulting product is dried and decorticated.
techniques are used only to a limited extent at commercial
scale.
Decortication
Decortication improves the biological availability of nutrients
Milling
and consumer acceptability of minor millets. It reduces total
protein and lysine, but it also improves the utilization of the
After dehulling, the material separated from the endosperm by
remaining protein. Decortication is sometimes accomplished
aspiration comprises the hull, bran, germ, and also a portion of
by using rice dehullers or other abrasive dehullers. Technically,
the endosperm in the form of broken grains and powder,
there are three types of decortication techniques that can be
which is used as poultry feed. The types of mills used for
employed to minor millets:
pulverizing minor millets are hammer mills, burr mills, and
1. Abrasive decorticators: Abrasive decorticators abrade away plate or disk mills. Wet milling is not performed for the minor
the fibrous pericarp (Figure 5). If all parts of the grains millets. Most of the minor millet processing involves semiwet
could be abraded away at the same rate, abrasive decortica- or dry milling techniques. Hammer mills are probably the
tion would be an efficient way of removing the pericarp. cheapest and simplest mills used for reducing the particle size
However, different parts of individual grains are abraded of the decorticated grains. These mills are available in different
away at different rates, and there is some loss of endosperm sizes depending on the size of operations. They consist of
(particularly from damaged grains) even when the grain is hammers installed on the rotor rotating rapidly in an enclosed
only lightly abraded. casing. The outlet is covered by a screen that determines the
2. Rubbing technique: For this technique, decortication is particle size of the flour. Overheating and moisture built up
achieved by a steel rotor, rotating the grain mass within a during the hammer milling result in caking of the flour and
generally cylindrical chamber. When the grain is properly cause reduction in the throughput of the process. Roller flour
tempered, the pericarp is rubbed off by the movement of milling is rarely used for minor millets as it is designed for
one seed against another. However, when the grain is too wheat milling. If roller mills are used for separating the endo-
dry, abrasion of the internal components of the mill sperm from the offal, the particle size is usually reduced in
becomes severe. The hulls and the endosperm fragments roller mills with smooth rollers. In cereals, the functional
are separated in a cyclone, and the endosperm particles are properties of flours are defined by the particle size distribution
milled in a proprietary mill. and damaged starch contents. However, there is not much
3. Metal friction machines: This is when attrition mills are used. research to evaluate the differences in the functionality of the
Attrition usually means particles getting smaller due to their millet flours owing to different milling procedures. Also, there
206 THE CEREAL GRAINS | Millet Minor: Overview
appears to be a difference between the flour produced by the (2011), a thresher-cum-pearler for finger millet and barnyard
wooden pestle and mortar and that produced by mechanical millet has been designed and developed. It works on the prin-
methods. The former seems to have better nutritional qualities. ciple of impact and shear on the grains. The threshing drum is
The biggest challenge is to develop a dehulling and milling provided with six leather flats, which provide gentle impact
technology according to specific physical characteristics of mil- and shear and a mild rubbing action for threshing, dehusking,
let grains that can yield flour with uniform quality. It is impor- pearling, and polishing of the grains. A sieve assembly is also
tant to note that some efforts have been made in this regard. attached with different operations. The design is simple and
The growth of this industry will depend upon constant easily adaptable in rural areas for efficient postharvest proces-
demand of millet-based products and regular feeding of the sing of these crops.
mills with millet grains, which have been properly harvested
and stored in good conditions.
Utilization of Minor Millets
Figure 6 CIAE-Millet mill (models I and II). Reproduced from Balasubramanian S (2013) Processing of millets. In: National Seminar on Recent
Advances in Processing, Utilization and Nutritional Impact of Small Millets. Madurai Symposium, Thamukkam Grounds, Madurai, 13 September 2013.
THE CEREAL GRAINS | Millet Minor: Overview 207
inexpensive foods prepared with other cereals like wheat Diseases in the Major Cereal Species; Grains Around the World:
breads and biscuits has made minor crops like millets lose Grain Production and Consumption: Africa; Non-wheat Foods:
their significance even in the local regions where they are Sorghum: Utilization; Proteins: Protein Synthesis and Deposition;
produced. Marketing millets as nutraceutical grains, while Scientific Organizations Related to Grains: Research
highlighting their key health benefits to create awareness Organizations of the World: CGIAR; The Basics: Grain: Morphology of
among consumers, could trigger demand of the millet-based Internal Structure; Taxonomic Classification of Grain Species; The
products. Support from the local governments at financial and Cereal Grains: An Overview of the Family of Cereal Grains Prominent
policy framework levels and involvement of cooperatives may in World Agriculture; Cereals: Domestication of the Cereal Grains;
help establish millet processing industry as a model in selected Millet Pearl: Overview; Teff: Overview; The Legumes and
sites, which can be replicated in other parts of the world. Minor Pseudocereals: Amaranth: Overview; Wheat-Based Foods:
millets are early maturing and can be cultivated under severe Cereals: Overview of Uses: Accent on Wheat Grain.
drought conditions. Thus, there exists a great potential of
exploiting these underutilized genetic germplasms to address
the current and future issues of health and global food security.
Further Reading
Exercises for Revision Annor GA (2013) Millet Starches: Structural Characteristics and Glycemic Attributes.
Doctoral dissertation, Canada: University of Guelph.
Annor GA, Marcone M, Bertoft E, and Seetharaman K (2013) In vitro starch digestibility
Give a few examples of minor millets and their genera. and expected glycemic index of kodo millet (Paspalum scrobiculatum) as affected by
What is the nutritional significance of polyphenols in starchproteinlipid interactions. Cereal Chemistry 90(3): 211217.
minor millets? Balasubramanian S (2013) Processing of millets. National Seminar on Recent Advances
What are the factors responsible for the hypoglycemic prop- in Processing, Utilization and Nutritional Impact of Small Millets. Madurai
Symposium, Thamukkam Grounds, Madurai, 13 September 2013.
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Investigate the recent advances in processing of minor mil- distribution of minerals in red non-tannin finger millet using PIXE microanalysis.
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Explore how minor millets could be a solution to global feeding on the early responses during the process of wound healing in diabetic rats.
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Determine how the molecular structure of minor millet Ramanathan MK and Gopalan C (1957) Effect of different cereals on blood sugar levels.
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Acknowledgments Sripriya G, Chandrasekharan K, Murthy VS, and Chandra TS (1996) ESR spectroscopic
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We gratefully acknowledge the late Dr. Koushik Seetharaman Taylor JRN, Belton PS, Beta T, and Duodu KG (2014) Increasing the utilisation of
for the inspiration to write this article. We also appreciate the sorghum, millets and pseudocereals: Developments in the science of their phenolic
support from Canadian International Food Security Research phytochemicals, biofortification and protein functionality. Journal of Cereal Science
Fund International Development Research Center and the 59(3): 257275.
Taylor JRN and Duodu KG (2014) Effects of processing sorghum and millets on their
Department of Foreign Affairs, Trade and Development, Gov- phenolic phytochemicals and the implications of this to the health-enhancing
ernment of Canada. properties of sorghum and millet food and beverage products. Journal of the
Science of Food and Agriculture 95(2): 225237. http://dx.doi.org/10.1002/
jsfa.6713.
See also: Agronomy of Grain Growing: Sorghum: Production and Tovey FI, Jayaraj AP, and Clark CG (1975) The possibility of dietary protective factors in
duodenal ulcer. Postgraduate Medical Journal 51: 366372.
Improvement Practices; Food Grains and the Consumer: Cultural Upadhyaya HD, Dwivedi SL, Singh SK, Singh S, Vetriventhan M, and Sharma S (2014)
Differences in Processing and Consumption; Grain Harvest, Forming core collections in Barnyard, Kodo, and Little Millets using morph
Storage and Transport: The Nature, Causes, and Control of Grain agronomic descriptors. Crop Science 54: 26732682.
Teff: Overview
G Bultosa, Haramaya University, Dire Dawa, Ethiopia; Botswana College of Agriculture, Gaborone, Botswana
2016 Elsevier Ltd. All rights reserved.
Topic Highlights and recommendations for further research to improve the teff
crop and grain utilization are suggested.
Teff crop description, origin, cultivation, and production.
Teff grain production constraints and progress of
improvement.
Teff Crop Description and Cultivation
Teff grain morphology, anatomical structure, composition,
and physicochemical and functional properties of the Description
major constituents of teff grain.
Teff is a C4 self-pollinated tetraploid cereal plant with a chro-
Teff grain nutrient merits, always consumed as whole-grain,
mosome number of 2n 4x 40. Teff is also an allotetraploid
gluten-free alternative food for celiac patients, high in min-
plant. The teff plant and panicles of some varieties are shown
eral nutrients particularly in iron with well-balanced amino
in Figure 1. Teffs root system is fibrous and most stems are
acids.
erect, while others are bending or elbowing types. It has a
Processing of teff grain flour to injera (lactic acid bacteria
panicle type of inflorescence showing different forms, from
and yeast-fermented, soft, spongy, sour, circular flatbread,
loose to compact. Its spikelets have 212 florets. Each floret
staple food for majority of Ethiopians).
has a lemma, a palea, three stamens, mostly two ovaries (in
Prospect for future teff productivity improvement and
some exceptional cases three), and feathery stigmas. In most
utilization and diversification of teff products.
varieties, the plant height is 50120 cm. A single teff plant is
known to produce from 9000 to 90 000 grains depending
upon the variety and production conditions.
Learning Objectives
Name and Origin
Achieve understanding on teff origin, description, grain
production, and constraints associated to grain production Teff (Eragrostis tef (Zucc.) Trotter) belongs to the family of
systems. Poaceae, subfamily Eragrostoideae, tribe Eragrosteae, and
Understand and apply the knowledge of teff grain mor- genus Eragrostis. About 350 species are known in the genus
phology, anatomical structures, nutrient compositions, Eragrostis of which teff is the only cultivated species. Chloridoi-
and grain component functionalities for processing into deae is used synonymously for Eragrostoideae of teff. Vernac-
various food products such as for gluten-free and whole- ular names in different parts of the world are as follows:
grain foods.
Achieve understanding on teff injera processing stages, Tahf: Arabic
nature of fermentation involved, characteristic of injera for Tef, teff, and Williams love grass: English
global consumers, and highlights on other traditional teff Xaafi/tafi/taafi: Oromo (O)/Afar/Sodo; tafe-e: Had; and tef,
grain food products. teff, and taff: Amarinya (A) and Tigrinya (T) (Ethiopian
languages)
See teff as a traditional Ethiopian grain, strongly linked to
national dietary culture, but with nutritional and sensory Mil ethiopien: French
benefits that could lead to product development in new Chimanganga, ndzungula (Ch), and chidzanjala (Lo): Malawi
markets. Teff is indigenous to Ethiopia. Ethiopia is also considered the
leading world center for teff genetic diversity. As of 2013, the
Ethiopian Institute of Biodiversity had conserved a total of
5169 accessions and 10 000 teff genotypes for varietal improve-
Introduction
ment study and to reduce loss of genetic diversity. The exact
details on teff domestication are unclear. However, teff is
Teff (Eragrostis tef (Zucc.) Trotter), today generally written as
believed to have been first domesticated by the pre-Semitic
tef, is a tropical cereal. Teff cultivation and usage as human
inhabitants of Ethiopia between 4000 and 1000 BCE and is
food are mostly confined to Ethiopia. This article highlights the
assumed to have originated in northeastern Africa. Even
origin, cultivation of the teff plant, and production and storage
though teff is an allotetraploid plant, to date, its diploid puta-
of the teff grain. Teff grain structure, anatomy, chemical
tive ancestors are not exactly established. But, based on the
composition, and physicochemical properties of the major
morphological data and cytological evidence, the following
chemical components are assessed and described. Teff milling,
species have been suggested as the ancestors and contributors
processing of injera (the staple food of the majority of Ethio-
to teff origin:
pians, a fermented, pancake-like, soft, spongy, sour, circular
flatbread), and other food products made from teff are Eragrostis aethiopica, E. atrovirens, E. longifolia, E. macilenta,
described. Lastly, teff potential, grain production limitations, E. pilosa, and E. pseudo tef as ancestor species of teff
(a) (c)
(b) (d)
Figure 1 Typical teff field plant and panicles of some teff varieties: (a) teff field of DZ-01-196, (b) panicles of DZ-01-196 (compact), (c) panicles
of DZ-Cr-37 (less compact), and (d) panicles of DZ-01-99 (loose). Reproduced with permission from Tefera, H., Ayele, M., Assefa, K., 1995. Improved
varieties of tef [Eragrostis tef (Zucc.) Trotter] release of 19701995. Research Bulletin 1. Debre Zeit Agricultural Research Center, Ethiopia.
E. aethiopica, E. bicolor, E. cilianensis, E. curvula, E. pilosa, and and E. minor) was not successful. However, it was recently
E. mexicana as contributor species to the origin of teff reported that E. tef and E. pilosa can be crossed with fertile
E. aethiopica, E. barrelieri, E. bicolor, E. cilianensis, E. hetero- offspring, suggesting that E. pilosa or an ancestor closely related
mera, E. mexicana, E. minor, E. papposa, and E. pilosa as very to E. pilosa is the most probable putative ancestor of teff.
closely related species to teff
E. aethiopica 2; E. barrelieri 6 ; E. cilianensis 2 , 4 , and
6; E. mexicana 6 ; E. minor 2 and 4 ; and E. pilosa 2 Cultivation and Production
based on cytological evidence as closely related species
Teff is one of the major cereals in Ethiopia with annual grain
to teff
production estimated to be ca. 3.5 106 t (ca. 21% of the
An attempted interspecific cross between teff and some wild cereal grain production). In Idaho, the United States, some
Eragrostis species (E. curvula, E. cilianensis (4 ), E. pilosa (4), teff grain is produced for the health-food market and for injera
THE CEREAL GRAINS | Teff: Overview 211
making. In South Africa, teff is widely grown as a fodder crop small and of light mass and can be easily blown away with
during the summer-rainfall season. However, the production the chaff. Use of mechanical threshers has recently shown an
of a combination of grain and fodder varieties has been improvement in grain yield and quality. Teff productivity may
introduced only recently. In Australia, India, Kenya, and the be improved in the future by adaption of this technology. In
United States, it is cultivated as a forage crop. Teff has been Ethiopia, the grain is traditionally stored in gotera (A, T) or
started to be cultivated in the Netherlands. Teff can adapt to a gotaara (O) (small hut-like stores) or pots or sacks. In compar-
wide range of environments, that is, moisture stress, high ison with other common cereals, teff grain is less prone to
rainfall, different soil types, and a wide range of altitudes attacks by weevils and other storage pests. Thus, it can be safely
from near sea level to over 3000 m. However, the best condi- stored under traditional storage conditions.
tions are 18002100 m above sea level, a temperature range
of 1027 C during the cultivation period, an annual rainfall
of 750850 mm, and rainfall of 450550 mm during the Teff Grain Marketing
growing season. Teff is known to have fewer disease and In local markets of Ethiopia, several small-scale grain traders
pest problems in the field as compared with maize, sorghum, distribute teff grain from major growing areas to the urban
wheat, and barley. However, the productivity is low. Lack of consumers and to regions of shortage in teff grain production.
high-yielding cultivars, lodging, weeds, waterlogging, low At Addis Ababa, teff grain is marketed in a place called Ehil
moisture, low-fertility conditions, loss of grains on harvest- berenda (Markato). Even though teff grain fetches premium
ing, threshing, and cleaning are major factors that contribute price in the global cereal grain markets, at present, interna-
to the low grain yield. The yields of improved varieties under tional teff grain export from Ethiopia is not practiced because
improved technologies (fertilizer, weed control, appropriate of limited production. But when permitted, wherever the
harvesting, and grain threshing) on farms are in the range of market is, companies (government-owned, joint ventures,
1.42.7 t ha1, and for some under research-managed farms, farmers union associations, and private firms), such as Ethio-
the yield is as high as 3.6 t ha1. Reduction of seeding rate pian Grain Trading Enterprise and Oromo Development Asso-
from traditionally recommended 2530 kg ha1 to ciation, which export oil and pulses, have the potential to
2.5 kg ha1 had recently demonstrated a substantial grain export teff grain as well.
yield increase as compared with previous grain yield
recorded. Because of adaption of some of the improved tech-
nologies by teff farmers, the average national yield in Ethiopia
Physicochemical Properties of Teff Grain
has increased annually from about 0.84 t ha1 in 2003 to
about 1.26 t ha1 in 2011. Morphology
Teff grain is hull-less (naked) and comes in a range of colors
Harvesting and Storage from milky white to almost dark brown. The most common
colors are white, creamy white, light brown, and dark brown
Teff grain is harvested when the vegetative and reproductive
(Figure 3). The grain is oval-shaped with size 0.91.7 mm
parts (pedicel, lemma, palea, and glumes) turn yellow or
(length) and 0.71.0 mm (diameter). The individual grain
straw color (4560 days for very early maturing, 60120
mass is generally 2 mg, 0.60.8% of the wheat grain
days for early maturing, and 120160 days for late maturing).
mass. The thousand kernel weights and hectoliter weight of
If harvested late, the grain loss will be significant due to
teff grain are in the range 0.190.42 g and 8587 kg hl1,
shattering and the natural grain color can also fade. However,
respectively.
if the grain is harvested early, it may become vitreous or
translucent. In Ethiopia, traditional harvesting is done man-
ually using a sickle (Figure 2(a)). The harvested panicles are Anatomy
gathered in batches, either on the day harvested or after 13
days, and temporarily stacked (in a conical shape, with the Pericarp
panicles toward the center) (Figure 2(b)) outdoors, under the The outer pericarp is thin and membranous and is equivalent
shade, till it is ready for threshing. Such storage duration will to the beeswing bran of wheat. In the pericarp, beneath the
help the teff grain to undergo the ripening process and dor- cuticle toward the nucellar epidermis, teff grain is known to
mancy breaking stages. bear slime layer rich in pectins, which form uniform layers on
At present, teff grain pre- and postharvest losses are high the inner surface of the cell wall. The ability of this slime layer
(2530%), largely incurred by lodging, shattering on harvest to absorb and maintain moisture around the grain is impli-
and harvest handling, threshing, and cleaning by traditional cated as a contributor to teff adaptive features to moisture
practices. Threshing is usually done by oxen feet trampling or stress. In the inner surface of the pericarp, the mesocarp and
manually by beating with sticks after spreading the dried pan- endocarp are fused and appear as a single layer. In this fused
icles on a dry circular threshing floor. The threshing floor is layer, some starch granules are observed, as in the pericarp of
prepared by smearing with cow dung, cement, or other suitable sorghum grain.
materials. The grain is traditionally winnowed by wafting in
the open air with the help of a rectangular flat piece of dried Seed coat (testa)
leather called an afarsa or hafarsaa (O) (ca. 0.4 m width - Next to the endocarp is the testa, which is adjacent to the
0.8 m length). Threshers or combine harvesters can be also aleurone layer. In some teff varieties, the testa is reported to
used. However, grain loss is large because teff grain is very contain tannins and is thus presumed to be thick. However, in
212 THE CEREAL GRAINS | Teff: Overview
Aleurone layer
The aleurone layer is one cell thick and is rich in protein and
lipid bodies.
Germ (embryo)
Like in other small-grain cereals, the germ occupies a relatively
large proportion of the grain and is rich in protein and lipids.
Endosperm
The endosperm is the largest component of the grain and
consists of outer and inner layers. The outer layer is vitreous
and contains most of the protein reserves of the endosperm and
a few starch granules. The inner layer is mealy consisting mainly
of thin-walled cells containing mostly starch granules with a
few protein bodies. Teff has compound-type starch granules
(Figure 4(a)), representing the contents of one amyloplast-
like rice, oats, Amaranthus, and quinoa starches. On milling,
individual starch granules are released along with small groups
of protein bodies. The protein bodies are individual entities in
nature and spherical in shape (Figure 4(a)), and unlike those of
wheat, they do not coalesce to form a matrix.
Carbohydrate
Carbohydrate content of teff grain is 73%, of which virtually
all is starch. The teff starch properties are given in Table 2.
Individual starch granules are very small (26 mm in diameter)
(Figures 4(b)4(f)), similar in size to rice starch granules, but
larger than Amaranthus and quinoa starch granules. The shape
is polygonal, smooth with no surface pores (Figure 4(f)). A few
granules are essentially cubic, and at high magnification, some
appear tortoise shell-shaped (Figure 4(e)). The composition of
teff starch granules is similar to other normal native cereal
starches, with 2530% amylose. Gelatinization temperature is
high, similar to other tropical cereal starches. The x-ray diffrac-
tion pattern of teff starch granule is A-type with crystallinity of
37% similar to rice starch granule. Pasting temperature is
similar to that of maize starch, but cooking time for peak
viscosity is longer. Peak, breakdown, and setback viscosities
are lower than those of maize starch. The paste clarity of
teff starch is opaque. The gel texture is short and smooth.
a-Amylase degradation of teff starch granules is by surface
erosion and endocorrosion in nature.
Because teff starch granules are very small, smooth, and of
uniform size, they offer good functionality as a fat substitute
Figure 2 (a) Harvesting of teff plant by the traditional method, (b) stacked and a flavor and aroma carrier, similar to other small-granule
teff panicles after harvest, and (c) teff grain after cleaning by winnowing and starches. Teff starch has good resistance to shear breakdown,
wafting, where TF threshing floor top layer smeared with cow dung and and thus, it may find good application in high-shear processed
dried, GS grass straw made of traditional bucket, and IM impurities. foods. Also, because of its slow retrogradation tendency, it
THE CEREAL GRAINS | Teff: Overview 213
Figure 3 Grains of different teff varieties: (a) DZ-01-196 (white), (b) DZ-Cr-37 (creamy white), (c) DZ-01-99 (light brown), and (d) South African brown
(dark brown).
Table 1 The proximate composition (db) of teff grain, Osborne protein fractions, and food energy
cw
sg pg
sg
sg
pb
pb pb
cb
(a) (d)
pg
tr
cb
(b) (e)
f cb nsp
pg
(c) (f)
Figure 4 Compound (a) and individual (bf) starch granules from different teff varieties: (a, b) South African brown, (c) DZ-01-1681, and (df)
DZ-01-196 (sg starch granules, pg polygonal, cb cubic, Tr tortoise-shell, pb protein, f fiber, and nsp no surface pores). Reproduced from
Bultosa, G., 2003. PhysicoChemical and Functional Properties of Grain tef [Eragrostis tef (Zucc.) Trotter] Starch. PhD thesis, University of
Pretoria, Pretoria, South Africa.
could have attractive applications where starch staling is pre- Protein and amino acids
ferred to be reduced (i.e., in baked and in refrigerated foods). Typical teff grain protein content (N 6.25) is 11%, with a
normal range of 913%. Thus, the protein content of teff
Fiber grain is similar to other common cereals. The major amino
The fiber content of teff grain (Table 1) is apparently higher acids are glutamic acid, alanine, proline, aspartic acid, leu-
than most other common cereals, because the grain is very cine, and valine (Table 3). Methionine, phenylalanine, and
small and the bran is proportionally large. histidine are slightly higher than in most other cereals, but
THE CEREAL GRAINS | Teff: Overview 215
Individual granule diameter (mm) and shape 26 (majority are 35), polygonal shape, Individuals are from compound granules
smooth surface
Amylose (%) (db) 28.4 2.8 Concanavalin A method
28.2 0.8 Iodine-binding method
Amylose weight-average molar mass, g mol1 (1.53.0) 106
Amylopectin weight-average molar mass, g mol1 (10.116.5) 107
Ash (%) (db) 0.16 0.04
Protein (%) (N 6.25) (db) 0.19 0.03
Lipids (db): total (mg g1) 8.9 0.7 24% HCl hydrolysis followed n-hexane and
in situ generated ethyl formate extract
Internal lipids (mg g1) 7.8 0.4 Hot (90 C) water-saturated butanol extract
Phosphorus (db) (mg g1) 0.65 0.08
Kofler gelatinization temp. range ( C): ToTpTc 68.074.080.0 To is onset, Tp is peak, and Tc is conclusion
gelatinization temperatures
DSC gelatinization endotherms range: To, Tp, and Tc in 63.865.4, 70.271.3, and 81.381.5 and DH is gelatinization enthalpy
C and DH in J g1, respectively 2.287.22
Pasting properties: Ti ( C), PV (RVU), BV (RVU), Rst 74.0 1.1, 269 13, 79 17, 8.4 1.8, Where Ti, PV, BV, Rst, and SBV are pasting
(RVU min1), SBV (RVU) 101 11 temperature, peak viscosity, breakdown
viscosity, rate of shear thinning, and
setback viscosity, respectively
Peak viscosity Mediumlow
Resistance to shear Mediumhigh
Retrogradation tendency Mediumlow
Crystallinity (%) 37
x-Ray diffraction: d-value (A) and intensity (%) 5.85 (83.8), 5.16 (97.0), 4.89 (99.4), 4.41 A-type starch crystalline polymorph
(36.4), and 3.84 (80.2)
Paste clarity Opaque
Gel texture Short
In vitro digestibility with porcine pancreatic a-amylase Surface erosion and endocorrosion type
Acid hydrolysis Gradual surface degradation (etching)
Source: Bultosa, G., 2003. PhysicoChemical and Functional Properties of Grain tef [Eragrostis tef (Zucc.) Trotter] Starch. PhD thesis, University of Pretoria, Pretoria, South Africa;
Bultosa, G., Hamaker, B.R., BeMiller, J.N., 2008. An SEC-MALLS study of molecular features of water-soluble amylopectin and amylose of tef [Eragrostis tef (Zucc.) Trotter] starches.
Starch/Starke 60, 822.
serine and glycine are lower. Lysine and arginine are essen- because teff grain bran is proportionally large. In particular,
tially higher in teff than in most other cereals, except rice and calcium, copper, iron, and zinc (Table 4) contents are higher
oats. The balance among essential amino acids is similar to compared with those in barley, wheat, and sorghum. The iron
the whole edible portion of egg protein, except for its lower content of traditionally harvested teff grain is especially high
lysine content. The overall amino acid profile of teff can be ( 15.7 mg per 100 g and in some reports as high as 27 mg per
regarded as well balanced. 100 g), in part because of grain contamination with the soil
Osborne protein fractions of teff grain are shown in Table 1. during harvest. However, when cleaned (with water and/or
Glutelins, albumins, and globulins are major fractions. The teff dilute acid), the level ( 5.7 mg per 100 g) is similar to other
prolamin fractions are reported variable (low and high) cereals. Most teff foods such as injera are fermented. Fermen-
depending on solvent type used in the extraction. The major tation for injera processing is known to destroy phytic acid to
prolamins of teff are somewhat similar to the a-prolamins of the extent of 90%, and such destruction is known to contrib-
maize, sorghum, and Coix. Teff is different from other cereals ute to high iron bioavailability in diets where fermented teff
in having higher albumins and globulins. Teff protein is essen- foods are the staples. Because of these two factors, iron-
tially free of the type of gluten found in wheat. Because of this, deficiency disease anemia is rare among regular teff injera
teff grain foods are today increasingly became important for consumers in Ethiopia.
consumers allergic to wheat gluten (e.g., for celiac patients). As
the main protein fractions (albumins and globulins) are the
most digestible types, teff protein digestibility is also presumed Fat and fatty acids
high. Amino acid compositions of the various Osborne protein Teff grain fat (Table 1) is lower than, for example, in maize and
fractions are shown in Table 3. oats. Thus, teff is different from other small-grain cereals in
having low fat even though the germ is large. As in most other
Ash and minerals cereal grains, palmitic, oleic, and linoleic acids are the major
The ash content in teff grain (Table 1) is apparently higher fatty acids (Table 5). Linolenic acid in teff is higher than in
than in wheat, rye, maize, barley, oats, rice, and millets, in part maize, sorghum, and wheat.
216 THE CEREAL GRAINS | Teff: Overview
Table 3 Amino acid composition of the whole teff grain protein and of the Osborne protein fractions (g per 100 g protein)
Source: Bultosa, G., Taylor, J.R.N., 2004. Teff. In: Wrigley, C., Corke, H., Walker, C.E. (Eds.), Encyclopedia of Grain Science, vol. 3. Elsevier, Oxford, UK, pp. 281290; Ketema, S.,
1997. Tef Eragrostis tef (Zucc.) Trotter. Promoting the Conservation and Use of Underutilized and Neglected Crops, Institute of Plant Genetics and Crop Plant Research, Gatersleben/
International Plant Genetic Resources Institute, Rome, Italy, Booklet 12; Assefa, K., Chanyalew, S., Tadele, Z. (Eds.), 2013. Achievements and Prospects of Tef Improvement:
Proceedings of the Second International Workshop, 79 November, 2011, Debre Zeit, Ethiopia. Ethiopian Institute of Agricultural Research/Institute of Plant Sciences, University of
Bern, Addis Ababa, Ethiopia/Bern, Switzerland; Hundera, F., 1998. Variations of morphoagronomic characters and grain chemical composition of released varieties of tef [Eragrostis
tef (Zucc.) Trotter]. J. Genet. Breed. 52, 307311; Bekele, E., 1995. Essential and non-essential amino acids in a free state and in the major protein fractions of teff seeds. SINET:
Ethiop. J. Sci. 18, 7999.
Table 5 Fatty acid composition of teff grain fat plasticity losses after a day. The rice flour injera-making ability
is similar to teff probably because of somewhat similar starches
Fatty acids (%) Range Mean in both grains. The hydrophobic and less polymerized nature
Palmitic (C16:0) 14.016.4 15.9 of teff grain protein prolamins is in part implicated as contrib-
Palmitoleic (C16:1) 0.10.6 0.3 utor to the making of superior semileavened flatbread injera
Stearic (C18:0) 3.03.7 3.3 when compared with sorghum injera. Teff grain flour is also
Oleic (C18:1) 23.324.9 24.0 used to make sweet unleavened bread called kitta (A), bixxille
Linoleic (C18:2) 41.346.5 44.2 (O), and daguwalo (T). Kitta can be consumed as bread, or it
Linolenic (C18:3) 6.99.9 7.9 can be used as an adjunct in traditional opaque beer (tella (A)
Arachidic (C20:0) 0.60.9 0.7 or farsoo (O)) or local spirit (katikalla (A) or araqii (O)). Por-
Arachidonic (C20:1) 0.51.2 0.8 ridge (genefo (A) or marqaa (O)) can also be made from teff
Behenic (C22:0) 0.31.1 0.5
flour. Thin, fermented teff flour batter is used to prepare soup
Erucic (C22:1) 0.00.9 0.4
(muk, (A)). Unfermented teff flour dough is also used in the
Source: Hundera, F., 1998. Variations of morphoagronomic characters and grain preparation of traditional snacks (dabo kolo (A) or hunkuroo
chemical composition of released varieties of tef [Eragrostis tef (Zucc.) Trotter]. J. (O)), where the dough is rolled into small balls and then
Genet. Breed. 52, 307311. roasted on a hot griddle. In the United States, teff has been
promoted as a thickener for soups, stews, and gravies probably
because teff flour paste gives the product a short and stiff
texture. Teff grain flour imparts a slight molasses-like sweetness
Table 6 Vitamin and antinutrient composition of teff grain (db)
to food products, making its inclusion in porridges, pancakes,
Component Typical value biscuits, cookies, cakes, stir-fry dishes, casseroles, soups, stews,
and puddings desirable. Processing of teff grain in Ethiopia has
Vitamins been limited to the household level. To date, technologies for
Vitamin A (mg per 100 g as RE) 8 large-scale commercial processing of teff grain, for the prepa-
Thiamine (mg per 100 g) 0.3 ration of foods like injera, are not well advanced. However,
Riboflavin (mg per 100 g) 0.2
apart from traditional usage, recent reports indicate that teff
Niacin (mg per 100 g) 2.5
grains, along with soybean, chickpea, and other grains, are
Vitamin C (mg per 100 g) 88
Antinutrients being used in the baby food industry. Teff grain flours are
Phytate (mg per 100 g) 707 used to improve dietary fiber, starch, protein, and mineral
Trypsin inhibitor activity (TIU per gram) 5584 content supply in gluten-free food products and in wheat
bread to improve the iron and antioxidant contents of the
RE is retinol equivalent; TIU is trypsin inhibitor unit. bread.
Source: Assefa, K., Chanyalew, S., Tadele, Z. (Eds.), 2013. Achievements and Prospects
of Tef Improvement: Proceedings of the Second International Workshop, 79
November, 2011, Debre Zeit, Ethiopia. Ethiopian Institute of Agricultural Research/ Teff Grain Injera Making
Institute of Plant Sciences, University of Bern, Addis Ababa, Ethiopia/Bern, Switzerland;
Bultosa, G., 2003. PhysicoChemical and Functional Properties of Grain tef [Eragrostis In Ethiopia, injera is regarded as the national staple food.
tef (Zucc.) Trotter] Starch. PhD thesis, University of Pretoria, Pretoria, South Africa; NRC Flowcharts of the injera-making process and injera-baking pro-
(National Research Council of the USA). 1996. Tef. In: Lost Crops of Africa, vol. 1, cess are shown in Figures 5 and 6, respectively. The process
Grains, pp. 214338. Washington, DC: National Academy Press. involves fermentation and then baking of the batter.
Fermentation
Flour is mixed with water and the dough is kneaded, usually by
hand. Fermentation for injera making involves two phases that
Usage
can last a total of 2472 h. The first phase starts spontaneously
Food made from teff grain is a staple diet for many Ethiopians. when the flour is wetted, due to contaminating microorgan-
Teff is considered to have a better food value than major grains, isms. Or it can be initiated by addition of irsho (A) (T) or
namely, wheat, barley, sorghum, and maize, as it is normally raacitii (O) (yellowish liquid saved from the previous batch
used as a whole grain, that is, the germ and bran are consumed fermentation). The initial 1824 h is notable for vigorous gas
along with the endosperm. Teff flour is used primarily for evolution and maximum dough expansion. At about 3033 h,
making of injera (A), caabitaa or budeena (O), and tayeta (T). an acidic yellowish liquid appears on the dough surface. This
Teff injera is regarded to have pre- and probiotic potentials due phase is characterized by a decrease in gas evolution up to 31 h,
to dietary fiber from whole grain and fermentation, an increase in liquid volume up to 48 h, and a decrease in pH
respectively. Although the lactic acid bacteria (LAB) and yeast to below 5.8.
involved in the fermentation of injera are killed during baking, The first phase of fermentation results in a liquid/solid
their dead cells and metabolic products are consumed as part separation after 24 h. The layer of liquid is then removed.
of the diet, and these were implicated as promoters of gut About 10% of the fermenting dough is mixed with water (1:3
health. In injera-making features, teff grain flours are superior ratio) and boiled (25 min), and as a result of starch gelatini-
to any other cereal grain flours used. The Ethiopian diaspora is zation, a dough binder, called absit, is formed. The absit is
using injera from rice or blend of rice and wheat flour, cooled and added to the fermentation vat signaling the second
although the injera suffers from staling, resilience, and phase of fermentation. The second phase (0.52 h) is
218 THE CEREAL GRAINS | Teff: Overview
Teff grain
Cleaning
Millings
characterized by a short duration of dough expansion and gas to the Enterobacteriaceae family are thought to initiate the
formation. fermentation. During the first 18 h of fermentation, the activ-
During the first phase of fermentation, the yellowish liquid ities of these bacteria reduce the dough pH to 5.8. A group of
that is removed contains water-soluble nutrients (amino acids, LAB of which major ones are Leuconostoc mesenteroides, Strepto-
sugars, and minerals) and a large number of microorganisms coccus faecalis, Pediococcus cerevisiae, P. pentosaceus, Lactobacillus
involved in the fermentation. This has negative nutritional brevis, L. plantarum, and L. fermentum are involved at the later
consequences. Thus, injera baked from a batter at 31 h of phase of fermentation (1872 h) in reducing the dough pH
fermentation without discarding the liquid is recommended as from 5.8 to 3.8. During the later phase of fermentation (22
being more nutritious. 24 h), yeasts of two genera Saccharomyces and Torulopsis are
A complex group of microorganisms is known to be reported to be involved. In the later phase ( 48 h), yeasts
involved in teff fermentation. A total of 107 LAB (some hetero- belonging to the genera Candida and Pichia are the dominant
fermentative and some homofermentative) and 68 yeast types isolated from the yellowish liquid removed from the
strains are reported in the teff fermentation. Bacteria belonging dough. In addition to amylases present in the grain, the
THE CEREAL GRAINS | Teff: Overview 219
DZ-01-196
DZ-01-787 DZ-01-99
Figure 6 Teff injera baking from fermented batter on electrically heated baking griddle (a). Portion of baked injera from white (b and c) and brown (d)
teff grains.
bacteria species Bacillus sp. A-001 involved in the fermentation nutritious. The thick batter used for aflegna injera is also used
has been characterized as one of the amylase-producing bacte- to prepare a slightly concave, thick flatbread called cumboo (O).
ria involved in partially attacking the starch granules. Cumboo is traditionally baked on the preheated surface of a
In the traditional teff fermentation for injera making, com- small-size concave griddle called bedde, which is placed upside
mercial yeast is not added externally. The source of the yeast is down on a flat larger griddle called elle chumbo. After pouring
either from irsho or from the endogenous microflora of teff the 1224 h fermented thick batter onto the upside down
grain and its flour that grows in the batter after the flour is concave bedde surface and covering with the lid, gentle heat is
wetted with water. Therefore, the yeast in the fermentation of applied underneath of the larger griddle, and depending on the
teff for injera making can be regarded as symbiotic yeast. batter size and heat intensity applied, the baking time for
cumboo is between 1 and 3 h.
Baking
Injera is usually baked after 24 h of fermentation. After the
fermentation, the batter is diluted slightly with water and then Prospects, Problems, and Recommendations
poured using circular motion from the outer perimeter toward
the center onto a hot, round, smooth griddle called a metad (A) Teff can be cultivated under harsh environmental conditions
or eelee caabitaa (O). It is then covered with a metad lid called where most other cereals are less viable. It has relatively few
akambalo (A), qadaada eelee caabitaa (O) to prevent steam from pest and disease problems in the field. The grain is less attacked
escaping. The griddle is traditionally made from clay. Before by weevils. The nutrient composition of teff grain indicates that
pouring the batter, the metad surface is swabbed with ground it has good potential to be used in various diets and as func-
oilseeds, commonly rapeseed or with animal fat in a piece of tional foods in health-food markets worldwide. However, the
cloth. This prevents the injera from sticking to the metad sur- grain yield of teff is low. The mechanized farming technologies
face. Depending on the batter thickness, heat intensity applied, that are used for the production of other cereal grains can be
and steaming, with the current electrically heated baking grid- problematic for teff because the plant stems are very thin and
dle, injera can be baked in 23 min. short and the grain is very small. Manipulation of teff plant
Based on the duration and nature of the fermentation genes through research on germplasm genetic diversity,
involved, three common types of injera are prepared:(1) injera improvement of yield using molecular breeding approach,
made from dough that does not contain absit, characterized by and a continuous teff production technology improvement
a soft, thin, fine appearance and a sour taste without the eyes for maximum grain production are required.
of injera (surface air cells); (2) injera made from partially At present, the milling of teff grain is limited to cottage-type
fermented paste (1224 h fermentation) called aflegna (A) millers. Processing of teff for different foods is usually by
(O) or bekuo (T), characterized by a sweet flavor, a pleasant traditional ways and mostly limited to the household level.
odor, and a rusty red underside; and (3) injera made from Over the last decade, utilization of teff grain as whole grain
overfermented paste called komtata injera (A) (T) or qomxoxaa supplements and for gluten-free markets is increased. More
caabitaa (O) that tastes very sour and is regarded as less research on teff large-scale milling and processing of teff grain
220 THE CEREAL GRAINS | Teff: Overview
for different commercial foods is required to promote world- Overview of the Family of Cereal Grains Prominent in World
wide teff product utilization. Agriculture; Millet Minor: Overview; Oats: Overview; Rice: Overview;
The Legumes and Pseudocereals: Amaranth: Overview.
What major features distinguish teff grain (particularly its Further Reading
starch and protein) from other common tropical cereals Adebowale AA, Emmambux MN, Beukes M, and Taylor JRN (2011) Fractionation and
like sorghum, rice, and maize? characterization of tef proteins. J. Cereal Sci. 30: 17.
Why is the productivity of the teff plants so low compared Assefa K, Chanyalew S, and Tadele Z (eds.) (2013) Achievements and Prospects of Tef
with that of maize and rice? Could teff be grown near where Improvement: Proceedings of the Second International Workshop, November 79,
2011, Debre Zeit, Ethiopia, Addis Ababa, Ethiopia/Bern, Switzerland: Ethiopian
you live?
Institute of Agricultural Research/Institute of Plant Sciences, University of Bern.
Why are teff grain products nutritionally and functionally Bekele E (1995) Essential and non-essential amino acids in a free state and in the major
attractive for health-food markets (e.g., for celiac patients)? protein fractions of teff seeds. SINET: Ethiop. J. Sci. 18: 7999.
Can you design and develop a novel food product utilizing Bergamo P, Maurano F, Mazzarella G, Iaquinto G, Vocca I, Rivelli AR, et al. (2011)
teff grain? Immunological evaluation of the alcohol-soluble protein fraction from gluten-free
grains in relation to celiac disease. Mol. Nutr. Food Res. 55: 12661270.
How is injera made? Is teff grain essential for injera manufac- Bultosa, G., 2003. PhysicoChemical and Functional Properties of Grain tef [Eragrostis
ture? Does teff injera have prebiotic and probiotic effects? tef (Zucc.) Trotter] Starch. PhD thesis, University of Pretoria, Pretoria, South Africa.
Bultosa G and Taylor JRN (2004) Teff. In: Wrigley C, Corke H, and Walker CE (eds.)
Encyclopedia of Grain Science, vol. 3, pp. 281290. Oxford, UK: Elsevier Ltd.
Bultosa G, Hamaker BR, and BeMiller JN (2008) An SEC-MALLS study of molecular
Exercises for Readers to Explore the Topic Further features of water-soluble amylopectin and amylose of tef [Eragrostis tef (Zucc.)
Trotter] starches. Starch/Starke 60: 822.
Value chain efficiency for teff productivity improvement Desiye A and Abegaz K (2013) Isolation, characterization and identification of lactic acid
such as high-yielding and lodge-resistant variety develop- bacteria and yeast involved in fermentation of teff (Eragrostis tef) batter. Adv. Res.
ments coupled with mechanized or semiautomated pro- Biol. Sci. 1(3): 3644.
Ebba T (1969) Tef (Eragrostis tef): the cultivation, usage and some of the known
duction technologies that suit very thin, short teff plant diseases and insect pests. Part I. Debre Zeit Agricultural Experiment Station. Dire
stems and very small grain size of light mass that would Dawa, Ethiopia: Addis Ababa University, College of Agriculture.
boost teff grain production and minimizes grain posthar- Hundera F (1998) Variations of morpho-agronomic characters and grain chemical
vest losses. composition of released varieties of tef [Eragrostis tef (Zucc.) Trotter]. J. Genet.
Breed. 52: 307311.
Development of teff grain-based composite flours for injera
Ketema S (1997) Tef Eragrostis tef (Zucc.) Trotter. Promoting the Conservation and Use
and other teff products. of Underutilized and Neglected Crops. Gatersleben/Rome: Institute of Plant Genetics
Efficiency of teff injera processing stages and cottage and and Crop Plant Research/International Plant Genetic Resources Institute Booklet 12.
large-scale injera processing and preservation. Kreitschitz A, Tadele Z, and Gola EM (2009) Slime cells on the surface of Eragrostis
Teff grain-based ready to eat foods for health-food markets seeds maintain a level of moisture around the grain to enhance germination. Seed
Sci. Res. 19: 2735.
such as gluten-free foods for celiac patients. NRC (National Research Council of the USA) (1996) Tef. In: Lost Crops of Africa, vol. 1.
Grains, pp. 214338. Washington, DC: National Academy Press.
Tefera H, Ayele M., Assefa, K., 1995. Improved varieties of tef [Eragrostis tef (Zucc.)
See also: Beverages from Grains: Fermentation: Foods and Trotter] release of 19701995. Research Bulletin 1. Debre Zeit Agricultural Research
Nonalcoholic Beverages; Food Grains and the Consumer: Cultural Center, Ethiopia.
Umeta M and Parker ML (1996) Microscopic studies of the major macro-components of
Differences in Processing and Consumption; Grains and Health; Food seeds, dough and injera from tef (Eragrostis tef). SINET: Ethiop. J. Sci.
Grains and Well-being: Functional Foods: Dietary Fibers, 19: 141148.
Prebiotics, Probiotics, and Synbiotics; Functional Foods: Overview; Umeta M, West CE, and Fufa H (2005) Content of zinc, iron, calcium and their
Food Grains: Intolerance, Allergy and Diseases: Celiac absorption inhibitors in foods commonly consumed in Ethiopia. J. Food Comp.
Anal. 18: 803817.
Disease; The Gluten-Free Diet; Appendix 1: Nutrient-Composition
Tables for Grains and for Grain-Based Products; Grain Composition
and Analysis: The Composition of Food Grains and Grain-Based
Products; Grains Around the World: Grain Production and Relevant Websites
Consumption: Africa; The Basics: Grain and Plant Morphology of
http://www.ars-grin.gov USDAs GRIN database.
Cereals and how characters can be used to identify varieties; The Grain http://mansfeld.ipk-gatersleben.de Mansfelds world database of agricultural and
Crops: An Overview; Grain: Morphology of Internal Structure; horticultural crops.
Taxonomic Classification of Grain Species; The Cereal Grains: An http://www.tef-research.org/-Tef Research project.
The Oilseeds
Contents
Oilseeds: Overview
Soybean: Overview
Canola: Overview
Cottonseed: Overview
Sunflower: Overview
Overview of the Oilseed Safflower (Carthamus tinctorius L.)
Linseed: Overview
Oilseeds: Overview
RJ Mailer, Australian Oils Research, Lambton, NSW, Australia
2016 Elsevier Ltd. All rights reserved.
The major oilseed crops of the world include soybean, Oilseed crops are generally grown for the oil in their seeds and
cottonseed, rapeseed, sunflower, groundnut, sesame seed, vary considerably in oil content, quality, and composition
linseed, safflower, and mustard seed. factors that rely heavily on the crop species or cultivar and
The global oilseed market is dominated by soybean with a upon the environmental conditions in which the crop is
total of 242 Mt produced in 2012. grown. The seed meal left after the oil has been extracted can
Rapeseed is now rated in second position in world produc- provide a high-protein product for use in either animal or
tion of oilseeds with 65 Mt produced in 2012. human food. In some crops, the seed meal or some other
Plant-breeding programs are aimed at increasing oil pro- factors are more important than the oil, and the oil is a by-
duction and quality. product. For example, soybean is primarily grown for its high-
World oilseed production is forecast to expand by 20% for protein meal and cotton for its fiber. Consequently, in
the period 201221. oil-producing crops, plant-breeding programs are aimed at
The use of edible oil for biodiesel is forecast to increase by increasing both oil production and quality with the additional
76% or 30 Mt by 2021. aim of increasing meal quality. In some oil crops, antinutri-
Meal is utilized as stock feed, in particular pork and poultry tional components are present (such as gossypol in cottonseed
and, more recently, aquaculture. and glucosinolates in rapeseed); they also need to be consid-
Palm oil fruit production in 2012 exceeded 50 Mt, second ered and may limit the use of the meal.
only to soybean.
The biggest changes will come in the future from genetic
modification.
Oilseeds and Their Uses
been a trend in human diets toward reduction of saturated fats Table 1 Approximate oil content of seed and protein content
such as palmitic acid (C16:0) and an increase in polyunsatu- of oil-free meal and the main uses of selected oilseed crops
rated (C18:2 and C18:3) and monounsaturated (C18:1) fats.
Oilseed Oil (%) Protein (%) Main use
Plant breeders have successfully developed more nutritionally
favorable products from traditional oilseed crops by altering Soybeans 20 46 Food
their fatty acid profiles and other constituents. Cottonseed 16 37 Fiber
There is strong demand for food, feed, and biofuel. According Peanut 41 46 Food
to the OECD-FAO Agricultural Outlook 201221, world oilseed Rapeseed 41 34 Oil
production is forecast to expand by 20% for the period, exceed- Sunflower seed 40 28 Oil
ing 529 Mt by 2021. This is a marked slowdown compared with Sesame seed 4060 42 Food
that of the last decade and is as a result of reduced area expan- Linseed 40 3234 Oil
Safflower seed 34 23 Oil
sion, environmental constraints, and competing crops. Vegeta-
Mustard seed 2050 35 Condiment/oil
ble oil production, however, will increase by about 28% due to
the contribution of palm and coconut oil. By 2021, palm oil is Source: Adapted from Australian Oilseed Federation Technical and Quality
expected to make up one-third of world vegetable oil production. Standards 2014 and other literature values.
World consumption of oils is forecast by FAO to grow by only
2% per year, half the growth rate of the last decade. China and complete and involved the use of hazardous flammable
the European Union are the dominant importers of oilseeds and solvents. In addition, only a few samples could be processed
their products. Although demand for food use is slowing down at a time reducing the capacity of the laboratories to service the
in developed countries as they reach saturation levels, consump- needs of plant-breeding programs. Kjeldahl nitrogen analysis
tion in developing countries is steadily increasing. There has to determine the nitrogen content of the meal also required the
been and continues to be a significant increase in biodiesel use use of dangerous chemicals and strong acids. More recent
in developed and developing countries. The use of edible oil for advances in oil content determination include nondestructive
biodiesel is forecast to increase by 76% or 30 Mt by 2021, raising seed analysis, such as the use of nuclear magnetic resonance
the share of vegetable oil for biodiesel from 12% to 16%. Non- spectroscopy. The rapid and accurate nondestructive testing
food uses of vegetable oils in developed countries are estimated of seed for oil content allows the breeder to retain the seed of
to continue to increase due to developments in the oleochemical potentially suitable lines. The introduction of near-infrared
industries with a shift away from petroleum-based products to reflectance spectroscopy has further enhanced the ability of
environmentally friendly oleochemicals. In addition, there is an laboratories to determine a wider range of seed components
increase in technological developments, particularly with the including oil, protein, moisture, fiber, and numerous others on
genetic modification of oilseeds to create a new range of prod- relatively small samples in a short time. Consequently, hun-
ucts. For cooking oils and salad dressings, plant breeders have dreds of seed samples can now be analyzed each day and the
selected cultivars with lower levels of polyunsaturated and satu- intact seed returned to the breeder for further growth and
rated fats and an increase in monounsaturates for increased evaluation.
oxidative stability. New developments include oils with
increased stearic acid to provide margarine type fats without
the need for hydrogenation and the subsequent production of New Variations on Old Crops
trans-fatty acids. The uses of vegetable oils in paints, lubricants,
cosmetics, and pharmaceuticals have been well documented. The major oilseed crops of the world include soybean, cotton-
Meal production globally will increase by 23% or about seed, rapeseed, sunflower, groundnut (or peanut), sesame
315 Mt by 2021, much of this as a result of expansion of seed, linseed (from which the name linolenic acid is derived),
rapeseed for biodiesel in the European Union. World con- safflower, and mustard seed. Many other crops can be used for
sumption of oilseed meal is also expected to increase by 23%, oilseed production including castor beans, grape seed, tobacco
which compared with the past decade reflects a slowing down seed, flax, corn oil, tung beans, and okra. The oil and protein
with livestock industries growing at a slower pace. Soybean content of the major oilseed crops is shown in Table 1.
meal dominates the market although it is expected that future From these crops, plant breeders have developed specialty
meal production from sunflower and rapeseed will increase oil types within the different species. Sunflower oil, for exam-
relative to soybean. Much of the meal is utilized as stock feed ple, is traditionally high in linoleic (polyunsaturated) acid and
and in the production of livestock products, in particular pork is promoted as such for its health benefits. With further plant
and poultry and, more recently, aquaculture. Meal products, breeding, there are now several grades of sunflower oil, from
such as protein extracts, are used for edible purposes, and high-linoleic acid, to high-oleic acid (monounsaturated), to
consumption of whole seed, including roasted soybeans, pea- intermediate types. Oleic acid, being monounsaturated, has
nuts, and sesame seed, is common. The meal also has indus- higher oxidative stability. In addition, the reduction in linoleic
trial applications in cosmetics, paints, and adhesives. acid means that the need for hydrogenation to reduce polyun-
saturated oils and the formation of trans-fatty acids is reduced.
Trans-fatty acids are nutritionally undesirable and similar to
Oil Analysis saturated fats in their effects. Examples of plant species, which
have been bred with increased oleic acid, include sunflowers
The traditional methods of oilseed analysis made plant breed- (high oleic acid and NuSun), linseed (solin, Linola), cot-
ing and selection a slow and tedious process. The Goldfisch or tonseed, soybean, rapeseed (canola, Monola), safflower, and
Soxhlet methods of oil extraction required several hours to mustard.
THE OILSEEDS | Oilseeds: Overview 223
Table 2 The fatty acid components and quantities in the major oilseed crops (amounts in rounded percentages)
Fatty acid Name Palmitic Palmitoleic Stearic Oleic Linoleic Linolenic Arachidic Gadoleic Behenic Erucic
Symbol C16:0 C16:1 C18:0 C18:1 C18:2 C18:3 C20:0 C20:1 C22:0 C22:1
Oilseed Type
Soybean Normal 814 <0.2 26 1730 4859 411 <0.6 <0.5 <0.7 <0.3
Soybean High oleic acid 6 3 84 2 4
Cottonseed Normal 2126 <1.2 23 1522 4758 <0.5 0.20.5 <0.1 <0.6 <0.3
Peanuts Normal 8.3 3.1 56 26 2
Rapeseed Low erucic acid 27 <0.6 13 5170 1530 514 0.21.5 0.15 <0.6 <2.0
High erucic 1.56 <3 0.53 860 1123 513 <3 315 <2 260
acid
High oleic acid 34 23 6376 1325 23 12 <0.6 <0.2
Sunflower Linoleic acid 58 <0.3 27 1440 4874 <0.3 0.10.5 <0.3 0.31.5 <0.3
Mid-oleic acid 45 34 5075 2030 <1
Oleic acid 25 <0.1 37 7591 217 <0.3 0.20.5 0.10.5 0.51.5 <0.3
Sesame Normal 812 <0.2 56 3642 4248 0.30.4 0.30.6 <0.3 <0.3
seed
Linseed Normal 57 <0.3 34 1920 1417 5261 <0.5 <0.6
Low linolenic 6 34 15 73 23
acid
Safflower Linoleic acid 58 <0.2 23 822 6883 <0.1 0.20.4 <0.3 <1.0 <1.8
Oleic acid 36 <0.2 13 7084 920 <1 0.30.6 <0.5 <0.4 <0.3
Mustard B. carinata 410 <2 823 1522 1827 <2 2050
seed B. nigra 27 <2 1027 1522 1127 3345
B. juncea 310 13 1564 1428 924 13 <40
Source: Adapted from Codex Alimentarius Commission, Codex Stan 210-1999, Amend. 2013 and other literature values.
Meal protein is high at 46%, but the amino acid profile is acid cultivars are also available, such as NuSun, with satu-
lacking in sulfur-containing amino acids particularly methio- rated fatty acid levels of 8% and only 2030% linoleic acid. The
nine. The components of whole peanuts are protein (26%), oil meal has high protein content and is used in animal feed for
(41%), and carbohydrates (24%). The main product is the oil livestock and poultry. A small percentage of the crop is used for
that has 80% unsaturated fatty acids making it a nutritionally nonoilseed production for confectionary purposes.
favorable oil. Additionally, the level of oleic acid and palmitic
acid makes it stable with good keeping qualities for cooking
Sesame Seed
and in food mixtures. Plant selection has seen the development
of new cultivars with high oleic acid levels of up to 60% and a Sesame seeds are used intact or as oil and meal. The seeds
subsequent reduction in linoleic acid content. Fatty acid pro- generally have high oil content of 50% and 25% protein,
files are influenced significantly by environmental conditions although the oil content can vary between 40% and 60%. The
with high temperatures favoring high oleic acid contents. fatty acid composition also shows a large range (Table 2).
A high oleic acid (40%) and linoleic acid (45%) content
makes the oil nutritionally beneficial. The unsaponifiable frac-
Rapeseed
tion of the oil contains sesamin and sesamolin that during the
Traditional rapeseed oil characteristically contains high levels refining process form sesamol and sesaminol. These are strong
of erucic acid, found to have detrimental effects on the myo- antioxidants that give the oil exceptional resistance to oxida-
cardial muscle of rats fed the oil. The value of the meal was also tion and rancidity. The oil-free meal is high in protein
low due to the presence of sulfur-containing compounds called (3450%), depending on the variety, and has a favorable
glucosinolates, found to have deleterious effects on the thyroid amino acid profile with high methionine and low lysine con-
gland of monogastric animals. Through plant breeding, major tent. Today, the seed is used for human consumption on bread
changes have been achieved to traditional rapeseed by increas- rolls, health food, and confectionary.
ing yield and oil while simultaneously increasing meal protein
content. The development of cultivars with very low levels of
Linseed
glucosinolate in the meal has also been achieved. With the
dramatic change in rapeseed from traditional types to those Due to its high iodine value, linseed oil has been used primar-
grown today, new terms have been used to discriminate ily for industrial purposes, such as linoleum floor covering,
between the types. Canola is recognized under ISO 5725, with a high level of unsaturated fatty acids making the oil very
Codex Alimentarius, and ISTA as cultivars of rapeseed with reactive and resulting in a short shelf life. Low linolenic acid
<2% erucic acid in the oil and < 30 mmol of aliphatic glucosi- cultivars have introduced linseed to the edible food market. In
nolates in the meal. European nomenclature to describe 1994, the Flax Council of Canada developed the term solin to
cultivars of rapeseed includes low-erucic acid rapeseed and describe linseed with < 5% linolenic acid. The original hybrid-
Colza, which is the French name for B. napus rapeseed in ization work was carried out by the CSIRO in Australia with the
general. The seed oil is the main value of the crop yielding release of two Linola cultivars in 1992 under the Plant Variety
42% oil, while the meal contains 35% protein. Current canola Rights scheme. Linola 947 was the first solin cultivar registered
cultivars have very low erucic acid and saturated fatty acid in Canada. Solin cv. Linola 989 has been reported as 46% oil
levels and a good proportion of oleic (C18:1), linoleic (dry basis) and 34% protein. Linseed meal has a high crude
(C18:2), and o-3 linolenic (C18:3) fatty acids. The balanced protein value but low lysine levels. It also has a high level of
fatty acid profile makes rapeseed ideal for mayonnaise and soluble fiber, called mucilage, that is indigestible to nonrumi-
salad dressings and a wide range of applications in the bakery nants and reduces the energy value of the meal. Linseed is
and confectionary industries. New high-oleic acid/low- traded at 40% oil although the oil can range significantly
linolenic acid types have improved oxidative stability for cook- depending on growing conditions.
ing and longer shelf life. Alternatively, high-erucic acid oil is
utilized for industrial purposes such as cutting oils. Erucic acid
Safflower
is extracted from high-erucic acid rapeseed and converted to
erucamide for plastic manufacture. Safflower was originally domesticated for its flowers: red
carthamin dye was extracted for coloring food and cloth.
Early cultivars were unsuitable for commercial development
Sunflower
due to low oil content of 30%. Breeding programs have since
Sunflower is grown for the seed oil, which is 80% of the seed increased oil contents and altered the fatty acid profiles to take
value. The dehulled meal has 2842% protein. The oil is highly it from an industrial product to the edible oil market with
considered due to its low linolenic acid value and therefore modern cultivars containing 34% or more oil. Traditional
oxidative stability for cooking, salad oil, and margarines. Her- high-linoleic acid types contain 6883% linoleic acid and
itability of high oil is reliable, and it has been possible to 822% oleic acid (Table 2). Through plant breeding, high oleic
increase oil contents from 30% in early types to over 50% in acid levels (7084%) have been achieved. Due to the wide
recent years. Sunflower cultivars with a range of fatty acid pro- range of environmental conditions from countries as diverse
files have been developed including high oleic acid (7591%) as Australia, India, and China, fatty acids may vary consider-
with reduced levels of linoleic acid (217%) and high levels of ably. The meal from safflower seed is also valued as a stock
alpha-tocopherol (vitamin E), providing the market with very feed. The protein content of the meal remains relatively low in
stable monounsaturated oil for cooking purposes. Mid-oleic relation to other oilseeds at only 23%.
THE OILSEEDS | Oilseeds: Overview 225
1000
100
Production (Mt)
10
0
05
06
07
08
09
10
11
12
20
20
20
20
20
20
20
20
Year
Figure 1 Global production of major oilseed crops (log10) from 2005 to 2012 in million tonnes (Mt). Adapted from Food and Agricultural Organization
of the United Nations Statistics.
Mustard Seed
Typically, mustard oil is high in erucic acid (2050%) and the
meal is high in glucosinolate compounds (50150 mM g 1), Table 3 World production of the major oilseed crops for 2011 and
2012
providing pungency to mustard for condiments but reducing
the palatability of the meal for stock feed. With plant breeding, Area harvested (Mha) Production (Mt)
very low levels of erucic acid have been achieved, and current
aims are to develop cultivars that are more closely related to 2011 2012 2011 2012
LEAR cultivars, with increased oleic acid content.
Soybeans 104 107 262 253
Rapeseed 34 34 63 65
Cottonseed NA NA 49 47
Production and Trade Groundnuts 25 25 40 41
Sunflower seed 26 25 41 38
For the purpose of this article, fruit oils will not be discussed Sesame seed 9 8 5 4
although production is significant. Palm oil fruit (from which Linseed 2 3 2 2
palmitic acid derives its name) production in 2012 exceeded Safflower seed 0.8 1.0 0.7 0.8
50 Mt, second only to soybean. Oilseed production from 2005 Mustard seed 0.7 0.6 0.6 0.5
to 2012 is shown in Figure 1 and the area grown and yields of NA, not available.
the various crops in 2011 and 2012 in Table 3. Source: Adapted from FAOSTAT | FAO Statistics Division 2014 | 03 February 2014.
226 THE OILSEEDS | Oilseeds: Overview
Future Developments Are the same countries always the largest producers or is
this changing? Why are some countries reducing
Oilseeds play an increasingly important role in society, both as production, while others are increasing?
an edible food product and for industrial uses. The benefits How are the uses of oil, meal, and oilseeds changing? Look
include the valuable oil component and the secondary but also at the new applications for these products, which differ
useful meal that provides high-energy and nutritionally impor- from the original use for food.
tant food. Breeders have been able to optimize the characteris- What will be the impact of industrial uses, particularly
tics of many of these crops to achieve higher levels of biodiesel to the food supply in the future?
production and to alter the products to suit specific require-
ments. These changes include improvements in nutritional
value, particularly with changes in fatty acid profiles and the See also: Agronomy of Grain Growing: Canola: Agronomy;
reduction in antinutritional components. Undoubtedly the Soybean: Agronomy; The Legumes and Pseudocereals: Peanuts:
biggest changes will come in the future from genetic modifica- Overview; The Oilseeds: Cottonseed: Overview; Sunflower: Overview.
tion in which long-term breeding programs can be dramati-
cally reduced to relatively short periods of time. Traditional,
breeding techniques rely on natural variability in plant popu- Further Reading
lations to achieve change. Genetic engineering can create new
products previously out of the reach of established breeding Appelqvist L-A and Ohlson R (eds.) (1972) Rapeseed: Cultivation, Composition
Processing and Utilization Amsterdam: Elsevier.
techniques. Engineering plants for specific aims or more suit- Benedict JH, Treacy MF, and Kinard DH (eds.) (1994) Vegetable Oils and Agrichemicals
able to environmental conditions can increase production. Memphis, TN: Cotton Foundation and National Cottonseed Products Association.
Nutritionally, fatty acid profiles can be dramatically altered to Bennett M (1998) Sesame seed. In: A Handbook for Farmers and Investors Australia:
reduce saturated and trans-fatty acids and simultaneously Rural Industries Research and Development Corporation.
FAO (2003) FAO Statistical Database. Rome: Food and Agriculture Organization of the
improve oxidative stability. Vegetable oils are already being
United Nations.
substituted, in many instances, for less environmentally Hui YH (ed.) (1996) Baileys Industrial Oil and Fat Products, 5th edn., Series 3. New
friendly petroleum products. The major competition to oilseed York: Wiley ISBN: 047159427X.
crops in the future will only be from the ever-growing range of Karleskind A (1996) Oil and Fats Manual. France: Lavoisier Publishing.
additional oilseed types. KeShun L (1997) Soybeans: Chemistry, Technology and Utilization. New York:
Chapman and Hall, ISBN: 0-8342-1299-4.
Kimber DS and McGregor DI (eds.) (1995) Brassica Oilseeds, Production and
Utilisation Wallingford: CAB International.
Robbelen G, Downey RK, and Ashri A (1989) Oil Crops of the World. New York:
Exercises for Revision
McGraw-Hill.
Weiss E (2000) Oilseed Crops, 2nd edn. Malden, MA: Iowa State Press.
Discuss the major oilseed crops of the world and their uses. Woodroof JG (1983) Peanut Production, Processing, Products, 3rd edn.
Which are the major producing and importing countries for Westport, CT: AVI.
each species of oilseed?
Consider and discuss what changes have been made to
make oilseed crops more useful? Relevant Websites
Evaluate the importance of fruit oils and how do they rate
http://www.amsoy.org American Soybean Association.
against oilseed crops? http://www.aocs.org American Oil Chemists Society.
Compare oilseeds that are used for oil with the ones that are http://www.apps.fao.org Food and Agriculture Organization.
grown for their high-protein meal and consider why? http://www.australianoilseeds.com Australian Oilseed Federation.
http://www.australianoilseeds.com 7th International Safflower Conference.
http://www.canola-council.org Canola Council of Canada.
http://www.codexalimentarius.net Codex Alimentarius Commission.
Exercises for Readers to Explore the Topic Further http://www.cottonseed.com National Cottonseed Products Association.
http://www.flaxcouncil.ca Flax Council of Canada.
http://www.grainscanada.gc.ca Canadian Grain Commission.
Consider the changes in oilseed production over years and www.sunflowernsa.com National Sunflower Association.
the ability of breeders and producers to keep pace with the www.regional.org.au 10th International Rapeseed Congress.
demand of an increasing population. http://www.rirdc.gov.au Australian Rural Industries and Research Foundation.
Soybean: Overview
K Liu, Agricultural Research Service, US Department of Agriculture, Aberdeen, ID, USA
2016 Elsevier Ltd. All rights reserved.
Topic Highlights During the same period, soybean production in the United
States increased from about 9 to 106 MMT and in Brazil and
Soybean is an ancient Chinese grain and now a top oilseed Argentina from virtually negligible amounts to 94 and
crop, with world annual production reaching more than 55 MMT, respectively, while soybean production in China
300 million tonnes. went up and down within a small range of 1015 MMT.
Soybean is also a legume, having the ability to fix nitrogen These differential changes have made the United States, Brazil,
from air through its roots. and Argentina the top three soybean producers and China
Soybean has the highest protein content (40%) and the from the worlds number 1 prior to the 1950s to the worlds
second highest oil content (20%) among food legumes. It number 4 soybean producer at the present time.
also contains an array of phytochemicals. There are several reasons for the rapid and sustainable
Soybean has a versatile use as food, feed, and industrial worldwide growth in soybean production. First, soybeans
material. have many possible end uses, including as human food, ani-
Soyfoods come in various types: traditional, modern mal feed, and industrial materials. Second, advances in plant
(Westernized), fermented, and nonfermented. breeding and production management have led to a steady
Soyfoods are not only nutritious but also healthy since increase in production yields in the field and improvements
regular consumption of soyfoods can help prevent certain in the seed quality. This is particularly true in recent years when
chronic diseases. application of crop biotechnology has resulted in the develop-
The majority of soybean produced annually is currently ment of new soybean varieties that are resistant to certain
crushed into oil for edible consumption and defatted diseases and pests, tolerant to herbicides, and/or improved in
meal mostly for animal feed. Only a small portion of seed composition. Third, the soybean is capable of producing a
annual production is used for whole soy food or soybean higher amount of edible protein than any other crops per
protein ingredients. hectare of land while producing a significant amount of edible
oil. Fourth, like other members of the legume family, soybeans
have an ability to fix nitrogen due to its characteristic symbiosis
Learning Objective with root bacteroids. This feature makes the soybean a good
rotational crop. Fifth, recent medical discoveries have linked
To have a basic understanding of soybeans with respect to the consumption of soyfoods to the prevention and treatment
its history, current production, agronomic traits, chemistry, of chronic diseases such as heart diseases, cancer, and bone
nutrition, processing, and various end uses diseases. This new development helps promote soybean utili-
zation as food. Finally, growth and globalization of world
economy and trades have been another important driving
force. We are seeing a rapid interchanging trend with respect
Introduction
to soybean utilization between China and the rest of the world.
In China, where soybeans originated and traditional soyfoods
From one of the five sacred grains in ancient China to one of
are popular, rapid economic development in recent years has
top cash crops, the soybean (Figure 1) has played a very
led to increased demand for soybeans as refined oil and animal
important role in nourishing humans and animals. For the
feed. In the rest of world, particularly in North America and
past several decades, progress has been made on many fronts
Europe, where historically soybeans are crushed for edible oil
of soybean research, from production management to variety
and animal feed, interest in soyfoods and soy-enriched food
development, from genome mapping to compositional modi-
has experienced a phenomenal rise over the past 2 decades.
fication, from processing to new product development, from
health benefits to identification of key responsible compo-
nents, and from animal nutrition to new industrial uses.
Since there are so many topics to be covered on soybeans, it Origins
is impossible to give a detailed account in this article. Readers Historical and geographic evidence indicates that soybeans
are advised to refer to several specific, topical chapters on originated in northern China and that its cultivation in the
soybeans as well as further reading under each article. region started approximately 5000 years ago. The soybean was
considered one of the five sacred grains, along with rice, wheat,
barley, and millets. It was repeatedly mentioned in later
Production
records. The Chinese gradually developed many nutritious
For the past six decades, the soybean has emerged as one of the foods out of soybeans, including soy milk, tofu, soy sprouts,
most important agricultural crops in the world, with a steady fermented soy paste, and soy sauce (Figure 3). Soybean culti-
increase in annual production. In 1955, the world soybean vation and soyfood preparation methods were gradually intro-
production was only about 20 million tonnes (MT). In 2014, duced into Japan, Korea, and other neighboring countries
global production increased to about 311 MMT (Figure 2). about 1100 years ago. People in these countries not only
accepted the Chinese way of preparing soyfoods but also cre- Agronomic Characteristics
ated new soyfoods. Japanese natto and Indonesian tempeh are
two good examples. All these foods are now known as tradi- The soybean belongs to the family Leguminosae. The culti-
tional soyfoods. vated form, Glycine max (L.) Merrill, grows annually. Its plant
The soybean and soybean products were introduced into is bushy, with a height between 0.2 and 2.0 m, and produces a
Europe during the Age of Discovery (sixteenth to seventeenth fruit having three to five hairy pods in a cluster (Figure 4). Each
centuries) and into the North America during the mid- pod, 38 cm long, contains two to four seeds that are spherical-
eighteenth century. Yet, large-scale official introduction into to oval-shaped. Most seeds are yellow, but they can also be
the United States did not occur until late nineteenth century green, dark brown, purplish black, or black.
and early twentieth century, during which the US Department Most mature plant seeds are made of three basic parts: the
of Agriculture developed a national soybean germplasm col- seed coat, embryo, and one or more food storage structures
lection, and thousands of new varieties were collected, mostly (such as the endosperm). The soybean seed, however, like
from China. Currently, the collection contains over 20 000 other legume seeds, is essentially devoid of endosperm and
strains, serving as a primary genetic source for soybean research consists of only a seed coat and a large, well-developed
and variety development.
Figure 3 Traditional soyfoods: soy milk, soy sauce, soy milk film, soy
Figure 1 Soybean seeds. sprouts, tofu, and okara (soy fiber).
USA
250
Brazil
Argentina
200
Million tonnes
China
150 India
Paraguay
100 Canada
Others
50
0
1955 1960 1970 1980 1990 2000 2010 2014
Production Year
Figure 2 Soybean production around the world. Data extracted from oil crops outlook, Economic Research Service, US Department of Agriculture,
Washington, DC.
230 THE OILSEEDS | Soybean: Overview
structural components found mainly in cell walls. Soluble soy-based feed has been a practical way to address the problem.
carbohydrates include measurable amounts of di- and oligo- Excess P due to unavailability or oversupplementation in ani-
saccharides, with sucrose in the range of 2.58.2%; raffinose, mal feed is excreted in feces, raising environmental concerns.
0.10.9%; and stachyose, 1.44.1%, and trace amounts of Among soybean carbohydrates, raffinose and stachyose gar-
monosaccharides, such as glucose and arabinose. The oligosac- ner the most attention, mainly because human and monogas-
charides are nonreducing sugars, containing fructose, glucose, tric animals cannot digest these sugars well due to the lack of a
and galactose as three or four units, linked by beta-fructosidic specific, endogenous alpha-galactosidase. As a result, their
and alpha-galactosidic linkages. presence has been linked to flatulence and abdominal discom-
The content of major minerals in soybeans ranges from fort associated with soybean consumption.
0.2% to 2.1%. Among them, potassium is in the highest con- Phospholipids are polar lipids, constituting 0.51.5% of
centration, followed by phosphorus, magnesium, sulfur, cal- the soybean seed and 13% of crude soybean oil. The com-
cium, chloride, and sodium. The minor minerals present in mercial product of phospholipids is known as lecithin, a major
soybeans are silicon, iron, zinc, manganese, copper, molybde- by-product of the soybean oil refining process. Phospholipids
num, fluorine, chromium, selenium, cobalt, cadmium, lead, are good emulsifying agents, soluble in alcohol and insoluble
arsenic, mercury, and iodine. Their content ranges from 0.01 in acetone. In living tissues, they are the major components of
to 140 ppm. Soybeans also contain both water-soluble and cell membranes.
oil-soluble vitamins. The water-soluble vitamins present in Isoflavones belong to a group of compounds that share a
soybeans include thiamin, riboflavin, niacin, pantothenic basic structure consisting of two benzyl rings joined by a three-
acid, and folic acid. The oil-soluble vitamins present in soy- carbon bridge that may or may not be closed in a pyran ring.
beans are vitamins A and E with essentially no measurable The soybean is unique in that it contains the highest amount of
vitamins D and K. isoflavones, being up to 3 mg g 1 dry weight. Among all the
health-promoting components, isoflavones are thought to be
most responsible for many of the hypothesized health benefits
Biological Active Components
of soyfoods and thus have gained the most attention in the
There are an unusually large number of heat-stable and heat- scientific community.
labile substances that are naturally found in soybeans. These Soybeans contain 0.10.3% saponins, which are composed
compounds, although present in a trace amount, are capable of of sugars bound to alkaloid, steroid, or triterpene compounds
eliciting diverse nutritional, biological, and physiological and have detergent surfactant properties. Saponins have blood
responses in humans and animals. These biological and phys- cholesterol-lowering properties rendered by their binding of
iological responses can be adverse or beneficial to human and cholesterol. The bound cholesterol is then passed to the colon
animals, and as such, some substances are known as anti- and excreted.
nutritional factors, while others are referred to as phytochem- Phytosterols are lipid-like compounds found in plants.
icals or nutraceuticals. Sometimes, the same substances may Soybeans, rapeseeds, and coniferous trees are the three major
exert both adverse and beneficial effects, depending on the commercial sources of phytosterols. The total phytosterol con-
dosage and specific application. Due to page limitations, only tent of soybeans is estimated at 0.30.6 mg g 1. Soybean ste-
a few key compounds are briefly described in the succeeding rols are obtained during oil processing as by-products of
text. vitamin E manufacturing. Although phytosterols are structur-
Proteinase inhibitors present in soybeans fall into two ally related to cholesterol, they have been clinically proven to
types: Kunitz trypsin inhibitor and BowmanBirk inhibitor. reduce blood cholesterol in our bodies.
The significance of soybean trypsin inhibitors lies in their
nutritional implications toward both humans and animals.
Nutritional Values
Unless inactivated by heat, these inhibitors can cause growth
depression by reducing the digestibility of proteins. In addition to their high content in soybeans, soybean oil and
Lectin, also known as hemagglutinin, is another antinutri- protein are both of high quality. Like other edible oils, aside
tional factor that is of protein nature found in soybeans. It from culinary roles, soybean oil provides us with calories,
possesses a remarkable ability to agglutinate erythrocytes and essential fatty acids, and fat-soluble vitamins. Since soybean
other types of cells. As demonstrated by animal studies, soy- oil contains very high percentages of unsaturated fatty acids, it
bean lectin is linked to an enlargement of the pancreas, a is an excellent source of essential fatty acids.
lowering of blood insulin levels, an inhibition of disacchari- Soybean protein contains all the identified essential amino
dase and proteases in the intestines, and degenerative changes acids for animals: isoleucine, leucine, lysine, methionine,
in the liver and kidneys, in addition to growth inhibition. cysteine, phenylalanine and tyrosine, threonine, tryptophan,
Phytate is the calciummagnesiumpotassium salt of ino- valine, and histidine. Most of these amino acids are present in
sitol hexaphosphoric acid, which is commonly known as phy- amounts that closely match those required for humans and
tic acid or phytin in the literature. As in most seeds, phytate is other animals. The only potentially limiting amino acids are
the principal source of phosphorus in soybeans. Its content sulfur-containing ones, primarily methionine and cysteine.
ranges from 1.00% to 1.47% on a dry matter basis. This value Yet, such deficiency is increasingly considered less problematic,
represents a little more than 50% of the total phosphorus in with respect to human nutrition. Protein quality depends on
soybean seeds and 6070% of total phosphorus in defatted the amino acid composition of the protein, amino acid
soybean meal. Phytate P is nutritionally unavailable to mono- requirements for humans or animals fed with the protein, the
gastric animals. Supplementation with inorganic P into active amounts of antinutritional factors, digestibility of the
232 THE OILSEEDS | Soybean: Overview
components. It is the final step after general processing. The Traditional Soyfoods
final product is refined, bleached, and deodorized (RBD) oil,
Over the course of several thousand years, long before the
commonly known as RBD oil. Yet, due to its high concentra-
modern processing emerged in the twentieth century, soybeans
tions of polyunsaturated fatty acids, RBD oil is prone to oxida-
had been processed into various soyfoods, known as tradi-
tion and off-flavor formation. It also lacks certain physical and
tional soyfoods. Since these foods were created in China and
functional properties. To overcome these limitations, addi-
neighboring countries, they are also known as Oriental soy-
tional processes (collectively known as soybean oil modifica-
foods. In general, traditional soyfoods are classified as non-
tion), such as hydrogenation, interesterification, winterization,
fermented and fermented. Nonfermented soyfoods include soy
and/or fractionation, may also be used.
milk, tofu, soy sprouts, yuba (soy milk film), okara (soy pulp),
immature soybeans, soynuts, and toasted soy flour (Figure 3).
Traditionally, soy milk is made in China by soaking, rins-
Alternative Processes ing, and grinding soybeans into slurry. This is followed by
filtering the slurry to separate the residue (okara) and cooking
There has been increased concern over the past several decades
the extract to become edible. As shown in Figure 7, the pro-
with the safety and environmental impact of hexane solvent-
cesses for making tofu, soy milk film, and okara are directly
based extraction. Several alternative oilseed milling processes
linked to soy milk production; tofu and soy milk film all start
have been available or under development. Screw pressing or
with soy milk making while okara is a by-product of soy milk
expelling is an old way of processing soybeans and other
making. To make tofu, soy milk is boiled for about 10 min,
oilseeds, which existed long before solvent extraction was
and a coagulant suspension is added to form curds, which is
developed. The method, also known as mechanical extraction,
then shaped and pressed into tofu.
has relatively low oil removal efficiency (leaving 48% residual
There are four major fermented soyfoods fermented soy
oil in the meal) and low production efficiency. The advantages
paste (miso), soy sauce, tempeh, and natto and two minor
are low initial cost and no solvent requirements.
fermented soyfoods: sufu (fermented tofu) and soy nuggets
Dry extrusion is another alternative processing method for
(fermented whole soybeans). They vary greatly in the methods
soybeans. It was developed in the 1960s to enable Midwestern
of preparation, microorganisms involved, and utilization.
US soybean growers to cook soybeans for use as livestock feed
While it takes less than a week to prepare tempeh and natto,
right on the farm where soybeans were produced. The process
the preparation of the remaining types of fermented soyfoods
generates friction as the sole source of heat to deactivate anti-
generally requires several months to complete. Except for
nutrients in soybeans. It does not remove any oil, though. Yet,
natto, which results from bacterial (Bacillus natto) fermenta-
when dry extrusion is combined with expelling, the through-
tion, all others are fermented mainly by fungi: soy paste, soy
put of the mechanical extraction is greatly improved. The pro-
sauce, and soy nuggets are all fermented by Aspergillus oryzae;
cess, known as extrudingexpelling, was developed in the
tempeh by Rhizopus oligosporus; and sufu by Mucor sp., Actino-
1980s and has become the primary choice for alternative pro-
mucor sp., and others.
cessing of soybeans.
Almost all nonfermented soyfoods are consumed primarily
Another alternative method for processing soybeans is
for nourishment. In contrast, the majority of fermented
aqueous extraction process. The method differs radically from
standard oil milling processes in that an aqueous system is
used to extract oil. Although the concept and general features
of the aqueous method appear attractive compared to the Whole soybeans
conventional hexane-based method, the comparatively low
oil yield and relatively high content of oil in the remaining Soaking
fractions have discouraged its commercial application.
Water Rinsing
Grinding
The soybean is the most versatile grain in terms of its end use. Cooking
Broadly speaking, it can be used as human food, animal feed,
and industrial materials. At the present, the majority of the Soymilk
annual soybean production is crushed, and its oil is solvent-
extracted with hexane for edible use. The defatted soybean Coagulant Coagulating
meal is generally toasted and serves as feed for various farm
animals and fish. A very small proportion of annual soybean Pressing (optional) Whey
production is processed into many types of traditional and
modern soyfoods. In addition, both the oil and protein meal Tofu
have been explored for a broad range of industrial uses, includ- Figure 7 Outline of soy milk and tofu making. Adapted from Liu K
ing paints, plastics, and adhesives. Most notably, soybean oil (1997) Soybeans: Chemistry, Technology, and Utilization. New York:
has been used as a feedstock for producing biodiesel to replace Chapman & Hall, Figure 4.2, with kind permission of Springer Science &
petroleum-based transportation fuels. Business Media.
234 THE OILSEEDS | Soybean: Overview
soyfoods contain salt and/or by-products (such as lactic acid) targeted for specific applications for their functional advan-
from a desirable fermentation. Both inhibit, delay, or slow tages over common soybean oil. Such effort will provide ben-
down spoilage of these products and allow them to have efits to food and industrial users and could increase the value
a relatively long shelf lives. They are generally served as sea- of soybeans as an oilseed.
sonings or condiments in cooking or making soups and con-
tribute more in flavor than in nutrition to the diet. Tempeh and
Soybean Protein Products
natto are two exceptions, which have short shelf lives, are often
consumed as a part of the main meal, and contribute protein The largest nutritionally valuable component present in soy-
and oil to the diet in addition to their characteristic flavor. beans is protein. Unlike soybean oil, which is used mainly for
In commercial processing, most traditional ways of making human consumption, soybean protein is used largely as feed-
soyfoods are now modernized. For example, in making mod- stuff. Only a small portion is for direct human consumption as
ern soy milk, several key steps are modified or added to reduce either traditional soyfoods or protein ingredients. Therefore,
beany flavor, increase production yield, and improve overall soybean protein has been severely underutilized in terms of
product quality and consumer acceptance, even though the human consumption.
basic principle is very similar to the traditional Chinese Soybean protein products are mostly made from defatted
method. The new steps include dehulling, blanching, fine soy flakes, the main coproduct of modern soybean processing.
milling, centrifuging, formulation, fortification, homogeniza- They are not consumed directly as food, but as versatile ingre-
tion, thermal processing, and aseptic packaging. dients used in virtually every type of food system, including
bakery, dairy, meat, breakfast cereal, beverages, infant formu-
las, and dairy and meat analogs. In these food systems, they not
Soybean Oil
only boost protein content but also provide many functional
Nearly all soybeans are processed for their oil and defatted properties, including gelling, emulsifying, water-holding, and
meal. With rare exceptions, crude soybean oil will have been fat-absorbing properties. There are four major types of soybean
refined, bleached, and deodorized to produce a high-quality protein products: flour, concentrates, isolates, and textured
RBD oil. About 85% of soybean oil is used for human con- soybean protein.
sumption. The rest is sold for biodiesel production or indus- Soy flour is one of the least-processed soybean protein
trial uses. products. Many types are available in the market, including
Soybean oil is the leading edible oil and widely used in defatted, full-fat, and low-fat, as well as enzyme-active, toasted,
various food products. Among them are salad and cooking oil, and textured. Defatted soy flour has been the most common
shortening, margarine, mayonnaise, and salad dressing. Other type. It is produced by simply grinding defatted soy flakes after
applications include uses in canned foods, confectionery coat- modern soybean processing, containing about 50% protein.
ings, pudding mixes, pancake and waffle mixes, macaroni and Full-fat soy flour is made by grinding dehulled whole soy-
cheese mixes, spaghetti sauces, dry breakfast cereals, frozen- beans. Low-fat soy flour can be made by extrudingexpelling
fried seafood, meat patties, pizza mixes, and others. soybeans, followed by milling the cake. In general, soy flour is
Since soybean oil contains relatively high proportions of primarily used as an ingredient in the bakery industry.
unsaturated linoleic and linolenic acids, it is susceptible to off- Soybean protein concentrate is traditionally made by aque-
flavor development. It also lacks some physical properties for ous alcohol extraction of defatted soy flakes. The resulting
certain applications, such as shortening and margarine. To product has about 70% protein with the remaining portion
increase oxidative stability and the melting point of soybean being mainly insoluble carbohydrates. The product may be
oil, hydrogenation was commonly used until a few years ago. further processed by thermal processing and homogenization
The process chemically adds two hydrogen atoms to a double for better functionality. Soybean protein concentrate is widely
bond of an unsaturated fatty acid in the oil. As a result, lower used in the meat industry to bind water and emulsify fat and as
unsaturated fatty acids are converted to saturated acids and a key ingredient for many meat alternatives. It is also used for
higher unsaturated fatty acid to lower unsaturated acids in protein fortification of various types of food.
the oil. However, hydrogenation causes the formation of Soybean protein isolate is produced by alkaline extraction
trans-fatty acids. In recent years, the adverse health effects of followed by precipitation at an acid pH. The process removes
trans fat has been well established, causing US regulatory agen- both soluble and insoluble carbohydrates. The resulting prod-
cies to call for reducing or even totally eliminating trans fat uct has a protein content of 90% and is light in color and bland
from foods commercially prepared for human consumption. in flavor. It has many functional properties, including gelation
This has created a great challenge for the food industry to find and emulsification. As the most refined soybean protein prod-
alternatives for meeting different application needs. uct, soybean protein isolate is used in processed meat, meat
One effective way to solve the trans fat issue, while keeping analogs, soup and sauce bases, nutritional beverages, infant
the stability and functional properties of soybean oil required formulas, and dairy replacement.
for many edible applications, is through plant breeding and Textured soybean proteins are made by a process that
genetic engineering. Already, several genotypes possessing imparts a structure, such as fiber, to a proteinaceous material.
large variation in fatty acid composition have been successfully The process is known as protein texturization. It is a unique
developed or released as commercial varieties with yields com- way to make vegetable proteins palatable. Among several pro-
parative to other soybean varieties in the United States. Among cesses, thermoplastic extrusion has been the most popular one
them are soybeans with linolenic acid as low as 3%, stearic as for soybean protein texturization. In a typical thermoplastic
high as 20%, and oleic as high as 80%. Oils from these lines are extrusion process, dry proteinaceous materials, predominantly
THE OILSEEDS | Soybean: Overview 235
defatted soy flour or soybean protein concentrate, are mixed livestock, representing two-thirds of the total world output of
with water, salts, flavorings, and/or colorants and then fed into protein feedstuffs, which include all other major oilseed meals
an extruder. Under high temperature and low moisture and fish meal. Over half of soybean meal processed for live-
(<30%) conditions, the extrudate expands rapidly upon stock feed is fed to poultry, about one-quarter is fed to swine,
emerging from the die. The final product, having porous tex- and the rest is used for beef cattle, dairy cattle, and pet food.
ture, comes in a variety of shapes, sizes, and colors. Its use Soybean protein is increasingly found in fish food, for both
ranges from meat extenders to meat analogs. It must be rehy- home aquariums and commercial aquaculture. Furthermore,
drated with water before use. limited availability and increasing cost of fish meal have led to
the use of soybean meal as a dietary protein source for a variety
of cultured marine species as well.
Western Soyfoods
Traditional soyfoods, along with soybeans, were officially
introduced to the West during the nineteenth and twentieth Soybean as Industrial Materials: The New Use
centuries. Since then, some of them, such as soy sauce, have While the soybean has been traditionally used as food for
been accepted by Westerners, whereas many others have been human consumption and is primarily used as animal feed at
modified to suite local food culture and tastes. Furthermore, present, its versatility and chemical composition make it an
many new soyfoods have been developed. The modified and attractive material for industrial uses as well. In recent years,
new soyfoods are known collectively as second-generation or the desire for biodegradable and less toxic polymers from
modern soyfoods. They can also be called Western or Western- renewable resources instead of fossil-based chemicals has
ized soyfoods, simply because they were developed in the West been intensified. This has stimulated research on using soy-
and may look or taste like Western foods (Figure 8). Examples bean as an industrial material. About 15% of soybean oil is
include soy ice cream, soy yogurts, soy cheese, soy butter, soy now used as feedstock for biodiesel, emulsifiers, lubricants,
puddings, tofu spreads, soy burgers, meatless meatballs, imita- plasticizers, surfactants, plastics, solvents, resins, and more.
tion bacon bits, and imitation chicken strips. Soybean and other vegetable oils have desirable environmental
qualities, such as being inherently biodegradable and renew-
able, having low ecotoxicity and low toxicity toward humans,
Soybean as Animal Feed and contributing no volatile organic chemicals. For example,
The majority of soybeans are used as animal feed mainly in the soy-based engine oil emits fewer harmful emissions than
form of soybean meal, which is the coproduct of modern petrochemical-based oil for increased worker safety. Soy ink
soybean processing. On a historical note, soybean meal has is superior to petroleum-based inks, because it is not toxic,
been an accepted part of livestock and poultry diets in the renewable, and environmentally friendly. When soybean oil
United States since the mid-1930s. Soybean feed production is converted into fatty acid methyl ester through transesterifica-
expanded rapidly in the mid-1970s and further accelerated in tion with methanol, the product is biodiesel. Soy biodiesel is
the early 1990s due to a growing demand for animal feed from nontoxic, renewable, and environmentally friendly.
developing countries. Prohibiting the use of slaughterhouse Soybean protein products, particularly defatted soybean
by-products in animal feeds and expanding aquaculture indus- meal, also have industrial applications, mostly as adhesives
try have also fueled the demand for this high-quality protein and paper coatings. These applications are essentially based
source. on such functional properties of soybean protein products,
Soybean meal is characterized by high protein content, low including adhesive strength, texture modification, film form-
crude fiber, and high palatability. It contains high amounts of ing, heat resistance, water retention, emulsification, and pro-
lysine, tryptophan, threonine, and isoleucine that are often tective colloid properties. For example, biocomposites are
lacking in cereal grains. Because of these features, soybean building materials made from recycled newspaper and soy
meal is the most important protein source used to feed flour. They replace other products traditionally made from
wood, such as furniture, flooring, and countertops. Particle-
board, laminated plywood, and finger-jointed lumber are
made with soy-based wood adhesives. Soy-based foams are
currently being developed for use in coolers, refrigerators,
automotive interiors, and even footwear. Soybean products
are also found in many popular brands of home and commer-
cial carpet and in auto upholstery applications.
protein for humans and animals and an important renewable Food Grains and the Consumer: Genetically Modified Grains and
source of industrial materials in the twenty-first century. the Consumer; Food Grains and Well-being: Nutrition: Soy-Based
Foods; Grain Marketing and Grading: Soybean: Grading and
Marketing; Grains Around the World: Grain Production and
Exercises for Revision Consumption: Oilseeds in North America;Non-wheat Foods:
Soybean: Soy-Based Fermented Foods; Soybean: Soymilk, Tofu, and
Why is soybean considered an oilseed? What differentiates Okara; Oilseed and Legume Processing: Soybean: Processing;
it from other oilseeds? Soybean: Soy Concentrates and Isolates; Scientific Organizations
What features make soybean a legume? Related to Grains: Research Organizations of the World: CGIAR;
Why has soybean become a top cash crop in the world since The Basics: Taxonomic Classification of Grain Species; The Grain
the middle of the last century? Chain: The Route from Genes to Grain-Based Products; The Grains that
If the typical soybean oil is healthy, why do we need to Feed the World; The Legumes and Pseudocereals: Grain
modify it through processing and/or genetic manipulation? Legumes and Their Dietary Impact: Overview; The Oilseeds: Oilseeds:
Why are the majority of traditional soyfoods not favored in Overview; The Basics: The Grain Crops: An Overview.
the West, and why do soyfoods have to be Westernized or
localized?
Why is soybean prized for its use as animal feed at the
present time? Further Reading
Do you think that genetically engineered soybeans are the
Anderson JW, Johnstone BM, and Cook-Newell ML (1995) Meta-analysis of the effects
future for improving soybean production and quality?
of soy protein intake on serum lipids. New England Journal of Medicine 333: 276.
Which type of soybean uses do you think will increase in Campbell KA, Glatz CE, Johnson LA, et al. (2011) Advances in aqueous extraction
the future: food, feed, or industrial material? processing of soybeans. Journal of the American Oil Chemists Society
88: 449465.
Chen KI, Erh MH, Su NW, Liu WH, Chou CC, and Cheng KC (2012) Soyfoods and
soybean products: From traditional use to modern applications. Applied
Exercises for Readers to Explore the Topic Further Microbiology and Biotechnology 96: 922.
Erickson DR (ed.) (1995) Practical Handbook of Soybean Processing and Utilization.
Currently, the food industry is making an effort to ban or Champaign, IL: AOCS Press.
eliminate trans fats from food. What are the best solutions Isanga J and Zhang GN (2008) Soybean bioactive components and their implications to
health A review. Food Reviews International 24: 252276.
for the soybean industry to meet this challenge?
Johnson LA, White PJ, and Galloway R (eds.) (2008) Soybeans Chemistry, Production,
Soyfood consumption in the West has been increasing Processing and Utilization. Champaign, IL: AOCS Press.
significantly for the past 23 decades. However, compared Liu K (1997) Soybeans: Chemistry, Technology, and Utilization. New York: Chapman &
with other foods, the room for growth is still huge. What Hall.
efforts need to be undertaken to further increase soyfood Liu K (ed.) (2005) Soybeans as Functional Foods and Ingredients. Champaign, IL:
AOCS Press.
consumption? Messina M and Lane B (2007) Soy protein, soybean isoflavones and coronary heart
Industrial use of soybeans has been considered new. What disease risk: Where do we stand? Future Lipidology 2: 5574.
research needs to be carried out to further increase this type Sikora S, Blazinkov M, Babic K, Sudaric A, and Redzepovic S (2008) Symbiotic nitrogen
of use? fixation and sustainable soybean production. Cereal Research Communications
36: 14831486.
Stupar RM and Specht JE (2013) Insights from the soybean (Glycine max and Glycine
soja) genome: Past, present, and future. Advances in Agronomy 118: 177204.
See also: Agronomy of Grain Growing: Soybean: Agronomy; Wang X, Cai X, and Zhang J, et al. (eds.) (1997) Zhong Guo Da Dou Zhi Ping (Chinese
Breeding of Grains: Soybean: Germplasm, Breeding, and Genetics; Soybean Food Products) (in Chinese). Beijing: China Light Industry Publisher.
Canola: Overview
VJ Barthet, Canadian Grain Commission, Winnipeg, MB, Canada
2016 Elsevier Ltd. All rights reserved.
Topic Highlights sunflower). Oil uses became easier since work on rapeseed oil
processing, refining, bleaching, and deodorizing showed that
Development of rapeseed into Canola rapeseed oil could be used as an edible oil, as soybean oil was.
Canola production in Canada since 1943 This led to the use of rapeseed oil as an edible oil in Canada.
Canola, countries exporting and importing low-erucic-acid However, work done in the early 1950s by Caroll on rats
(LEA) rapeseed showed that rapeseed oil, and erucic acid in particular, had a
Canola major compound contents oil, protein, and negative effect on the heath of the rats. This led, in 1956, the
glucosinolate Food and Drug Directorate of the Department of National
Main fatty acid present in Canola and LEA rapeseeds Health and Welfare (now Health Canada) to prohibit rapeseed
Canola processing and health effects oil in human food. Even though this directive was canceled
within months, it gave a very strong push to the Canadian
rapeseed breeding programs to develop a low-erucic-acid
Learning Objective (LEA) rapeseed. These breeding programs were located in the
prairies, Winnipeg with Dr. Stefansson (University of Mani-
In the following article, the reader will learn about canola, toba) and Saskatoon with Dr. Downey (Dominion Forage
its production, composition, and processing. Crop Laboratory, now Agriculture and Agri-Food Canada).
In 1958, a first variety trial was conducted with six varieties
(Argentine, Arlo, Golden, Gute, Polish, and Regina II) of rape-
Introduction seed (B. napus and B. campestris) grown in 22 locations
throughout the western provinces. This led in 1961 to the
Canola is a Canadian crop and the name canola is a trademark; first paper showing an effect of variety and environment on
it came from the contraction of Canada and ola for oil. the fatty acid composition of the oil for all fatty acids except
Canola plants belong to the Brassicaceae (Cruciferae) family linolenic acid. The paper also showed an inverse relationship
that comprises about 350 genera and 3000 species including between oleic acid content and erucic acid content for both B.
Brassica rapa (syn. B. campestris subsp. oleifera (turnip rape)), B. napus and B. campestris. The author concluded by saying sin-
napus (rutabagas), B. oleracea (cabbage and cauliflower), and B. gles plant selection and crosses should yield valuable informa-
juncea (mustard). B. campestris originated in the foothills of the tion of individual fatty acid and the development of gas liquid
Himalayan mountains; B. napus most likely originated in the chromatography (GC) makes it possible to do such studies.
Mediterranean region and was the result of natural crosses Truly, the development of LEA rapeseed became linked to the
between B. campestris and B. oleracea plants. development of GC, since precise and short-time analysis was a
requirement to analyze accurately a large number of samples.
To look for LEA rapeseed (and high), single-plant analyses
Development of Low-Glucosinolate Low-Erucic-Acid of Liho (a variety of B. napus from Germany), known for its LEA
Rapeseed content compared to other varieties, were conducted and large
variations in erucic acid content were observed (650%). This
Rapeseed culture was introduced in western Canada during led to the first LEA rapeseed plants; progeny of selected LEA
World War II (WWII). The seeds were crushed and the oil plants, grown in both Winnipeg and Saskatoon, gave plants
was used principally as a lubricant. At the time, two types of with seeds containing 0% erucic acid in their oil in both
rapeseeds were grown: B. napus, also called Argentine canola, locations. In 1963, Downey and Harvey developed a technique
and B. campestris (rapa), referred to as Polish seed. To make where only half of the seed was analyzed for the fatty acid
lubricants, the iodine value of the oil should be in the 103105 composition while the other half, formed by half of the coty-
range; it was found that B. napus was ideal, with an iodine value ledon attached to the radicle, was germinated and planted. This
close to 105, whereas Polish seeds (B. campestris) were not as innovative technique allowed growing plants with a better
suitable since the iodine value of the oil was close to 115. A knowledge and control of the oil composition. Subsequent
breeding program was begun to develop rapeseed varieties work showed that the fatty acid composition was controlled
with improved agronomy such as uniformity and lodging by the genotype of the developing seeds and not by the mater-
and an increased oil content while lowering the iodine value nal plant, and several genes were involved and their actions
of the oil. were additive. Later on, it was shown that five genes (e, Ea, Eb,
Later on, rapeseed almost disappeared from the prairies Ec, and Ed) controlled the level of erucic in B. napus in an
because the oil demands changed and other oilseed crops additive manner.
were available. However, rapeseed had shown that its culture Work was also conducted with B. campestris (rapa) to get
was quite favorable for western Canada since it needed fewer LEA B. campestris varieties. In 1964, Downey reported on the
heat units to grow than the other oilseeds (soybean or selection of inbred seed from a Polish LEA variety of
B. campestris having no erucic acid in its oil. Then, Dorrel and acid, these varieties are not canola varieties; they are low-
Downey showed that fatty acid composition (erucic acid con- glucosinolate HEAR. An important breeding program to
tent) of B. campestris was as in B. napus, controlled by the develop varieties with very high erucic acid and low glucosino-
genotype of the developing seed; however, they could not lates is still present at the University of Manitoba.
confirm their hypothesis that a single gene controls the erucic In order to increase variation and create new potential for
level. Instead, they showed that the fatty acid composition was breeding, ethyl methanesulfonate was tested on B. napus to
affected by the stage of the seed development. promote mutations. As a result, in 1987, Stellar was registered.
Simultaneously, work was started to reduce the glucosino- It was the first low-linolenic (3.3%), high-linoleic (28.9%)
late content of the seeds since several studies had shown that canola (0.1% erucic acid in the oil and 8 mmol g1 glucosino-
glucosinolates had a negative effect on animal health. All these late in the meal) variety registered in Canada.
nutritional studies were summarized in 1965 by Bowland et al.
in a report. In 1959, the first Canadian report showing signif-
icant variations of isothiocyanate contents between varieties
but not locations was published. Important differences were Production
reported between Argentine-type (B. napus L.) rapeseed and
Polish-type (B. campestris (rapa) L.) rapeseed. In 1970, Kondra Rapeseed has been an important source of edible vegetable oil
and Downey reported that, opposite to erucic acid, glucona- in Asia for almost 4000 years, used as a lighting oil and edible
pin, glucobrassicanapin, and progoitrin (the main glucosino- oil in Europe since at least the middle ages. However, in
lates) contents were determined by the genotype of the Canada, rapeseed production started during WWII as a source
maternal plant. The work was done using Bronowski, a variety of high-quality lubricant for marine engines. It was a very small
from Poland containing low levels of the three glucosinolates, production about 1000 metric tonnes in 1943 (Figure 1). The
and Target, a Canadian variety with high levels of glucosino- development of the first canola variety in 1974 led to an
lates. Work was also done in Europe with winter rape to increase in production and a switch from rapeseed to canola.
develop low-glucosinolate rapeseeds also using the Bronowski By 1977, almost 2 million tonnes of canola was produced in
variety. western Canada. Canola production increased steadily up to
This led in 1974 to the registration of Tower, the first 1995; then, the introduction of the first herbicide-resistant
summer rape (B. napus) with LEA content in the oil and low- varieties led to another jump in production (Figure 1). This
glucosinolate content of the meal. In 1977, the first B. campes- was followed by the development of new varieties, mostly
tris low in erucic acid and in total glucosinolates, called Candle, hybrid, with better agronomic traits allowing a rapid produc-
was registered in Canada by the Agriculture and Agri-Food tion increase since 2000. Starting in 2004, the demand in
Canada Saskatoon Research Centre. canola increased due to a sharp increase in the world biodiesel
In 1978, the Rapeseed Association of Canada decided to production. This demand was matched by an increase in pro-
name this new crop (low-glucosinolate LEA or double-low duction due to varieties with better yields. In 2013, western
rapeseed) canola, a name made from the condensation of Canadian canola production reached 17.96 million tonnes
can from Canada to represent the Canadian origin of the (Figure 1). Canola is considered as the Cinderella crop in
new crop and ola to follow some famous brand name exam- Canada, from a forgotten crop to a major cash crop for Cana-
ples (cola, Mazola, etc.). Canola is now defined as a rapeseed dian farmers. Today, almost all canola varieties grown in Can-
that produces an oil that must contain less than 2% erucic acid, ada are herbicide-tolerant. Three types are found, two were
and the solid component of the seed must contain less than obtained by insertion of a gene and are GMO (Roundup
30 mmol of any one or any mixture of 3-butenyl glucosinolate,
4-pentenyl glucosinolate, 2-hydroxy-3-butenyl glucosinolate,
and 2-hydroxy-4-pentenyl glucosinolate per gram of air-dry, 18,000
oil-free solid (http://www.canolacouncil.org/oil-and-meal/
16,000
what-is-canola/).
Production (metric tonnes 1000)
Hero 50.2% erucic acid in the oil and 15 mmol g1 of total Figure 1 Canadian production of rapeseed and canola in metric tonnes
glucosinolate in the meal. Due to their high levels of erucic (1000) from 1943 to 2013.
THE OILSEEDS | Canola: Overview 239
Ready and LibertyLink), and one was obtained by mutation seeded in August and harvested the next year from early July to
and is a non-GMO (Clearfield). August. Australia also grows canola, but opposite to Canada,
The world canola and LEA rapeseed production is con- Australian canola is a winter crop seeded at the end of
centrated in the European Community, Canada, and Aus- summerearly autumn and harvested the following year end
tralia (Table 1). China and India are important rapeseed of springearly summer.
producers; however, most of the rapeseed crops grown in
these countries are not low in erucic acid and low in glu-
cosinolate (Table 3). Composition
Canola/LEA rapeseed trade data are presented in Table 2.
The European Community, China, Japan, and Mexico are the Canola is marketed for its oil. Therefore, canola quality can be
main importing countries, whereas Canada, Australia, and defined by its oil content and the fatty acid composition of the
Ukraine are the main, if not the only, exporting countries. oil. Canola meal (what is left of the seed after crushing) is used
Canada is the main canola exporter; since 2004, Canada has for animal feed so the glucosinolate content is also an impor-
been responsible of 70% of the world total canola exports. The tant parameter in determining canola seed quality for feed.
United States is the main canola customer for Canada, import- Table 3 presents the proximate analysis and the relative fatty
ing an equivalent of 5 million tonnes of seeds; however, less acid composition of canola and rapeseed grown in various part
than 0.5 million tonnes of seeds is exported to the United of the world.
States from Canada. Canadian canola export to the United
States is in the form of oil, about 1.5 million tonnes of canola
oil. There is also an important export in the form of canola
meal well over 2.5 million tonnes mainly used by the US Processing and Uses
dairy industry.
Canola seeds (LEA rapeseed and rapeseed) are processed by
crushing plant to produce oil and meal. When the seed is
delivered to the crushing plant, they are stored in bins to be
Growing Conditions processed. Processing steps might depend on the crushing plan
and the country. In Canada, the weather is usually dry and
Canola, in Canada, is a summer crop. It is seeded in spring cold, so the first step of the processing is to pass the seed
(May) and harvested from August to October. In Europe and through a conditioner with heat and moisture allowing better
China, LEA rapeseed and rapeseed are winter crops; they are flaking and to heat the seed, making the oil more fluid and
Table 1 Top five canola/rapeseed producing countries and world production for the last 10 years (1000 tonnes)
Country
Year European Community China Japan Mexico Pakistan World total Canada Australia Ukraine World total
200405 224 362 2277 1196 672 5507 3756 927 160 5507
200506 712 713 2333 1259 820 7521 5846 827 357 7521
200607 433 954 2170 1065 806 7106 5426 230 755 7106
200708 1388 855 2210 1317 535 8648 5916 508 1496 8648
200809 2897 3085 2061 1077 596 11 584 7660 1111 2157 11 584
200910 2284 2059 2287 1403 940 11 146 7315 1172 1910 11 146
201011 2829 1100 2337 1492 811 10 484 7249 1507 1116 10 484
201112 3485 2487 2391 1653 932 13 071 8680 2486 1341 13 071
240 THE OILSEEDS | Canola: Overview
Table 3 Oil, protein, and total glucosinolate content ranges (dry basis) and relative fatty acid composition of canola/LEA rapeseed and rapeseed
grown in various parts of the world
easier to extract. The second step is flaking where the seeds are shown to increase milk production when compared to diets
placed into big rollers in order to break the seed structure to containing cottonseed or soybean. It is also used in poultry and
allow an easier crush and facilitate the oil extraction. A cooking swine feeds.
step follows to ease the oil extraction and to inactivate endog-
enous enzymes of the seeds. Canola contains low levels of
glucosinolates; however, it is important to ensure that they Food Uses and Health Effects
are not hydrolyzed during the processing to guarantee good
quality for the meal. The myrosinase responsible for the glu- Canola oil is considered a very healthy oil due to its fatty acid
cosinolate hydrolysis should be inactivated. Enzymes (lipases) composition. It averages about 60% oleic acid (C18:1), 20%
able to degrade the triglycerides must also be inactivated to linoleic acid (C18:2), and 10% ALA (C18:3) (Table 1). This
ensure no oil degradation occurs during the processing. Then, makes canola oil a good source of ALA with the ideal ratio
the seeds are submitted to a screw press (expeller press) that (2:1) of omega-6 (o-6) to omega-3 (o-3). When compared to
squeezes the oil out of the seeds. Not all oil is recovered at this the other vegetable oils, it has the lowest amount of saturated
step. A solvent extraction, usually with hexane, is performed on fatty acid (7% or less) compared to sunflower (12%). Studies
the expeller press cake. After solvent extraction, depending on with diets containing canola oil suggested that canola could
the crushing plan, up to 99% of the oil could be recovered reduce blood cholesterol levels and reduce the risk of cardio-
from the seeds. The last step of canola processing is a refining vascular disease. Canola oil can be used by food manufactures
process during which undesirable substances are removed to produce various kinds of salad dressing. It is also found at
from the crude oil to give the yellowish oil found in local food stores as a salad oil, which can also be used as
supermarkets. cooking oil. Margarines with canola can also be found at
Chlorophyll is present in canola seeds when the seeds are food markets and used by processors for baking.
harvested before reaching full maturity. Canadian canola seeds In response to the mandatory labeling of trans-fatty acids in
usually contain higher levels of chlorophyll because the grow- food products, canola varieties with high-stability oils
ing season is very short May to September compared to (ALA < 3%) were grown. These new oils have higher levels of
canola grown in Europe, Australia, or China, September to oleic acid (6574% vs. 60%) and lower levels of ALA (13% vs.
June. Chlorophyll is an undesirable substance in canola oil 10%) than the conventional canola oil. These high-stability
and has to be removed. Chlorophyll with its heme structure oils are used by the food industry for frying, allowing the
and the magnesium ion in the center is highly reactive and can reduction/elimination of trans-fatty acids from fried foods
oxidize unsaturated fatty acids in the oil. Canola oil contains (e.g., potato chips) making these products a healthier choice.
about 20% linoleic acid (C18:2) and 10% a-linolenic acid In North America, some chip brands and other fried snacks are
(ALA) (C18:3); these fatty acids can oxidize 10 (linoleic acid) processed using high-stability canola oil.
and 50 times (ALA) faster than oleic acid (C18:1). To protect
the canola oil from oxidation and to allow further processing
of the oil, chlorophylls have to be removed during refining. Exercises for Revision
Refined canola oil can be further processed to give other
food products such as margarines and shortenings or industrial What are the three Brassica species that can be called canola?
products such as biodiesel. What are the major producing and importing countries for
Canola meal, the leftover from the seed once the oil is canola seeds?
removed, can also be further processed. It is then used by the What is the main fatty acid in canola oil?
feed industry as pellet or mash product. Canola meal is con- What were the two compounds reduced from rapeseed to
sidered a premium ingredient in dairy diets where it has been give canola?
THE OILSEEDS | Canola: Overview 241
What is the ratio of omega-6/omega-3 for conventional Jahries G and Schafer U (2011) Rapeseed (Brassica napus) oil and its benefits for
canola? human health. In: Preedy V, Watson R, and Patel V (eds.) Nuts and Seeds in Health
Why must chlorophyll be removed from canola crude oil? and Disease Prevention, 1st ed., Chapter 114, pp. 967974. ISBN 9780123756893.
Krzymanski J and Downey RK (1969) Inheritance of fatty acid composition in winter
forms of rapeseed Brassica napus. Canadian Journal of Plant Science 49: 313319.
Le MHA, Landero JL, Beltranena E, and Zijlstra RT (2014) The effect of feeding
Exercises for Readers to Explore the Topic Further increasing inclusion of extruded Brassica juncea canola expeller on growth
performance and nutrient digestibility in weaned pigs. Animal Feed Science and
Technology 192: 7380. http://dx.doi.org/10.1016/j.anifeedsci.2014.02.006.
What are the advantages of modifying the fatty acid com- Li P, Ding X, and Zhou H (2011) Rapeseed quality improvement and monitoring plan in
position of canola seeds? China from 2000 to 2010. In: Proceeding of the 13th International Rapeseed
Why can changes in weather patterns affect the quality Congress, Prague, Czech Republic, 59 June 2011.
parameters (chlorophyll, oil, and fatty acid composition) Mag TK (1983) Canola oil processing in Canada. Journal of the American Oil Chemists
Society 60(2): 380384.
of Canadian canola seeds? Martineau R, Ouellet DR, and Lapierre H (2013) Feeding canola meal to dairy cows: A
meta-analysis on lactational responses. Journal of Dairy Science 96(3):
17011714.
See also: Breeding of Grains: Canola/Rapeseed: Genetics and Mulrooney CN, Schingoethe DJ, Kalscheur KF, and Hippen AR (2009) Canola meal
Breeding; Food Grains and the Consumer: Genetically Modified replacing distillers grains with solubles for lactating dairy cows. Journal of Dairy
Science 92(11): 56695676.
Grains and the Consumer; Grain Harvest, Storage and Transport: Scarth R, McVetty PBE, Rimmer SR, and Stefansson BR (1988) Stellar low linolenic-
Canola: Harvest, Transport, and Storage; Grains Around the World: high linoleic acid summer rape. Canadian Journal of Plant Science 68: 509511.
Grain Production and Consumption: Oilseeds in North America; Scarth R, McVetty PBE, Rimmer SR, and Stefansson BR (1991) Hero summer rape.
Oilseed and Legume Processing: Canola: Processing; The Canadian Journal of Plant Science 71: 865866.
Slominski BA, Simbaya J, Campbell LD, Rakow G, and Guenter W (1999) Nutritive value
Basics: The Grain Crops: An Overview.
for broilers of meals derived from newly developed varieties of yellow-seeded
canola. Animal Feed Science and Technology 78(34): 249262.
Stefansson BR and Hougen FW (1964) Selection of rape plants (Brassica napus) with
seed oil practically free of erucic acid. Canadian Journal of Plant Science 44(4):
Further Reading 359364.
Stefansson BR, Hougen FW, and Downey RK (1961) Note on the isolation of rape plants
Abraham V and de Man JM (1986) Hydrogenation of canola oil as affected by with seed oil free from erucic acid. Canadian Journal of Plant Science 41(1):
chlorophyll. Journal of the American Oil Chemists Society 63(9): 11851188. 218219.
Anand IJ and Downey RK (1981) A study of erucic acid alleles in digenomic rapeseeds Stefansson RR and Kondra ZP (1975) Tower summer rape. Canadian Journal of Plant
(Brassica napus L.). Canadian Journal of Plant Science 61: 199203. Science 55: 343344.
Aukema H and Campbell L (2011) Oil nutrition and utilization. In: Daun JK, Eskin NAM, Stricker H, Duchini F, Facchini M, and Mombelli G (2008) Canola oil decreases
and Hickling D (eds.) Canola, Chemistry, Production, Processing and Utilization, cholesterol and improves endothelial function in patients with peripheral arterial
pp. 245280. Urbana, IL: AOCS Press, Chapter 9. occlusive disease A pilot study. Artery Research 2(2): 6773.
Bowland JP, Clandinin DR, and Wetter LR (1965) Oil and Meal from Canadian Tautorus CL and Low NH (1993) Chemical aspects of chlorophyll breakdown products
Rapeseed. Ottawa, Canada: Department of Agriculture, Publication # 1257. https:// and their relevance to canola oil stability. Journal of the American Oil Chemists
archive.org/details/oilmealfromcanad00bell. Society 70(9): 843847.
Busch H and Robbelen G (1981) Neidriger glucosinolatgehalt als zuchtzeil fu
winterraps. Agew. Botanik 55: 361371.
Craig BM (1961) Varietal and environmental effects on rapeseed III. Fatty acid
composition of 1958 varietal test. Canadian Journal of Plant Science 41: 204210. Relevant Websites
Downey RK and Harvey BL (1963) Methods of breeding for oil quality in rape. Canadian
Journal of Plant Science 43: 271275. http://www.australianoilseeds.com/oilseeds_industry/quality_of_australian_canola
Fowler DB and Stefansson BR (1972) Effects of the mutagenic agent ethyl Australian Oilseeds Federation.
methanesulfonate on the M1 generation of rape (Brassica napus). Canadian Journal http://www.cetiom.fr/index.php?id15446 Centre Technique Interprofessionnel des
of Plant Science 52(1): 5362. Oleagineux et du Chanvre.
Fowler DB and Stefansson BR (1975) Ethyl methanesulfonate-induced mutations in http://grainscanada.gc.ca/canola/hqcm-mqrc-eng.htm Canadian Grain Commission.
rape (Brassica napus). Canadian Journal of Plant Science 55(3): 817821. http://www5.statcan.gc.ca/cansim/a26?langeng&retrLangeng&id0010010&
Harvey BL and Downey RK (1964) The inheritance of erucic acid content in rapeseed paSer&pattern&stByVal1&p11&p231&tabModedataTable&csid
(Brassica napus). Canadian Journal of Plant Science 44: 104111. Statistics Canada.
Cottonseed: Overview
E Hernandez, Texas A&M University, College Station, TX, USA
2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition, volume 1, pp 343348, 2004, Elsevier Ltd.
Topic Highlights The glands in the seed are ovoid structures containing 3550%
gossypol and are 0.0250.178 mm in diameter. These gossypol
Traditional varieties of cottonseed contain gossypol glands are difficult to break by mechanical means, but
(0.62.0% in raw cottonseed kernels). heat generated in extraction of oil by pressing, binds gossypol
Gossypol is a polyphenolic compound that is toxic to man to protein, turning it nontoxic. Switching to solvent extraction
and monogastric animals, presumably functioning as a with hexane, where no appreciable heat is generated, increases
natural insecticide for the plant. the free gossypol content in the meal over ten times.
Gossypol is located in glands, appearing as black specks Raw cottonseed kernels may contain 0.62.0% free gossy-
in the stems, leaves, and seed. pol. The Food and Drug Administration in the US (FDA) limits
The breeding of glandless cotton varieties (since the free gossypol in human food products and ingredients at
1960s) means that cottonseed is available without gossypol. 450 ppm, and the Protein Advisory Group of the United
Meal and oil from glandless cottonseed are similar in com- Nations Food and Agriculture Organization (FAO) and
position to these products from the traditional glanded World Health Organization (WHO) has set maximum guide-
seeds. lines of 600 ppm for free gossypol and 12 000 ppm total gos-
Glandless cottonseed offers the advantages for processing sypol. The feed industry in the US set limits for free gossypol
of reduced energy costs, elimination of prepress operations, levels in poultry diets at 100 ppm maximum for broilers and
and the absence of color in the resulting oil. 40 ppm for laying hens.
Food products from glandless cottonseed include whole Removal by mechanical separation of the gossypol-contai-
roasted seeds, flour, protein concentrates, and isolates. ning glands has also been reported. Also addition of iron
Glandless cottonseed is suitable to feed monogastric ani- salts, such as ferrous sulfate, which bind the gossypol in
mals, not only due to the absence of gossypol but also feeds and render it biologically inactive, is practiced in some
because of higher available energy. countries.
Solvent extraction with hexane of cottonseed oil, com-
monly used commercially, removes only small amounts of
Learning Objectives gossypol with the oil. However, extraction with more polar
solvents is effective in removing gossypol. Examples of polar
Understand the important differences between glandless solvents used include aqueous acetone, a mixture of acetone
and the traditional glanded cotton varieties. and hexane, and sequential extraction with hexane, aqueous
Become familiar with the processing involved in producing acetone, and anhydrous acetone. Blends of methylene chloride
the various products from cottonseed for food and feed and hexane-acetic acid have also been used. These solvents are
uses. not widely used commercially.
Techniques have also been reported that mill cottonseed in
the presence of hexane and then remove the intact, heavier
Introduction gossypol glands by the liquid cyclone process. Also, an air
classification process has been developed to separate intact
Most varieties of cotton (Gossypium hirsutum L., G. aboreum L., gossypol glands from solvent-extracted ground flour.
G. barbadense L., or G. herbaceum L.) are grown mainly in warm
climates around the world. Over 98.5 million tonnes (Mt) of
cotton were produced worldwide in 200102 and over 33.6 Mt Glandless Cottonseed
of cottonseed. Cotton is grown for its fiber (over 80% of its
value) and the seed is used mostly for oil recovery and feed. Some of the first studies on the development of glandless
Whole cottonseed can be fed to dairy cattle, and the meal cottonseed in the US were done in the early 1950s. These
resulting from oil extraction is fed primarily to ruminants studies described that selection of plants from the Hopi
and, in limited amounts, to poultry and swine. Moencopi variety could result in almost complete elimination
Traditional varieties of cottonseed contain gossypol, a of pigment glands from leaves and bolls. When crossing Moen-
yellow-green polyphenolic compound considered toxic to copi with upland cotton varieties (G. hirsutum), it was found
man and monogastric animals (Figure 1), reportedly affecting that glandless seed appeared in later segregating generations.
the heart, liver, and reproductive organs. It has been used in Production of glandless cotton was first conducted in Iguala,
China as a male contraceptive but the practice was abandoned Mexico in 1960.
because of permanent side effects. Gossypol is dispersed in Basically two genes, gl-2 and gl-3, in the plant are found to
the plant as deposited structures or glands, which can be control the production of gossypol pigments in the seed. When
seen as black specks in the stems, leaves, and seed (Figure 2). present in the homozygous recessive condition (gl2gl2gl3gl3),
all parts of the plant aboveground, including the seed, have Other than gossypol content very few differences are found in
no pigment glands. The presence of pigmented glands can glandless compared with glanded cottonseeds. Evaluations of
be easily seen as black specks by cross-sectioning a seed eight varieties each of glanded and glandless cottonseed were
(Figure 2). reported (Table 1). Amino acid profiles were similar for
Recently, the value of the whole seed as cattle feed has glanded and glandless seed. The storage and processing char-
increased making it less profitable to extract the oil and sell acteristics of glandless and glanded cottonseed are also
the meal. In some cases cottonseed is left at the gin in exchange reported to be essentially identical with regard to development
for ginning and baling services. Sometimes glandless cotton- of free fatty acids, moisture levels, and refining losses in the
seed is considered to produce fiber of lower yield and quality crude oils.
than traditional glanded varieties. Some studies have shown No major differences were encountered either in solvent
that the glandless factor in itself does not decrease fiber yield or extraction of glanded versus glandless cottonseed. Extraction
quality as long as care is taken to prevent weed and insect rates of oil from glandless cottonseed flakes are about the same
infestation. as from glanded flakes, using commercial hexane. It has also
Gossypol is regarded also to be a natural insecticide. Reports been suggested that glandless cottonseed can also be a source
show that glandless cotton attracts more insects than glanded of a new type of lecithin. Cottonseed has the highest content of
varieties. Entomology studies have shown that cotton bollworm phospholipids after soybeans, present at about 2.2% in the oil,
(Heliothis zea Boddie), tobacco budworm (H. virescens F.), pink but due to gossypol presence it was not commercially viable.
bollworm (Pectinophua gossyprella Saunders), and lygus insects Currently, the major commercial source of lecithin is soybean
prefer glandless cotton. It is generally recognized that glandless oil. Composition of cottonseed phospholipids consists of
varieties require closer supervision to intercept and control phosphatidylcholine, phosphatidylethanolamine, and phos-
insect infestations. phatidylinositiol (33%, 22%, and 37%, respectively) and is
Processing and utilization of glandless cottonseed was considered more oxidatively stable than soybean phospho-
reported in the early 1960s in the US. Glandless cottonseed lipids due to lower content of unsaturated fatty acids.
oil and meal were evaluated in poultry broiler and layer One of the main advantages of milling glandless cottonseed
rations. Glandless cottonseed meal was found nearly as effec- is that it simplifies direct solvent extraction. Current commercial
tive as soybean meal in achieving broiler gains and did not practice of refining glanded cottonseed oil requires that the oil
244 THE OILSEEDS | Cottonseed: Overview
Table 1 Comparative analysis means of eight varieties each Advantages of processing glandless cottonseed over glanded
of glanded and glandless cottonseed and their productsa seed include:
Whole glandless cottonseed kernels are prepared either for separated by centrifugation. The solids in the liquor are
direct use as whole nut substitutes or for food ingredients. also precipitated at pH 7, and the storage protein curd
These are also used in the production of high-quality flours removed by centrifugation and then dried. The liquor is
and food protein concentrates or isolates. The use of glandless further acidified to pH 4 to precipitate the NSP, also
cottonseed kernels for the production of nutlike products by removed by centrifugation and then dried. The soluble
means of various roasting methods has been reported, includ- matter remaining in the whey is then removed by either
ing dry roasting at several temperatures, roasting under vac- the selective extraction or selective precipitation method.
uum followed by steam injection, pressure steaming followed A relatively pure storage protein fraction, containing over
by oven roasting, and deep fat frying in various oils (including 90% protein (dry weight basis), can be prepared.
corn, cottonseed, peanut, safflower, soybean, and sunflower 3. The selective extraction procedure consists basically of
oils). When the main product is to produce high-protein leaching the proteins soluble with water at neutral pH,
flours, less care needs to be taken to minimize breakage during where the solids are then centrifuged out. The liquor is
de-hulling. then acidified to pH 4; this results in a precipitated protein
Glandless cottonseed flour can be prepared by extracting de- curd. The curd is dried to produce an NSP isolate. The solids
hulled glandless seed flakes with hexane. The extracted flakes are from the original water leaching are then solubilized in
then desolventized and ground to a desired mesh size. In the alkali at pH 10 and centrifuged to remove insolubles. The
production of cottonseed concentrates, the defatted flakes are re- liquor is then precipitated at pH 7 and the resulting curd is
extracted with acidified water or ethanol to remove the soluble concentrated and dried to produce an SP isolate.
sugars and flavor compounds, and dried to produce protein
A patented process for the preparation of glandless cottonseed
concentrate. A spray dryer can also be used in preparing con-
protein concentrate from solvent-extracted flour using industrial
centrates where the defatted flakes are ground prior to extrac-
membranes has also been used. Defatted flour is sieved through
tion. Concentrates have been prepared from flour by a dry air
an 80-mesh screen to break up any agglomerates and remove
classification method and by acidic water extraction. Figure 3
hull particles, and suspended in acidified water at pH 4.04.5.
shows a simplified diagram of the process to be used to generate
This solution is passed through an ultrafiltration membrane
protein concentrate from glandless cottonseed.
(100 000 MW cutoff) and the solids retained are either dried
Three major techniques have been reported for making
directly or neutralized before drying. The liquid fraction perme-
glandless cottonseed isolates. These isolates can be conven-
ate may then be passed through a reverse osmosis membrane to
tional protein isolates, storage proteins (SP) (soluble at high
concentrate the soluble solids and to recover water which may
and low pH), and nonstorage proteins (NSP) (soluble at near
be reused in the process.
neutrality):
Figure 3 Protein concentrate production from glandless cottonseed. (Data from Lusas EW and Jividen GM (1987) J. Am. Oil Chem. Soc. 64(6):
839854.)
246 THE OILSEEDS | Cottonseed: Overview
used with glanded cottonseed. Commercial glanded cotton- See also: Agronomy of Grain Growing: Soybean: Agronomy;
seed meal intended for poultry feeding usually receives consid- Barley, Rice and Maize Processing: Oil from Rice and Maize;
erable heat during processing to intentionally bind the Fats: Healthy Fats and Oils; Lipid Chemistry; Food Grains and the
gossypol. Also in order to increase the amount of glanded Consumer: Genetically Modified Grains and the Consumer; Grains
meal gossypol, addition of iron to the meal is used to inactivate Around the World: Grain Production and Consumption: Oilseeds in
gossypol. In the cases of glandless cotton meal, it would be North America; Non-food Products from Grains: Biodiesel; Cereal
expected that lower heat treatment needed for glandless cotton Grains as Animal Feed; Oilseed and Legume Processing:
would have beneficial effects in the protein availability and Soybean: Processing; Soybean: Soy Concentrates and Isolates; The
quality of the meal. Some heat pretreatment of the meal is still Basics: Grain and Plant Morphology of Cereals and how characters
recommended, however, before hexane extraction, to improve can be used to identify varieties; Taxonomic Classification of Grain
protein efficiency. Heating glandless cottonseed with 12% Species; The Grain Crops: An Overview; The Oilseeds: Oilseeds:
added water for 10 min at 82 C, followed by 105 C for Overview; Soybean: Overview.
20 min before hexane extraction, was reported to appreciably
increase broiler weight gains when compared with glanded
cottonseed feed. It has also been shown that meal from gland-
less cottonseed had an equal protein availability to soybean
meal in supporting chick growth. Feed trials with glanded Further Reading
cottonseed meal with swine, catfish, fish, and shrimp have
Berot S, Chenu M, Coutret J, and Gueguen J (1995) Cottonseed protein-rich products
also been described. from glandless African varieties: 1. Pilot-plant scale production of protein
concentrates, Sciences-des-Aliments 15(3): 203215.
Camire ME, King CC, and Bittner DR (1991) Characteristics of extruded mixtures of
Exercises for Revision cornmeal and glandless cottonseed flour. Cereal Chemistry 68(4): 419424.
Cater CM (1968) Processing glandless cottonseed. Oil Mill Gazeteer 72(7): 815.
Cherry JP, Gray MS, and Jones LA (1981) A review of lecithin chemistry and glandless
What is the meaning of Glandless in relation to cottonseed as a potential commercial source. Seed-oil content of glanded and
cottonseed? glandless, cottons. Journal of the American Oil Chemists Society 58(10): 903913.
What is gossypol? Glandless cotton: Its significance, status, and prospects: Proceedings of a conference,
December 1314, 1977, Dallas, Texas/sponsored by Agricultural Research Service,
What natural advantage does gossypol offer to the
United States Department of Agriculture and National Cottonseed Products
cotton plant? Association, Inc., Agricultural Research Service Southern Region, US Dept. of
Where is gossypol located in the cotton plant? Agriculture, 1978.
What is the toxicity of gossypol to humans, to animals Kohel RJ and Lee JA (1984) Genetic analysis of Egyptian glandless cotton. Crop
and to insects? Science 24(6): 11191121.
Lawhon JT, Cater CM, and Mattil KF (1977) Journal of the American Oil Chemists
Describe the ramifications of breeding glandless cotton for Society 54: 75.
the cotton-production industry, for the cotton-oil industry La-Rue DC, Knabe DA, Izard RS, and Tanksley TD (1986) Glandless cottonseed kernels
and for the food and feed industries. for starter, grower and finisher swine. Journal of Animal Science 64(4): 10511059.
Explain the relative advantages (and disadvantages) of Lusas EW and Jividen GM (1987) Glandless cottonseed: A review of the first 25 years of
processing and utilization research. Journal of the American Oil Chemists Society
glandless cotton (versus glanded cotton) to all parts of the 64(6): 839854.
cotton industry. Miravalle RJ (1972) The plant geneticists contribution toward changing the lipid and
List the food uses of cottonseed and compare these with the amino acid composition of cottonseed. Journal of the American Oil Chemists
food uses of the soybean. Society 49: 2426.
Percival AE, Wendel JF, and Stewart JM (1999) Taxonomy and Germ plasm resources.
In: Smith CW and Cothren JT (eds.) Cotton. New York: Wiley.
Robinson EH, Rawles SD, Oldenburg PW, and Stickney RR (1984) Effects of feeding
Exercises for Readers to Explore the Topic Further glandless or glanded cottonseed products and gossypol to Tilapia aurea.
Aquaculture 38(2): 145154.
Find out about the differences between the three major Ryan JR, Kratzer FH, Grau CR, and Vohra P (1986) Glandless cottonseed meal for laying
and breeding hens and broiler chicks. Poultary Science 65(5): 949955.
species of cotton (Gossypium hirsutum L., G. aboreum L.,
Simmons RG and Golightly NH (1981) Cottonseed Cookery. College Station, TX: Food
G. barbadense L., G. herbaceum L.). What potential advan- Protein R&D Center, Texas A&M University.
tages might accrue from the transfer of genes from one of Silva-Ramos LC and Kohel RJ (1987) Seed-oil content of glanded and glandless
these to the more common cotton species? cottons. Journal of the American Oil Chemists Society 64(9): 13371340.
Consider the possibility of extracting gossypol from glanded
cottonseed for use as an insecticide for growing glandless
cotton.
Research the uses and value of lecithin, and examine the Relevant Website
possibility of producing lecithin from glandless cottonseed.
cottonseed.com http://www.cottonseed.com National Cottonseed Products
Find out about genetically modified (GM) cotton, Association (NCPA) is a trade association for the cottonseed processing industry.
the nature of its consequent tolerance to insect attack, its Besides commercial information, it has further links to technical information on
suitability for producing cotton, oil and meal, and the cottonseed products.
possible public resistance to cotton as a genetically modi-
fied organism (GMO).
Sunflower: Overview
GJ Seiler and TJ Gulya, USDA-ARS, Northern Crop Science Laboratory, Fargo, ND, USA
2016 Elsevier Ltd. All rights reserved.
Topic Highlights Midwest and as far as northern Mexico. The Native Americans
also used sunflower hulls as a source of dye, leaves for herbal
Sunflower history origin and crop development medicines, and pollen in religious ceremonies.
Sunflower botany Historical records indicate that the Spanish were the first to
Sunflower production practices in North America introduce sunflower to Europe in the early 1500s. Sunflower
Sunflower agronomic problems: insect, disease, and preda- was initially grown as an ornamental plant. Early English and
tors (birds) French explorers also introduced it to their respective coun-
Sunflower processing and uses tries. From Western Europe, sunflower spread along the trade
routes to Egypt, Afghanistan, India, China, and Russia. By the
early 1700s, sunflower seeds were eaten as a snack, and in
Learning Objectives 1716, the first patent for the use of sunflower oil (for industrial
purposes) was filed in England. The most significant boost for
To understand the basics of the origin of sunflower and its sunflower as a crop, however, came from the Russian Ortho-
agronomy, production, processing, and uses dox Church. Lenten regulations prohibited the consumption
To understand the diversity of sunflower types and where of many oily foods, but since sunflower was not specifically
sunflower is produced listed, the seed and oil became a staple diet item in Russia.
To understand the variation in fatty acid composition and Efforts by Russian scientists led to significant crop improve-
its significance in sunflower utilization ments with oil contents soon exceeding 40%. While sunflower
was grown throughout Western and Eastern Europe, Russia
historically was the largest producer with in excess of 3 million
hectares in the early twentieth century, compared to one-half
Overview
million hectares for the rest of Europe. Russian immigrants are
credited with introducing sunflower to North America. In fact,
Sunflower (Helianthus annuus) is grown as an oilseed crop world-
the open-pollinated variety Russian Mammoth still sold by
wide in temperate and subtropical climates in 72 countries.
garden seed firms traces its lineage back to the same-named
Among oilseeds, sunflower generally ranks fifth behind soybean,
variety initially introduced in the 1880s. Early cultivation of
rapeseed, cottonseed, and peanut, with an average annual world
sunflower in North America was primarily for livestock silage
production of 3244 million tonnes (Table 1). Unlike soybean,
and seed for poultry. By the second half of the twentieth
sunflower is primarily an oil crop, with high-protein meal being
century, improved Russian varieties with oil levels of 4555%
a by-product. Sunflower is grown on every continent, except
were available. Increased US production of these high-oil sun-
Antarctica, with Ukraine being the largest producer, followed
flower varieties spurred interest by oil crushers, which led to
by the Russian Federation, Argentina, China, France, Romania,
expanded US production, especially in the Northern Great
Bulgaria, Turkey, Hungary, and the United States in 201112
Plains (North Dakota and Minnesota). The discovery of cyto-
(Table 2). US production accounts for 35% of the world out-
plasmic male sterility (CMS) by French scientists laid the foun-
put. Nutritionally, sunflower oil is somewhat superior to other
dation for the development of sunflower hybrids in the early
vegetable oils due to the greater proportion of the unsaturated
1970s. Hybrid sunflower, with higher yields and oil content,
fatty acids (oleic, linoleic, and linolenic) and lower saturated
more uniformity, and disease resistance, in comparison with
fatty acids (palmitic and stearic), especially in the recently devel-
open-pollinated varieties, provided the last great impetus in
oped midoleic content NuSun hybrids (Table 3). Sunflower
establishing sunflower as a worldwide crop (Figure 1).
oil contains zero trans fats, which have been implicated in ele-
vated cholesterol levels and increased risk of coronary heart
disease.
Botany
Table 1 World production (million tonnes) of major oilseed crops over the last 5 years
Source: http://apps.fas.usda.gov/psdonline/circulars/oilseeds.pdf.
Table 2 World sunflower seed production (in 1000 tonnes) over the last 5 years
Source: http://faostat.fao.org/.
Table 3 Fatty acid composition of traditional, high oleic and NuSun sunflower oil in comparison with other vegetable oils
Source: http://www.sunflowernsa.com.
freely hybridize with each other in nature, giving rise to inter- improved oil quality. All annual species, except H. agrestis,
mediate forms, which has led to confusion in taxonomy. While can be crossed with cultivated H. annuus. There has been
H. annuus is a geographically diverse species occupying a wide more success in crossing tetraploid and hexaploid perennial
range of habitats, other Helianthus species are very habitat- species with cultivated H. annuus than with the diploid peren-
specific and thus are endemic in a limited number of locations. nial species.
For example, H. niveus ssp. tephrodes is only found in active
sand dunes in southern California and in northern Sonora,
Mexico. H. paradoxus is only found in saline, marshy sites in Production Practices in North America
western Texas and New Mexico, and H. exilis is only found in
serpentine sites in Northern California whose soils are nearly Sunflower production in the United States is concentrated in
toxic to many nonadapted plants. Wild Helianthus species, the Northern Great Plains (North Dakota, South Dakota, and
both annual and perennial, have contributed many useful Minnesota) with a secondary production area in the Central
traits to domesticated sunflower, including disease resistance, Great Plains (western Kansas, Nebraska, and eastern Colorado)
drought and salt tolerance, herbicide resistance, CMS, and (Table 4), but there is some amount of sunflower grown in
THE OILSEEDS | Sunflower: Overview 249
Source: http://www.sunflowernsa.com/.
Figure 1 Sunflower field in bloom. Courtesy of Dr. Brady Vick (Retired),
USDA-ARS Sunflower and Plant Biology Unit, Fargo, ND, USA.
Sunflower is typically planted when soil temperatures reach
10 C. Below 10 C, germination is very slow. Sunflower is gen-
erally planted in row widths of 5075 cm to match the available
equipment, although solid seeding (widths < 30 cm) is also
practiced. Sunflower is planted from 4 to 7 cm deep at rates of
3762 000 seeds ha1 for oilseed and 3550 000 seeds ha1 for
confection. Lower plant populations for confectionery sunflower
produce larger-sized heads and a greater proportion of large seed,
which is desirable. Conversely, high plant populations will result
in smaller heads, which not only dry down faster but also have
thinner stalks that are more prone to lodging.
Sunflower has a root system characterized by both a deep
taproot and an extensive fibrous root system, which makes it
efficient at both water uptake and nutrient uptake. While its
water requirements are similar to that of corn, sunflowers greater
efficiency can result in higher returns under moisture-limiting
conditions. Sunflower responds well to nitrogen fertilization,
Figure 2 Common annual sunflower, Helianthus annuus, the with a general rule of 5 kg ha1 nitrogen required for 100 kg
progenitor of cultivated sunflower, with multiple flowers on a branched seed ha1. Thus, for a yield target of 2000 kg seed ha1, the soil
stem. Courtesy of Dr. Brady Vick (retired), USDA-ARS Sunflower and nitrogen content plus added nitrogen would be 100 kg ha1.
Plant Biology Unit, Fargo, ND, USA. Early weed control is important in sunflower with several
preplant, preemergence herbicides available for grass control
and now postemergent ones for broadleaf weed control.
Hybrids have been developed through conventional breeding
each of the 48 contiguous states (especially for birdseed). Most with tolerance to both imidazolinone and sulfonylurea herbi-
of the production recommendations have come from research cides, allowing for postemergent applications. There are no
conducted in North Dakota, but as interest in sunflower glyphosate-resistant hybrids in any country, nor are there
increases in other areas, new research generated from other currently any other commercially available traits introduced
states addresses these different environments. into sunflower from other organisms beyond the genus
Sunflower is traditionally planted between May 1 and the Helianthus (i.e., sunflower is currently still a non-GMO crop).
middle of June in the Northern Great Plains, with hybrid Sunflower in the Northern Great Plains typically flowers in
maturities of 100120 days to fit the short, 120-day average August, about 6070 days after planting. Four to five weeks
frost-free growing season of this region. Longer maturing post-pollination is required for seed maturation, at which
hybrids are used where short growing seasons are not a restric- point seed moisture is 3040%. If the crop is killed by frost,
tion. Similarly, there are early maturing hybrids that would be seeds may be harvested without use of a chemical desiccant.
suitable for a double-crop situation. Double-crop sunflowers Any conventional grain combine can be used to harvest sun-
are typically planted in July, either following the failure of an flower, with a variety of attachments to facilitate harvesting of
initial crop or after harvest of a small grain crop. Yields of the sunflower heads. Chemical desiccants can be used to
22003500 kg ha1 are attainable under optimal, irrigated reduce seed moisture, minimizing losses due to birds, lodging,
conditions, while the average dryland yield of sunflower in and severe disease attack of the heads. Seed moisture must be at
the United States ranges from 1516 to 1750 kg ha1. The phe- 10% or less to retard fungal deterioration in storage, with 8%
nology of sunflower hybrid maturation is depicted in Table 5. moisture preferable for long-term storage.
250 THE OILSEEDS | Sunflower: Overview
Table 5 Sunflower growth stages and approximate time and growing degree days (GDD) to reach each stage
VE Emergence 93 10
V4 4 True leaves 194 20
V8 8 True leaves 303 28
V12 12 True leaves 383 34
V16 16 True leaves 429 38
V20 20 True leaves 484 44
R1 Miniature terminal bud 511 46
R2 Bud <2 cm from last leaf 695 61
R3 Bud >2 cm from last leaf 774 67
R4 Bud open, ray flowers visible 829 71
R5.1 Early flower 859 73
R5.5 50% flowered 902 77
R6 Flowering complete 989 84
R7 Back of head pale yellow 1140 86
R8 Bracts green, head back yellow 1228 104
R9 Bracts yellow, head back brown 1372 119
Pest Problems The main head rot pathogens are Sclerotinia sclerotiorum, Rhizopus
(three species), Botrytis cinerea, and Erwinia carotovora.
Cultivated sunflower has a number of pest problems, including Control of most diseases is accomplished primarily with
diseases, insects, and birds. Since the genus Helianthus is indig- resistant hybrids and, to a lesser extent, with cultural practices.
enous to North America and coevolved with the pests, there are Single dominant genes are used to confer resistance to rust,
a great diversity of insects and pathogens adapted to sunflower, Verticillium wilt, and downy mildew. At least two dominant
plus a natural reservoir of these pests on the wild progenitors. genes are required for resistance to Phomopsis stem canker,
On other continents, the diversity of sunflower pests and path- while resistance to Sclerotinia head rot and stalk rot is con-
ogens is generally less, although international seed movement trolled polygenically, making it the most difficult disease to
has dispersed most sunflower pathogens around the globe. select for resistance. Very few diseases are managed with fun-
gicides, primarily because of economics. Seed treatments for
control of downy mildew had been very effective until the
Diseases pathogen developed resistance to the fungicide metalaxyl. In
Sunflower is subject to a number of fungal, bacterial, and viral some countries, foliar fungicides are used to manage Phomopsis
diseases, with fungi the most numerous and economically disease and rust. The use of foliar fungicides is more practical
serious. Diseases causing the most losses worldwide are Scler- with confection sunflower where the profit margin is greater
otinia head rot and stalk rot, Phomopsis stem canker, rust, and than with oilseeds.
downy mildew. Some diseases are serious in only a few coun-
tries, such as Verticillium wilt in Argentina or white rust
Insects
(Albugo) in South Africa. Most sunflower diseases are caused
by pathogens specific to sunflower, such as Phomopsis helianthi, In the United States, there is a wide variety of sunflower-
Alternaria helianthi, and Plasmopara halstedii. Some of the most specific, native insects that preferentially feed on stems, leaves,
serious diseases, however, are caused by pathogens with wide roots, or seeds. Stem-feeding insects on sunflower include at
host ranges, such as Sclerotinia sclerotiorum that occurs on 137 least three different cutworms (Euxoa species and Feltia jaculi-
different genera of plants. fera), the sunflower stem weevil (Cylindrocopturus adspersus), the
Disease organisms may be grouped based on plant parts black sunflower stem weevil (Apion occidentale), the sunflower
affected. The only significant seedling disease is downy mildew maggot (Strauzia longipennis), and the long-horned sunflower
(Plasmopara halstedii). Foliar diseases on sunflower include rust stem girdler (Dectes texanus). Root-feeding insects include the
(Puccinia helianthi), blights caused by Alternaria (five species) and carrot beetle (Ligyrus gibbosus), the sunflower root weevil (Baris
Septoria helianthi, powdery mildew (Erysiphe cichoracearum), bac- strenua), and occasionally wireworms. Leaf-feeding insects
terial blight (Pseudomonas syringae pv. helianthi), apical chlorosis include grasshoppers (four Melanoplus species and Camnula
(Pseudomonas syringae pv. tagetis), sunflower mosaic virus, and pellucida), aphids (primarily Aphis helianthi and Masonaphis
sunflower chlorotic mottle virus. The main stem diseases of masoni), the painted lady caterpillar (Vanessa cardui), the sun-
sunflower are Sclerotinia stalk rot (S. sclerotiorum), stalk rots flower beetle (Zygogramma exclamationis), and the palestriped
caused by S. minor and Sclerotium rolfsii, Phomopsis stem canker flea beetle (Systena blanda). Seed- and head-feeding insects
(P. helianthi), charcoal rot (Macrophomina phaseolina), and bac- include the sunflower moth (Homoeosoma electellum), the sun-
terial stalk rot (Erwinia carotovora). Head rots are probably the flower midge (Contarinia schulzi), the red sunflower seed weevil
most devastating because of their direct impact upon seed yield. (Smicronyx fulvus), the gray seed weevil (Smicronyx sordidus), the
THE OILSEEDS | Sunflower: Overview 251
banded sunflower moth (Cochylis hospes), the sunflower head nutritionally than polyunsaturated fatty acids, like linoleic
moth (Gymnocarena diffusa), the sunflower seed maggot (Neote- acid, and far superior to saturated fatty acids. While oleic acid
phritis finalis), the sunflower headclipping weevil (Haplorhynch- levels of 8090% are attainable, the frying industry prefers an
ites aeneus), and the tarnished plant bug (Lygus lineolaris). In oleic acid content of 5560%, primarily because of the taste
other continents, insects are generally considered minimal or associated with the remnant linoleic acid. High and mid-oleic
infrequent problems, and when they occur, they are generally sunflower oil also has a very high oxidative stability, almost
caused by omnivorous insects such as aphids, plant bugs (Lygus 10 that of traditional sunflower oil, extending product shelf
spp.), and other non-sunflower-specific insects. Management life and longevity of the frying oil. For a complete comparison
of sunflower insect pests usually concentrates on cultural and of the fatty acid composition of various vegetable oils (see
chemical strategies. To date, selection for effective insect resis- Table 3).
tance has been limited in cultivated sunflower. Oil extraction from sunflower seed is a relatively straight-
forward process. Seed is first thoroughly dried, dehulled, and
then flaked or rolled and heated to 8590 C. The crushed seed
Birds
is then subjected to one of three extraction methods: (1) a
Sunflower is vulnerable to damage by birds due to its easy mechanical screw press, (2) extraction with an organic solvent,
accessibility and high nutritional value of the seed. Ranging usually hexane, followed by distillation to remove the solvent,
from tiny sparrows (Passeridae) to large parrots (Psittacidae), and (3) a combination of the screw press and solvent extrac-
birds are a constant problem to sunflower on all continents. In tion. In the United States, the latter is the preferred method and
commercial birdseed mixtures, sunflower is the preferred seed, generally recovers about 99% of the total seed oil. The crude oil
so it is not surprising that the crop is beset by depredation by a is further processed to remove gums, waxes, free fatty acids,
wide variety of birds. In the United States, the migratory red- and odors. The refined oil is now suitable as a vegetable oil or
winged blackbird (Agelaius phoeniceus) causes the most dam- can be hydrogenated and blended with soybean or canola oil
age. In Europe, many different sparrows (Passer species) and for margarines.
doves (Streptopelia spp.) are the major problems, while in Sunflower oil can be used as an alternative or additive to
South America, parakeets (Psittacidae) and doves (Columbidae) diesel fuel to create biodiesel, a clean burning alternative fuel
predominate. produced from a renewable resource. This could help farmers
Seed losses due to bird feeding can easily exceed 10% in decrease their dependence upon petroleum fuels by substituting
fields planted close to nesting areas, and losses of 100% are not farm-grown fuel. For diesel engine use, sunflower oil requires
unheard of. In the United States, yield loss due to birds is more extensive purification including removal of waxes and
significant enough that many different management strategies gums. Minor engine modifications, including improved fuel
have been investigated. There has been limited success in filters, are also necessary to burn any vegetable oil. Since the
breeding for sunflower varieties that have physical barriers energy content of sunflower oil is less than that of diesel, fuel
(long necks, tight bracts, and seed coloration) to bird feeding. consumption would be greater and power output less. As the
Most research efforts have explored cultural methods, habitat price of diesel increases and the world supply diminishes, the
management, and mechanical frightening devices (noise feasibility of vegetable oil use in diesel engines becomes more of
makers). Various registered avicides have been tested, but a reality.
none have been successful in reducing bird depredation. How-
ever, one novel approach under investigation is a commercial
product, which contains the FDA-approved grape flavor com- Sunflower By-products
ponent, which ironically many birds find repellent.
Sunflower Meal
After sunflower seeds are crushed, the remaining meal is a high-
Sunflower Oil, Processing, and Uses protein product commonly used in livestock and poultry feeds.
The meal contains 260500 g kg1 protein, 120350 g kg1
Sunflower is grown worldwide, primarily as an edible oil crop. fiber (from remnant hulls), and 1090 g kg1 fat. Sunflower
Traditional sunflower oil (high linoleic) is used mainly as salad meal is lower in lysine content and higher in methionine than
oil and frying oil, although it can be hydrogenated for use in soybean meal and thus is usually combined with soybean meal
margarines. In terms of world production, sunflower usually in livestock feeds. Feeding trials with both dairy and beef cattle
ranks fifth among oilseed crops behind soybean, rapeseed or have adequately demonstrated the utility of sunflower meal
canola, cottonseed, and peanut. The oil content of oilseed since the meal is more ruminally degradable than either soybean
sunflower varies from 40% to 50% by weight. There are cur- or canola meal.
rently three categories of sunflower oil: traditional, NuSun,
and high oleic. Traditional sunflower oil is characterized by a
Hulls
high concentration of linoleic acid (70%) and a moderate
amount of oleic acid (20%). Plant breeders, using conven- Sunflower hulls comprise 2130% of the total seed weight and
tional methods, have selected lines in which oleic acid is the are often a waste by-product. In oil extraction factories, the
major fatty acid component (65%), and this oil has been hulls are often burned as a source of heat, both for the plant
given the trade name of NuSun. High oleic oil has an oleic and for the distillation process to remove the hexane solvent.
acid content of 8590%. Oleic acid, a monounsaturated fatty Hulls, mixed with appropriate binders and glue, have also been
acid, is considered by health and dietary experts to be better molded into cylinders and sold as ersatz fireplace logs. Public
252 THE OILSEEDS | Sunflower: Overview
buildings in the vicinity of processing plants have also used may be 25100% greater than with oilseeds, extra management
sunflower hulls for heating, with only minor modifications of practices to control insects and diseases are more feasible.
their boilers.
A red dye, suitable as an FDA-approved food colorant, can
be extracted from the hulls of purple-seeded sunflower. With Future Trends
the trend away from the use of synthetic dyes, many foods high
in anthocyanin content have been viewed as dye sources. The Sunflower production in the United States is in direct compe-
anthocyanin content of purple-hulled seed ranges from 6 to tition with soybean and rapeseed, which currently have the
16 g kg1 of hulls, which compares favorably with that of other advantage of glyphosate resistance for weed management. Sun-
sources (grape skins and beet pulp). Natural red dyes lack the flower is also confronted by disease and insect problems that
chemical stability of synthetic dyes and thus may only be are lesser concerns in soybeans and rapeseed. The challenge to
usable in certain applications. the sunflower industry is to make the crop as easy and profit-
Pectin, used in jellies and as a general food thickener, can be able to grow as its competitors. The development of
extracted from sunflower heads. The pectin content, after seed imidazolinone- and sulfonylurea-resistant hybrids gives
removal, is from 150 to 250 g kg1, and thus, a hectare of growers new tools to control postemergent weeds, similar to
sunflower could yield 200350 kg pectin. Sunflower pectin is the glyphosate resistance technology. Finding management
chemically a low-methoxyl pectin, as contrasted to high- schemes to control insects and diseases in sunflower is a con-
methoxyl pectins in apples and citrus. Low-methoxyl pectins tinuing challenge. Worldwide, Sclerotinia and Phomopsis
are used in making jellies low in sugar and thus would be used diseases are the two most devastating diseases. Both diseases
primarily in the diet food market. can be managed to some degree with fungicides, but genetic
Sunflower butter, a relatively new product marketed as resistance would be more economical and safer for the envi-
SunButter, is a delicious and healthy alternative to peanut ronment. The many native Helianthus species offer untapped
butter. Packed with nutrition, it is an excellent choice for resources for resistance genes for Sclerotinia and other diseases.
people with peanut allergies. Made from specially roasted sun- Another option to make sunflower more competitive is to
flower seeds, it is completely peanut-free, tree nut-free, and change its fatty acid composition to make the oil nutritionally
gluten-free. A single serving of SunButter Natural Omega-3 better than other oils, the so-called designer-specialty oils.
provides 461 mg of omega-3, with 1/3 less saturated fat than Moves in this direction include the NuSun hybrids, with
peanut butter. oil nearly comparable to olive oil. The global sunflower market
is trending toward high oleic (8590%) oil, which has zero
trans fats, low saturated fats, and high monounsaturated fatty
acid with high oxidative stability. The transformation is already
Confection Sunflower under way but will probably take 10 years to totally convert the
industry. As the world public becomes more health-conscious
Sunflower hybrids grown for human consumption are termed and with the trend to increase label information, consumers
confectionery sunflower. The seeds are generally roasted, are demanding to know what in a product makes it a healthy
salted, and eaten as a snack food, either in-shell or dehulled choice. Healthy sunflower oil will benefit from these demands
to produce nutmeats. The nutmeats are also used in breakfast and in turn generate more interest among growers. Commit-
cereals, trail mixes, and baked goods. Sunflower kernels have ments by large snack food processors and fast-food chain
been chocolate- or candy-coated as a snack. In shell, they have restaurants in the use of NuSun have been the first step in
also been coated with various flavorings, such as bacon, bar- increasing domestic usage of sunflower oil.
beque, salt, and pepper to name a few. Confectionery
sunflower comprises 1822% of US total sunflower produc-
tion, with the United States the largest producer in the world Exercises for Revision
market, followed by Argentina. Confectionery sunflower is
characterized by its larger seed size and by having white or From which continent did the sunflower originate?
gray stripes on a black or brown background, as compared to Who introduced the sunflower to Europe?
uniform black color of oilseed sunflower. Which are the main sunflower-producing countries?
Confectionery seed is graded on the basis of size. The largest Who were the first to increase sunflower oil content?
seeds, referred to as in-shell, are those passing over a 7.9 mm How many days are needed for sunflower to reach maturity
round-hole screen and generally make up 1525% of the in North America?
harvested seed. The medium-sized seed, destined for dehulling What are the various uses of sunflower?
for use as nutmeats, comprises 3050% of the crop and is that What are NuSun sunflowers?
fraction recovered between a 7.9 and 7.1 mm round-hole Which birds are the main problems in North and South
screen. The smallest seeds, passing through the 7.1 mm Americas?
round-hole screen, are used primarily as birdseed and com-
prise 1520% of the crop.
Confectionery sunflower is genetically unique from oilseed Exercises for Readers to Explore the Topic Further
sunflower, primarily because seed size and hull coloration are
more important than oil content. As a group, they are much more Propose novel technological or plant breeding solutions to
susceptible to diseases and insects. Since the profit to the grower the problem of bird damage in sunflower production.
THE OILSEEDS | Sunflower: Overview 253
Compare the productivity of sunflower with that of other Salunkhe DK, Chavan JK, Adsule RN, and Kadam SS (1992) Sunflower. In: World
oilseed crops in a region of interest to you. Oilseeds: Chemistry, Technology, and Utilization. New York: Van Nostrand
What scope is there for further value-added development of Reinhold, 554 pp.
Schneiter AA (1997) Sunflower Technology and Production. Madison WI: American
specialty oils from sunflower? Society of Agronomy Monograph #35.
Seiler GJ (ed.) (1992) Sunflower: Special Issue. Field Crops Research 30(34):
191448.
Seiler GJ and Jan CC (2010) Basic sunflower information. In: Hu J, Seiler G, and Kole C
(eds.) Genetics, Genomics, and Breeding of Sunflower, pp. 150. Enfield, NH:
Further Reading Science Publishers Chapter 1, 335 pp.
Skoric D, Seiler GJ, Zhao L, Jan CC, Miller JF, and Charlet L (eds.) (2012) Sunflower
Berglund DR (2007) Sunflower Production. Fargo, ND: North Dakota State University Genetics and Breeding. Novi Sad, Serbia: Serbian Academy of Science and Arts,
Extension Service Bulletin 25 (revised), 119 pp. 520 pp.
Fick GN (1989) Sunflower. In: Robbelen G, Downey RK, and Shri A (eds.) Oil Crops of Weiss EA (ed.) (2000) Sunflower. In: Oilseed Crops, 364 pp. Malden, MA: Blackwell
the World. New York: McGraw-Hill, 553 pp. Science.
Heiser Jr. CB (1976) The Sunflower. Norman, OK: University Oklahoma Press, 198 pp.
Heiser Jr. CB, Smith DM, Clevenger SB, and Martin Jr. WC (1969) The North American
sunflowers (Helianthus). Memoirs of the Torrey Botanical Club 22(3): 1218.
Jan CC and Seiler GJ (2007) Sunflower. In: Singh RJ (ed.) Genetics Resources,
Chromosome Engineering, and Crop Improvement. Oilseed Crops, vol. 4. Relevant Websites
New York: CRC Press, Taylor and Francis Group Chapter 5, 304 pp.
Maiti RK, Singh VP, Purohit SS, and Vidyasagar P (eds.) (2007) Research Advances in www.ars.usda.gov/main/site_main.htm?modecode54-42-05-20 U.S. Department
Sunflower. Jodhpur, India: Agrobios International, 512 pp. of Agriculture Agricultural Research Service, Sunflower and Plant Biology
Miller JF (1987) Sunflower. In: Fehr WR (ed.) Principles of Cultivar Development, vol. Research Unit.
2. New York: Macmillan Publication, 761 pp. http://www.ext.nodak.edu/extpubs/plantsci/rowcrops/eb25w-12.htm North Dakota
Pappalardo J (2008) Sunflower, the Secret History. Woodstock, NY: The Overlook State University Extension Service, with individual bulletins on many aspects of
Press, 256 pp. sunflower production.
Rogers CE, Thompson TE, and Seiler GJ (1982) Sunflower Species of the United States. www.isasunflower.org International Sunflower Association.
Bismarck, ND: National Sunflower Association, 75 pp. www.sunflowernsa.com National Sunflower Association.
Overview of the Oilseed Safflower (Carthamus tinctorius L.)
C Hall III, North Dakota State University, Fargo, ND, USA
2016 Elsevier Ltd. All rights reserved.
content. Seed planting density is another agronomic practice common. Lutein and zeaxanthin levels are approximately 0.8
used to enhance yield of many crops. However, safflower tends and 1.5 mg g 1 oil, respectively, as determined by HPLC. How-
to branch when sown in wide seed spacing (i.e., less seed ever, photometric (445 nm) analysis indicated levels as high as
density) and thus compensates for the lower plant numbers 7 mg g 1 seed. The underestimations of carotenoids could
by producing more seeds per plant compared to greater seed occur for oils that contain predominantly carotenes. In the
densities. As a result, oil yield per hectare and oil content per case of safflower, trans- and cis-b-carotene levels were only
seed are not significantly different among plants grown from 0.140.16 mg ml 1 oil, while a-carotene accounted for less
low- and high-density seeding rates. than 0.04 mg ml 1 oil. Therefore, the total carotenoid level
Harvesting of safflower usually begins 3540 days after close to 2.5 mg g 1 oil is realistic. Contrary to expectations,
flowering. These harvesting dates also coincide with maximum the yellow pigments in the flowers of safflower are not carot-
oil content. Thus, harvesting date can influence oil yield. The enoids. Although carotenoid levels as high as 650 mg g 1 dried
maturity of seed also contributes to the final composition of flower petals have been reported, hydroxyl safflower yellow A,
the seeds. Over the course of 100 days, palmitic and oleic acids safflower yellow B, and anhydrosafflor yellow B are the pre-
increased, while stearic acid decreased. In contrast, linoleic acid dominant yellow pigments. These pigments are glycosylated
reached a maximum at day 75 and then decreased when mea- flavonoids and thus have high water solubility in contrast to
sured at day 100. Safflower can grow on marginal cropland, carotenoids.
including soils of moderate salinity. Under hydroponic condi- A significant variation in literature among total, free,
tions, 50 mM sodium chloride was found to inhibit safflower and esterified sterol exists. The phytosterol content in saf-
growth and increase the oxygenated essential oils. In general, flowers is approximately 20004500 mg g 1 seed. The highest
7% and 29% reductions in oil content and oil yield, respec- sterol content was obtained in seeds at 24 days after anthesis
tively, in safflower subjected to saline stress have been (i.e., flowering) and stabilized at 2400 mg g 1 seed at day 47
reported. Also, oleic acid tended to increase under saline (i.e., end of maturity). Therefore, the variation in phytosterol
conditions, while linoleic decreased. data may be due to seed maturity. Phytosterol contents of
15205745 mg g 1 of oil have been reported. In general, the
free (48%) sterol form accounts for the majority of the phy-
Nonsaponifiable Components tosterol content followed by esterified (39%) form. Steryl
glycosides and acyl steryl glycosides collectively make up
Tocopherols, carotenoids, and phytosterols make up the non- about 13% of the total sterols. Regardless of the sterol form,
saponifiable lipid components of safflower oil. Traditional and b-sitosterol accounts for the largest percentage (5070%) of
high-oleic acid safflower oils are composed of approximately the phytosterols. Campesterol, D7-stigmastenol, D5-
250 mg tocopherol per gram of oil or 50100 mg tocopherol per avenasterol, and stigmasterol each make up 612% of the
gram of seed. However, tocopherol contents as high as 684 and total phytosterol content. Reductions in total sterols due to
941 mg g 1 of oil have been reported for traditional safflower maturity and refining have been documented; however, the
and high-oleic acid safflower oil, respectively. Mean tocoph- percentage of each phytosterol remains relatively consistent,
erol concentrations from seven geographic regions in the range for example, b-sitosterol accounting for the greatest percentage.
of 676827 mg g 1 of oil have been reported. No differentia-
tion of safflower oil type was provided, but the lowest and
highest tocopherol contents were from East Africa and South- Phenolic Components
west Asia geographic locations, respectively.
In traditional safflower, a-tocopherol accounts for 9398% The majority of the phenolic compounds of safflower are
of the total tocopherols, while b-, g-, and d-tocopherols located in the flower petals and meal component of the seed.
account for the remaining in varying concentrations. The meals contain 38005700 mg g 1 of phenolic glycosides.
a-Tocopherol accounted for 8398% of the tocopherols in Although humans find these compounds bitter, animals
high-oleic acid safflower oil; however, no b analog was appear to be less impacted by the bitterness, and thus, saf-
found. In addition to tocopherols, 5 and 12 mg of a- and flower is often used as a protein source in animal rations.
g-tocotrienols per gram oil, respectively, were reported in cul- Total phenolic contents of water extracts were 40 and
tivated safflower. In contrast to cultivated species, g-tocopherol 126.0 mg gallic acid equivalence (GAE) per gram flowers and
contents as high as 36% were reported in oil obtained from seed, respectively. In contrast, only 26 mg GAE per gram was
wild safflower. A mutant safflower obtained through breeding observed in the oil. Flavonoid contents of safflower water
contained a unique tocopherol profile where g-tocopherol extract obtained from the seed were 62.2 1.9 mg quercetin
accounted for 96% of the total tocopherols. equivalence per gram, while an extract from dried flowers
Approximately 29% of the tocopherols are removed during contained approximately 10 catechin equivalences per gram
refining, with the greatest removal occurring during the bleach- extract.
ing and deodorization steps of the process. Thus, adjustments Further characterization of the water extract of the seed
to the aforementioned numbers should be considered when indicated that the main phenolic compound was
refining safflower oil. Roasting of the safflower at temperatures ( )-epigallocatechin (109.6 mg g 1). Other phenolics includ-
between 140 and 180 C did not negatively impact tocoph- ing 4-hydroxy-benzhydrazide derivative (18.2 mg g 1), 2-
erols or tocotrienols. amino-3,4-dimethylbenzoic acid (16.8 mg g 1), and gallocate-
The safflower carotenoids are present in trace amounts in chin (17.0 mg g 1) account for the majority of the remaining
the oil. Total carotenoid levels of less than 2.5 mg g 1 oil are phenolic compounds. The phenolic acids trans-ferulic,
THE OILSEEDS | Overview of the Oilseed Safflower (Carthamus tinctorius L.) 257
chlorogenic, p-coumaric, and syringic acids are present at 3.0, and candy. The stability of the pigments can be problematic
2.4, 0.5, and 0.2 mg g 1 extract, respectively. Additional flavo- under the presence of light. Carthamin in particular has poor
noids included naringin, rutin hydrate, quercetin dehydrate, stability when added to aqueous-based food systems. The sta-
luteolin, and kaempferol at concentrations of 6.0, 3.7, 2.2, 1.6, bility improves in a 75% sucrosecholine solution compared
and 0.8 mg g 1, respectively. Chlorogenic acid, quercetin-3- to water. A 90% glucosecholine chloride solution had the
galactoside, and gallic acid are present at concentrations of greatest stabilizing effect on carthamin. Although degradation
3742, 1516, and 1012.1 mg g 1 dried flower petals, respec- occurred, carthamin had greater stability in a xylitolcholine
tively. Other phenolic compounds were present at concentra- solution than in water under thermal degradation conditions.
tions less than 9 mg g 1. Although differences in concentrations In contrast to carthamin, the yellow pigments tend to be more
of phenolics were observed, the general trend in individual stable but tend to degrade slightly under acid conditions. Solu-
phenolics remained consistent regardless of growing location tion containing lactic acid and ethanol facilitated the rapid
and cultivar. degradation of carthamin and to a lesser extent the yellow
pigments. Multiple studies have demonstrated the stability
issues of safflower pigment. The importance of the food matrix
Uses in stabilizing/destabilizing the colorants must be considered if
safflower pigments are to be used in foods.
Although considered a modern use, safflower as a vegetable oil Traditional Chinese medicine has used safflower as a cure
dates to around 260 BC. The use of safflower as a vegetable oil for many ailments. However, until relatively recently, evidence
is limited due to the availability of inexpensive commodity supporting the use of safflower as a treatment for many
oils. However, high-oleic acid safflower has promise as frying ailments has been lacking. A summary of the pharmacological
oil due to the low level of polyunsaturated fatty acids. Many and medicinal benefits can be found in the literature.
oxidative stability index evaluations indicate that high-oleic However, a few examples include the antiatherogenic effect
acid safflower oil has comparable oxidative stability to that of of N-(p-coumaroyl)serotonin and antiadipogenic effects of
high-oleic acid sunflower oil. The oxidative stability of high- N-feruloylserotonin of safflower seed extract. The serotonin
oleic acid safflower was further enhanced in a safflower line derivatives are believed to inhibit the activation of the vascular
that contained high g-tocopherol content. The non- smooth muscle cells, thereby preventing atherosclerosis.
saponifiable fraction of safflower was found to inhibit frying Alcohol extracts of the flowers had matrix metalloproteases-2
oil polymerization. (MMP-2) inhibitory activity. MMP-2 are important for many
In countries such as Korea, safflower oil is used as condi- biological functions but if not properly controlled can contrib-
ment oil and is therefore roasted prior to oil extraction. Mild ute to the development of diseases such as cancer and athero-
roasting of the safflower seed does not affect fatty acid compo- sclerosis. Therefore, the MMP-2 inhibitory activity of hydroxyl
sition, but increases in tocopherols and oxidative stability have safflower yellow A and safflower yellow B could be important
been documented by several researchers. However, excessive treatments for disease affected by MMP-2.
heat treatments such as boiling and microwaving decrease The use as a clothing dye and a coloring for food has been
tocopherol concentration and oxidative stability. In addition documented. Safflower as a potential source of health-
to tocopherol and Maillard browning compounds, formed promoting oil (i.e., high g-linolenic acid) and as a biodiesel
during roasting, phytosterols and phenolic compounds are feed stock has attracted attention from commercial entities.
thought to stabilize the safflower oil. Safflower is also a common bird food. Recently, pharmacolog-
Extracts of safflower have been shown to exhibit antioxi- ical and medicinal benefits of safflower have attracted the
dant activity and thus could potentially serve as a source of attention of health researchers. Therefore, the author invites
natural antioxidants. Water extracts of safflower seeds showed readers to review these source materials.
dose-dependent activity in several radical scavenging assays.
The ORAC value of 63 mM Trolox equivalence per gram of
seed extract was comparable to that of tomatoes. Radical scav-
enging properties were also found for extracts of the flower Future Prospects
petals. Roasting increased slightly the Trolox equivalent antiox-
idant capacity (TEAC), while microwaving of the seed signifi- A significant body of knowledge exists regarding the produc-
cantly increased TEAC compared to the control or raw seed. The tion and composition of safflower. However, additional pro-
list of studies supporting the antioxidant activity of safflower duction research related to stressed environments such as high
continues to grow. The diverse phenolic composition, yellow salinity and low moisture conditions are needed, specifically
pigments, and tocopherol are just a few examples of compounds improving yield under stress conditions. Furthermore, the use
that likely contribute to the antioxidant activity. of molecular techniques to alter or enhance composition is
In addition to their antioxidant activity, the pigments of the needed. The genetic manipulation of safflower to produce
flowers have been used extensively as a food, cosmetic, and high g-linolenic acid safflower oil represents an opportunity
textile dye. Hydroxyl safflower yellow A, safflower yellow B, to produce safflower oil with eicosapentaenoic acid and doc-
and anhydrosafflor yellow B are the yellow pigments of osahexaenoic acid, that is, long chain omega-3 fatty acids are
safflower, while carthamin is the red color. Unlike the yellow important for health and are normally found in fish and algae.
pigments, carthamin has limited water solubility and is there- Safflower is unique among many oilseeds in that temperature
fore used in lipid-based products such as chocolate. The yellow during growing and seed filling does not significantly affect
pigments have been used to color cheese and sausage, juice, unsaturated fatty acid content. Therefore, greater production
258 THE OILSEEDS | Overview of the Oilseed Safflower (Carthamus tinctorius L.)
area could be achieved in contrast to oilseeds that have See also: Bioactives and Toxins: Bioactives: Antioxidants; Fats:
temperature-dependent regulation of unsaturated fatty acids. Healthy Fats and Oils; Lipid Chemistry; Food Grains and Well-
Expanding the understanding of the functional characteris- being: Functional Foods: Overview; Genetics of Grains:
tics of safflower seed and flower is needed. Several studies have Development of Genetically Modified Grains; The Oilseeds: Canola:
documented the color stability of flower extracts in various Overview; Linseed: Overview; Oilseeds: Overview; Soybean: Overview;
model systems. However, additional investigation is needed in Sunflower: Overview; Cottonseed: Overview.
food-based systems. The bitterness associated the meal, while
not a deterrent in animal feed, is not well suited for human
consumption. Identifying ways to reduce or eliminate the bit-
terness at a commercial level is needed. Alternatively, methods
to extract and purify the bitter compounds could be justified if Further Reading
sufficient bioactives could be isolated. Some of the bitter com-
Asgarpanah J and Kazemivash N (2013) Phytochemistry, pharmacology and medicinal
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control of the linoleic acid content in safflower (Carthamus tinctorius) seed oil.
acterized. Although a number of studies have been completed,
Journal of Agricultural and Food Chemistry 52: 332336.
there is still a need to assess the bioactivity of safflower. Hamrouni-Sellami I, Salah H, Kchouk M, and Marzouk B (2007) Variations in
phytosterol composition during the ripening of Tunisian safflower Carthamus
tinctorius L. seeds. Pakistan Journal of Biological Sciences 10(21):
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Kisha T and Johnson R (2012) Safflower. In: Gupta SK (ed.) Technological Innovations
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biofuel, coloring, and bioactive compounds? (Carthamus tinctorius L.) in semi-arid conditions. Biotechnology and
Why is safflower grown worldwide? Biotechnological Equipment 22(4): 947953.
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inhibitor from Carthamus tinctorius L. using ultrafiltration liquid
Why is the taproot of safflower important? chromatographymass spectrometry. Journal of Liquid Chromatography and
How does growing environment affect safflower seed Related Technologies 37: 23272336.
composition? Luterottia S, Franko M, Sikovec M, and Bicanic D (2002) Ultrasensitive assays of trans-
Why is safflower unique among oilseeds in terms of fatty and cis-b-carotenes in vegetable oils by high-performance liquid
chromatographythermal lens detection. Analytica Chimica Acta 460: 193200.
acid composition?
Mariod A, Ahmed S, Abdelwahab S, et al. (2012) Effects of roasting and boiling on the
What bioactive compounds are present in safflower and chemical composition, amino acids and oil stability of safflower seeds. International
what part of the plant can these compounds be found? Journal of Food Science and Technology 47: 17371743.
What type of safflower oil can be used as frying oil? What Mayerhofer M, Mayerhofer R, Topinka D, Christianson J, and Good A (2011)
Introgression potential between safflower (Carthamus tinctorius) and wild relatives
makes this oil well suited for frying applications?
of genus Carthamus. BMC Plant Biology 11: 47. http://dx.doi.org/10.1186/1471-
Why is safflower a potential source of omega-3 lipid? 2229-11-47.
Why is the yellow pigment not commonly used in food Nykiforuk C, Shewmaker C, Harry I, et al. (2012) High level accumulation of gamma
systems? linolenic acid (C18:3D6.9,12 cis) in transgenic safflower (Carthamus tinctorius)
seeds. Transgenic Research 21: 367381.
Ortega-Garca J, Gamez-Meza N, Noriega-Rodriguez J, et al. (2006) Refining of high
oleic safflower oil: Effect on the sterols and tocopherols content. European Food
Exercises for Readers to Explore the Topic Further Research and Technology 223: 775779.
Singh V and Nimbkar N (2006) Safflower (Carthamus tinctorius L.). In: Singh RJ (ed.)
How does safflower differ from other oilseeds in terms of Genetic Resources, Chromosome Engineering, and Crop Improvement, Vol. 4.
Oilseed Crops, pp. 167194. Boca Raton, FL: CRC Press, Taylor & Francis Group.
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Smith J (1996) Safflower. Champaign, IL: AOCS Press.
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Identify several countries that produce safflower and com- in Carthamus (Asteraceae): RAPD markers and sectional classification. Botanical
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produced seeds? Do any countries use wild safflower? Weiss EA (2000) Safflower. In: Weiss EA (ed.) Oilseed Crops, pp. 93129. Oxford:
Blackwell Science.
Why is defatted safflower meal used only for livestock feed
and not human food? How does this compare to other
oilseed meals? Cross-reference to oilseed articles.
Linseed: Overview
S Cloutier, Agriculture and Agri-Food Canada, Ottawa, ON, Canada
2016 Elsevier Ltd. All rights reserved.
Topic Highlights The term linseed often refers to the seed destined for oil
extraction with industrial end uses while the term flaxseed is
Linseed is an ancient crop that has been useful and versatile more commonly associated with nonprocessed seed used
through the ages. directly in human food or animal feed. In reality, the two
Linseed is a minor oilseed crop with a unique fatty acid terms represent flaxseed with the same oilseed quality and
profile. they can be used interchangeably. Currently cultivated linseed
Linseed oil is rich in alpha-linolenic acid (ALA) that gives varieties contain 4050% oil, of which 5557% is ALA, an
the oil unique drying properties. essential fatty acid. LinolaTM is a trademark name of the
Many health benefits have been attributed to the consump- solin (generic term) type of flax that contains < 5% linolenic
tion of linseed. acid, resulting in oil similar in composition to sunflower. In
Low yield relative to other major oilseeds limits the expan- Canada, LinolaTM or solin varieties were required to have a
sion of the crop. yellow seed coat to make them visually distinguishable from
brown seeded linseed varieties, although there are also yellow
linseed varieties. High linolenic acid content varieties charac-
Learning Objectives terized by more than 60% ALA are generically referred to as
high-lin such as the European variety Omega or under the
registered name Nulin.
In the following article, the reader is expected to understand
the uniqueness of linseed compared to other oilseed crops.
An overview of the agronomic practices and production
limitations is presented to illustrate the challenges associ- End Uses
ated with the development of improved cultivars and of
cultivating this crop. Emphases have been placed on not An overview of the end use versatility of fiber flax and linseed is
only the composition of linseed as it relates to its multiple illustrated (Figure 1). For the purpose of this article, the focus
end uses but also the multiple health benefits associated will be strictly on the seeds. Linseed is used as human food in
with the consumption of linseed. baked goods and other products, either whole or ground.
Whole seeds can also be fed to poultry because they can
grind the seeds with their gizzard. The processing of linseed
Introduction to extract oil produces a by-product called meal. Rich in pro-
tein and energy, the meal is a high-value feed for livestock. The
The species epithet of the Latin name of flax, Linum usitatissi- extracted oil can be utilized as human food, mostly as a sup-
mum L., means most useful, an appropriate name because this plement, but its main application is as industrial oil in, for
crop has been highly versatile and esteemed through the ages. example, stains, paints and linoleum flooring.
Flax has two morphotypes: fiber and oilseed. Tall and
branched only in the top quarter of the plant, fiber flax is
grown for its stem fibers that are extracted and processed into Composition
linen, nonwoven mats, paper, and various other products.
Oilseed flax plants, also called linseed or flaxseed, are shorter Expressed on a dry basis, typical linseed contains 4143% oil,
and more branched and have larger seeds than the fiber type. 28% dietary fiber, 2123% protein, 6% carbohydrate, and
Flax is one of the worlds oldest cultivated crops, praised by 4% ash but these can vary substantially among genotypes,
ancient civilizations for its fiber and seed uses. Today, varieties particularly oil content. In genebanks, oil contents ranging
are grown for either stem fiber or seed, although value can be from 31.4% to 45.7% were reported while levels up to 52.3%
extracted from the seeds of fiber flax and, conversely, the stem were recorded in field plot trials. Most modern linseed cultivars
fiber of linseed. Over the last 50 years, the worlds production field-grown in temperate climates contain 4549% oil.
of linseed fluctuated between 2 and 4 million tonnes (Mt), but The fatty acid composition of linseed oil is unique among
in the last decade, it has stabilized at a little more than 2 Mt plants because it is particularly high in ALA, the parent fatty
with the largest producers being Canada, the Russian acid of the omega-3 family. The major oilseeds of the world
Federation, and China. In 2012, world oil production from contain much less ALA than linseed (Figure 2). Rapeseed
the top 12 major oilseed crops amounted to 157 Mt, of which (canola) and soybean oils have 10% and 8% ALA, respec-
linseed oil production at 544 t accounted for a mere 0.35%. As tively, while sunflower, cottonseed, corn, and peanut oils only
such, linseed is a minor oilseed crop worldwide, but the have traces (<1%). Conventional linseed varieties typically
unique characteristics of its oil combined with its high content contain 57% ALA, 16% linoleic acid (LA), 18% oleic acid, 4%
of seed dietary fibers and lignans respond to specific markets stearic acid, and 5% palmitic acid. With 73% polyunsaturated
not addressed by the major oilseeds. fatty acid content, linseed oil has unique drying properties.
Linseed
Rapeseed
Soybean
Solin
Sunflower
Cottonseed
Corn
Extraction
Peanut
incidence of cardiovascular diseases. Recently, Health Canada Animal Feed and Omega-3-Enriched Products
has granted a health claim regarding the effectiveness of
ground flaxseed on lowering cholesterol allowing health Linseed meal, ground or whole flaxseed, can be used as feed for
claims such as the following to be made on flaxseed labels: poultry, swine, and cattle because it is a good source of protein,
16 g or 2 tablespoons of ground flaxseed supplies 40% of the oil, and energy although whole flaxseed can only be used with
daily amount shown to help lower cholesterol. On the other poultry. There are several reports on the health benefits asso-
hand, LA can be converted into omega-6 AA, which has been ciated with feeding linseed-derived products to livestock. How-
shown to be detrimental by promoting inflammatory reac- ever, as outlined in the earlier-mentioned section, livestock
tions. The enzymes delta-6, delta-5, and delta-4 desaturases producers must follow recommended guidelines to avoid the
are shared between the omega-3 and omega-6 pathways. For- side effects associated with deficiencies, antinutritional effects,
tunately, the high omega-3 to omega-6 ratio of linseed favors or ability to process the ration.
the omega-3 pathway over the omega-6 pathway. The Western A main driver for using linseed as feed is for the production
diet is generally considered to have an omega-6 to omega-3 of omega-3-enriched eggs and meat. The feeding of 10% lin-
ratio that is too high (20:1) due to the types of fats and oils seed ration to laying hens increased the omega-3 content in
consumed (e.g., soybean; Figure 2) and hence favoring the eggs by a factor of 4. In Canada, a minimum of 300 mg of
omega-6 pathway. The replacement of some of these fats and omega-3 fatty acid per serving is required to allow the labeling
oils with whole and ground linseed or linseed oil would rep- of a product as a source of omega-3. Such labeling commands a
resent a healthy alternative to meet the recommended 2:1 to premium that health-conscious consumers are willing to pay.
6:1 ratio. Omega-3-enriched chicken and pork products are also avail-
Linseed contains both soluble and insoluble dietary fibers. able to consumers.
The predominant fraction of the soluble fiber is the mucilage,
which has been associated with lowering of blood cholesterol
and glucose. Lignans are phytoestrogens that are also part of
the dietary fiber component. Linseed is one of the best
Agronomy
sources of these diphenolic compounds. Secoisolariciresinol
In temperate climates, linseed is grown as a summer annual
diglucoside, the major linseed lignan, has been associated
crop that matures in 90150 days including a vegetative, a
with reduced risks of hormone-sensitive cancers such as pros-
flowering, and a seed maturation period. In some areas of the
tate, colon, and breast cancers. Lignans also have antioxidant
world, winter-type linseed is grown. These varieties do not
properties leading to their positive role in reducing the activity
require vernalization per se to flower but they have the ability
of free radicals.
to endure some cold temperatures in the early developmental
stages. While each boll may contain a maximum of ten seeds,
linseed and fiber flax accessions in western Canadian field
trials produced only 3.58.1 seeds per boll. The flat, oval,
Antinutritional Factors
and pointed seeds are small with thousand-seed weight rang-
ing from 2.7 to 8.4 g, but the majority of Canadian and US
There are two main well-documented antinutritional compo-
linseed varieties have thousand-seed weight of 55.5 g.
nents in linseed: cyanogenic glucosides and linatine. The two
Ideally, linseed is sown in firm, moist soil 1.53.0 cm deep
main cyanogenic glucosides contained in mature seeds are
in rows 1520 cm apart at a seeding rate of 3045 kg ha1.
linustatin and neolinustatin. Following ingestion, hydrolysis
Early seeding is recommended to avoid some diseases (see
can lead to the release of hydrogen cyanide that can impede
succeeding text) and drought and heat during flowering and
the essential process of cell respiration. Fortunately, the
seed filling, which translates into higher yield, oil content, and
amount of cyanogenic glucosides in linseed is low, and
quality. Heat and drought stresses at flowering time and during
because the acute lethal dose of hydrogen cyanide in human
seed filling result in significant yield losses and lower oil con-
is 0.53.5 mg Kg1 of body weight, an average adult would
tent. Seeding practices for linseed differ greatly from those of
have to consume 1000 g of linseed in a day to receive a fatal
fiber flax where a high plant density is critical to fiber produc-
dose. The recommended daily intake of linseed of 3040
tion and quality. In general, the utilization of agronomic best
grams contains well below the toxic dosage. Moreover, the
management practices such as early seeding, crop rotation, and
acidic environment of the human stomach is not conducive
weed and disease control has a major effect on linseed yield.
to cyanogenic glucoside hydrolysis. As a consequence, these
toxic degradation products are more of a concern in
ruminants.
Linatine is an antagonist of vitamin B6. Symptoms associ- Genetics
ated with B6 deficiencies were observed in young poultry;
hence, their ration should be limited to <10% linseed meal. Linum usitatissimum L. is a diploid species with 15 pairs of
As mentioned earlier, the mucilage comprises soluble dietary chromosomes (2n 30). Its genome is composed of an esti-
fibers well known for their positive role in laxation. In animal mated 373 Mbp of nucleotides for which an assembled
feed, the outcome of rapid passage in the intestinal tract is poor sequence is available (www.phytozome.net). There are 48 000
nutrient absorption and its consequent performance reduc- accessions of cultivated flax preserved in genebanks around the
tion, particularly in poultry, a consequence that also limits world but only 10 000 are estimated to be unique because of
the linseed meal proportion of the ration. the redundancy between collections. Nevertheless, these
262 THE OILSEEDS | Linseed: Overview
collections represent a tremendous amount of diversity for A core collection of flax including more than 300 linseed
genetic enhancement (Figure 3). Other important genetic types and 92 fiber types was extensively characterized in the
resources include a large collection of expressed sequences field at two locations during four consecutive years. Agronomic
(genes), a consensus genetic map, and a physical map that traits such as yield and its components (thousand-seed weight,
were used to order and annotate the genome sequence. seeds per boll, and bolls per area), lodging, maturity, flowering
Figure 3 Genetic diversity in cultivated flax, Linum usitatissimum. (a) Flower color varies from white to shades of pink or blue and petals overlap,
margin, and folding are also indicative of the breadth of the genetic diversity. (b) At maturity, some accessions have completely closed bolls (or
capsules) while others are slightly or completely dehiscent. The latter is a useful trait to disseminate the seeds in the wild but is undesirable and selected
against by breeders for usage as a crop. (c) The seed coat color of linseed can vary from bright yellow to olive to dark brown and even include variegated.
Photos, reproduced with permission, are courtesy of Dr. Axel Diederichsen, Plant Gene Resources of Canada, Saskatoon, SK, Canada.
THE OILSEEDS | Linseed: Overview 263
time, height, and plant branching were measured. Seed quality to keep the soil inoculum low are effective ways of controlling
traits such as fatty acid composition, iodine value, oil content, the disease. The host disease resistance mechanism seems to be
and protein content were also evaluated. Finally, disease rat- controlled by a few major genes meaning that the development
ings for rust, pasmo, and powdery mildew were recorded. The of resistant varieties should be highly feasible in a reasonable
collection was genotyped with molecular markers and a time frame. However, the large environmental component
genome-wide association study revealed several marker-trait involved in the expression of wilt tolerance challenges plant
associations. These important studies provided background breeders and continues to impede breeding progress.
information on the flax genome and the genetics of important Rust spores are airborne and can infect flax plants when
traits that, combined together, will assist and accelerate temperature and moisture conditions are conducive. Visual
improvement of linseed by plant breeders. symptoms are the presence of pustules with or without chlo-
rosis or necrosis on the aerial parts of the plants. Interestingly,
it was while studying flax rust in the 1950s that Harold Henry
Yield Flor came up with the gene-for-gene concept, stating that for
each gene conditioning rust reaction in the host, there is a gene
In western Canadian field plots, yield of linseed accessions as counterpart in the fungus conditioning pathogenicity. This
high as 2.5 tonnes per hectare (t ha1) was reported, but higher concept has since been found to be applicable to many plant
yields have been measured (Paul Dribnenki, personal diseases. In flax, several rust resistance genes (R genes) have
communication, January 2014). However, this yield potential been cloned in the last two decades assisting in their introgres-
is not achieved for the crop at large and the yields of linseed sion in breeding lines, an important asset to breeders because
worldwide remain at < 1 t ha1. Even in the main producing rust-resistant varieties are the best way to counter this disease.
countries, it is still only 1.4 t ha1 and the rate of yield increase Powdery mildew infection appears as a white mass of myce-
remains below 1.5% per year, far less than its oilseed compet- lium and spores on the leaves. Heavy infection can occur under
itors canola (1.62 t ha1) and soybean (1.62.6 t ha1). Yield high humidity conditions. Similarly to fusarium wilt, resis-
improvement is therefore the main objective of linseed breed- tance seems to be controlled by a few major genes. As with
ing programs because it is the single most important trait to the other two previous fungal diseases, genetic resistance exists
ensure competitiveness. In recent years, plant breeders have and planting resistant cultivars is the best practice to control
realized that the narrow genetic base of the germplasm has the disease in areas where it is prevalent.
limited yield improvement. To overcome this limitation, lin- Pasmo is the most prevalent disease on the Canadian Prai-
seed breeders are now incorporating exotic material, using ries. Early infection of pasmo has a major detrimental impact
modified breeding schemes, and applying molecular markers on yield and quality. The early brown spots can evolve into
into breeding programs. For example, backcrosses of molecular complete leaf desiccation. The disease girdles the stems and
markers associated with thousand-seed weight to introgress the increases the prevalence of lodging, which has also been shown
trait into adapted elite germplasm are being implemented. to cause significant yield reduction in flax and negatively affect
Genomic selection could also prove useful toward yield seed quality. Early seeding of clean seeds, crop rotation, and
improvement, although there are no reports yet on its usage fungicide application are the best control methods because
in linseed improvement. commercial cultivars released to date seem to lack resistance
Many oilseed crops such as corn, canola, and soybean have to this disease.
well-established hybrid systems that can take advantage of
heterosis, also known as hybrid vigor. As mentioned earlier,
linseed is highly self-pollinated, and to date, a single hybrid Transgenic
has been released in China. Although hybrid production of
linseed bears promise and is theoretically possible, the devel- In 1996, CDC Triffid was the first and only genetically modi-
opment of an economically viable and stable system poses fied (GM) linseed variety to be registered in Canada. CDC
major challenges. Triffid was the result of the transformation of NorLin with a
modified Arabidopsis acetolactate synthase gene that conferred
resistance to residual soil activity of sulfonylurea herbicides.
Diseases Breeder seed of the variety was distributed to seed growers for
pedigreed seed production. In 199798, in light of the Euro-
Several diseases can affect the crop during its growing season. pean Union (EU) moratorium on GM crops and foods, the
The most important are fungal diseases that are prevalent in seed company owning the right to CDC Triffid voluntarily
most growing areas of the world: fusarium wilt caused by halted its seed multiplication and coordinated its retrieval
Fusarium oxysporum, rust caused by Melampsora lini, pasmo and destruction with the Flax Council of Canada to prevent
caused by Septoria linicola, and powdery mildew caused by potential harm to the industry. CDC Triffid pedigreed seed
Oidium lini. The crop can also be affected by other fungal, production was recalled and destroyed. In 2009, a shipment
phytoplasma, and viral diseases but these tend to either be of linseed from Canada to the EU tested positive for a well-
localized or have a negligible impact on yield or quality. known GM marker. Further testing confirmed the low-level
Fusarium wilt, a soil-borne pathogen, can wreak havoc on a presence of CDC Triffid. While this variety had received
crop because heavy infection of susceptible varieties results in approval for feed and industrial uses in Canada and the United
complete wilting of the plants and major yield reduction. The States, it constituted a nonapproved event in the EU and there
combination of moderate resistant varieties and crop rotation is zero tolerance for such. Canada and the EU entered into
264 THE OILSEEDS | Linseed: Overview
discussions to mitigate the crisis that led to the establishment sequencing, is proving viable in many major crops such as
of a stewardship program and testing protocols. In 2013, one wheat, corn, barley, and rice and bears promise for other
can conclude that the monitoring has been a success because crops such as linseed that has the advantage of a small
the low-level presence of the CDC Triffid gene construct in genome. Genetic transformation has the potential to bring
Canadian linseed has been substantially reduced. Unfortu- about novel products because linseed is readily amenable
nately, exports to the EU are only half what they were prior to this type of technologies. As a health food, the anti-
to 2009 indicating that market confidence remains to be fully nutritional compounds of linseed remain and will continue
regained. to be an important research target and the topic of various
research activities.
Conclusion
Linseed is a unique oilseed crop that combines high oil content Acknowledgments
with high content of ALA giving the oil distinctive drying
properties sought in the manufacturing of biodegradable The author sincerely thanks Drs. Helen Booker, Axel
industrial products. ALA, lignans, and high fiber content of Diederichsen, Paul Dribnenki, and Gordon Rowland and
linseed are also contributing to making linseed highly suitable Mr. Andrzej Walichnowski for their comments and edits
for the food and feed markets because these seed components made to this article. The author also thanks Ms. Michelle Beaith
have been linked to several health attributes in human and and Drs. Allan Green and Paul Dribnenki for the information
animals. While linseed remains a minor oilseed crop world- on the status of LinolaTM around the world and Ms. Kelley
wide, its versatility and uniqueness continue to be unmatched Fitzpatrick for the information on the regulatory affairs of
by the main oilseed crops. flax health claims.
Contents
Grain Legumes and Their Dietary Impact: Overview
Pseudocereals: Overview
Lupin: Overview
Amaranth: Overview
Beans: Overview
Buckwheat: Overview
Chickpea: Overview
Pea: Overview
Peanuts: Overview
Quinoa: Overview
Table 1 Names and regions of greatest genetic diversity for major grain legume species
Adapted from Sinha SK (1977) Food Legumes: Distribution, Adaptability and Biology of Yield. Rome, Italy: Food and Agriculture Organization of the United Nations. Based on Sinha
(FAO) and Adams MW and Pipoly JJ III (1980) Biological structure, classification and distribution of economic legumes. In: Summerfield RJ and Bunting AH (eds.) Advances in
Legume Science, pp. 116. Kew, UK: Royal Botanic Gardens; Zohary D, Hopf M, and Weiss E (2012) Domestication of Plants in the Old World, pp. 98100. Oxford, UK: Oxford
University Press; Grabiele M, Chalup L, Robledo G, and Seijo G (2012) Genetic and geographic origin of domesticated peanut as evidenced by 5S rDNA and chloroplast; DNA
sequences. Plant Systematics and Evolution 298: 11511165.
Table 2 Protein and other nutritional constituents of grain legumes rice). Oilseed legumes, including soybean (Glycine max) and
and other reference foods (g constituent 100 g food1) peanut (groundnut, Arachis hypogaea), in contrast to pulses,
accumulate higher protein, lower carbohydrate, and higher
Food Protein Lipid Carbohydrate Water lipid. Because of the high lipid content, they are often utilized
as a source of vegetable oil for culinary and industrial uses, but
Pulses
they are also important as pulse-like ingredients in prepared
Common bean 22.2 1.1 61.5 11.5
Lima bean 21.5 0.7 63.4 10.7 foods. For example, soybean is the base of many nonfermented
Chickpea 19.3 6.0 60.7 11.5 and fermented foods such as tofu and tempeh, while peanut is
Green gram 23.9 1.1 62.6 9.0 the main ingredient in some snacks and sandwich spreads. Lupin
Cowpea 23.8 2.1 59.6 11.1 (Lupinus L.) seeds protein, lipid, and carbohydrate composition
Lupin 36.7 11.5 45.4 is intermediate to that of pulses and oilseeds, and its dominant
Oilseeds use in many areas of the world is as an animal feed. However, in
Soybean 36.5 19.9 30.2 8.5 Europe and South America, it has a long, pulse-like history of
Peanut 25.8 49.2 16.1 6.5 human consumption and now enjoys increasing popularity as a
Cereals
food crop in North America and Australia.
Wheat, hard white 11.3 1.7 75.9 9.6
Grain legumes are ancient foods that help fulfill the ageless
Brown rice, raw 7.5 2.7 76.2 12.4
Animal products need for human sustenance. The word pulse arrived in mod-
Beef, lean ground 20.0 10.0 0.0 69.5 ern English from the Greek poltos meaning porridge of meal,
Pork, ground 16.9 21.2 0.0 61.0 by way of Latin (puls and pultis), Old French (pols), and
Fish, cod 17.8 0.7 0.0 81.2 Middle English (puls).
Milk, cows 3.25% fat 3.3 3.3 4.7 88.0 Gathering plant parts for food is a reliable way (in compar-
Egg, hens 12.5 10.0 1.2 75.3 ison with episodic hunts for animal prey) of feeding oneself in
a hunting and gathering society. Gathering pulse seeds is par-
Based on data from FAOSTAT Database http://www.fao.org/ag/; Mohamed AA and
ticularly convenient compared with parts of other plants. Pods
Rayas-Duarte P (1995) Composition of Lupinus albus. Cereal Chemistry 72: 643647;
from grain legume plants are handy and easily grabbed com-
Kohajdova Z, Karovicova J, and Schmidt S (2011) Lupin composition and possible use
in bakery A review. Czech Journal of Food Sciences 29: 203211. pared with subterranean roots and tubers. No tools other than
fingers and palms are needed to grab and hang on to the large
seeds. Keeping both feet firmly on the ground is less daunting
(Tables 1 and 2). Pulses also contain high levels of dietary fiber than climbing trees for aerial fruits and nuts. The dense seeds
(20 g fiber per 100 g compared with 12 g fiber per 100 g for are more filling than leaves and will satisfy hunger in the
wheat, 9 g fiber per 100 g for soybean, and 3 g fiber per 100 g for absence of meat if hunting is unsuccessful. As a bonus to
THE LEGUMES AND PSEUDOCEREALS | Grain Legumes and Their Dietary Impact: Overview 267
those concerned about times of scarcity, the seeds can be dried (Aztec and Mayan New World civilizations), and those where
and stored for a period of months or more without loss of food predominant religious practices excluded animal protein con-
value when eaten or viability when planted. Pulse seeds are sumption (Hindu veneration of the cow in South Central
large relative to cereals, so they are convenient to handle for Asia). Even today, those regions of the world consume more
food preparation and planting. Because of their convenience, pulses than elsewhere.
availability, and composition, wild pulses were an accessible, Besides influencing the amount of pulse contribution to the
nutritional contributor to early hunting and gathering societies diet, cultural inertia expresses a parallel effect on the pulse
across the world. species consumed within a region. Those pulses with long
The wild progenitors from which pulses were domesticated records of use in a particular country or region of the world
have been identified, with the exception of Vicia faba. The continue as dietary preferences today (Table 3). The common
archeological record for some pulses, particularly Phaseolus bean, Phaseolus vulgaris, appears to be the most universally
species and Lens culinaris, is more complete than for others consumed pulse species.
such as Vigna unguiculata and Cajanus cajan. In whatever man- Early in the Common Era, the 40-day fast of Jesus in the
ner they were developed, the domesticates quickly spread desert became the standard reference for monks belonging to
across regions and continents, particularly to regions lacking the Roman Church for the custom of mortification of the body.
a wide range of highly desirable, locally domesticated food The fasting monks were required to remain engaged in con-
crops such as Europe and North America. Over time, domesti- ducting the monasterys daily functions; thus, the elders of the
cation, selection, and trade resulted in a wide variety of pulses monastic community began, in the fourth century, to standard-
grown, contributing protein to human diets across the globe. ize the fast diet in order to ensure that their workforce received
Pulse and cereal grain domestication appear to have taken at least the nutrition required to keep the body and soul intact
place at roughly the same time in human history, which sug- but without arousing earthly sensuality in monks eating
gests that these two foods may be ancient partners in providing flavorful food. A satisfyingly dull puree of pulses, accompanied
human nutrition. While cereals provided most of the energy perhaps by a vegetable soup, became typical of the single meal
and much of the protein requirements of humans, pulses allowed during fasts. Monks certainly had their fill of pulses as
played a strong supporting role as a contributor of concen- they contemplated eternity; the fifth-century monasteries
trated dietary protein. Concurrent pulse and cereal production scheduled 230 fast days per year.
and their dietary consumption resulted in more sustainable Before widespread adoption in seventeenth-century Europe
farming systems, as well as better nutritional balance, than of cuisine innovations such as leavened bread, people of medi-
would have been derived from production and consumption eval societies typically ate warm servings of porridge twice
of cereals alone. daily. These plates or bowls of boiled gruel incorporated read-
ily available and affordable cereal grains such as barley and
wheat and also pulses such as the wild-growing or domesticated
lentils first found in the Eastern Mediterranean and West Asia.
Consumption Meat consumption declined with increasing population,
decline of wild game stocks, and movement of people into
Current grain legume consumption in many parts of the Old towns and cities. World trade, natural sciences, and the influ-
World and New World appears to be affected by cultural iner- ence of aristocratic culture began to influence diets in the late
tia; past consumption patterns in those regions are echoed in 1600s. Three-a-day meals became common across socio-
the present. While pulses are common components of diets economic groups, and cookbooks spread food culture through-
around the world, the extent to which they contribute to the out the urban and rural societies. New foods including spices,
diet correlates closely with the historic availability and accep- potatoes, and a thickener made by browning flour in fat known
tance of animal protein. Ancient civilizations that consumed as roux made their appearance and elbowed the cereal-and-
more pulses were those where animal protein was less available pulse-based porridge out of popularity. The Protestant Refor-
or acceptable. They included those that maintained high mation deemphasized fasting, which further reduced the
human population densities and overwhelmed local supplies consumption of sturdy but unglamorous pulse porridges.
of wild and domestic animal protein (East Asian Old World Two vastly different problems influence contemporary
civilizations), those where few animals were domesticated grain legume consumption: malnutrition in food-deficit
Adapted from Smart J (1990) Grain Legumes: Evolution and Genetic Resources. Cambridge: Cambridge University Press.
268 THE LEGUMES AND PSEUDOCEREALS | Grain Legumes and Their Dietary Impact: Overview
regions and diet-related diseases such as obesity, diabetes, and trend in developed countries between the 1960s and now is, if
heart disease in food surplus regions. In food-deficit regions, anything, to include slightly less pulse in diets. In contrast, the
increased consumption of pulse crops could provide a concen- increased protein and energy consumption between the 1960s
trated source of protein, vitamins, and micronutrients that and now reported in the least developed countries (as defined
complement traditional low-protein, high-carbohydrate grain by the Food and Agriculture Organization of the United
and root-based diets. Additionally, because of the legumes Nations) results from inclusion of both more animal protein
ability to support biological nitrogen fixation and its contribu- and more plant protein in the diets, including a small increase
tion to increased crop diversity, local farms could benefit from in pulse consumption. One aggravating reason for low pulse
agricultural systems that integrate pulse crops adapted to the consumption is flatulence resulting from intestinal fermenta-
regional environment. In food surplus regions, substitution of tion of complex sugars and dietary fiber. If gassiness causes the
plant-based diets for excessive meat consumption and con- consumer discomfort or embarrassment, pulse foods will be
sumption of foods rich in dietary fiber such as pulses are avoided when other choices are available.
among the strategies promoted to combat the dramatic As one might expect, food use of pulses is higher in least
increase in diet-related diseases. developed countries when compared with developed countries
(Table 6). However, the percentage of total domestic pulse
supply used for food in the least developed countries declined
Composition substantially over the past 40 years, while in developed
countries, it increased slightly. Feed use and food use are mutu-
Pulses, with roughly double the protein content of cereals, are ally exclusive, so in the least developed countries, there is a
damned with faint praise as the protein of the poor. They relatively low percentage of supply used for feed, while in devel-
contribute excellent nutrition to all, but because they are inex- oped countries, there is a high percentage used for feed. Table 6
pensive relative to meat, they are particularly beneficial to indicates that on average both the least developed and the devel-
subsistence farmers and those with low disposable income in oped countries exported more pulse seed than they imported,
food-deficient countries. In such circumstances, pulses can with the exception of developed countries in the 1960s. Trade
account for 11% of daily protein intake and 6% of the
daily energy in the human diet (Tables 4 and 5). The praise, Table 5 Energy per capita consumption in two decades by country
however, is faint and damning because this association with groupings (kcal per person per day)
poverty, a connection perhaps firmly established with the por-
ridges of the Middle Ages, stigmatizes pulse foods among those Least developed
with or aspiring to have higher income who increasingly favor countries Developed countries
meat protein. In developed nations, protein consumption has
Protein sources 196271 200211 196271 200211
increased from the 1960s to now and is associated with
increased consumption of animal products. Scarcely, 2% of Total plant 1818 (93)a 2055 (92) 2176 (71) 2339 (71)
dietary protein in developed nations, and 1% of energy, Pulses 79 (4) 94 (4) 34 (1) 28 (1)
comes from pulses. In more affluent settings, pulses remain Cereals 1248 (64) 1324 (59) 1030 (34) 954 (29)
an underused alternative to high-priced animal protein. Inter- Starchy roots 171 (9) 202 (9) 146 (5) 132 (4)
estingly, as noted in the preceding text, at least one segment of Oilseeds 43 (2) 50 (2) 19 (1) 30 (1)
affluent society, lifestyle- or health-conscious eaters such as Total animal 142 (7) 180 (8) 886 (29) 935 (29)
Meat 54 (3) 67 (3) 277 (9) 352 (11)
vegetarians or those seeking the nutritional benefits of high
Eggs 3 (0) 5 (0) 42 (1) 44 (1)
dietary fiber, still values these plants. Even so, the consumption Milk 53 (3) 70 (3) 296 (10) 313 (10)
Fish 13 (1) 19 (1) 35 (1) 48 (1)
Table 4 Protein per capita consumption in two decades by country Grand total 1961 2236 3061 3274
groupings (g protein per person per day)
a
kcal capita1 day1 (% of grand total).
Least developed nations Developed nations Based on data from FAOSTAT Database http://www.fao.org.
movement of pulses, however, is much more complex than can at least 2820 mg lysine. This requirement could be satisfied by
be summarized using these aggregates of countries grouped by consumption of 288 g of wheat (Triticum aestivum) and 124 g of
economic development. The reader is encouraged to examine bean (Phaseolus vulgaris) (Tables 2 and 7). To meet the lysine
trade data at the regional and country levels in order to discern requirement through consumption of wheat alone, the same
how pulses are moving around the globe. adult would need to eat 960 g wheat containing 108 g total
protein. The intentional combination of cereals and pulses at
meals offers inexpensive (in comparison with meat) plant-based
Protein Complementation diets with balanced amino acid composition.
While the central theme of this overview is the enormous
Understanding amino acid balance is key to recognizing the contribution made by dry grain legume seeds, and in particular
special contribution made by grain legumes to protein comple- pulse seeds, to the human diet, these crops have other food
mentation in cereal-based diets. Different sources of plant pro- uses. Some species are used as vegetables, as either green mature
tein are made up of characteristic concentrations of or immature seeds or pods, as sprouted seeds, or even as stocks
indispensable (essential) amino acids and therefore contribute for fermentation. As noted earlier, soybean is the base of many
upon digestion a different balance of amino acids for human nonfermented and fermented foods such as tofu and tempeh.
nutrition. Amino acid balance in a particular protein source can Fermented pulse seeds primarily involving black gram (Vigna
be scored against that of a reference protein, typically hens egg mungo), Bengal gram (Cicer arietinum), and green gram (Vigna
(regarded as well balanced in relation to adult human metabolic radiata) are components of diets particularly in India as idli,
needs) or milk (Table 7). No plant-based protein mimics the dhokla, and khaman. Fermentation, sometimes in combina-
indispensable amino acid balance present in eggs, so it follows tion with cereal grains, causes desirable reduction in beany
that each type of plant protein must exhibit a characteristic set of flavor, aroma, and consistency and improves digestibility.
limiting amino acids. For example, whole wheat flour and white- Nonfood uses include dry seeds used as animal feed and
seeded common beans generally have lower proportions of the forage from pulse vegetation, in aquaculture feeds, and even
indispensable amino acids than eggs or milk does. In particular, as ornamentals. In a Southeast Asian setting, well-established
the amino acid lysine limits efficient utilization of wheat protein shrubs of pigeon pea, a short-term perennial grown on field
in human metabolism, while the indispensable sulfur- banks or as an intercrop, as well as dry season-grown horse
containing amino acid methionine limits bean protein. The gram and cowpea, can be intermittently grazed or cut for hay.
indispensable amino acid balance of wheat and bean is charac- N concentration of the hay varies between 2.5% and 4.5%,
teristic of other cereal grains and pulses. Cereal grains are typi- but livestock may need a few days to be accustomed to the
cally deficient in lysine but high in methionine, while the feed. Dairy cattle are reported to similarly consume and uti-
opposite is the case for pulses: high in lysine and low in methi- lize faba bean silage and grasslegume silage. From a cool
onine. Cysteine, a dispensable sulfur-containing amino acid season perspective, the seeds of soybean, pea, lentil, faba
synthesized from methionine and serine, is in many references bean, and chickpea can be used as a protein source in live-
listed along with methionine as a limiting amino acid of pulses. stock feed. Oil and fiber contents of the seeds affect the
Plant protein complementation, accomplished by consum- metabolizable energy (MEn) values of pulses when fed to
ing protein from more than one plant source, can provide various types of livestock. For example, the relatively low-
humans with a balanced diet of amino acids approximating oil, high-fiber pea has a low (2570 kcal kg1) MEn when
that found in hens egg. Grain legumes, with their high concen- fed to poultry, while for swine, the pea MEn is much higher
tration of lysine relative to lysine-deficient cereals, and cereals, (3740 kcal kg1). The pulse seed amino acid profile for
with their high concentration of methionine and cysteine rela- monogastric feeds, while good, is inferior to soybean. Pea
tive to that found in pulses, combine as ideal dietary partners, and faba bean protein is highly digested, but some livestock
because the amino acids making up their proteins are very good feeding studies have shown an increased excretion of endog-
complements. For example, at 0.7 g kg1 body mass per day, an enous protein, which may be due to seed antinutrients. Dry
85 kg adult would require 60 g of protein per day containing roasting the whole seeds has been shown to improve
Table 7 Indispensable amino acid requirements in dietary protein and the amino acid composition of selected foods (mg amino acid per g protein)
Amino acid Infant requirement Adult requirement Hens egg Cows milk Whole wheat flour White bean flour
Histidine 23 17 24 28 22 28
Isoleucine 57 23 63 60 40 42
Leucine 101 52 88 98 63 76
Lysine 69 47 70 79 26 72
Methionine cysteine 38 23 56 34 35 19
Phenylalanine tyrosine 87 41 98 96 81 77
Threonine 47 24 49 45 27 39
Tryptophan 18 6 16 14 11 10
Valine 56 29 72 67 43 46
Based on data from Anonymous (2002) Dietary Reference Intakes for Energy, Carbohydrates, Fiber, Fat, Protein and Amino Acids (Macronutrients), Chapter 10, p. 71. Washington,
DC: National Academies Press (online version available on p. 535 at the URL http://www.nap.edu/books/0309085373/html/).
270 THE LEGUMES AND PSEUDOCEREALS | Grain Legumes and Their Dietary Impact: Overview
digestibility and protein utilization. Pulses have great poten- phytohemagglutinins) of different animal species depending
tial for use in aquaculture diets, but in this application, they on the specific receptors on the cell membrane surface. This
are hampered by their carbohydrates. However, field peas agglutination specificity provides a method of rapid detection
(Pisum sativum), faba beans (Vicia faba), chickpeas (Cicer and classification. Lectins differ in the severity of their impact.
arietinum), and vetch (Vicia sativa) seeds can be used in Some (mainly found in Phaseolus vulgaris) are classified as
aquaculture feeds for juvenile silver perch following de- toxic, others are considered only growth inhibitory (Phaseolus
hulling, which removes much of the undesirable carbohy- lunatus), and still others essentially nontoxic (Pisum sativum,
drate fraction. Lens culinaris, and Vicia faba). The role of lectins in the plant is
not well established, although they appear to help the plant
defend itself against specific bacteria, fungi, and insects. They
Antinutritional Factors may also aid processing and movements of glycosylated stor-
age materials during seed maturation, assist in the establish-
Low digestibility hampers full utilization of grain legume pro- ment of the symbiotic relationship of legumes and rhizobia,
tein. In part, the problem may result from the more rapid and promote cell growth and division. Or, like protease inhib-
discharge, relative to other foods, of digesting pulses from the itors, lectins may simply serve as storage proteins.
intestinal tract and reduced protein hydrolysis by gut enzymes. The effect of ingested lectins on human and other animal
However, antinutritional factors, compounds in pulses that metabolisms varies according to the lectin type, the species
interfere with their digestion and metabolism, also play a ingesting the protein, and the age, nutritional, and health
major role in restricting dietary utilization in some pulse spe- status of that human or animal. To affect human metabolism,
cies. These compounds generally include proteinaceous mole- plant lectins first must bind to epithelial cells in the gut. This
cules such as protease inhibitors (particularly trypsin binding requires the presence of particular carbohydrate
inhibitors) and lectins and also nonproteinaceous compounds groups on the lumen of the gut epithelial cells. Dietary lectins
such as tannins. strongly resist proteolytic degradation in the gut. Toxic dietary
Most of the wild relatives of pulses contain toxins and lectins can modify hormone balance and lipid and muscle
antimetabolites. One of the key features of pulse domestication metabolism. These modifications deplete stores of lipid, glyco-
from wild forms is human selection for lower levels of these gen, and protein and lead to weight loss. Some nontoxic lectins
compounds. Traditional food preparation techniques used for bind to the gut epithelium but have no deleterious effect on
pulse seeds result in reduction or elimination of these meta- metabolism, while others exhibit only limited binding capac-
bolic impediments. Some modern food pulses today, in par- ity. Deleterious lectin effects appear to be reversible.
ticular the Phaseolus species, can still contain sufficiently high Tannins can form strong cross-linked complexes with die-
levels of antinutritional factors in their dry seeds to cause tary proteins and enzymes. The antinutritional impact of tan-
digestive difficulties if eaten without proper processing. For nins on digestion and metabolism is not fully understood.
this reason, pulse seeds are not eaten raw, but processed in a Protease inhibitors and lectins are heat labile and
way (moist heat >100 C, sprouting, and fermentation) that rendered innocuous by usual methods of cooking. Moist heat
inactivates the antinutritional factors. treatment such as the traditional soaking of beans, followed
Protease inhibitors, a major class of antinutritional factors by cooking in boiling water is generally considered to be
in pulses, often inhibit the digestive enzyme trypsin but may sufficient to inactivate protease inhibitors. Soaking may help
act more broadly by inhibiting chymotrypsin and other serine leach out some tannins, but heat treatment will not reduce the
proteases. The role of these inhibitors in the plants that pro- impact of that remaining in the seeds.
duce them is uncertain but may involve defense against disease Particular grain legume species accumulate additional clas-
and insects. An alternative explanation is that these com- ses of antinutrients. For example, Lupinus albus seeds may
pounds might simply serve as protein reserves high in sulfur- contain quinolizidine alkaloids that give the seeds a bitter
containing amino acids such as cysteine and coincidentally flavor and, if not successfully removed prior to ingestion, can
inhibit human digestion. The primary concern regarding raw result in poisoning. A. hypogaea cotyledons contain low-
pulse consumption is with very young humans and other molecular-weight glycoproteins to which some individuals
young animals that are more susceptible to the effects of pro- exhibit allergic reaction upon contact and ingestion. Neuro-
tease inhibitors than adults. In vitro studies suggest that prote- and osteolathyrism, the former a reversible paralysis and the
ase inhibitors can impair nutrient utilization and reduce latter a skeletal abnormality, have been linked to excessive
growth rates in these groups. In vivo animal studies indicate consumption of grass pea (Lathyrus sativus) seeds. Broad bean
that interference with growth occurs primarily when the level (Vicia faba) antinutrient levels are low and do not require moist
of dietary protein itself is limiting. Pancreas activity can be heating or fermentation to inactivate but can induce favism, a
affected when dietary protein digestion is inhibited and high hemolytic anemia, in individuals with congenital deficiency of
levels of undigested protein interfere with appropriate pancre- glucose-6-phosphate dehydrogenase. Pulses of Mediterranean
atic regulatory control. Since pulses are used in some human origin such as peas (Pisum sativum), chickpeas (Cicer arieti-
infant diets, cooking must be done with particular care to num), and lentils (Lens culinaris) generally have low toxic and
ensure that protease inhibitors are inactivated. antinutrient levels. While protease inhibitors have been found
Lectins, another major class of antinutritional factors in in cowpeas (Vigna unguiculata), they are apparently not present
pulses, are proteins that bind to carbohydrates or to molecules in serious levels, nor have harmful quantities of cowpea lectins
containing carbohydrates. This binding capacity allows them been reported. Vigna mungo and Vigna radiata seeds likewise
to agglutinate red blood cells (lectins are sometimes called contain little in the way of antinutritional factors.
THE LEGUMES AND PSEUDOCEREALS | Grain Legumes and Their Dietary Impact: Overview 271
Agronomy similar growth period will benefit directly from the pulse crops
nitrogen fixation, but perennials, crops with longer growth
The Leguminosae family is enormous, with an estimated periods, or subsequent crops may benefit from the nutrients
16 00019 000 species in 750 genera. If the raw number of released through the decomposition of pulse crop litter. The
species were to determine the greatness of a family, Legumino- agronomic impact of a legume crop on the intercrop will
sae would be eclipsed only by Orchidaceae and Compositae. If depend on the crop species (and even the cultivar) in the
instead the economic value of a family were considered, intercrop, the extent of its inclusion in the plot, tillage, and
legumes would be second only to Gramineae. Taxonomists soil type.
place pulses in the legume subfamily Papilionoideae, mostly Pulses can assist in soil improvement, particularly when
in tribe Phaseoleae, but a few in the tribes Vicieae, Cicereae, used as green manure crops when the whole plant is plowed
and Genisteae. down into the soil. Fixed nitrogen and in some cases phospho-
Cropping systems develop in response to the demand for rus are made available to subsequent crops, and the added
particular crops, soil nutrient status, soil physical structure, and organic matter contributes to improved soil structure. Also,
biotic competition and are constrained by available resources the deep tap roots of some crops such as pigeon pea reduce
(including climate) and knowledge. Pulses in temperate crop- soil erosion and mine deep soil for nutrients that are returned
ping systems are commonly planted in monocultures as part of to the surface through leaf drop.
a rotational sequence with maize (Zea mays), small grain In sustainable shifting cultivation systems of intercropping,
cereals, or occasionally forages or periods of fallow in dryland a few years of crop production are followed by a decade or
areas. Since in these cropping systems pulses are typically cash more of bush fallow where cropping is suspended and native
crops and occasionally feed crops, their inclusion in the rota- vegetation regrows. Cultivated pulses grown during the crop-
tion and total acreage from year to year depends on the status ping years provide concentrated protein for human diets using
of cash markets and economic benefits of growing that pulse nitrogen fixed from the atmosphere and help maintain the soil
crop relative to other crops, rather than with maintaining a set nitrogen status by not depleting fixed soil nitrogen. During the
crop rotation, maintaining particular soil characteristics, or bush fallow stage, indigenous legumes, including trees, con-
addressing biotic competition. Higher-value crop alternatives tribute to the restoration of soil nitrogen. When the human
may be readily substituted for pulses in the rotation in some population density is low, relative to available land, and bush
climates. National farm strategies with strong economic fallow periods are sufficiently long to restore soil nutrients,
impact, such as government subsidies or deficiency payments, shifting cultivation can be sustained. With increased popula-
can affect pulse production by inclusion or exclusion of partic- tion or decreased land base, the cropping period must be
ular pulse crops from the programs. For example, in the United extended, bush fallow period reduced, and as a result, soil
States, the Agricultural Act of 2014 identifies specialty crops, nutrient status and crop productivity decline.
including pulse crops, for indirect support such as assistance Since in most cases pulse crops are not alternate hosts for
with marketing, promotion and trade, crop insurance, and cereal crop diseases, their use in crop rotations will help reduce
research and research-based information extended to growers. cereal disease inocula. However, pulses are certainly not
While not a big producer of any of these pulse crops, the immune to pests; they are attacked by many fungal, bacterial,
United States is an exporter, and additional production stimu- and viral diseases and a range of sucking, boring, and chewing
lated by the Agricultural Act could affect prices in other pulse- insects. Also, pulses are typically poor competitors against
exporting or pulse-importing countries. weeds, except perhaps under stress conditions such as drought
Leguminosae will collaborate with genus Rhizobium bacte- conditions where annual weeds are also checked.
ria in an ecologically unique symbiotic relationship. This sym- At harvest time, pulses often disappoint and frustrate
biosis supports biological nitrogen fixation and frees legumes growers by failing to meet yield potential. Pulses are often
from the need to access soil sources of fixed nitrogen. Rhizobia considered to be extremely sensitive to environmental stress,
invade the legume root hairs and reside in cortical root swell- relative to the more robust cereals, a sensitivity which may in
ings (nodules) where nitrogen from the atmosphere is fixed for part be associated with maintaining the symbiotic relationship
eventual transport within the plant and assimilation into pro- with Rhizobium. The RhizobiumLeguminosae symbiosis
tein. Even though a legume can support the symbiotic relation- requires energy supplied by the host plant to support the
ship, it will not occur unless the rhizosphere contains metabolic needs of the bacterial colonies in return for assimi-
compatible rhizobia. Strains of rhizobia differ according to lated nitrogen. In the case of the pulses, which are typically
the particular legume hosts that they will inoculate and are grown as annual crops, the symbiotic relationship must be
classified according to this host range. reestablished with every new crop. Farmers watching the
Cropping systems in developing countries typically rely on color of the growing annual pulse crops in their fields can
intercropping where pulses are planted along with other clearly see the lag time between planting and the establishment
species, commonly cereals and vegetables. The intercropping of functioning nodules. For example, a temperate common
strategy can provide subsistence farmers with several essential bean crop planted in a soil with compatible rhizobia will tend
foods from one plot of land, produce higher protein yield per to have a deeper shade of green about a month after planting, as
hectare, combine complementary canopy structures (low- fixed nitrogen becomes available to the plant. During the early
stature pulses with high-stature sorghum (Sorghum bicolor) portion of the growing season, the host plant pays the costs of
and maize (Z. mays)), balance utilization of soil moisture establishment and only later receives the benefits. Plant-
following rainy periods, or reduce crop competition by weeds generated energy used to support the symbiotic relationship is
and insects. It is unlikely that a nonlegume companion crop of not available to the plant for dry matter production, so some
272 THE LEGUMES AND PSEUDOCEREALS | Grain Legumes and Their Dietary Impact: Overview
amount of yield penalty should be anticipated. Effective nodu- Which pulse crop is the most universally consumed?
lation and nitrogen fixation depend on favorable interactions A porridge made of grain legumes and cereal grains was a
between the host and appropriate Rhizobium genotypes and the common food in medieval societies. What events and
environment. If these interactions falter and nitrogen fixation cultural shifts resulted in the decline of porridge in diets?
lags behind demand, dry matter production will suffer. When did this transition take place?
Review the change in protein consumption from pulses and
Final Words from meat in the 1960s compared with recent years. How
do the trends differ between developed and developing
Declining per capita pulse consumption, particularly given the nations?
nutritional value of pulses and their affordability, is a cause for Identify examples of protein complementation in your diet.
concern. Although some regions in developed countries now Do any of your examples involve combinations of cereal
benefit from new pulse export opportunities and are increasing grains and grain legumes?
production, this expansion reflects changing markets and What are examples of nonfood uses of grain legumes?
increased population, not increased dietary utilization. Much Identify a reason for and a reason against using grain
more global impact could be accomplished by addressing con- legumes for animal feed.
straints to consumption, including the social stigma associated Why are pulse seeds not eaten raw? What types of proces-
with eating poor pulses compared with rich meat, flatulence, sing are necessary and how do they change the grain?
yield variation, and agronomic constraints to product quality How would you integrate grain legumes into a cropping
and stable high yield. Further increasing pulse consumption in system? What benefit would they provide to the system?
developing and low-income countries may be impossible Describe the symbiosis that is key to the role of grain
given the strength of the human drive toward diets associated legumes in the cropping system.
with affluence. Perhaps, the greatest initial gains could occur What constraints to consumption must be addressed before
among consumers in developed countries currently with very grain legumes can play a larger role in global nutrition?
low pulse consumption. The keys to higher consumption in
those countries may involve increasing the availability of
value-added pulses such as those produced by certified organic
methods or food quality assurance programs and highlighting Exercises for Readers to Explore the Topic Further
their inclusion in national and regional cuisine initiatives. In
tandem, these efforts to add value and visibility could impart Can you outline a climate change scenario that would
some cache to pulses and improve their appeal. influence the production and consumption of grain
legumes?
Then said Daniel to Melzar, whom the prince of the eunuchs had set Genetic transformation is not a common strategy for the
over Daniel, Hananiah, Mishael, and Azariah, prove thy servants, I improvement of pulse crop cultivars. Why is this? What
beseech thee, ten days; and let them give us pulse to eat, and water to factors might encourage or discourage the development of
drink. Then let our countenances be looked upon before thee, and GMO pulse cultivars?
the countenance of the children that eat of the portion of the kings
meat: and as thou seest, deal with thy servants. So he consented to What are the key challenges in producing pulse crops in
them in this matter, and proved them ten days. And at the end of ten organic production systems? What are the benefits?
days their countenances appeared fairer and fatter in flesh than all Few fast-food outlets include pulse-based foods on their
the children which did eat the portion of the kings meat. Thus menus. Why is this? What types of meals made from pulses
Melzar took away the portion of their meat, and the wine that they
might have promise as a fast and nutritious food?
should drink; and gave them pulse.
Daniel 1:1116 (RSV) What types of research have been conducted on the impact
of pulse-based foods on cancer and obesity? What have
been the outcomes of those research projects?
How can the social stigmas associated with pulse-based
foods (poverty and flatulence) be overcome?
How could the legumeRhizobium symbiosis be improved?
Exercises for Revision
What seed constituents are similar between pulses and See also: The Basics: Taxonomic Classification of Grain Species;
oilseed legumes? What seed constituents differ? Which The Oilseeds: Oilseeds: Overview; Soybean: Overview; Grains
grain legumes seed constituents are roughly intermediate Around the World: Grain Production and Consumption: Overview.
between pulses and oilseed legumes?
How do seed constituents of grain legumes compare with
those of cereal grains?
Did domestication of grain legumes generally occur in the Further Reading
same region of the world or in vastly different regions?
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Expanding the Production and Use of Cool Season Food Legumes, pp. 5376.
Dordrecht, The Netherlands: Kluwer Academic.
Jansman AJM, Hill GD, Huisman J, and van der Poel AFB (eds.) (1998) Recent
Advances of Research in Antinutritional Factors in Legume Seeds and Rapeseed.
Wageningen, The Netherlands: Wageningen Pers. Relevant Websites
Labuza TP and Erdman Jr. JW (1984) Food Science and Nutritional Health: An
Introduction. St. Paul, MN: West Publishing. http://www.fao.org FAOSTAT Database, a comprehensive resource for agricultural
Marquardt RR and Bell JM (1988) Future potential of pulses of use in animal feeds. production data, including than on pulse crops.
In: Summerfield RJ (ed.) World Crops: Cool Season Food Legumes, pp. 421444. http://www.icarda.org The website of the International Center for Agricultural Research
Dordrecht, The Netherlands: Kluwer Academic. in the Dry Areas, based in Syria, has useful information on pulse agronomy and
Reddy NR, Pierson MD, and Salunkhe DK (eds.) (1986) Legume-Based Fermented utilization.
Foods. Boca Raton, FL: CRC Press.
Pseudocereals: Overview
RJ Fletcher, University of Queensland, Gatton, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition, volume 2, pp. 488493, 2004, Elsevier Ltd.
Learning Objectives
Current Role of Pseudocereals with sugar to make confectionery items, rolled into balls,
cooked as porridge or popped.
The pseudocereals are relatively unimportant on a global scale Grain amaranth has enjoyed a resurgence in recent years,
today. However, they have been significant contributors to the especially through the health food market, since its crude
human diet in certain defined regions in the past and could be protein is high (1419%), with a high lysine (up to perhaps
developed again as important new crops. For example, pseu- 6% of the protein) and tryptophan content. It appeals to the
docereals may play a role in human nutrition for those who modern consumer since these essential amino acids are low in
have allergies to traditional cereals; for the primary producer, cereals.
pseudocereals can play a role in cereal rotations, reducing the Grain amaranth is being developed as an energy food to be
buildup of grass weeds, pests, or diseases. combined with traditional cereal grains in breakfast food,
The three best-known pseudocereal crops are grain ama- bread, multigrain crackers, pastes, pancake mixes, or popped
ranth (Amaranthus caudatus; A. cruentus; A. hypochondriacus; as a snack food product. Popping can, however, affect its
family: Amaranthceae), quinoa (Chenopodium quinoa subsp. nutritional quality.
quinoa; Chenopodiaceae), and buckwheat (Fagopyrum The grain is easy to digest and heat processing improves its
esculentum; Polygonaceae). Each is briefly described below, digestibility.
from the viewpoint of functionality. A range of potential Amaranth starch (up to 69% of the grain) is principally
pseudocereals from other plant families is then presented, amylopectin; granules are relatively small (13 mm) compared
with dicotyledonous families followed by monocotyledonous with cereals (330 mm) and have a higher solubility and gela-
families. tinization temperature, rendering a distinctive gel. The seed can
comprise as much as 10% oil; this oil contains squalene, which
is used in cosmetics manufacturing. Antinutritional trypsin
Weeds inhibitors occur in concentrations up to twice that observed
in wheat.
Many of the species listed as potential pseudocereals below are
currently regarded, in some areas, as weeds; one of them,
Amaranthus retroflexus, could be regarded as the worlds most Potential pseudocereals in the Amaranthaceae family
cosmopolitan weed. Amaranthus dubius (spleen amaranth, khada sag), Amar-
A weed is defined as a plant which is growing in the wrong anthus frumentaceous (poong keeray, tola kura), Amaranthus
place. If some of the potential pseudocereals listed below can tricolor (tampala), Amaranthus graecizans (spreading pig-
be commercialized, they would no longer be regarded as weed), Amaranthus blitum (livid amaranth), Amaranthus
weeds. quitensis (yuyo colorado, sangorache), Amaranthus retro-
flexus (redroot, pigweed), Amaranthus spinosus (spiny ama-
ranth): the seeds are eaten raw in India and other places or
Warning mixed with other grains or processed into flour or other
products.
The historic use of any obscure plant by a culture for medicinal
or similar personal use does not mean that the plant is safe for Achyranthes aspera (devils horsewhip, prickly chaff-flower):
the seeds have been used in desert areas in India as a famine
use as a human food. Poisonous or antinutritional factors exist
food.
in many plants and their historic use was often complemented
by elaborate preparation to reduce the effect of any poison. As Celosia argentea var. argentea (red-fox), Celosia argentea var.
cristata (cockscomb): the seeds of these species have been
an example, Australian aborigines were able to consume cycad
used as famine foods.
seeds and yams but only after extensive preparation, which
reduced the toxicity of the material.
Regular use of a plant over a lifetime by a group of people
may also render a plant relatively safe for that group of people, Chenopodiaceae
whereas a newcomer who has never eaten the food can suffer Chenopodium quinoa subsp. quinoa (quinoa)
serious effects on eating it for the first time. Chenopodium quinoa subsp. quinoa is native to the border areas
between Peru and Bolivia. It held a special position in the
agricultural and ceremonial life of the Inca people and was
Amaranthaceae
grown in Colombia, Peru, the southern areas of Bolivia, the
Amaranthus caudatus, A. cruentus, A. hypochondriacus northern areas of Argentina, and the central areas of Chile.
(grain amaranth) Quinoa grain has a protein content ranging as high as
Amaranthus caudatus (Inca wheat, love-lies-bleeding) is native 22.8%, a carbohydrate content up to 77.4%, a fat content up
to the northern higher-altitude regions of Bolivia, Peru, and to 9.5% (and relatively stable), and a fiber content up to 5.8%.
Ecuador, whereas Amaranthus cruentus (Purple amaranth) and The grain is higher in protein, fat and fiber, and lower in
Amaranthus hypochondriacus (Princes feather) are native to carbohydrate content than most comparable cereals, due prin-
Guatemala and Mexico. The species were grown by pre- cipally to the proportional size of the embryo within the grain
Colombian civilizations, Aztecs, and indigenous US tribes (up to 30% of the grains gross weight, compared with 1% for
but have mostly been replaced by cereal crops. most cereals).
Depending on the species, grain amaranth seeds are The proteins in quinoa are principally albumins and glob-
sprouted, parched, toasted, ground into flour, baked, mixed ulins, which are at higher concentrations than is normally
276 THE LEGUMES AND PSEUDOCEREALS | Pseudocereals: Overview
found in the major cereals (wheat, rice, or maize). The amino or a granulated form to be boiled, steamed, or baked. Buck-
acid balance (higher in histidine, lysine, isoleucine, methio- wheat flour itself is used in soba-style Japanese noodles,
nine, and cysteine content) and mineral content (calcium, polenta (a form of porridge), desserts, ice cream cones, dietetic
magnesium, phosphorus, potassium, and iron) of quinoa foods, and canned meat or vegetable products and as a com-
grain is superior to most cereals. However, there are saponins ponent (3040%) of the flour used in pancakes, breads, pasta
in the seedcoat which have antinutritional properties. products, cakes and dumplings, often combined with wheat
flour.
Potential pseudocereals in the Chenopodiaceae family It is expected that buckwheat flour could be used more
Chenopodium album (fat hen, lambs quarters). The seed is readily in specialty breads, pasta, extruded snack food, and
sprouted or ground into flour for breads, pancakes, muf- ready-to-eat cereals.
fins, and biscuits in Russia and China; the grain is reported The proteins in buckwheat grain are principally globulins
to have 16.1% protein, 6.9% fat, 48.9% carbohydrate, and with a small proportion of prolamins; buckwheat protein has
5.8% ash. twice the lysine content of wheat or white rice. The grains are
Chenopodium ambrosioides (wormseed, Mexican tea). In the relatively high in potassium, magnesium, phosphate, iron, and
Himalayas, the seed is ground into flour and mixed with vitamins B1 and B2.
warm water or roasted or added to alcoholic beverages; care
needs to be taken with its preparation and use. Potential pseudocereals in the Polygonaceae family
Chenopodium berlandieri subsp. nuttalliae (southern huauzon- Fagopyrum cymosum (perennial buckwheat) or Fagopyrum
tle). The seed is ground to a meal and used for bread or tataricum (Tartary buckwheat). In the Himalayas, the seeds
gruel (a light form of porridge produced by boiling meal in are eaten or ground into flour.
water or milk). Calligonum polygonoides (phog, phogalli).The seeds are eaten
Chenopodium murale (nettle-leaf goosefoot, sowbane). The seed raw in India.
is popped or parched and ground to make gruel. Polygonum aviculare (knotgrass, gooseweed). The seeds are
Chenopodium pallidicaule (canihua). The seed is toasted and eaten or ground into flour for use in cookies or pancakes;
ground into a flour and eaten as a breakfast food, mixed they are sometimes mixed with wheat; care needs to be
with wheat flour in baked products or drunk as a beverage. taken with their preparation and eating.
Chenopodium sp. (California Yokuts lambs quarters). This Polygonum glabrum (sauri arak, jioti, aatlaria). The seeds are
selection was used as a semi-domesticated grain crop by parched and made into a kind of Indian sattu (a tradi-
Yokuts Indian villages, in the Central Valley, California. tional weaning food mix often eaten by adults, usually
Atriplex canescens (fourwing saltbush). The seed is ground based upon roasted Bengal gram, wheat, and sugar with
into meal and used as flour, or as a beverage. spices added and served in a number of ways, including as a
Atriplex confertifolia (shadscale). The seed is ground into meal porridge).
and used for bread or mush (a heavier form of porridge than Polygonum orientale (Princes feather).The seeds are eaten in
gruel, produced by boiling meal in water or milk). China.
Atriplex hortensis (orache). Flour is prepared from the seeds Polygonum plebeium (chimtee sag, raniphul, machichi). The
and used in soup and muffins; the seed is reportedly rich in seeds are crushed on stones, cooked, and eaten in the
vitamin A. form of a damper; the dry seed contains 17.4% protein,
Atriplex lentiformis (quail bush) and Atriplex polycarpa (all 50.4% carbohydrate, 3% fat, and 16.2% fiber.
scale).The seeds are eaten by the Native American Pima Rumex acetosa (sorrel dock). The seeds are used as a base for
group, located in Arizona. bread in Scandinavian countries; care needs to be taken
Atriplex patula (halberd-leaved saltbush). The seed is ground with their preparation and eating.
and mixed with corn and steamed as meat balls by the Zuni Rumex acetosella subsp. acetosella (sheep sorrel). The seeds are
Indians. ground into flour to make a flat bread called sygrasbrod.
Atriplex prostrata (hastate saltplant). The seed is ground into Rumex crispus (curled dock). The seeds are ground into meal
flour for use in baked products. or flour and used in pancakes; care needs to be taken with
Haloxylon sadicornicum.The seeds are mixed with other preparation and eating.
grains for bread making in India.
Suaeda corniculata and Suaeda heteroptera. The seeds are
eaten raw in Manchuria. Potential Pseudocereals in Other Dicot Families
Tecticornia verrucosa. The seeds are ground to flour by
Australian aborigines and baked. Other dicotyledonous families with potential pseudocereal
species producing starchy grains include: Bixaceae, Cactaceae,
Cannabidaceae, Caryophyllaceae, Chloranthaceae, Cistaceae,
Portulacaceae, and Trapaceae. Species of potential interest are
Polygonaceae
named in the following.
Fagopyrum esculentum (buckwheat) There are many other minor dicotyledonous families with
Buckwheat originated from south central China. plants producing starchy grains which will not be referred
This crop is harvested with its green pericarp intact; once the to again; these include: Ancistrocladaceae, Basellaceae, Buddle-
pericarp (hull) has been removed, the groat can be coarsely jaceae, Desfontainiaceae, Dioncophyllaceae, Droseraceae, Ery-
milled into grits for breakfast food or roasted and sold in whole throxylaceae, Frankeniaceae, Gelsemiaceae, Leitneriaceae,
THE LEGUMES AND PSEUDOCEREALS | Pseudocereals: Overview 277
Bixaceae
Bixa orellana (annatto). The seeds are used as coloring and Trapaceae
flavoring for meat, poultry, or fish; the seedcoat contains the
carotenoid, bixin, which is used as a tasteless coloring for
Trapa natans (water caltrop, water truffle).The seeds are eaten
raw, roasted, boiled, fried, or ground into flour for bread
cheese, butter, margarine and chocolate and for color in soap
and sweet puddings.
and skin products.
Trapa bispinosa. The seeds are cooked and eaten in India.
Cactaceae
Carnegiea gigantea (giant cactus). The seeds are ground into
flour and used to make soup, paste, and other products by Potential Pseudocereals in Monocot Families
the Native American Papago group.
Monocotyledonous families with potential pseudocereal spe-
Ferocactus wislizeni (fishhook cactus). The seeds are ground,
parched, and used for bread and gruel. cies include: Araceae, Cannaceae, Commelinaceae, Cyperaceae,
Marantaceae, Musaceae, Palmae, Pandanaceae, Pontederiaceae,
Opuntia clavata (ishikana). The seeds are roasted by Native
Americans of the Acoma and Laguna groups; however, the Typhaceae, and Zingiberaceae. Species of potential interest are
mucilage in the fruit can render the food objectionable. named in the following.
There are also many minor monocotyledonous families
Opuntia phaeacantha (bastard fig). The seeds are dried,
parched, and ground into meal for gruel and cakes. with plants that produce starchy grains which will not be
referred to again; these include: Alismataceae, Anarthriaceae,
Opuntia soehrensii (ayrampo). A violet dye is extracted from
the seeds and used as a food coloring. Aponogetonaceae, Bromeliaceae, Burmanniaceae, Butoma-
ceae, Centrolepidaceae, Costaceae, Cyanastraceae, Cyclantha-
Pachycereus pecten-aboriginum (cardon hecho hecho). In Mex-
ico, the seeds are ground and made into cakes. ceae, Eriocaulaceae, Flagellariaceae, Haemodoraceae,
Hydatellaceae, Hydrocharitaceae, Joinvilleaceae, Juncaceae,
Pachycereus pringlei (cardon). The seeds are toasted, ground,
and fashioned into balls. Juncaginaceae, Lemnaceae, Mayacaceae, Melanthiaceae, Naja-
daceae, Philydraceae, Potamogetonaceae, Rapateaceae, Rhipo-
Stenocereus stellatus (xoconochtli, joconostle). The seeds have
been eaten. gonaceae, Scheuchzeriaceae, Sparganiaceae, Triuridaceae, and
Velloziaceae.
Cannabidaceae
Cannabis sativa (hemp). The seeds are parched, fried as cakes, Araceae
used in asanomi (in Japan), in the spice mix shichimi, in ale or
Peltandra virginica (arrow arum, tuckahoe). The seed produces a
are eaten as sprouts.
bread similar to corn bread, with a cocoa flavor.
Caryophyllaceae
Lychnia segetum. The seeds have been eaten as a famine food Cannaceae
in France. Canna edulis (Queensland arrowroot). Immature seeds are
Spergularia arvensis (corn spurrey). The seeds are used for cooked in tortillas or cakes; the plant is used as a source of
bread flour in Norway and Sweden. arrowroot in Colombia.
Stellaria media (chickweed). The seeds are used as bread or as
a soup thickening in India.
Commelinaceae
Chloranthaceae
Commelina benghalensis (keng, bokna, mu-Kengeria), Comme-
Sarcandra glabra (tea scent). The seeds are roasted and used as a lina forskailii (kansura, kanshura). The seeds are ground into
sesame substitute. flour for bread.
Commelina communis (spider wort, day flower), Commelina
obliqua (kena, keni, kana, kanjura). The seeds are eaten.
Cistaceae
Cyanotis axilaris (vichaka, narido, damro, soltra). The seeds
Cistus ladanifer (labdanum). The seeds are ground and used for are ground into flour for bread or bhadku, a mixture of
cakes and breads. flour and salt.
278 THE LEGUMES AND PSEUDOCEREALS | Pseudocereals: Overview
fiber.
Contrast and compare the characteristics (grain yield and
utilization value) of the three main pseudocereals: ama-
ranth, quinoa and buckwheat.
Pandanaceae Consider a further six potential pseudocereal species (e.g.,
chia) and indicate what beneficial attributes they offer.
Pandanus tectorius (Nicobar breadfruit, screwpine). The seeds are
edible but only after careful preparation.
Pontederiaceae
Exercises for Readers to Explore the Topic Further
Pontederia cordata var. cordata (lance-leaf pickerel weed). The Find out the potential situations in which some of the less
seeds are eaten raw, boiled or parched, and ground into flour common pseudocereals might become useful, considering
for bread. growth vigor and seed usefulness (including anti-
nutritional properties, such as saponins).
Typhaceae
Amaranthus retroflexus is considered a weed in some situa-
tions, but it appears to have some potential for animal feed,
Typha angustifolia (narrow-leaf cattail). The seeds are roasted possibly even for human food. Investigate the situations in
and eaten. which its potential value might be harnessed for good.
Zingiberaceae
See also: Food Grains and the Consumer: Grains and Health;
Aframomum angustifolium (Madagascar cardamom). The
Food Grains and Well-being: Functional Foods: Dietary
dried seeds are used like pepper or added to coffee.
Fibers, Prebiotics, Probiotics, and Synbiotics; Food Grains:
Aframomum melegueta (Melegueta pepper). The seeds are used
Intolerance, Allergy and Diseases: The Gluten-Free Diet;
as flavoring for beverages, meats, ice cream, candy, and
Grain Composition and Analysis: The Composition of Food
bread.
Grains and Grain-Based Products; Non-food Products from
Amomum compactum (round cardamom). The seeds are used
Grains: Cereal Grains as Animal Feed; The Basics: Grain and
in cakes in Malaysia.
Plant Morphology of Cereals and how characters can be used to
Amomum subulatum (Indian cardamom). The seeds are used
identify varieties; Taxonomic Classification of Grain Species; The
for flavoring in char masala, in Afghanistan.
Grain Crops: An Overview; The Legumes and Pseudocereals:
Amomum xanthioides (bastard cardamom). The seeds are used
Amaranth: Overview; Buckwheat: Overview; Quinoa: Overview.
as flavoring in liqueurs in China.
THE LEGUMES AND PSEUDOCEREALS | Pseudocereals: Overview 279
Further Reading Wiersema JH and Leon B (1999) World Economic Plants. Boca Raton, Florida: CRC
Press.
Belton PS and Taylor JRN (2002) Pseudocereals and Less Common Cereals. Berlin: Williams JT (1995) Cereals and Pseudocereals. London: Chapman and Hall.
Springer.
Crawford M (2000) Edible Plants for Temperate Climates. Totnes, Devon: Agroforestry
Research Trust.
Facciola S (1998) Cornucopia II. A Source Book of Edible Plants. Vistas, CA: Kampong Relevant Websites
Publications.
Hegarty MP, Hegarty EE, and Wills RBH (2001) Food Safety of Australian Plant Biodiversity Research Collection, University of Connecticut - The families of flowering
Bushfoods. Canberra, ACT: Rural Industries Research and Development plants: descriptions. http://www.biodiversity.uconn.edu.
Corporation. http://www.pfaf.org/ Plants for a future.
Mabberley DJ (1997) The Plant-Book. A Portable Dictionary of the Vascular Plants, 2nd hort.purdue.edu http://www.hort.purdue.edu Center for new crops and plant products,
edn Cambridge: Cambridge University Press. Purdue University and Famine Foods database.
Lupin: Overview
DS Petterson, Department of Agriculture, Nedlands, WA, Australia
2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition, volume 2, pp 166174, 2004, Elsevier Ltd.
Learning Objectives
To understand the structural and compositional aspects of Grain Morphology and Composition
lupins
To understand the origin and uses of various types of Lupins have a typical dicotyledon structure. Their thick seed
lupins coat (hull or testa) composes 30% of the seed weight for L.
To appreciate the requirements for and types of lupin luteus, 25% for L. angustifolius, 15% for L. albus, and 12% for L.
processing mutabilis. This is considerably higher than for most domesti-
To understand the various antinutritional factors present in cated grain species. Within the cotyledons (kernels), energy is
some lupins mostly stored in the form of thickened cell wall material
( 25% of the cotyledons) and oil bodies ( 614%). There
is virtually no starch in any of the lupin species. This is in
Definition and Origins of Lupins marked contrast to crops such as field peas and chickpeas,
which can have 5070% of the cotyledon weight as starch
The seed or grains of domesticated Lupinus species are generi- and have low protein and oil content, and the soybean with
cally called lupins. The same term is used widely in the com- 1520% oil, some starch, and a high protein content. Their
munity in referring to wild and domesticated plants and the crude protein content ranges from 28% to 42%. Proximate
seed/grain of all Lupinus species. The genus Lupinus belongs to analyses for the whole grain of the major domesticated species,
the Fabaceae, a family of legumes. There are 12 lupin species, and the Andean lupin, are shown in Table 1. There are some
all large-seeded, native to Europe and the Mediterranean wild and partly domesticated Lupinus lines containing up to
regions. Three of these are now fully domesticated. Lupinus 45% crude protein and up to 21% oil; others have as little as
albus, the European white lupin or albus lupin has a white 20% crude protein and 3% oil.
flower. Wild types of L. angustifolius, the narrow-leafed lupin, The thick seed coat, which is mostly cellulose and hemicel-
have a blue flower: however, modern cultivars bred for low lulose, of Lupinus species means that it is important to also
alkaloid content in Australia have a white flower and are consider the composition and nutritional value of their coty-
known widely as the Australian sweet lupin (ASL). L. luteus, ledons: these data are in Table 2.
the yellow lupin (YL), has a yellow flower. Their grain is widely The major lupin proteins are a group of globulins, called
used in the animal feeds industry and to a lesser extent as a conglutins. The three major ones are conglutin a (which sed-
food (or food ingredient). There are 100 lupin species native iments at 1112S and is analogous to the legumin of peas
to the Americas. They are mostly small-seeded, and only L. and glycinin of soybeans), conglutin b (which sediments at
mutabilis, the pearl or Andean lupin, has been used as a food 7S and is analogous to the vicilin of peas and conglycinins a
source. Worldwide, several species are being domesticated for and b of soybeans), and conglutin g (a sulfur-rich protein that
possible use in agriculture. The main value of lupin crops to sediments at 2S and is analogous to the conglycinin of
farmers is their replenishment of soil nitrogen, providing a soybeans). They compose 85% of the total protein and
disease break for cereals in crop rotation systems and a mar- have similar sizes and physical properties to the storage pro-
ketable grain. Lupins tend to grow relatively better on poorer teins of other grain legume species. The remaining 15% of
soils than most other crop species. proteins are albumins, which are soluble at pH 5 and vary in
Lupins are also well known for their spectacular flowers. size from 6000 to 117 000 Da. There is evidence for the 2S
The Russell lupin, a hybrid derived from L. polyphyllus, is widely globulin actually being an albumin: depending on the species,
used as an ornamental flower, particularly in Europe. Parts of this fraction consists of four (L. luteus), five (L. albus), or six
the southern highlands in New Zealand are a blaze of color (L. angustifolius) proteins. For more details on lupin and other
from the alpine lupin, a variant of the Russell lupin, in legume proteins, see CEREALS | Protein Chemistry.
Table 1 Chemical composition of the major lupin species (g kg1 as Table 3 Essential amino acid profile for major lupin species
received)a (% in grain)a
ADF, acid detergent fiber; NDF, neutral detergent fiber; TDF, total dietary fiber.
a water, about eightfold by weight for lupins, and maintain
Average of worldwide data.
normal gut motility. The profiles for various species are
shown in Table 4. The oil content of lupins varies from
Typical amino acid profiles for the domesticated lupin spe- 15% in the pearl lupin down to 4% in the YL. There are
cies, compared with the FAO standards for infants and others, differences in individual fatty acid profiles; they are typically
are shown in Table 3. The deficiencies in lysine and methio- high in oleic and/or linoleic acids and contain a total of 75%
nine are common to most grain legumes. unsaturated fatty acids (cf. 85% in olive oil). The oil is heat-
The seed hulls and cotyledons contain different types of stable and has a high antioxidant capacity.
carbohydrate. Lupin hulls are predominantly composed of Variation in the content of major essential minerals in
structural polysaccharides: cellulose, hemicelluloses, and pec- lupins, up to 30% of the mean, is typical of legume species,
tins. The main carbohydrate reserves of the cotyledons are the probably reflecting both genotype and environmental factors
nonstructural polysaccharides of the cell walls, with the main (Table 5). The essential trace mineral content of lupin species
components being galactose, arabinose, and uronic acid. These not only is influenced by genotype but also tends to reflect the
complex compounds are referred to as nonstarch polysaccha- soil types on which they are grown. For example, the selenium
rides (NSPs). Their structure, based on a rhamnogalacturonans content of lupin seeds grown in Western Australia varies
backbone, is not yet fully defined. The water-soluble portion, according to species and rainfall zone. Where the species
5% of the seed, is considered to have an antinutritional effect were grown together, the selenium content of L. albus > L.
due to its viscous nature and effect on intestinal transit time. luteus > L. angustifolius. In all cases, the higher the rainfall, the
The insoluble NSP, 30% of the seed, has a minimal effect on lower the selenium content. Similar variations apply to the
nutrient utilization by monogastric species. An important attri- content of copper, cobalt, and zinc in lupins. The accumula-
bute of insoluble NSP is their ability to hold large quantities of tion of manganese by L. albus is well documented with most of
282 THE LEGUMES AND PSEUDOCEREALS | Lupin: Overview
Table 5 Mineral content of major lupin species (range of data sets) Table 6 Antinutritional factors in the major lupin speciesa
bacterial fermentation in the colon to produce carbon dioxide, gramine, an indole alkaloid, occur in some lines of L. luteus,
methane, and hydrogen. This causes abdominal discomfort but the significance of these levels is not known.
and cramps and leads to flatulence, which seems to be a The only mycotoxin associated with lupins to date is the
major reason for the low interest in consuming grain legumes family of phomopsins, a group of linear hexapeptides with an
in many societies. The oligosaccharides are a rich source of antimicrotubule effect, which is most pronounced in the liver
nutrients for bifidobacteria in the colon. This can have a ben- and kidneys. They are capable of disrupting mitosis and of
eficial effect, as the bifidobacteria counter the activity of putre- reducing the activity of motile cells and organisms, rendering
fying bacteria and reduce their production of harmful, and their host prone to secondary infections. There is no relation-
possibly carcinogenic, fermentation products. One further ship between lupinosis and lupin alkaloid poisoning that can
source of confusion about the significance of oligosaccharides occur when sheep graze bitter lupins and their stubble or graze
is the inclusion by some authors of sucrose in their total for the perennial lupins of North America. Grains that contain
oligosaccharides. This is nutritionally confusing as sucrose can phomopsins are smaller and less dense than normal-colored,
be absorbed from the stomach and upper digestive tract of nontoxin-containing grain and can be easily removed from the
humans and nonruminant animals. rest of the harvested grain by grading, using screens, gravity
Isoflavones have generally been regarded as ANFs because tables, and aspirators. A rapid enzyme-linked immunosorbent
of their negative effects on fertility in ruminants grazing pas- assay for the phomopsins is readily available to the food
ture medics and subterranean clovers. However, they are now industry. The most recent cultivars of L. angustifolius are resis-
recognized as preventative agents against some forms of cancer tant to invasion by the responsible fungus (Diaporthe toxica), so
and osteoporosis in women. Isoflavones are present in green- that the risk of phomopsin ingestion is now extremely low and
leaf material and in the hypocotyls of germinating seeds likely to be even less in the future.
(sprouts). There is conflicting evidence as to their presence in
mature grain.
The lupin alkaloids are usually bicyclic, tricyclic, or tetra- Grain Processing
cyclic derivatives of quinolizidine (Figure 1). The individual
Dehulling
alkaloids and their concentrations vary widely between,
and sometimes within, species (Table 7). Grain of modern The simplest processing of lupin grain is repetitive abrasion of
domesticated lupin cultivars typically contains less than the testa in a ripple flow mill, tangentially abrasive dehulling
200 mg kg1 alkaloids. In contrast, grain from bitter wild device, or similar device. (The thickness and hardness of the
types that still exist in many countries may contain from testa make it harder to dehull lupins than most grains.) Alter-
5000 to 40 000 mg kg1 alkaloids. There have been human natively, the grain may be conditioned by the addition of
deaths associated with the consumption of grain from bitter water, to swell the grain, followed by a mild heat treatment
wild types of lupin. One would need to eat 10 kg of the to help break the bonding between the testa and cotyledons. It
modern varieties of lupin grain in a very short time to absorb is then possible to dehull using the same roller mills as used for
sufficient alkaloids to be at risk. The quinolizidine alkaloids milling wheat and other cereals. The major components are
have a short half-life in humans and are excreted largely separated by air aspiration and grading across gravity tables.
unchanged in the urine of >90% of humans. Traces of The germ fraction usually separates with the hulls and is
mostly lost.
The main reason for dehulling is that the cotyledons are
much more nutritious than whole grain for nonruminant ani-
mals such as pigs, poultry, and fish. It is then possible to mill
N the cotyledons to make flour or to separate it into protein-rich
and fiber-rich fractions for the feed, human food, and cosmetic
Quinolizidine
industries.
Figure 1 Structure of quinolizidine.
Milling
Table 7 Quinolizidine alkaloids in the major lupin species (% in Hammer milling of whole grain to make a coarse meal is
alkaloid fraction)a common in the animal feed industry, simply to make the
constituents more available to ruminant animals by increasing
L. albus L. angustifolius L. luteus L. mutabilis
the surface area of material for digestion, and in some cases, it
Albine 15 may be uneconomic to dehull for nonruminant feeds. Ham-
Lupanine 70 70 70 mer cutting and roller milling of the splits are used to make a
Multiflorine 3 meal or fine flour for nonruminant feeds. The particle size
13-Hydroxylupanine 8 12 2 range can be from 10 to 800 mm, which is far greater than for
4-Hydroxylupanine 1 7 cereal, pea, and soy flours.
Angustifoline 10 <1
Lupinine 60
Sparteine 30 20 Fractionation
a
Data are for modern cultivars or developing lines of these species. Up to 10% gramine, Several reports in the scientific literature and patents describe the
an indole alkaloid, is present in L. luteus. separation of protein, fiber, and oil fractions from cotyledons.
284 THE LEGUMES AND PSEUDOCEREALS | Lupin: Overview
Some processes cater for the recovery of alkaloids, for use as Fermented Foods
biological agents against insect pests and some microorganisms,
Tempeh is a traditional Indonesian food, made by two suc-
and low-molecular-weight carbohydrates for use in the chemical
cessive fermentations using soybeans as substrate. A bacterial
industry. These mostly involve wet grinding of the cotyledons
fermentation during the soaking of cooked dehulled soybeans
and separation of fractions according to solubility at different
is followed by a solid-state fermentation of the bean mass by
pH or in different solvents. There may be an initial dry milling
the mold Rhizopus oligosporus. Tempeh has been successfully
and separation of particles by air classification.
made from L. albus, L. angustifolius, and blends of both with
soybeans. Indonesian consumers like the taste of lupin-based
Extrusion tempeh but frequently comment that the texture is too firm.
The production of second-generation products, such as burgers
Extrusion can improve the quality of lupin-based formulated
and patties, from lupin-based tempeh seems to overcome this
feeds. There is no expansion of the pellets as with starch-
objection.
enriched feeds, but there are improvements in feed digestibil-
Miso is a fermented paste made from soybeans, usually
ity, which makes the process cost-effective.
mixed with rice. A traditional process involves preparing a rice
koji by fermenting cooked rice with a culture of Aspergillus
oryzae (tane-koji) and then adding cooked soybeans and salt
Food Uses
to the koji. Lupins have been used experimentally to make
Traditional Uses miso of comparable quality with soy miso. Natto, another
Japanese fermented food, can also be made from lupin grain.
Lupins have been used as a food for as much as 6000 years in
Shoyu is the traditional soy sauce of Japan. Some Japanese
the Andean highlands and over 3000 years around the Medi-
people now use lupin shoyu as an alternative. Lupins can be
terranean. The pearl lupin of the Andean highlands of South
used to make sauces similar in flavor and texture to the tradi-
America, known locally as tarwi or chocho, was extensively
tional soy sauces of China, Korea, and Indonesia.
cultivated, and consumers would soak the seed in running
Several thousand tonnes of ASLs have been used commer-
water to remove most of the bitter alkaloids before toasting
cially in Indonesia for tempeh production, and small commer-
or boiling and drying to make kirku. Andean people have
cial batches of lupin-based miso have been sold in Japan.
long recognized the benefits of consuming lupins, associating
their use with religious festivals. In South America, the tradi-
tional use of tarwi as staple food continues in many Andean
communities. Around the Mediterranean, lupini beans, large-
Flour Additive
seeded bitter grains of L. albus, have been extensively used as a
snack food and as a bean substitute in times of drought. They Lupin kernel flour can be mixed in with wheat or wholemeal
are still commonly used as a snack bar. The beans (grains) are flours to make bread more nutritious, by giving a better bal-
substantially debittered before being pickled and bottled or ance of essential amino acids. Legume grains contain relatively
canned for use as a snack food. In the Middle East, the grain more lysine and less methionine than cereal grains; so, adding
of L. albus is used to make snack foods, most commonly at the 10% lupin flour to wheat flours will give a product with
time of Ramadan (several thousand tonnes are imported from similar properties to the full wheat product but with an
Australia each year for this). They are usually boiled whole and improvement in amino acid score from under 40% to over
spiced, with the testa being removed before eating. Another use 70% relative to egg albumin. Loaf volume is compromised
is to make a paste similar to the traditional falafel from faba with increasing lupin content but fortifying with gluten or
beans. using harder wheat in the blend can overcome this. In practice,
the limit of inclusion is 10%. There is an increase in water-
holding capacity. The texture, flavor, and golden color of the
Bean/Sprout
lupinwheat flour are appealing to many consumers. In
Immature seeds do not yet contain any alkaloids and have a Australia, some bread manufacturers use lupin hull flour to
similar taste and nutritional value to immature (green) peas. provide bulk in high-fiber breads. Ultrafine kernel flour has
They can be used as an alternative to soybeans to make an attractive yellow color, good dispersion in aqueous systems,
edamame. Lupins can be germinated to make a sprout suit- and good emulsifying properties. Albus lupin flour can be
able for vegetable or salad use. Germination lowers not only added into wheat flour to make the traditional Chilean breads
the content of alkaloids, phytate, and oligosaccharides on a dry hallulla and marraqueta. Up to 10% micronized flour from
weight basis but also the overall content of the protein fraction. L. albus cotyledons could be included in breads, biscuits, and
The lupin sprout compares favorably with soy and mung cakes, enhancing protein quality, color, and taste. The added
sprouts for taste and texture, although it has a slight beany lupin flour retards staling, probably by increased water reten-
and bitter aftertaste. Unlike most species, there is no vitamin C tion and better emulsification properties. Studies have shown a
production on germination. There is, however, considerable high acceptability of pasta enriched with L. angustifolius kernel
production of the isoflavones, genistein and daidzein. flour, and up to 15% substitution of L. albus flour for semolina
Whole seeds may be deep-fried or made into spreads sim- or durum flours enhanced the protein quality and appearance
ilar in nature to hummus (traditionally made from chickpeas) of spaghetti noodles without affecting sensory qualities. Flour
and falafel (traditionally made from faba beans) and used with from L. albus has been blended with noodle wheat flour to
salads or breads. make unsalted white noodles with improved protein and
THE LEGUMES AND PSEUDOCEREALS | Lupin: Overview 285
dietary fiber contents and color and texture. Up to 50% lupin Feed Uses
flour can be incorporated into a range of cakes and biscuits.
There are several suppliers of lupin flour and other products Ruminants
in Europe, but there are no available data on sales. Almost all of the testa can be fermented in the rumen, so there
is no need to dehull lupins for ruminant feeds. The whole grain
is highly fermentable with a minimal risk of acidosis. It is best
Lupin Kernel Fiber to crack the grain open as a minimum action, to increase access
for the rumen flora, but any grinding should not be too fine
The purified cell wall fraction from lupin kernels is virtually
because there can be a loss of material to the hindgut where it is
colorless, odorless, and tasteless and can hold up to eight times
used less efficiently. A significant amount of protein degrada-
its own weight of water. The fiber has cholesterol-lowering
tion occurs in the rumen, but some protection from this is
properties and acts as a fecal bulking agent, thus reducing
possible by treating the grain with formaldehyde before feed-
stool transit time and benefiting bowel health. It also enhances
ing. Other methods include flame roasting the grain, steaming
satiety, probably due to its high water-holding capacity, and
and rolling the grain into flakes, and coarsely grinding the
has potential as an antiobesity agent. The soluble fiber fraction
grain and then extruding the meal.
can function as a fat replacer. The fiber can be used to supple-
Lupins are a valuable component of diets of feedlot cattle
ment breads, pastas, biscuits, mousses, and jellies.
(up to 40% of the diet), and milking cows can readily consume
large amounts of (cracked) grain while in the milking sheds.
Rangeland farmers often supplement their grazing sheep with
Protein Fractions 50250 g grain per head per day in dry periods when pastures
Lupin kernel proteins have some valuable functional proper- are low. There is ample evidence for an enhancement of the
ties, combining good emulsifying, foaming, and water- and reproductive performance of both rams and ewes when their
oil-holding properties but with less thermal stability than soy diets are supplemented with lupin grain prior to mating. No
proteins. Nevertheless, lupin protein concentrates and isolates specific factor has been attributed to this.
are used as meat extenders and refined fractions used to Several hundred thousand tonnes of grain are retained on-
enhance the foaming properties of dairy desserts and other farm in Australia each year for grazing supplements for sheep.
foods. The greatest consumption of traded lupin grain is by cattle,
either in feedlots or under intensive housing, when fed with
formulated diets containing lupin grain.
Pigs rank second to cattle in the overall consumption of Exercises for Revision
lupin grain, almost entirely as a component of formulated
feeds. How can lupins be used in fermented foods?
What antinutrients can be present in lupins?
Characterize the major lupin proteins.
Poultry
The apparent metabolizable energy content of both ASL and YL
grains is 8.6 MJ kg1. It is not practical to use more than Exercises for Readers to Explore the Topic Further
100 g kg1 feed because of the high water-holding capacity of
the undigested carbohydrates, which can cause wet droppings. Increase the applications of lupin flours in foods
For layer birds, it is possible to include up to 300 g kg1 in the Increase the industrial-scale production of traditional
diets, as the wet droppings are less of a problem. Albus lupins lupin-containing foods
can be readily included in poultry diets. There is some benefit
from adding carbohydrase enzymes to poultry diets. The oli-
gosaccharides in lupin meals seem to have a beneficial effect on See also: Agronomy of Grain Growing: Lupin: Agronomy;
the osmotic stability of the gut enhancing nutrient absorption. Breeding of Grains: Lupin: Breeding; Food Grains and the
Poultry ranks third as a consumer of lupins in formulated Consumer: Grains and Health; Non-food Products from Grains:
feeds. Cereal Grains as Animal Feed; Processing of Grains: Extrusion
Technologies; Proteins: The Protein Chemistry of Dicotyledonous
Grains; The Basics: Grain: Morphology of Internal Structure; The
Finfish and crustaceans Legumes and Pseudocereals: Grain Legumes and Their Dietary
Lupins can be used in both high nutrient density (HND; high
Impact: Overview.
protein and oil content) and low nutrient density (low protein
and oil content) diets for finfish. When included in HND diets
Further Reading
fed to rainbow trout (Oncorhynchus mykiss), the DE value of
lupin kernel meals ranges from 12.8 MJ kg1 for L. angustifolius Gladstones JS, Atkins C, and Hamblin J (eds.) (1998) Lupin as Crop Plants: Biology,
to 14.8 MJ kg1 for L. albus. Protein digestibility of all lupin Production and Utilization. Oxon, UK: CAB International. This book is the definitive
varieties when fed to fish is high, typically greater than 95%. work on all aspects of the Genus Lupinus, and was drawn on for this article.
Nahrung 45(6) (2002) A special edition covering the processing of lupins and other
The low essential amino acid content is less of a problem for grain legumes.
finfish fed with HND diets than for pigs and poultry because of Petterson DS (2000) The use of lupins in feeding systems: Review. Asian-Australasian
the high level of protein required in the diets restricting the Journal of Animal Sciences 13(6): 861882.
practical inclusion levels of the meal and also that so much of Van Barneveld RJ (1999) Understanding the nutritional chemistry of lupin (Lupinus sp.)
seed to improve livestock production efficiency. Nutrition Research Reviews
the dietary protein is metabolized as an energy source. Good
12: 130.
acceptance and palatability of HND diets containing L. angu- Wink M, Meissner C, and Witte L (1995) Patterns of quinolizidine alkaloids in 56
stifolius, L. albus, and L. luteus have been reported, with accept- species of the genus Lupinus. Phytochemistry 38(1): 139153.
able inclusion levels up to 500 g kg1.
Lupin meals are used commercially in diets for freshwater,
estuarine, and marine finfish and crustaceans. This is a small
Relevant Websites
but emerging market, as the relative value of the splits seems http://www.agric.wa.gov.au Department of Agriculture, Western Australia.
high for finfish. http://www.rala.is The International Lupin Association.
Amaranth: Overview
H Corke and YZ Cai, Hubei University of Technology, Wuhan, China; University of Hong Kong, Hong Kong, China
HX Wu, Wuhan Chortle Biotechnology Company Limited, Wuhan, China
2016 Elsevier Ltd. All rights reserved.
Cotyledons
Procambium
Starchy
Cotyledons perisperm
Seed Vascular
coat bundles
Endosperm
Shoot
apex
Perisperm
Procambium
Root
Procambium Radicle
(Root)
(a)
Cross-section Longitudinal section
(b)
Figure 2 (a) Illustration of A. cruentus seed in cross and longitudinal sections; (b) scanning electron micrograph of A. cruentus seed. Adapted
from Irving DW, Betschart AA, and Saunders RM (1981) Morphological studies on Amaranthus cruentus. Journal of Food Science 46(4): 1170; Becker R
(1994) Amaranth oil: Composition, processing, and nutritional qualities. In: Paredes-Lopez O (ed.) Amaranth: Biology, Chemistry, and Technology,
p. 135. Boca Raton, FL: CRC Press.
There has also been much information on other compounds in and amylopectin. Table 4 shows that amylose makes up from
amaranth (tannin, phytic acid, oxalate, saponins, nitrates, tryp- 0% (waxy) to 34.3% of total starch in amaranth grain with the
sin inhibitor, etc.) with antiphysiological activity in animal balance being the amylopectin, showing that there is consid-
performance. Tannins are of importance from the nutritional erable variation in amylose/amylopectin ratio among and
point of view, since they may influence protein digestibility and within the Amaranthus species. Light-colored grains normally
reduce mineral bioavailability. Tannins in different amaranth contain more starch (mean 69%) than dark-colored grains.
species range between 0.1% and 0.25% and seem to be low Starches isolated from A. hypochondriacus genotypes contain
enough not to have nutritional impact. The phytate content the lowest amount of amylose (7.8%), whereas A. retroflexus
(0.340.61%) in amaranth is higher than that found in rice genotypes have the highest amylose (34.3%). The average
(0.10.14%), but lower than that detected in maize and wheat. amylose content of 124 genotypes is 19.2%.
Amaranth starch granules come in polygonal, lenticular,
circular, and elliptical shapes. Amaranth starch granules are
Amaranth Starch extremely small, ranging from 0.8 to 2.5 mm in diameter. Com-
paratively, diameters of the granules in commercial starch range
Starch is the main component of amaranth grain, composing from 3 to 8 mm for rice starch to 15100 mm for potato starch.
4869% of its total dry weight. Starch generally consists of Amaranth starch granule size and its size distribution are char-
straight-chain and branched-chain molecules, that is, amylose acteristics that influence the functional properties of starch.
290 THE LEGUMES AND PSEUDOCEREALS | Amaranth: Overview
Table 1 Comparison of proximate composition between amaranth Since the 1970s, some interesting findings on physico-
grains and some cerealsa chemical and functional properties of amaranth starch have
been reported, for example, stable paste properties and a wide
Composition Amaranthb Wheat Corn Rice
range of viscosity, resistance to shear thinning, and gelatiniza-
Carbohydrate 59.2 66.9 67.7 75.4 tion temperatures. The focus of such research was restricted to
Crude protein 16.6c 14.0d 10.3e 8.5e very few Amaranthus species or genotypes and sometimes gave
Fat 7.2 2.1 4.5 2.1 contradictory results due to the genetic variation of the prop-
Crude fiber 4.1 2.6 2.3 0.9 erties of amaranth starch. A. hypochondriacus starch had a
Ash 3.3 1.9 1.4 1.4 higher amylograph viscosity, a higher gelatinization tempera-
Moisture 9.6 12.5 13.8 11.7 ture, a lower swelling power, and a higher solubility than
a wheat starch. A. paniculatus (synonym: A. cruentus) starch had
Percentage on dry weight basis.
b
Mean values of four Amaranthus species (A. cruentus, A. caudatus, A.
moderate swelling power, higher solubility, and lower pasting
hypochondriacus, and A. hybridus). viscosity than foxtail millet starch. Compared to maize starch,
c
N 5.85. A. paniculatus starch had a higher paste viscosity, lower paste
d
N 5.7. clarity, and higher freeze/thaw stability.
e
N 6.25. A large number of Amaranthus germplasm (26 species)
Data from Saunders RM and Becker R (1984) Amaranthus: A potential food and feed accessions were introduced into China for the evaluation of
resource. In: Pomeranz Y (ed.) Advances in Cereal Science and Technology, vol. 6, pp. amaranth starch properties. Compared to the reference maize,
357396. St. Paul, MN: American Association of Cereal Chemists; Singhal RS and rice, potato, and wheat starches, Amaranthus starch tended to
Kulkarni PR (1988) Composition of the seeds of some Amaranthus species. Journal of
have more stable paste, that is, lower pasting viscosities (lower
the Science of Food and Agriculture 42(4): 325331; Yue SX, Sun HL, and Tang DF
shear thinning and lower retrogradation), higher gelatiniza-
(1993) The Research and Development of Grain and Amaranth in China (in Chinese).
Beijing: Chinese Agricultural Science and Technology Press; Segura-Nieto M, Barba de
tion temperature, and higher energy of enthalpy for gelatiniza-
la Rosa AP, and Paredes-Lopez O (1994) Biochemistry of amaranth proteins. In: tion. Also, Amaranthus starch paste was more resistant to cold
Paredes-Lopez O (ed.) Amaranth: Biology, Chemistry, and Technology, pp. 75106. storage and had softer gel texture and lower changes of hard-
Boca Raton, FL: CRC Press. ness, cohesiveness, modulus, and adhesiveness. Figure 3
Amaranthus speciesb
Component 1 2 3 4 5 6 Spinach
displays the Rapid Visco Analyzer (RVA) pasting profiles of plastics. The unique properties of A. caudatus starch, namely, its
Amaranthus starch compared with maize starch. A wide range pasting properties and high susceptibility to amylase, might
of variation was found in the several tested properties of starch represent potential uses as food, feed, and fuel. A. hypochon-
among the Amaranthus species and among genotypes within driacus starch had relatively low gelatinization temperature and
the same species, suggesting that the wide genetic diversity food freeze/thaw stability, indicating its use in gravies, sauces,
necessitates specific choices for specific uses. Interestingly, it and soups. A. paniculatus starch could be used in salad dress-
was observed that there were significant effects of environment ings and custard preparations. There were also reports on
on the tested properties of Amaranthus starch, indicating that chemical modification (e.g., succinylation and hydroxypropy-
starch of the same genotype might have different starch prop- lation) of amaranth starches and their properties.
erties if grown in different locations. Additionally, obvious Many studies demonstrated that starches isolated from
effects on pasting and rheological properties from genotypic, many Amaranthus species could be used as good thickeners
pH, and salt factors were found. and stabilizers in salad dressing, canned food, sauce, frozen
Recently, amaranth starch has attracted increasing attention food, etc., because of their stable paste, low retrogradation,
because of its unusual properties. Possible product uses of freeze/thaw resistance, and heating and cooling resistance.
amaranth starch, capitalizing on its small granule size, could Also, there has been a report that describes amaranth starch
include food thickeners, dusting powders for cosmetic as well as behaving very poorly in breads and cakes from the stand-
as food uses, laundry starch, and even possible biodegradable point of color and appearance, texture, and volume, probably
because of its low amylose content. During starch application,
Table 4 Starch and amylose content of amaranth grains (%)a it is worth paying attention to a fairly wide variation of ama-
ranth starch properties among different species and genotypes.
Species Starch Amylose
K112
Viscosity (RVU)
scores higher than other seeds (e.g., wheat, barley, soybean, proteins showed similar effect, but to a much lesser extent.
and maize) in the FAO/WHO Nutritionists Protein Value Furthermore, amaranth protein concentrates were examined
Chart: a score of 100 is the ideal, whereas amaranth protein for direct application in certain food systems (wheat dough
received the highest score of 75, compared to cow milk (score and noodle). It was found that the mixing properties of wheat
72), soybeans (68), and peanuts (52), indicating that its bal- flour were improved by the addition of the protein concen-
ance was closer to the optimum required in the human diet trates. Addition of protein concentrates to a noodle formula-
(FAO/WHO). tion also showed some positive influence on final product
Seed proteins are generally classified into four types based quality. A few amaranth protein concentrates were also evalu-
on solubility: albumin, globulin, prolamin, and glutelin. Most ated as fillers for an emulsion-type meat product. Only one of
studies on amaranth proteins suggested that albumin was the the amaranth protein concentrates (genotype K112) gave some
major fraction (4965%), followed by glutelin (2242%), favorable results.
globulin (1418%), and prolamin (1.03.2%). However,
there was variation in amaranth protein fractions because of
protein extraction and fractionation procedures and due to Amaranth Oil
genetic variation. For instance, it was reported that the frac-
tionated proteins from five different Amaranthus species aver- Oil is another chemical component of much interest in ama-
aged albumin (65%), globulin (17%), prolamin (11%), and ranth grain, because it contains relatively higher levels of oil
glutelin (7%). Also, it was reported that the isolated proteins (58%) than cereal grains; moreover, there is a considerable
from A. hypochondriacus consisted of 4649% albumin plus amount of squalene in amaranth oil (5%) compared with that
globulin, 3% prolamin, and 3033% glutelin, and in another in oils of cereals and other oil crops (0.010.4%) (Table 7).
study of same species, similar percentages of albumin, globu- Palmitic, oleic, and linoleic acids are the major fatty acids in
lin, prolamin, and glutelin were 51.0, 15.9, 2.0, and 31.1, amaranth oil, with the saturated/unsaturated fatty acid (S/U)
respectively. ratios ranging from 0.12 to 0.50. Published data showed that
In the 1980s and 1990s, many researchers isolated major the fatty acid profiles in amaranth oil from different species
fractions (albumin, globulin, and glutelin) of proteins from were as follows: palmitic (1225%), oleic (1935%), linoleic
main Amaranthus species and examined their biochemical and (2562%), stearic (28.6%), and linolenic (0.32.2%) acids.
physicochemical properties and molecular weights. Based on The squalene levels range from 5% to 8% in A. cruentus oil.
their measurements of its emulsion activity, foaming stability, Lower amounts of squalene (25%) are tested in some
and surface hydroscopicity, along with its relatively high heat
stability, it was concluded that amaranth proteins could be Table 7 Oil and squalene content of amaranth grains and other seeds
useful as effective emulsifying and foaming agents. On the
other hand, there were reports that A. cruentus and A. hypochon- Species Oil (%) Squalene (%)
driacus protein fractions had poor emulsifying and foaming
A. cruentus (1) 5.577.72 4.525.44
properties, limited water absorption, and good gel-foaming A. cruentus (2)a 5.08.0
properties. A. hybridus 6.40 5.23
It was found that protein concentrates made from ama- A. hypochondriacus 5.357.05 3.625.01
ranth grains of five genotypes exhibited better solubility, emul- A. tricolor 5.08 6.14
sification, and foaming properties than the commercially Maize 4.0 0.03
available soy proteins (Table 6). The effect of proteins on Cottonseed 7.0 0.01
starch gelatinization and retrogradation was investigated Rice 1.03.0 0.3
using the hydrolyzed and native protein concentrates of ama- Olive 36.0 0.4
Peanut 47.0 0.03
ranth. The results suggested that native proteins tend to be
attracted toward the starch granule and contribute to the regu- a
Data from Lyon and Becker (1987), other data in the table from Becker R (1994) and He
lation of swelling and amylase leaching during gelatinization, HP, Cai YZ, Sun M, and Corke H (2002). Extraction and purification of squalene
therefore resulting in a weaker starch gel. However, hydrolyzed from Amaranthus grain. Journal of Agricultural and Food Chemistry 50: 368372.
Table 6 Functional properties of amaranth protein concentrations, compared to soy protein isolates
Species Genotypes Emulsifying activity (%) Solubility (%) Foam expansion (ml) Foam stability (ml)
Amaranthus species of West Africa. Other lipids in amaranth, because of safety and health concerns. Red-colored vegetative
such as triglycerides, sterols, methyl sterols, terpenic and ali- tissues of Amaranthus plants contain a high concentration of
phatic alcohols, tocopherols, and hydrocarbons, have also betalain pigments and are often produced in high biomass
been identified. Recent results indicated that the major fatty (Table 8). They have attracted considerable interest as potential
acids in amaranth oil from 11 Amaranthus genotypes included substitutes for the well-known betalains from beetroot (Beta
palmitic acid (19.123.4%), oleic acid (18.738.9%), and vulgaris), which are extensively used in the food industry world-
linoleic acid (36.755.9%), with S/U ratios of 0.260.32. The wide. A comprehensive systematic study on amaranth pigments
squalene content in amaranth oils from 11 genotypes of four has covered field evaluation and selection of betalain-
species ranges from 3.6% to 6.1% (Table 7). These reports producing genotypes/species; extraction, identification, and
reveal the significant variation in oil composition content quantification of betalain components; determination and
among various species and genotypes. evaluation of properties and antioxidant activity; and pigment
Squalene is a biosynthetic precursor to all steroids. It is an production and utilization in certain food systems.
important functional ingredient in skin cosmetics and also as an Betalains are divided into two major structural groups, that
excellent computer-disk lubricant. It has been reported that the is, red-violet betacyanins and yellow betaxanthins. Amaranth
decreased risk for some cancers might be associated with squalene pigments belong to betacyanins. They could be easily extracted
consumption, and the use of squalene alone is effective in decreas- with water or methanol. Sixteen kinds of betacyanins were
ing serum cholesterol levels. The traditional source of squalene is isolated and characterized from plants in the Amaranthaceae.
from shark and whale liver oil. Estimates in 2015 place the world Figure 4 shows HPLC profile of amaranthine-type betacyanins
market at 2600 tonne year1, with a growth in demand of per- and their molecular structures, which are predominant pig-
haps 10% per year. It is very expensive ($43 000 per tonne). ments in all Amaranthus species. Quantitative analysis data
Amaranth oil will attract more attention as an alternative plant showed that the screened amaranth species/varieties yield
source of squalene because of its abundant squalene content, higher betacyanin pigments than feasible in beetroot produc-
although efforts to further increase the level are necessary. tion. Colorant properties and stability of the betacyanins are
Squalene can be easily extracted from amaranth grain by the key factors of pigment quality that directly influence their
simple vacuum distillation. Recently, work to screen genotypes application in food systems. Assessment of color characteristics
containing a higher content of squalene in a wide range and stability of betacyanins from Amaranthus species showed
of Amaranthus species and to develop efficient large-scale that Amaranthus betacyanins exhibited bright red-violet color
separation technique has been conducted; a method was estab- and favorable stability under selected conditions (pH 57 and
lished to separate and purify squalene from amaranth oils. temperature <14 C), as compared to red radish anthocyanins.
After saponification, squalene content increased from 4.2% Dried Amaranthus pigments had good storage stability (t1/
to 43.3% in the unsaponifiables by the removal of the saponifi- 2 23.3 months) at room temperature. The presented data
ables. The unsaponifiables were fractionated by a chromato- support the potential development and utilization of Amar-
graphic method to obtain highly purified squalene. The final anthus betacyanins as new natural colorants for use in the food
purity of squalene reached 9498%. industry, particularly for low-temperature uses.
However, amaranth oil occurs in very low levels and at Aqueous extracts of natural pigments are commonly
higher cost than oils of peanut, olive, rapeseed, etc. Unless a unstable and sensitive to storage environmental factors. Spray
specialty starch or protein could also be produced from the drying can significantly improve storage stability. Amaranthus
defatted meal, a substantial decrease in raw product cost will betacyanin extracts were spray-dried using a range of 1025 DE
probably be required before amaranth oil goes commercial. maltodextrins as carrier and coating agents at different
inlet/outlet air temperatures and various feed solid contents.
The best dried pigment powder made was superior to commer-
Amaranth Pigments cial red beet powder in terms of physical properties (Table 9).
The results demonstrated the feasibility of the production
There has been an increasing trend toward replacement of of spray-dried Amaranthus betacyanin extracts as a food-grade
synthetic colorants with natural pigments since the 1980s colorant.
Table 8 Pigment content and fresh weight of 12 genotypes from seven Amaranthus species
Species Genotypes Fresh weight of colored parts (kg ha1)a Total betalain content (mg per 100 g FW)
mAU
1
1: amaranthine HOCH2
2500 HO O
1: isoamaranthine (C15 6 1 O H
epimer) HO 5 2
HOOC 7 + COO
2000 O O HO N
HO
2
HO 1
1500 OH
1 H 15
1000 N COOH
HOOC
H
500
Table 9 Comparison of amaranth pigment powder and commercial beet powder (No. 3600 or E162)a
Color parameters
Spray-dried Betacyanin content Hygroscopic moisture Pigment
powder (%)b (g/100 g) L* a* b* H0 retention (%)c
Amaranth powder 0.74 48.5 49.35 32.60 2.47 355.7 87.9 1.8
(4.3)
Beet powder 0.31 58.8 51.54 19.64 0.71 2.0 84.3 1.4
a
Amaranth pigment powder made by spray drying at 180C inlet temperature using 19.8% solids in feed mixture (15 DE maltodextrin).
b
Calculated as % amaranthine and % betanin.
c
Determined at 25 C and 32% RH after 16 week storage.
Data from Cai YZ and Corke H (2000) Production and properties of spray-dried Amaranthus betacyanin pigments. Journal of Food Science 65: 12481252.
Application of Amaranthus betacyanins in jelly, ice cream, Himalayas, laddoos in India, and sattoo in Nepal. These
and a model beverage was evaluated by comparison to com- amaranth-based foods are still consumed today in various
mercial colorants (red radish anthocyanins and synthetic areas of the world. Modern food applications of amaranth
colorant FD&C Red#3). The results revealed the feasibility grains include breakfast foods, infant/weaning food formula-
and potential of Amaranthus betacyanins for use in the model tions, breads, pastas, flakes, drinks, and beverages. Currently
food systems at low temperature. Amaranthus betacyanins in available consumer food products include popped foods, bak-
the foods examined exhibited better color characteristics than ing mixes, malted flours, breakfast cereals, snacks, baked
red radish anthocyanins at the same levels, but were not as goods, and whole amaranth products. In China, a wide range
bright as FD&C Red#3. The betacyanins had color stability of amaranth food products has been developed for retail sale.
comparable to red radish and FD&C Red#3 at lower tempera- These include instant flour for beverages or porridge, packaged
ture (<14 C) and also similar color stability at 25 C, but not dried noodles, cakes, biscuits, and soy sauce made from
as stable as the two commercial colorants at higher tempera- brown/black amaranth and other grains (wheat and soybean).
ture (37 C). Additionally, all betalains isolated from plants in China has a wide diversity of traditional food products made
the Amaranthaceae exhibited strong antioxidant activity. To from specialty grains, such as buckwheat, millets, sorghum,
date, there has been evidence suggesting that certain natural and legumes. By adapting products made from these, addi-
colorants as nutritional antioxidants in the diet may reduce the tional amaranth-based foods have been developed, such as
risk of some diseases. This gives the possibility of these beta- composite flour instant noodles, distilled spirits, and vinegar.
lains being used both as natural colorants and as potential Based on the properties of diverse fractions of amaranth grain,
natural antioxidants. and depending on the physical, nutritional, and sensory
requirements of specific products, there is continuing potential
for new product development based on amaranth. Examples of
Food and Feed Utilization of Amaranth recent studies include that of Martinez et al. who investigated
the effect of A. mantegazzianus flour on the technological and
Traditional foods are made from amaranth grains in many sensory quality of bread wheat pasta. Also, Diaz et al. drew
countries, for example, alegria and atole in Mexico, attention to the significance of amaranth as source of gluten-
alboroto in Guatemala, bollos in Peru, Chapati in free ingredients. They added 20% amaranth to a maize base
THE LEGUMES AND PSEUDOCEREALS | Amaranth: Overview 295
used in twin screw extrusion. Addition of amaranth conferred animal feed. Its red-colored vegetative tissue with high level
increased sectional expansion index. Also, in recent years, ama- of betacyanins is a potential alternative source of well-known
ranth grains have been developed and processed for many beetroot pigments (betalains). Amaranth has been attracting
innovative new food products in China, such as amaranth- worldwide attention and will be a high-potential new crop
flavored cake, amaranth yogurt, and amaranth Taosu (a type with multiple uses. Since the first edition of this article 12
of traditional Chinese sweet cake). years ago, more than 250 research papers in international
Furthermore, vegetable amaranth has been used as a tradi- scientific journals have been published on food science aspects
tional vegetable for several centuries in China. In sub-Saharan of amaranth, most regarding novel applications of nutritional
Africa and Central and South America, vegetable amaranth is fractions of the grain. This developing corps of technical
also an important leafy vegetable, consumed as a cooked green knowledge may help to increase demand to a level that will
vegetable or used in soup. justify investment in major production and processing sys-
Amaranth is a fast-growing, high-nitrogen, and high- tems. However, intense and continuous efforts are necessary
biomass plant that can be used fresh or as silage for animal in several aspects, such as amaranth breeding and field culti-
feed. Amaranth grain flour or powdered leaf meal can be used vation (for yield, shattering, and lodging), relevant food/feed
to make commercial pelleted or other compound feeds. Cur- processing and development, product industrialization, and
rently, feed application of amaranth has been successful and marketing.
popular in China and has played a role in developing Chinese
animal husbandry. Amaranth feeds made from leaves, stems,
panicles, and grains include green fodder, silage, leaf and seed- Exercises for Revision
meal feed, pellet feed, etc. The evaluation of feed use has
shown that amaranth feeds efficiently raise pigs, chickens,
What makes Amaranthus most suited for food or for feed
cattle, sheep, and fish to improve the quality and productivity production?
of animal production. Chinese peasant households are accus-
What are the general properties of Amaranthus starch and
tomed to raise livestock and poultry, and nowadays, in many protein?
areas, amaranth is the main source of silage for household pig
production. However, it is not widely used as feed in other
countries.
Exercises for Readers to Explore the Topic Further
Cai YZ and Corke H (2000) Production and properties of spray-dried Amaranthus Saunders RM and Becker R (1984) Amaranthus: A potential food and feed resource.
betacyanin pigments. Journal of Food Science 65: 12481252. In: Pomeranz Y (ed.) Advances in Cereal Science and Technology, vol. 6,
Cai Y, Sun M, and Corke H (2005) Development and application of betalains from the pp. 357396. St. Paul, MN: American Association of Cereal Chemists.
Amaranthaceae. Trends in Food Science and Technology 16: 370376. Segura-Nieto M, Barba de la Rosa AP, and Paredes-Lopez O (1994) Biochemistry of
Cai YZ, Sun M, Wu HX, Huang RH, and Corke H (1998) Characterization and amaranth proteins. In: Paredes-Lopez O (ed.) Amaranth: Biology, Chemistry, and
quantification of betacyanin pigments from diverse Amaranthus species. Journal of Technology, pp. 75106. Boca Raton, FL: CRC Press.
Agricultural and Food Chemistry 46(6): 20632070. Singhal RS and Kulkarni PR (1988) Composition of the seeds of some Amaranthus
Chen JZ (2011) Development and preparation of amaranth yogurt. Food Research and species. Journal of the Science of Food and Agriculture 42(4): 325331.
Development (in Chinese) 12: 7273. Sleugh BB, Moore KJ, Brummer EC, Knapp AD, Russell J, and Gibson L (2001) Forage
Corke H, Wu HX, Yue SX, and Sun HL (1997) Developing specialty starches from new nutritive value of various amaranth species at different harvest dates. Crop Science
crops: A case study using grain amaranth. In: Campbell GM, Webb C, and 41: 466472.
McKee SL (eds.) Cereals: Novel Uses and Processes, pp. 91102. New York: Teutonico RA and Knorr D (1985) Amaranth: Composition, properties, and applications
Plenum. of a rediscovered food crop. Food Technology 39(4): 4954.
Diaz JMR, Kirjoranta S, Tenitz S, et al. (2013) Use of amaranth, quinoa and kaniwa in Venskutonis PR and Kraujalis P (2013) Nutritional components of amaranth seeds and
extruded corn-based snacks. Journal of Cereal Science 58: 5967. vegetables: A review on composition, properties, and uses. Comprehensive Reviews
He HP, Cai YZ, Sun M, and Corke H (2002) Extraction and purification of in Food Science and Food Safety 12: 381412.
squalene from Amaranthus grain. Journal of Agricultural and Food Chemistry Wang YY, Wang HS, Zhang HT, and Cui YH (2011) Research advances of amaranth as a
50: 368372. high-quality feed plant. Feed Research (in Chinese) 1: 1920.
Irving DW, Betschart AA, and Saunders RM (1981) Morphological studies on Williams JT and Brenner D (1995) Grain amaranth (Amaranthus species).
Amaranthus cruentus. Journal of Food Science 46(4): 11701174. In: Williams JT (ed.) Cereals and Pseudocereals, pp. 129186. London: Chapman
Lamothe LM, Srichuwong S, Reuhs BL, and Hamaker BR (2015) Quinoa and Hall.
(Chenopodium quinoa W.) and amaranth (Amaranthus caudatus L.) provide Wu HX and Corke H (1999) Genetic diversity in physical properties of starch from a
dietary fibres high in pectic substances and xyloglucans. Food Chemistry world collection of Amaranthus. Cereal Chemistry 76: 877883.
167: 490496. Wu HX, Yue SX, Sun HL, and Corke H (1995) Physical properties of starch from two
Leon-Camacho M, Garcia-Gonzalez DL, and Aparicio R (2001) A detailed and genotypes of Amaranthus cruentus of agricultural significance in China. Starch/
comprehensive study of amaranth (Amaranthus cruentus L.) oil fatty profile. Starke 47: 295297.
European Food Research and Technology 213: 349355. Yue SX, Sun HL, and Tang DF (1993) The Research and Development of Grain
Lyon CK and Becker R (1987) Extraction and refining of oil from amaranth seed. Journal Amaranth in China (in Chinese). Beijing: Chinese Agricultural Science and
of the American Oil Chemists Society 64: 233236. Technology Press.
Martinez CS, Ribotta PD, Anon MC, and Leon AE (2014) Effect of amaranth flour Zhang FF, Jiang YP, Cao YJ, and Sun HN (2014) Feed value of amaranth and its
(Amaranthus mantegazzianus) on the technological and sensory quality of bread utilization in poultry. Poultry Science (in Chinese) 3: 68.
wheat pasta. Food Science and Technology International 20: 127135.
Paredes-Lopez O (1994) Amaranth: Biology, Chemistry, and Technology. Boca Raton,
FL: CRC Press.
Quiroga AV, Barrio DA, and Anon MC (2015) Amaranth lectin
presents potential antitumor properties. LWT Food Science and Technology Relevant Websites
60: 478485.
Sabbione AC, Scilingo A, and Anon MC (2015) Potential antithrombotic activity http://www.agmrc.org/commodities__products/specialty_crops/amaranth/ Website
detected in amaranth proteins and its hydrolysates. LWT Food Science and of the Agricultural Marketing Resource Center, Iowa State University.
Technology 60: 171177.
Beans: Overview
SK Sathe, Florida State University, Tallahassee, FL, USA
2016 Elsevier Ltd. All rights reserved.
Data from Deshpande SS and Srinivasan D (1990) Food legumes: Chemistry and technology. Advances in Cereal Sciences and Technology 10: 147241; Doughty J and Walker A
(1982) Legumes in Human Nutrition. Food and Nutrition Paper 20, 152 pp. Rome, Italy: Food and Agriculture Organization of the United Nations.
Morphology of the Pods/Seeds the seed coat, the cotyledons, and the embryo axis. In most dry
seeds, they account for 820%, 8090%, and 12% of the seed
Regardless of the fat content, most legume seeds have similar weight, respectively. The majority of the nutrients are present
structure. Mature legume seeds have three major components: in cotyledons. Typical seed structure and various anatomical
THE LEGUMES AND PSEUDOCEREALS | Beans: Overview 299
Table 2 Acreage, production, yield, and import/export data for pulses and dry beans
Tons $ Tons $
P D P D P D D D D D
World 5 903 158 28 780 377 8554 8040 5 049 539 23 140 276 3 321 726 3 227 563 000 3 393 895 3 074 549 000
Africa 1 890 425 7 198 019 6357 6585 1 201 768 4 740 016 408 551 331 881 000 311 174 212 970 000
Asia 2 617 155 14 499 325 6404 7359 1 675 921 10 670 688 1 397 795 1 343 293 1 681 843 1 523 401 000
Caribbean 2156 257 317 13 919 9195 3001 236 602 100 273 89 401 000 897 1 019 000
Europe 688 286 263 487 17 230 18 536 1 185 919 488 408 609 885 690 887 000 139 648 167 956 000
Central 8750 2 278 678 31 657 7240 27 700 1 649 772 230 600 220 369 000 104 363 117 170 000
America
North NA 804 482 NA 21 392 NA 1 720 950 225 709 258 776 000 603 769 513 547 000
America
South 1790 3 419 069 14 441 10 482 2585 3 583 840 332 041 272 811 000 455 509 444 047
America
Oceania 694 596 60 000 13 715 8333 952 645 50 000 15 872 20 145 000 96 692 94 439 000
Seedcoat (testa)
Cotyledon
Raphe
Plumule
Hilum Hypocotyl
Radicle
Micropyle
Outer
integument
Seedcoat
Inner
integument
Epidermis
Cotyledon
Storage
parenchyma
cells
(d)
Figure 1 Dry bean (Phaseolus vulgaris) seed: (a) external side view, (b) external face or edge view (viewing at hilum side), (c) cross section (one
cotyledon removed), and (d) detailed cross section across seed coat and cotyledon. Reproduced from Encyclopedia of Food Sciences and Nutrition,
2nd edn. (2003), p. 407, Elsevier Ltd.
parts of legume seeds are shown in Figure 1. The outermost is the small opening in the seed coat where originally the
layer of the seed is the seed coat or testa. The external seed pollen tube enters the valve. The raphe is the ridge at the side
structure includes the hilum, micropyle, and raphe. The hilum of hilum opposite to the micropyle and represents the base of
is a scar-like structure (usually oval shaped) near the middle the stalk that fuses with the seed coat upon seed maturation. In
edge where the seed breaks away from the stalk. The micropyle most legumes, the endosperm is short-lived and shrinks to a
300 THE LEGUMES AND PSEUDOCEREALS | Beans: Overview
thin layer surrounding the cotyledons (or embryo). On soak- are often used as key structures in the identification of different
ing the seeds, the endosperm is easily removed along with the plant types.
seed coat. The remainder (embryo) of the seed consists of
shoot (which contains two cotyledons) and a short axis
above and below the cotyledons and terminates in the shoot
tip. The plumule or embryonic stem is well developed in the Chemical and Nutritional Composition
resting seed and lies between two cotyledons.
The outermost layer of seed coat is the cuticle, which has The majority of nutrients in dry beans are primarily located in
papillae or papillae-like growth in some legumes (e.g., green the cotyledons and account for up to 90% of the total nutritive
gram), but in most legumes, it is a smooth structure. The value. Typically, dry beans provide 12551464 kilojoules (kJ)
thickness of the seed coat is quite variable depending upon per 100 g of dry seeds. The main constituents of cotyledons are
the type of bean. Generally, seeds containing thick seed coats proteins and carbohydrates and, respectively, account for
tend to have high fat content. Both the hilum and micropyle 1525% and 5075% of the total seed weight. The remaining
are important in water imbibition by the testa. Palisade cells components consist of lipids, minerals, fiber, and vitamins.
derived from the outer epidermis of the outer integument, With the exception of oilseeds, dry beans generally contain
which are next to the cuticle, are either loosely packed or low amounts (13% of seed weight) of lipids. Although most
densely packed, depending upon the seed maturity, and may minerals are present in cotyledons, some (such as calcium and
affect seed hydration. Next to palisade cells are the hourglass iron) may be present in the seed coat in significant proportion.
cells. The shape of hourglass cells can vary from bottle-shaped Typical nutrient composition of several Phaseolus beans is
(guar) to dumbbell-shaped (broad beans) to hourglass- shown in Table 3. Dry beans contain not only significant
shaped (soybeans). Only a few legume species (such as amounts of nutrients but also several undesirable components
Dolichos, Cajanus, and Vigna) have more than one layer of and attributes: inhibitors of enzymes such as trypsin, chymo-
hourglass cells. The remainder of the testa contains primarily trypsin, subtilisin, amylases, and elastase; lectins; phenolic
mesophyll cells. compounds including tannins; phytates; toxic amino acids
Legume cotyledons are primarily composed of parenchyma mimosine and djenkolic acid; cyanogenic glycosides, which
cells, which have variable size (70100 mM) and act as storage produce HCN; flatulence-causing oligosaccharides raffinose,
sites for most nutrients. Each cell of the cotyledon is bound by stachyose, and verbascose; lipoxygenases, which catalyze the
the cell wall and the middle lamella. Vascular bundles in cotyle- development of rancidity; and off odors often described as
dons generally are devoid of any filling material. Vascular bundles beany, grassy, painty, cardboard-like, and chalky.
Bean Moisture (%) Protein (%) Carbohydrates (%) Fat (%) Ash (%) Crude fiber (%)
(including agglutinating activity) with respect to chemical shown to be 50 000. Appropriate moist-heat treatment (such
composition. as cooking or autoclaving) can inactivate both the protease and
amylase inhibitors.
Other proteins
Phaseolus beans contain trypsin inhibitors (many of them also
Allergenic Proteins
inhibit chymotrypsin), amylase inhibitors, lipoxygenases, and
several other minor protein components. Most of these pro- With heightened concerns about food allergies and the
teins are a part of albumins. Trypsin and chymotrypsin inhib- increased frequency of food recalls due to actual or suspected
itors in Phaseolus beans typically account for up to 10% of the presence of allergen in food, food-induced allergens are of
total proteins and are generally rich in sulfur amino acids. The increased interest to consumers and regulatory agencies alike.
MWs of these inhibitors range from 2000 to 23 000. Most Since any food protein could be a potential allergen, it is
Phaseolus beans lack Kunitz-type (inhibitors with 170200 important to recognize that dry bean proteins may be aller-
amino acids with MW of 20 000) trypsin inhibitors. Amylase genic. Compared to aeroallergens, such as pollen, perhaps with
inhibitors in dry beans have been characterized from only a the exception of peanut and soybean proteins, very little is
few cultivars and therefore not yet extensively studied. The MW known about dry bean protein allergens. A few dry bean pro-
of kidney bean amylase inhibitor (a glycoprotein) has been teins have been identified to be allergens (Table 4) and even
MW (kDa) Allergen
Legume SDS-PAGE Identity designation Seq.a Accession number PubMed index
Dry beans
Chickpea
Cicer 10106 11527247,
arietinum 10705224
Cowpea
Vigna 41, 55 Albumins 12546052
sinensis
Grass pea
Lathyrus 21, 28, 46 11295670
sativus
Green pea
Pisum 20, 2030, Vicilin-like proteins 12589366
sativum 33, 50
1.8 Glycoprotein 991447
French 8.89 Nonspecific lipid transfer protein type 1 C GQ434007.1
(green) (ns-LTP-1)
bean Nonspecific lipid transfer protein 1b C GQ434008.1
precursor
Lentil
Lens 1284 11080720,
culinaris 10604559,
9893199
48 Vicilin-like protein Len c 1 C
66 Seed-specific biotinylated protein Len c 2 P 11080720
Mung bean 16 Pathogenesis-related protein, PR-10 C AY792956.1
Vigna radiata Bet v 1 family member
53 8S globulin (vicilin) C Q198W3, B1NPN8
30 Albumin C Q43680
18 Cytokinin-specific binding protein (CSBP), C AB012218
Bet v 1 family member
Oil seeds
Castor bean
Ricinus 11.2 2S albumin Ric c 1 C P01089 9430499
communis
12 2S albumin Ric c 3 P 9430499
11S crystalloid protein Ric c 2 3392372
47/51 Protein doublet 3392372
Lupin
Lupinus 30175 10359910
albus
Peanut
THE LEGUMES AND PSEUDOCEREALS | Beans: Overview 303
Table 4 (Continued)
MW (kDa) Allergen
Legume SDS-PAGE Identity designation Seq.a Accession number PubMed index
shape (round, oval, oblong, elliptical, spherical, kidney-shape- beans, phytate phosphorus accounts for up to 80% of the total
d, and irregular) and size (560 mm) and typically contain phosphorus. The amount of phytate in Phaseolus beans ranges
1045% (of total starch) amylose. The average degree of poly- from 0.6% to 2.1% (by weight) of the total seed weight.
merization for amyloses and chain lengths for amylopectins of Because phytates are chelating agents, they may interfere in
Phaseolus bean starches ranges from 1600 to 1900 and 22 to 26, mineral utilization. Germination, fermentation, and soaking
respectively. Based on x-ray diffraction spectra, Phaseolus dry followed by cooking (if both soak and cook waters are dis-
bean starches are mostly C type (mixture of A (typical of cereal carded) are effective methods of removing phytates (5080%
starches) and B (typical of root and tuber and high-amylose reduction). Tannins (especially condensed tannins) are heat-
cereal starches) types). These starches have restricted swelling, stable compounds and are present in Phaseolus seeds (espe-
gelatinization temperature range of 6089 C, high solution cially in colored varieties) up to 2% of total seed weight.
viscosities, and good thermal stabilities. Upon gelatinization, Because of their ionic character, they may interact with other
they produce opaque gels. Dry bean starches (especially if constituents (notably proteins) and adversely affect the nutri-
cooked) are well digested (digestibility is comparable to those tional bioavailability of that constituent. Tannins are thought
of many cereal and tuber starches) by humans. However, as dry to offer protection to the plant from insects and pests.
bean starches are digested slowly, they are hypoglycemic and
therefore useful in the diets of diabetics. The nonstarchy poly-
Lipids
saccharides are not digestible in many mammals due to the
lack of required enzymes in their digestive tracts and therefore Phaseolus beans contain 13% lipids (by weight) depending
contribute to the total dietary fiber. These nonstarchy poly- upon the species. Neutral lipids (3050% of total) and phos-
saccharides help increase viscosity and may thereby lower pholipids (2535% of total) are the major constituents, and
digestion rate of starch (i.e., lower glycemic index). Further, glycolipids may account for up to 10% of the total lipids.
they may assist removal of dietary lipids (i.e., hypocholester- Regardless of the variety, Phaseolus bean lipids primarily con-
olemic effect). tain palmitic, oleic, linoleic, and linolenic acids. Polyunsatu-
rated fatty acids and saturated fatty acids typically account for
5587% and 1228% of total lipids.
Vitamins and Minerals
Phaseolus beans are a good source of B vitamins, especially
Beneficial Bioactive Compounds
thiamin, riboflavin, niacin, and folacin. Typically, thiamine,
riboflavin, niacin, and folacin contents (on a dry weight basis) Continuing research on raffinose oligosaccharides, phytates,
of Phaseolus beans are, respectively, 0.51.14, 0.10.25, tannins, simple phenolic compounds, sterols, protease inhib-
0.43.14, and 0.0370.676 mg per 100 g. Vitamin E content itors, and several other major and minor constituents of dry
ranges from 0.72 to 1.97 mg per 100 g and B6 content ranges beans suggests that some of these compounds, under appro-
from 0.2 to 0.659 mg per 100 g. Phaseolus beans are not good priate conditions, may offer beneficial effects for human
sources of vitamin A and C. Phaseolus beans are excellent health. Numerous reports suggesting the beneficial effects of
sources of several minerals including Ca, Fe, Cu, Zn, P, K, one or more components (either alone or in combination)
and Mg. Typically, raw beans contain 70260, 0.51.40, have appeared. These investigations have used in vitro and
3.3413.5, 160320, 1.02.1, 380570, 13201780, in vivo model studies to decipher possible beneficial effect(s).
4.021.0, and 1.96.5 (all expressed as mg per 100 g, dry The primary goals of these investigations are to (a) derive
weight basis) of Ca, Cu, Fe, Mg, Mn, P, K, Na, and Zn, respec- nutritionally beneficial effect for long-term good physical and
tively. The low sodium and high potassium content of raw mental health and (b) treat/cure disease. Experimental dem-
beans makes them a desirable constituent of human food onstration of causeeffect relationship can be tenuous and
(especially for people with hypertension). difficult to prove/disprove and may depend on the experimen-
tal design. With the simultaneous presence of several com-
pounds in the same food source, establishing unequivocal
Antinutritional Factors
relationship between the compound(s) and their physiological
In addition to protease inhibitors and amylase inhibitors, Pha- effect(s) is, at best, very difficult. In recent years, attempts to
seolus beans contain several other antinutritional and investigate the underlying molecular and mechanistic basis for
antiphysiological factors such as phytic acid, tannins, cyano- the reported beneficial effect(s) of tested compound(s)/com-
genic glycosides, saponins, and allergens. Because these com- ponent(s) have therefore often resulted in outcomes with very
ponents are usually present in small quantities (less than 5% of limited success. Long-term detailed and fundamental research
the total seed weight), they do not pose a serious health hazard is needed to fully realize the potential health benefits of dry
under normal conditions (i.e., when beans are a part of total bean consumption.
diet and are properly processed prior to consumption). Of
these antinutritional factors, phytates and tannins are of par-
ticular concern because both of them are heat-stable and can- Grading, Handling, and Storage
not be easily removed from beans during normal home
processing. Phytate is a general term used for mono- to dodeca Dry beans are usually harvested at maturity. The seeds are
anions of phytate along with esters lower than hexaphosphate. removed from the pods either manually or mechanically and
CaMg salts of phytic acid (myoinositol 1, 2, 3, 4, 5, 6-hexakis cleaned to remove dirt, stalks, leaves, blemished, wrinkled, and
(dihydrogen phosphate)) are referred to as phytin. In most dry broken seeds and packaged prior to storage. The grading of
THE LEGUMES AND PSEUDOCEREALS | Beans: Overview 305
seeds is usually based on external characteristics such as color, preparation of idli, a breakfast food popular in India and Sri
gloss, seed size, seed soundness, seed firmness, and presence of Lanka, after fermenting it with rice. Industrial processing
contaminating substances. The seeds are stored at farmer, includes freezing for such beans as green French beans and
trader, or government levels. Typically, farmers hold up to snap beans, milling (production of flours and high-protein
8% of harvested seeds until next season so that they can be flours), baking (baked beans), cooking and frying (refried
used as planting seeds if crop failures occur. In developed and beans), and canning (alone in saltwater or tomato juice or in
developing nations, a majority of the seeds are stored by combination with meats such as beef and pork). In developing
traders and/or governments to protect against subsequent countries, dehusking and splitting to produce dal is also done
crop failures (or low yields), price fluctuations, and fluctua- at industrial scale.
tions in supply (change in demand, shortages, and famines). In developing and underdeveloped countries, Phaseolus
Losses in seeds occur both pre- and postharvest. The normal beans are used in numerous ways depending on the type of
preharvest losses are mainly due to birds and mammals feed- bean and regional preference. They may be eaten as raw,
ing on bean plant seeds. Drought/floods, insects, and rodents immature seeds; cooked as green vegetables (such as French
can also contribute to preharvest losses. In developed nations, beans); consumed as part of salads; used in making curry; used
preharvest losses are usually small (as low as 1% of the crop). A as a soup ingredient; cooked, mashed, and mixed with condi-
majority of the losses occur during postharvest handling and ments and spices and used as gruels and porridges; prepared as
storage and can range from 8% to as high as 50% of total crop. pastes to be extruded to prepare fried snack products; sprouted;
It is estimated that as much as 48% of food produced in the puffed or roasted and eaten as snack foods; and fermented to
world is lost (due to pre- and postharvest losses). Factors that prepare numerous fermented products.
influence postharvest losses of legumes include moisture, tem- In developed countries, Phaseolus beans are consumed as
perature, respiration rate, insect damage, microbial spoilage, salad and soup ingredient, sprouts, canned, frozen, and refried
and damage caused by mites and rodents. Properly packaged beans. They are also extensively used in the preparation of
dry beans should be stored at low relative humidity and tem- Mexican-style preparations, such as burrito, chimichanga,
perature conditions. High relative humidity and temperature taco, bean dips, and tamale, and often canned with meats
favor the hard-to-cook beans. These conditions also favor the such as beef and pork. In many South American countries,
growth of molds and insects. Three major insect genera that cooked black beans are a preferred part of breakfast. With
cause much of the damage to stored legumes are Bruchus, increased ethnic populations in the Western world, increased
Acanthoscelides, and Callosobruchus. Dehusked, split stored emphasis on vegetarian foods, and recommended reduction of
pulses are also damaged by Rhyzopertha, Trogoderma, and Tri- meats and animal foods in the human diet, opportunities need
bolium species. Usually, pests seem to have preference for the to be explored to incorporate dry beans in food products for
type of bean they infest although the basis for such preference improved and nutritious food products.
(or the lack of it) has not been elucidated yet. The major Although there is a good potential for the preparation of
microbial problem during bean storage is contamination by protein concentrates and isolates and the development of food
aflatoxin-producing molds (Aspergillus flavus and Aspergillus starches, Phaseolus beans have not been used on large scale for
parasiticus). Mites can consume food up to their own weight such purposes. In many countries, especially the developed
(68 mg) and because of their large numbers can cause serious ones, Phaseolus beans have been extensively used as animal
losses. Rodents cause twofold damage to stored legumes by not feed. In developing and underdeveloped countries, the green
only consuming but also contaminating (up to 20 times the foliage, deseeded pods, and roots and shoots of bean plants are
amount they would eat). Because rodents are carriers of many used as natural fertilizers especially after composting. Because
communicable diseases, they pose serious damage to stored the legume roots fix nitrogen, they help conserve soil quality.
beans. The species that most commonly cause damage include For this reason, in many developing countries, they are exten-
Rattus rattus, R. norvegicus, Bandicota indica, B. bengalensis, and sively used for soil quality conservation.
Mus musculus.
Phaseolus beans are processed and used in a variety of ways. The What distinguishes cereals from legumes?
processing of beans is mostly at the household level in devel- What is the difference between the terms legume and
oping and underdeveloped countries, while in most developed oilseed?
countries, the majority of the processing is done at the indus- Which type of beans are the most widely consumed in the
trial level. Home processing methods include milling, soaking, world?
cooking, frying, germination, fermentation (either alone or in Which compound(s) account for 80% phosphorus in dry
combination with cereals), roasting, puffing, parching, extru- beans?
sion and frying, and toasting. The method(s) used for home What is the advantage of using dry beans with cereals in
processing depends on regional preference for bean variety and foods?
the desired end product. For example, mung beans in sprouted What is the advantage of the nonstarchy polysaccharides
form are popular on global scale and therefore germination is present in dry beans in human nutrition?
one of the preferred household processing methods used. What is the difference between the terms pulse and
Black gram, on the other hand, is extensively used for the legume?
306 THE LEGUMES AND PSEUDOCEREALS | Beans: Overview
Exercises for Readers to Explore the Topic Further Doughty J and Walker A (1982) Legumes in Human Nutrition, p. 152. Rome, Italy: Food
and Agriculture Organization of the United Nations Food and Nutrition Paper 20.
FAO (2013) http://faostat.fao.org/site/567/DesktopDefault.aspx?PageID567#ancor
What are the possible beneficial actions of bioactive com- (accessed 22.02.2014).
ponents of dry bean? FAO (2013) Dimensions of Need Staple Food: What Do People Eat? http://www.fao.
What are the factors that result in pre- and postharvest org/docrep/u8480e/u8480e07.htm (accessed 22.02.2014).
losses? Geil PB and Anderson JW (1994) Nutrition and health implications of dry beans: A
review. Journal of the American College of Nutrition 13: 549558.
How does the ionic character of phytic acid influence its Guillon F and Champ MM (2002) Carbohydrate fractions of legumes: Uses in
interaction with dietary minerals and proteins? human nutrition and potential for health. British Journal of Nutrition
In what ways can the dry bean utilization be improved? 88: S293S306.
Kami JA, Velasquez VB, Debouck DG, and Gepts P (1995) Identification of presumed
ancestral DNA sequences of phaseolin in Phaseolus vulgaris. Proceedings
of the National Academy of Sciences of the United States of America
See also: Agronomy of Grain Growing: Lentil: Agronomy; 92: 11011104.
Breeding of Grains: Lentil: Breeding; Food Grains and the Muntz K (1998) Deposition of storage proteins. Plant Molecular Biology 38: 7799.
Osborn TC (1988) Genetic control of bean seed proteins. Critical Reviews in Plant
Consumer: Cultural Differences in Processing and Consumption; Sciences 7: 93116.
Proteins: The Protein Chemistry of Dicotyledonous Grains; The Patwardhan VN (1962) Pulses and beans in human nutrition. American Journal of
Basics: Taxonomic Classification of Grain Species; The Legumes Clinical Nutrition 11: 1230.
and Pseudocereals: Chickpea: Overview; Grain Legumes and Their Reddy NR, Pierson MD, Sathe SK, and Salunkhe DK (1984) Chemical, nutritional and
physiological aspects of dry bean carbohydrates: A review. Food Chemistry
Dietary Impact: Overview; Pea: Overview. 13: 2568.
Reddy NR, Pierson MD, and Salunkhe DK (1986) Legume-Based Fermented Foods,
p. 254. Boca Raton, FL: CRC Press.
Salunkhe DK and Kadam SS (1989) Handbook of World Food Legumes: Nutritional
Chemistry, Processing Technology, and Utilization. Boca Raton, FL: CRC Press
Further Reading vols. I (310 pp.), II (294 pp.), and III (323 pp.).
Sathe SK (2002) Dry bean protein functionality. Critical Reviews in Biotechnology
Broughton WJ, Hernandez G, Blair M, Beebe S, Gepts P, and Vanderleyden J (2003) 22: 175223.
Beans (Phaseolus spp.) Model food legumes. Plant and Soil 252: 55128. Sathe SK, Deshpande SS, and Salunkhe DK (1984) Dry beans of Phaseolus: A review.
Derbyshire E, Wright DJ, and Boulter D (1976) Legumin and vicilin, storage proteins of Part 1. Chemical composition: Proteins. CRC Critical Reviews in Food Science and
legume seeds. Photochemistry 15: 324. Nutrition 20: 146.
Deshpande SS (1992) Food legumes in human nutrition: A personal perspective. CRC Siddiq M and Uebersax MA (eds.) (2013) Dry Beans and Pulses: Production,
Critical Reviews in Food Science and Nutrition 32: 333363. Processing and Nutrition, pp. 398. Ames, IA: Wiley.
Buckwheat: Overview
YZ Cai, China National Seed Group, Beijing, China; Huazhong Agricultural University, Wuhan, China
H Corke, The University of Hong Kong, Hong Kong, Pokfulam, China; Hubei University of Technology, Wuhan, China
D Wang, Huazhong Agricultural University, Wuhan, China
WD Li, Shanxi University, Taiyuan, China
2016 Elsevier Ltd. All rights reserved.
Topic Highlights cereals. In addition, it also contains high levels of fiber, min-
erals, and vitamins and other bioactive compounds (e.g., fla-
Cultivated buckwheat comprises two species of the genus vonoids, phytosterols, D-chiro-inositol, myo-inositol, free and
Fagopyrum, namely, F. esculentum Moench (common buck- bound phenolic acids, and phenylpropanoid glycosides) with
wheat) and F. tataricum L. (Tartary or bitter buckwheat). positive therapeutic effects on the human body. Buckwheat
Originating in China and widely produced, mostly in Asia with both edible and pharmaceutical values has increasingly
and in eastern Europe, buckwheat produces a pseudocereal attracted attention because of its positive effects on some
grain that can be dehulled and milled to flour, for use in a chronic disease conditions, such as hypertension, hypercholes-
wide range of food products. terolemia, diabetes, and other cardiovascular diseases. Buck-
The composition and properties of buckwheat flour, wheat has further potential for use as an important functional
derived from its components of starch, protein, and various food source.
therapeutic components such as flavonoids and phytos-
terols, give a wide scope for development of healthful
grain-based products. General Aspects of Buckwheat Production
perennial species such as Fagopyrum cymosum Meissn and Fago- glycosides). These are distributed in different parts of the buck-
pyrum suffruticosum Fr. Schmidt. Among these species, only wheat grain, for example, protein mainly occurs in the aleu-
F. esculentum and F. tataricum are cultivated species while rone layer and embryo, starch in the endosperm, and fiber, ash,
others are wild species. Buckwheat has broad ecological adapt- and flavonoids are normally in the testa and pericarp. Milling
ability and can grow well in adverse soil and climatic condi- procedures, fineness, and flour yield influence chemical com-
tions; in particular, Tartary buckwheat can grow in poor soils position in the final products of buckwheat milling (Table 2).
even at low temperature and low precipitation, such as in high- Different buckwheat varieties (genotypes) and plant condi-
altitude mountainous regions. tions (soil and climate) also cause compositional differences.
Common buckwheat is characterized by petiolate leaves,
with terminal and axillary flowers, and corymbose or paniculate
Protein and Amino Acids
cyme. Tartary buckwheat has narrow-shaped leaves, more
branches, and smaller flowers than common buckwheat, with Buckwheat grain contains not only a high level of protein but
inconspicuous greenish-white sepals. Buckwheat flowers are also a good quality or nutritional balance of protein. Protein
indefinite inflorescences that are white, pink, or yellow. Because content of buckwheat grain and flour is similar to or slightly
of its flourishing flowers, buckwheat is cultivated in China to higher than that in wheat grain and flour (Table 2) but
support the production of honey, considering one of the highest- significantly higher than those in many other cereals (e.g.,
yielding honey plants. Buckwheat grain is actually a fruit rice, maize, and sorghum). Amino acid composition of buck-
(achene) whose color ranges from a glossy or dull gray to wheat protein is nutritionally well balanced and is rich in
brown or black. Buckwheat grain (achene) consists of hull (peri- essential types (e.g., lysine and arginine), compared with
carp) and kernel or seed (testa, aleurone layer, embryo, and most cereals. Table 3 shows that all amino acids are higher
endosperm). Such achenes are mostly triangular, 49 mm in buckwheat grain than in wheat grain, except for glutamic
long with 1000 grain weight ranging from 15 to 35 g. Common acid and proline.
buckwheat grain is generally bigger and huskier than Tartary
buckwheat grain. Common buckwheat has a softer hull than
Carbohydrate
Tartary buckwheat. Common buckwheat grain tastes harsh, and
Tartary buckwheat tastes slightly bitter. Carbohydrate (mainly starch) is the major component
(5974%) of buckwheat grain (Table 2). Amylose content in
starch generally ranges from 20% to 26%. Starch granules are
Chemical Composition of Buckwheat 215 mm in diameter, round, oval, or polygonal in shape with
a few holes and pits on the surface. Buckwheat starch exhibits a
Chemical components of buckwheat include protein/amino typical A-type x-ray diffraction pattern and the crystallinity
acids, starch, lipids, fiber, minerals, vitamins, and other func- varies from 38.3% to 51.3%. Buckwheat grains also contain
tional components (e.g., flavonoids, phytosterols, fagopyrins, 0.650.76% reducing sugars, 0.791.16% oligosaccharides,
D-chiro-inositol, myo-inositol, and phenylpropanoid and 0.10.2% nonstarchy polysaccharides.
Composition Buckwheat seeds Buckwheat groats Buckwheat crude flour Buckwheat fine flour Wheat grain Bread wheat flour
buckwheat grain generally has higher crude fiber than Tartary Vitamins
buckwheat. Content of crude fiber in buckwheat flour com-
Buckwheat grain contains higher levels of vitamin B1
monly depends on the milling process because fiber is mainly
(thiamine), B2 (riboflavin), E (tocopherol), and PP (nicotinic
distributed in the testa and pericarp. Crude flour contains more
acid/amide) than most cereals (Table 4). Generally, Tartary
fiber (3.46.9%) than fine flour (0.11.6%).
buckwheat has more vitamin B1, B2, and PP, but less vitamin
E than common buckwheat.
Minerals
Buckwheat grain is rich in minerals including Mg, Se, Fe, K, Ca, Therapeutic Components
Cu, Mn, and Zn. Some studies show that buckwheat flour
contains more Mg (1113 times), Sr (536 times), Li (57 Buckwheat is also rich in components with therapeutic effects,
times), Fe (34 times), and K (2.93.6 times) than wheat including flavonoids, phytosterols, fagopyrins, D-chiro-
flour. Additionally, a high level of Se is detected in Tartary inositol, myo-inositol, phenylpropanoid glycosides, free and
buckwheat flour (0.43 mg kg1). bound phenolic acids, and proanthocyanidins.
Table 4 Comparison of vitamin contents between buckwheat flour and some cereal flour
Table 5 Functional properties of common buckwheat protein concentrates, compared to other protein isolates
Protein concentrate Emulsifying activity at pH 7.0 (%) Solubility (%) Foam stability (ml)
antinutritional factors: trypsin inhibitors, tannin, and a high and noodles containing more buckwheat induced significantly
content of dietary fiber. Buckwheat trypsin inhibitors are rela- lower postprandial blood glucose and insulin responses than
tively thermostable, acid-resistant, and partially susceptible to with whole wheat bread.
pepsin action. In addition, allergens in buckwheat proteins Additionally, there are effects of processing treatments
have attracted attention. During buckwheat grain milling and (including steaming or autoclaving and extruding) on buck-
food processing, some workers have allergic reactions, such as wheat starch properties. The results showed that hydrothermal
nose itching, sneezing, rhinorrhea, dyspnea, asthma, and even treatment increases gelatinization temperature, swelling
more serious symptoms. It is necessary to know more about power, and solubility of buckwheat starch. Autoclaving/cool-
the behavior and processing properties of specific buckwheat ing treatment of buckwheat groats resulted in improved avail-
allergenic proteins. ability of buckwheat starch and led to more apparent amylose
and true amylose in the buckwheat starch and more resistant-
starch formation in comparison to the untreated samples. It
Starch was also reported that extrusion influenced the physico-
chemical properties of starch and formation of starchprotein
Buckwheat starch normally has higher peak viscosity, higher
and starchlipid complexes in extruded mixtures containing
gelatinization temperature, greater swelling power, and higher
buckwheat flour and starch. Starchprotein complexes formed
susceptibility to acid and enzymatic degradation when com-
during extrusion played an important role in the stability of the
pared with cereal starches (maize and wheat). Buckwheat
porous structure in the extruded products containing buck-
starch retrogradation is generally slower than that of maize
wheat starch.
and wheat starch. Buckwheat starch has lower syneresis when
stored at 4 C for 310 days and has better stability to syneresis
after freezethaw cycles at 12 to 25 C.
The starch properties of common and Tartary buckwheat Therapeutic Effects of Buckwheat
compared to wheat starch. Clear differences were found in
physicochemical properties between buckwheat and wheat, Buckwheat, especially Tartary buckwheat, has been regarded as
but less variation was observed between three common and an effective medicinal herb in Chinese medicine for more than
three Tartary buckwheat genotypes. Buckwheat starch had 1000 years. Over 80 prescriptions utilizing buckwheat in the
higher peak gelatinization temperature (average Tp 68.7 C) treatment of many diseases have been described in Chinese
than standard wheat starch (Tp 63.7 C). The important past- traditional medicine books. In modern times, buckwheat
ing properties of buckwheat starch were higher hot paste vis- products are popular as functional or therapeutic foods in
cosity and cool paste viscosity, better resistance to shear China and some other countries. Regular consumption of
thinning, and less effect of NaCl on peak viscosity than for buckwheat foods may reduce the occurrence of hyperlipid-
wheat starch. Starch swelling volume is 27.428.0 ml for com- emia, obesity, and diabetes. A clinical trial in Beijing Tongren
mon buckwheat and 26.530.8 ml for Tartary buckwheat, Hospital showed that Tartary buckwheat flour had hypolipi-
compared to the reference (20.1 ml) for wheat starch. demic and hypoglycemic effects in treating hyperlipidemia and
Investigations on the nutritional properties of buckwheat diabetes (Table 6). Table 7 shows that health foods made from
starch showed that it has potential use in the design of foods Tartary buckwheat have positive effects in reducing cholesterol,
with lower glycemic index properties. The in vitro rate of starch blood lipid, blood sugar, urinary sugar, and other indices.
hydrolysis and resistant-starch formation in boiled buckwheat These therapeutic effects should be attributed to unique pro-
groats and in a series of buckwheat/wheat breads and noodles teins, resistant starch, and dietary fiber as well as vitamins,
(3070% buckwheat flour) were evaluated. The highest level of minerals, flavonoids, and phytosterols in buckwheat.
resistant starch (6% total starch basis) and the lowest starch Protein and indigestible carbohydrates (fiber and resistant
hydrolysis index were found in boiled buckwheat groats. The starch) of buckwheat (particularly Tartary buckwheat) can be
rate of in vitro amylolysis was much lower in all buckwheat used as potential functional food additives to treat conditions
products compared to the reference whole wheat bread or such as hypertension, obesity, and constipation. Buckwheat
noodles. Consumption of boiled buckwheat groats or bread protein extract could lower blood cholesterol level more
Table 6 Clinical effects of tartary buckwheat flour on hyperlipidemia and diabetes (hyperglycemia) after 1-month dietary treatment
Hypolipidemic effect
Serum triglyceride (mg dl1) (n 18) 310 241 233 173
Serum cholesterol (mg dl1) (n 13) 247 57 197 43
Hypoglycemic effect
Blood sugar (mg dl1) (n 23) 202 46 157 40
Urinary sugar (g 24 h1) (n 27) 44 43 20 22
a
n is number of subjects.
Source: Data from Lin RF, Zhou MD, Tao YR, Li JY, and Zhang ZW (eds.) (1992) Proceedings of the 5th International Symposium on Buckwheat, 2026 August, Taiyuan, China.
Beijing: Agricultural Publishing House; Corke H and Lin RF (eds.) (1998) Proceedings of the 1st International Conference on Asian Food Product Development Focus on Specialty
Grains and Grain Products, 610 September, Taiyuan, China. Beijing and New York: Science Press.
312 THE LEGUMES AND PSEUDOCEREALS | Buckwheat: Overview
Treatment Triglyceride (mg dl1) Cholesterol (mg dl1) Lipoprotein (mg dl1) Blood sugar (mg dl1) Urinary sugar (nmol 24 h1)
efficiently, compared with soy protein and casein. Sulfur- medicinal herb for Chinese consumption, has been widely
containing amino acids and glycine and arginine and lysine grown for a long time. A rich array of traditional buckwheat
are also considered to have important influence on this foods are found in various places of China, such as buckwheat
cholesterol-lowering effect. These amino acids can conduct wantuo and helao (or hele), buckwheat cats-ear noodles,
downregulation of hepatic LDL receptors and improve the buckwheat noodles (vermicelli), pancake, griddle cakes, jelly
removal of cholesterol and its esters in outlying tissues and noodles, and porridge. These foods are famous and popular in
thus reduce the serum cholesterol level. Many studies have China, particularly for their long history, rich nutritional pro-
further indicated that consumption of buckwheat protein can file, special flavor, various processing methods, and functional
cause suppression in body fat, constipation, mammary carci- food role in maintaining human health. For example, buck-
nogenesis, and colon carcinogenesis in rats and in the forma- wheat helao (hele), wantuo, and cats-ear noodles are well-
tion of cholesterol gallstones in hamsters. Some researchers known local foods in the northern region of China (Shanx
explain that the cholesterol-lowering, obesity-treating, and and Shaanxi provinces).
diabetes-controlling effects are also possibly associated with In Japan, buckwheat noodles called soba and buckwheat
other components (dietary fiber and resistant starch) in buck- roasted groats have long been very popular, while in Korea, a
wheat. The metabolites, formed during fermentation of resis- favorite jelly-type food called mook is also made from buck-
tant starch and other indigestible carbohydrates in the large wheat. In India, buckwheat flour is used for making chillare,
intestine, contribute to the maintenance of colon health and an unleavened bread fried with ghee, and also for a crisp food
also have beneficial effects on glucose metabolism. Moreover, a called pakora. Honey produced from common buckwheat is
novel protein isolated from Tartary buckwheat water-soluble prized in both China and India. In eastern Europe, buckwheat
extracts exhibited antiproliferative effect and antitumor activity. grain forms an important part of the traditional diet. Buck-
The flavonoids from buckwheat (especially Tartary buck- wheat groat porridge and meals and soup made from flour are
wheat) have been found to be effective in reducing blood still favorite traditional foods in Poland, Russia, and some
cholesterol levels, keeping capillaries and arteries strong and other eastern European countries. In the United States,
flexible, improving blood microcirculation, and protecting although buckwheat is mainly used for animal feed, buck-
blood vessels from rupturing and forming clots. These flavo- wheat cake was once a special breakfast item and buckwheat
noids also demonstrate antioxidant, antimicrobial, and anti- flour is also an ingredient of some pancake mixes.
inflammatory activities. Because Tartary buckwheat leaves and Modern buckwheat food products include buckwheat dried
stems contain high concentrations of rutin and other flavo- noodles or vermicelli, instant noodles, spaghetti or macaroni,
noids at flowering, extracts of buckwheat plants are helpful for biscuits, breads, cakes, pastry, breakfast cereals and snacks,
treating keratitis, while dried powdered buckwheat plants are vinegar, drinks, and beverages (Tartary buckwheat tea, buck-
sometimes used for curing external wounds and ulcers. wheat beer, and liquor). Most of these products, particularly
The vitamins and minerals (including Mg, Fe, and Se) in made from Tartary buckwheat, are popular healthy foods in
buckwheat can also play a crucial complementary role in the China. It is estimated that commercial food products made
previously mentioned therapeutic effects. Phytosterols found from buckwheat have produced over 10 billion RMB of value
in buckwheat grain, although at a low level, also showed in China. In addition, tender buckwheat shoots are used as
positive effects on blood cholesterol level. Other functions of fresh vegetables and are also processed into canned vegetables.
buckwheat sterols are waiting for further investigation, because
some phytosterols from other plant sources were found to have
strong pharmaceutical effects, for example, antivirus and anti-
tumor activities. Also, there is a report that the fagopyrins Buckwheat Processing and Development
isolated in buckwheat grain are an effective component for
the treatment of type 2 diabetes. However, there is still some Buckwheat grains are dehulled and milled into flour for food
dispute on this issue. processing. Buckwheat flour milling is similar to wheat dry
milling. Buckwheat flour yield ranges from 58% to 75%,
mainly depending on variety and milling techniques. Buck-
Buckwheat Food Products wheat flour contains similar levels of starch and protein to
wheat flour. However, the behavior of buckwheat flour is
Buckwheat products are recognized as functional food in some different from wheat flour, and it is difficult to make wheat
Asian countries. Buckwheat, as a nutritional staple crop and flour-like foods because there is no gluten. Therefore,
THE LEGUMES AND PSEUDOCEREALS | Buckwheat: Overview 313
buckwheat flour is usually mixed with wheat flour and other make pancakes, flakes, biscuits, bread, noodles, spaghetti or
flours before use. macaroni, and ready-to-eat breakfast cereals. The processing
Chinese researchers and food factories have mixed buck- of these foods can be conducted using ordinary baking,
wheat and wheat flour along with other flours to successfully extruding, or expansion methods. The amount of buckwheat
make buckwheat pastry, sandwich cake, dried noodles, instant flour added to other cereal flours is generally < 50%. For
noodles, and spaghetti (Tables 8 and 9). The processing steps instance, acceptable bread contained 25% buckwheat flour
for dried noodles are shown in Figure 1. Suitable levels of with a formulation that included vital gluten, whey, or sour
buckwheat flour are 10% for crisp pastry and cakes, milk. A recipe for bread with 10% Tartary buckwheat flour is
2040% for noodles, and 3050% for palatable spaghetti. shown in Table 8. It was also reported that the buckwheat
The cooking quality of spaghettis mixed from different ratios corn flakes were produced by extrusion processing. The for-
of buckwheat and wheat flours is shown in Table 9. Buck- mulations with an addition of 18%, 22%, and 33% buck-
wheat/wheat spaghettis generally have higher water absorp- wheat flour were used. The addition of 30% buckwheat flour
tion, higher cooking loss, and higher protein loss rate than gave the best organoleptic properties.
pure wheat spaghetti, because buckwheat flour contains much Although buckwheat is still cultivated as a minor crop in the
more albumins (water-soluble) and globulins (soluble in salt world, buckwheat product development has quickened since
water) than wheat flour. These commercial foods are produced the 1990s, particularly in China. To date, there has been an
by modern food-processing equipment. However, most tradi- emerging trend in the research and development of high value-
tional buckwheat foods are still handmade in the countryside added buckwheat products, especially Tartary buckwheat
and even cities in China. An example of the basic hand- products. Tartary buckwheat has high potential in the health
preparation method for buckwheat helao is shown in Figure 2. food market. Since Tartary buckwheat contains a high level of
In Japan, there are many noodle-making factories produc- flavonoids and other therapeutic components, some cosmetics
ing soba noodles. Buckwheat flour used for making soba is and medicine products have successfully been developed and
commonly mixed with 1050% wheat flour. These noodles marketed using buckwheat materials in China, such as cream,
are sold in precooked, boiled, dried, or instant stages, as well shampoo, lotion, toothpaste, and flavonoid capsules. How-
as being made for direct sale to buckwheat noodle restau- ever, many such new products have not been accepted well in
rants, small shops, and stands. In Europe and the United the marketplace, because of their high cost and erratic quality
States, buckwheat flour is mixed with other cereal flours to control.
Table 9 Cooking quality of spaghetti made from common buckwheat and wheat flour at different ratios
Buckwheat/wheat flour ratio Water absorption (%) Cooking loss of dry materials (%) Cooking in water Cooking in salt water
Source: Data from Wei YM (1995) Quality and Processing of Buckwheat Grain (in Chinese). Xian, China: World Publishing Corporation.
314 THE LEGUMES AND PSEUDOCEREALS | Buckwheat: Overview
Nowadays, buckwheat functional foods and related prod- Figure 2 Traditional method of buckwheat helao preparation.
ucts have attracted widespread interest and have become more
popular. Much more attention should be paid to the biological
functions, palatability, and acceptability of buckwheat prod-
into added-value components that would be attractive to
ucts. It is necessary to enhance development and commercial-
the health food industry.
ization of traditional buckwheat foods and to improve modern
buckwheat food-processing techniques.
Compare Chinese and Japanese buckwheat noodles in
depth, showing how different traditional products achieve
their required quality outcomes. Suggest new product ideas
for buckwheat-based noodles.
Exercises for Revision
Describe the composition of buckwheat seed in relation to See also: Carbohydrates: Starch: Chemistry; Food Grains:
physical properties of the food product (e.g., the effect of Intolerance, Allergy and Diseases: Celiac Disease; Proteins:
starch on noodle texture). The Protein Chemistry of Cereal Grains; The Basics: Taxonomic
Outline the health claims made for the various fractions Classification of Grain Species; The Legumes and Pseudocereals:
and components of the buckwheat seed. Amaranth: Overview; Pseudocereals: Overview.
Try to make buckwheat noodles, experimenting with the
effect of varying the proportion of wheat flour to buck-
wheat flour.
Further Reading
Bejosano FP and Corke H (1998) Amaranthus and buckwheat protein concentrate effects
Exercises for Readers to Explore the Topic Further on an emulsion-type meat product. Meat Science 50: 343353.
Corke H and Lin RF (eds.) (1998) Proceedings of the 1st International Conference on
Design a breeding program for the development of nutri- Asian Food Product Development Focus on Specialty Grains and Grain Products,
tionally improved buckwheat, suitable for fractionation Taiyuan, China, 610 September. Beijing and New York: Science Press.
THE LEGUMES AND PSEUDOCEREALS | Buckwheat: Overview 315
Guo XN, Zhu KX, Zhang H, and Yao HY (2010) Anti-tumor activity of a novel protein Tao CJ, Yuan P, Wang X, and Liao B (2014) Advances in the development of buckwheat
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Chickpea: Overview
EJ Knights and KB Hobson, New South Wales Department of Primary Industries, Tamworth, NSW, Australia
2016 Elsevier Ltd. All rights reserved.
This article is a revision of the previous edition article by E.J. Knights, volume 1, pp 280287, 2004, Elsevier Ltd.
Chickpea is an annual legume belonging to the Cicer genus. The genus Cicer contains 43 species, 34 perennial and 9
Seed shape separates the species into three main groups: annual. All annual species are diploids and have 16 chromo-
desi, kabuli, and pea. somes (2N 16). Among these, C. reticulatum is the most
Chickpea is a key dietary component in many countries closely related to the cultivated chickpea and the presumed
where vegetarian diets dominate or animal protein is too progenitor. Domestication occurred about 10 000 years ago
expensive. in the region of southeastern Turkey, part of the Fertile Cres-
Chickpea is grown mainly as a sole crop in a range of cent. Recent evidence suggests this occurred as a single event
rotations. due to the limited geographic distribution of the wild species
World production averages 10 Mha annually. populations and the very low genetic variation of the cultigen.
India produces more than 67% of the worlds annual About 40005000 years ago, the domesticated forms, initially
production. desi types, commenced a westward movement along estab-
lished trade routes toward the Mediterranean. Subsequent
spread to southern Europe and North Africa, including Ethio-
pia, occurred by land and/or sea routes. Differentiation of the
Learning Objective
kabuli type is thought to have occurred in the Mediterranean
region in comparatively recent times. The eastward movement
To achieve understanding of the agriculturally important
of desi types to India commenced about 4000 years ago,
species in both farming systems and its end uses
whereas kabuli types did not arrive there until about 300
years ago. (The prominence of Kabul on the Silk Road sug-
gests both an overland introduction and the origin of the word
Introduction kabuli.) In the sixteenth century, Spanish and Portuguese
travelers carried kabuli types to South and Central America
Chickpea (Cicer arietinum L.) is a herbaceous, annual legume. where they came to be known by their Spanish name gar-
Plants are typically short (2050 cm) with a semierect to erect banzo. Chickpea industries have comparatively recent histo-
growth habit and a variable number of primary and secondary ries in the United States (1950s), Australia (1970s), and
branches. An imparipinnate leaf type, producing 1118 small Canada (1990s).
leaflets, gives the foliage its distinct fernlike appearance
(Figure 1). Chickpea is a long day plant; however, time to
flowering is accelerated by increasing temperature. Flowering Production
is indeterminate with single (rarely double), small purple or
white flowers produced on racemes arising from the leaf axil. During the 1990s, world chickpea production has averaged
Pods have a characteristic oval to rhomboid shape and inflate 8 million tonnes (Mt) from 10.5 million hectares (Mha). In
rapidly after (self-) fertilization (Figure 1). All aerial plant parts the 2000s, this increased to 8.5 Mt from a similar area, with the
excepting flowers have a dense covering of fine hairs, which increased production coming from increased yield. Crop area
secrete a mixture of organic acids. Roots are colonized by for the past 3 years (20112013) has averaged 13 Mha, which
Mesorhizobium ciceri, a nitrogen-fixing bacterium specific to is the highest 3-year average production (Figure 3). Histori-
the genus Cicer. Nodules formed by these bacteria vary in cally, Asia has been the major chickpea producer. For years
size, the largest approaching 3 cm in diameter. The seed is 20092013, it accounted for 85.1% of all production, followed
highly variable in appearance but invariably possesses the by Australia (5.3%), Africa (5.0%), Americas (3.8%), and
characteristic chickpea beak protruding over the embryo. Europe (0.9%). Desi types contribute 70% of production
Seed shape, color, and, to a lesser extent, size separate the worldwide. Chickpea is cropped in approximately 80 coun-
species into three main groups: desi, kabuli, and pea tries. India produces two-thirds of the world crop (Table 1),
(Figure 2). and after a steady decline in the 1990s, production has been
Desi (derived from the Hindi and Urdu word for native or increasing in recent years. A similar trend has occurred in
local) seeds have an angular shape and thick, colored (mostly Pakistan, the worlds second largest producer. In European
brown) seed coat. In contrast, kabuli seeds have a more countries, chickpea production has fallen dramatically, by
rounded shape and thin, white to cream-beige seed coat. 6097% since the 1960s. On the other hand, production has
There is considerable overlap between these two seed types in risen significantly in some countries. In Australia, the worlds
seed size; however, desi seeds are generally smaller (weighing fourth largest producer, chickpea area increased rapidly during
80350 mg) than kabuli seeds (100750 mg). The pea-type the 2000s as the demand increased for cereal break crops and
seed has little commercial significance. alternatives were sought for nitrogen fertilizer. Expanded chickpea
16000 1000
area
production
yield 900
800
Area (x1000 ha)/production (x1000 t)
12000
700
600
400
300
4000
200
100
0 0
1962
1965
1968
1971
1974
1977
1980
1983
1986
1989
1992
1995
1998
2001
2004
2007
2010
2013
Figure 3 World area, production and yield of chickpea, 19612013. Each datum is the mean for a 3-year period. Source: FAO.
Table 1 Mean area, production, and yield of chickpea in major chickpea-producing countries (20092013)
implements, although some regions are introducing mechani- mainly on growing season rainfall. Winter sowing occurs in
zed processes. Egypt and Sudan. Sowing is usually delayed until late February
to May to minimize losses from cold and/or ascochyta blight.
Rapidly increasing temperatures and moisture stress in summer
West Asia, North Africa, and Southern Europe hasten maturity and limit crop duration to 90120 days.
Chickpea is generally confined to areas having an annual
Production in this region is almost exclusively of kabuli types. rainfall above 400 mm. It is grown as a sole crop in rotation
The crop is grown at high altitudes in west Asia (Afghanistan, with wheat (bread or durum) or barley; a fallow phase or
Iran, and Turkey) but at comparatively lower altitudes in the summer or forage crop may be included in drier areas. There
Middle East (Syria, Yemen, Jordan, Israel, and Iraq), North is some mechanization of production, particularly sowing and
Africa (Egypt, Sudan, Algeria, Tunisia, and Morocco), and south- weeding, but harvest is still largely manual. High labor costs
ern Europe (Greece, Italy, Spain, and Portugal). It is grown as a have contributed to the decline in chickpea production in parts
spring crop in this Mediterranean-type environment, relying of the region, especially in Europe.
THE LEGUMES AND PSEUDOCEREALS | Chickpea: Overview 319
East Africa (Ethiopia, Eritrea, Kenya, Tanzania, Malawi, southeast and southwest, production in the southeast region
and Uganda) is mainly kabuli type. Chickpea is grown as a winter crop,
sown from May to July and harvested from October to January.
Mostly desi and some kabuli chickpeas are grown in this Most crops are produced under rainfed conditions; incrop
region, usually at high altitudes (up to 2400 m). The area of rainfall is a major yield determinant in most regions, especially
kabuli is increasing in some countries with the introduction of in the southeast and southwest. As in North America, produc-
improved varieties and is up to one-third of the crop in Ethi- tion is highly mechanized. Control of ascochyta blight, the
opia. Crops are mainly sown as pure stands in rotation with major industry problem in most regions, relies heavily on
wheat, barley, and teff, but to a lesser extent as mixtures with ground rig application of foliar fungicides. In northeast Aus-
sorghum, safflower, and maize. Crop growth relies mainly on tralia, Phytophthora root rot is a common biotic constraint,
residual soil moisture. Sowing follows the wet season, from particularly in seasons conducive to the disease.
July to early September in Ethiopia and from February to April
further south. Inputs of fertilizers and pesticides are minimal,
and although there is some use of tractors in sowing, harvest Production Constraints
and threshing remain manual operations.
There is a large discrepancy between the yield in farmers fields
and that commonly obtained under experimental conditions.
The low yields generally obtained by farmers reflect a plethora
Central and Southern America (Mexico and Argentina)
of problems that beset the crop. The major production con-
straints and their impacts are described below.
In Mexico, desi chickpeas are produced under rainfed condi-
tions in the west-central parts of the country and kabuli types
under irrigation in the drier northwest. Chickpea is grown as a
winter crop following maize in the west-central region and
Physical Constraints
following soybean or sesame in the northwest. Sowing is from Drought and Heat
October to December and harvest from March to May. There
Chickpea is mostly grown in low rainfall environments. Inad-
has been some recent transition to mechanized production,
equate soil moisture is therefore a critical production con-
particularly for kabuli types. Production in Argentina has
straint and may occur throughout the growth cycle. Winter/
recently increased to 40 000 ha of kabuli type. It is sown
spring rains in west Asia, or soil moisture remaining from the
from April to July.
wet season in the Indian subcontinent and East Africa, may not
be sufficient for farmers to sow; for many regions, variability in
main season rainfall explains much of the year-to-year varia-
North America (Canada and the United States) tion in chickpea area. Receding soil moisture, particularly in
low rainfall areas in India and Pakistan, may also be inade-
The North American chickpea industry began in coastal dis- quate to sustain seedlings. Intermittent moisture stress of
tricts of California where spring-sown kabuli crops were grown established crops can result from breaks in winter and early
on residual soil moisture. Later, winter crops were grown under spring rainfall, but a more serious problem is terminal mois-
irrigation in the drier San Joaquin Valley. During the 1980s, ture stress suffered by spring-sown crops (and winter crops in
both desi and kabuli chickpeas were introduced to the Palouse the Indian subcontinent) as podding coincides with rapidly
region of northwestern United States and, in the following declining soil moisture. Terminal drought is a major constraint
decade, to the brown and dark brown soil zones of Saskatch- in an estimated 80% of the global chickpea area.
ewan and Alberta in Canada. Current production areas are the Heat stress (temperature >35 C) during the reproductive
Pacific Northwest in the United States and the province of phase has become a serious constraint as chickpea moves from
Saskatchewan in Canada. The threat of cold and ascochyta cooler long-season environments to warmer short-season envi-
blight forces chickpeas to be grown as a spring crop in both ronments, an increased area under later sowing, and expected
these regions. Sowing commences in May and harvest, partic- increases in temperatures due to increased climate variability.
ularly in Canada, needs to be completed by September to avoid
freezing injury to immature seeds. In the United States and
Canada, all stages of chickpea production, including seed Cold
inoculation; sowing; weed, disease, and insect control; harvest; Low temperatures affect the plant in two main ways: tissue
and seed drying, are highly mechanized. In most cases, the damage through disruption of cell membranes and abortion of
machinery and processes have been modified from those flowers. Sensitivity to low temperatures (<5 C) is one reason
used in cereal production. for spring sowing in higher elevation areas of west Asia. How-
ever, genotypes have now been identified that can survive tem-
peratures as low as 25 C. Winter sowing of these cold-tolerant
Australia genotypes can increase yield potential by 70% through
increased biomass production and improved water-use effi-
National production (90%) is mainly desi type and this ciency. Where the crop is sown in autumn/winter, such as in
predominantly occurs in the summer rainfall-dominant northern India, Pakistan, and Australia, chilling damage in the
regions in the northeast of Australia. In the winter early flowering phase narrows the window for pod setting.
rainfall-dominant Mediterranean-type environments in the Sowing is normally delayed so that commencement of
320 THE LEGUMES AND PSEUDOCEREALS | Chickpea: Overview
Hostile Soils
A range of soil chemical and physical problems limit chickpea
growth. The most important of these is salinity, since chickpea is
comparatively intolerant of the ionic imbalances characteristic of
saline soils. Sodic soils, in which high levels of exchangeable
sodium are associated with increased bulk density, low water
infiltration, and poor aeration, are also common in northern
India and in other regions where soil pH is very high. Retarded
root growth and poor nodulation inhibit chickpea growth and
productivity under such conditions. Other locally important soil
factors are acidity (poor nodulation), deficiencies of iron and
zinc, and either a deficiency or toxicity of boron.
Ascochyta Blight
Fusarium Wilt
Ascochyta blight is the most important disease of chickpea
worldwide and has been recorded in nearly all producing coun- Fusarium wilt is the other major disease of chickpea and, like
tries. Recent epidemics in the United States (1980s) and Austra- ascochyta blight, causes economic damage in a large number of
lia (1990s) caused major industry disruptions, reflected by a producing countries. It is a soil- and seed-borne disease caused
sharp decline in the area sown. Elsewhere, for example, in west by different races of the fungus Fusarium oxysporum f.sp.ciceris
Asia and the Mediterranean region, the threat of ascochyta blight Matuo and K. Sato. The races are differentiated on their ability
causes sowing to be delayed, thereby significantly reducing yield to incite specific reactions causing two types of symptoms:
potential. The causal agent of ascochyta blight is Ascochyta rabiei yellowing syndrome and wilting syndrome. The wilting syn-
(Passerini) Labrousse (syn. Phoma rabiei and teleomorph Didy- drome is more destructive and economically more important
mella rabiei (Kovachevski)), a fungus spread by infected seed and than yellowing syndrome. Fusarium wilt is more frequently
crop residues, long-range dispersal of sexually produced asco- expressed under warm, dry growing conditions and is therefore
spores, or, within the crop, short-range dispersal of asexually a greater problem in the Indian subcontinent and in regions
produced pycnidiospores. Lesions are formed on leaves, stems, where the crop is spring-sown. Fungicidal seed dressings pro-
pods (Figure 4), and seeds. The disease progresses quickly under vide protection against seed-borne infection, but the longevity
favorable wet conditions and can cause total crop failure. Asco- of the pathogen in soil makes host resistance the most success-
chyta can be controlled by an integrated disease management ful way of combating the disease.
approach based on host resistance and, depending on location,
removal of infected plant debris, delayed sowing, seed dressing, Root and Collar Rots
and foliar fungicides.
A number of fungal root or collar diseases have the potential to
inflict significant yield losses. Dry root rot (Rhizoctonia batati-
cola (Taubenhaus) Butler) is the most important of these,
Other Foliar Diseases
especially on the Indian subcontinent. The disease generally
Botrytis gray mold (Botrytis cinerea Pres.) is the most important presents as a sudden drying of scattered plants at the podding
of the foliar diseases after ascochyta. The pathogen has a very stage and is favored by hot, dry conditions. Other diseases
broad host range; therefore, the disease is widely distributed. include wet root rot (Rhizoctonia solani Kuhn), black root rot
Most damage occurs under high humidity and generally at (Fusarium solani (Martius) Saccardo), collar rot (Sclerotium rolf-
temperatures higher than optimum for ascochyta. Botrytis sii Saccardo), and phytophthora root rot (Phytophthora medica-
has contributed to the decline of the Bangladesh chickpea ginis Hansen and Maxwell). The latter has been recorded in a
industry and has also caused significant or total crop losses number of countries, but economically significant damage has
in the higher rainfall zones of northeastern India and in Aus- only been observed in northeastern Australia where it is the
tralia. The following diseases are also regarded as locally sig- major production problem.
nificant: sclerotinia stem rot (Sclerotinia sclerotiorum (Lib.)
de Bary), phoma blight (Phoma medicaginis Malbr. and
Viruses
Roumeguere), alternaria blight (Alternaria alternata (Fries:
Fries) Keissler), stemphylium blight (Stemphylium sarciniforme Considerable crop losses have been attributed to virus disease,
(Cavara) Wiltshire), and rust (Uromyces ciceris-arietini Jaczewski especially in India, Pakistan, Iran, the United States, and
in Boyer & Jaczewski). Australia. In most cases, a complex of viruses has been
THE LEGUMES AND PSEUDOCEREALS | Chickpea: Overview 321
implicated. Aphids, particularly Aphis craccivora Koch, are almost cultivation. Options for pre- and postemergent control of
always the vector responsible for disease transmission. In India, both grass and broadleaf weeds are now available and used
chickpea stunt is the name given to a syndrome characterized by extensively in Australia, the United States, and Canada. Grass
foliage discoloration (red in desi and yellow in kabuli), stunting, herbicides are very effective, have no toxicity to chickpea, and
phloem browning, and plant death. The disease has been contribute to the productivity of ensuing cereal crops by elim-
ascribed to the leafhopper-transmitted chickpea chlorotic inating graminaceous diseases that can otherwise survive on
dwarf virus and some aphid-transmitted luteoviruses, including grass weeds. The development of herbicide-tolerant varieties
bean leafroll luteovirus. Other viruses known to cause disease will also provide further weed control options.
are alfalfa mosaic virus, cucumber mosaic virus, beet western
yellows virus, and subterranean clover red leaf virus.
Utilization
Nematodes Nutritional Status
Nematodes reduce yield by colonizing and damaging roots, Desi chickpea seeds are composed of the embryo (1.5%), seed
reducing the efficiency of nitrogen fixation and in some cases coat (15.5%), and cotyledons (83%); the seed coat fraction of
acting synergistically with fungal diseases such as fusarium kabuli seeds is much lower (6.5%), resulting in a higher cotyle-
wilt. Many nematode species are known to infect chickpea don fraction (92%). Protein averages 23% and is deficient in the
although only three are considered economically important. sulfur-containing amino acids methionine and cystine. Carbo-
Root-knot nematodes (Meloidogyne spp.) cause abundant for- hydrate, the main constituent, averages 63.5% of total seed and
mation of galls, whereas both cyst nematode (Heterodera ciceri is mainly starch. Soluble sugars comprise 10% of the starch
Vovlas, Greco & di Vito) and root-lesion nematodes (mainly and include oligosaccharides such as stachyose and raffinose
Pratylenchus thornei Sher and Allenand and P. neglectus that cannot be broken down by human digestive enzymes.
(Rensch) Chitwood and Oteifa) damage root cellular tissue Gases produced by bacterial degradation of these oligosaccha-
causing necrotic regions. Root-knot nematodes and root-lesion rides cause flatulence and intestinal discomfit. Oils comprise
nematodes have an extensive host range and are widely dis- 5% of the seed but are well below commercially extractable
tributed; cyst nematode has a very limited host range and its levels. Chickpea is generally low in antinutritional factors such
distribution is confined to the Middle East. as protease and amylase inhibitors, whose activities are further
reduced by cooking, and protein-binding phenolics, which are
largely removed with processing.
Insects
The small number of insect pests causing economic damage to
Uses and Processing
chickpea is attributed to the antibiotic effect of the plants
acidic (mainly malic acid) exudate. The most widespread and Chickpea is a key dietary component in many countries where
damaging pest is the pod borer (Helicoverpa spp.). Various animal protein is too expensive or in other countries where the
larval instars feed on leaves, flowers, and pods. Most yield crop has a long history of cultivation and consumption. The
loss is caused when late instar larvae enter the pod and partly complementarity of pulse and cereal proteins has given rise to
or fully consume the developing seed. There is little effective many traditional dishes in which chickpea is combined with
host resistance within chickpea so control is reliant on a com- wheat, rice, or some locally important cereal, such as teff, to
bination of chemical and biopesticides (such as nuclear poly- provide the bulk of dietary protein and calories. The use of
hedrosis virus). Genetic transformation has been used to chickpea as human food is hugely varied, reflecting both the
develop chickpea containing the toxins from the bacterium Bt crops antiquity and its broad cultural base (Table 2). The
and it is currently undergoing field testing. mature seed, either whole or processed, accounts for nearly
The other major field pest is leaf miner (Liriomyza cicerina all consumption. However, there are specialty uses for the
(Rondani)) whose distribution is confined to the Mediterra- green, immature plant. Fully podded plants are commonly
nean region. Larvae tunnel through the leaf parenchyma caus- sold in street markets in India and the Middle East. The seeds
ing loss of photosynthetic tissue and premature leaf drop. are either roasted in the pod, consumed raw as a snack, or
Infestation of stored seed is mainly by small bruchid beetles boiled; fresh leaves are also used as a vegetable. Ripe seeds (desi
(Callosobruchus spp.). Larvae emerge from eggs laid on the and kabuli) are presoaked and boiled, with or without the
(stored) seed surface and tunnel beneath the seed coat where addition of vegetables or meat, to produce a range of tradi-
they feed and pupate. Adults emerge from the seed, ready for tional dishes. Kabuli seeds are also canned after cooking, but a
the next life cycle and leaving a substantial cavity. more important use is as hommos bitihneh (pureed and mixed
with oil). Both desi and kabuli seeds are also roasted. This
process generally requires a quick soaking followed by high-
Weeds
temperature treatment (e.g., at 250 C) for 23 min in pre-
Chickpeas slow early growth and low total biomass predispose heated sand or an oven. Variations to the soaking, roasting,
the crop to severe weed competition. In most countries, weed and final processing (e.g., decortication, polishing, or sugar
control is done by manual weeding or cultivation by animal- coating) give this snack food its regional characteristics.
or tractor-drawn implements. These practices are feasible Most desi seeds are milled to produce dhal, a process that
where crop area is small, although yield potential is often involves removal of the seed coat and cleaving of the cotyle-
compromised where wide rows are employed to facilitate dons (Figure 5). Milling of chickpeas is a major industry in the
322 THE LEGUMES AND PSEUDOCEREALS | Chickpea: Overview
What is the most important disease of chickpea worldwide? Basics: Taxonomic Classification of Grain Species; The Grain Chain:
What are the main abiotic constraints of chickpea? The Route from Genes to Grain-Based Products; The Grain Crops: An
What is the average protein content of the grain? Overview; The Legumes and Pseudocereals: Grain Legumes and
Which two wild relatives of chickpea are being exploited by Their Dietary Impact: Overview.
breeding programs?
Further Reading
Exercises for Readers to Explore the Topic Further
Gaur PM, Jukanti AK, and Varshney RK (2012) Impact of genomic technologies on
The Cicer genus contains nine annual species. How diverse chickpea breeding strategies. Agron 2: 199221.
are they and what is their potential for use in chickpea Knight R (ed.) (2000) Linking Research and Marketing Opportunities for Pulses in the
21st Century. Dordrecht: Kluwer Academic.
improvement programs?
Loss S, Brandon N, and Siddique KHM (eds.) (1998) The Chickpea Book: A Technical
What are the products made from besan (chickpea flour) in Guide to Chickpea Production. Western Australia: Department of Agriculture and
the subcontinent? Food.
Maiti R and Wesche-Ebeling P (eds.) (2001) Advances in Chickpea Science. Enfield:
Science Publishers.
Saxena MC and Singh KB (eds.) (1987) The Chickpea. Wallingford: CAB International.
See also: Agronomy of Grain Growing: Chickpea: Agronomy;
Saxena NP (1984) Chickpea. In: Goldsworthy PR and Fisher NM (eds.) The Physiology
Appendix 1: Nutrient-Composition Tables for Grains and for Grain- of Tropical Field Crops, pp. 419452. Chichester: Wiley.
Based Products; Appendix 2: Lists of Standardized Testing Methods Sharma RN, Shrivastava GK, Rathore AL, Sharma ML, and Khan MA (eds.) (2003)
for the Analysis of Grain and Grain-Based Foods; Food Grains and Chickpea Research for the Millennium. Raipur: Indira Gandhi Agricultural
the Consumer: Consumer Trends in Grain Consumption; Cultural University.
Singh KB and Saxena MC (eds.) (1999) The Tropical Agriculturalist: Chickpeas.
Differences in Processing and Consumption; Fortification of Grain- London: MacMillan.
Based Foods; Genetically Modified Grains and the Consumer; Grains Smithson JB, Thompson JA, and Summerfield RJ (1985) Chickpea (Cicer arietinum L.).
and Health; Food Grains and Well-being: Functional Foods: In: Summerfield RJ and Roberts EH (eds.) Grain Legume Crops, pp. 312390.
Overview; Food-quality Testing: The Application of Sensory Science London: Collins.
van der Maesen LJG (1972) Cicer L, A Monograph of the Genus, with Special Reference
to the Evaluation of Grain-Based Foods; Grain Composition and
to the Chickpea (Cicer arietinum L.), Its Ecology and Cultivation. Wageningen: H
Analysis: Standardized Test Methods for Grains and Grain-Based Veenman and Zonen NV.
Products; The Composition of Food Grains and Grain-Based Products; Van Rheenen HA and Saxena MC (eds.) (1990) Chickpea in the Nineties. Patancheru:
Units of Grain Science and Trade: Equivalence between the US, ICRISAT.
Chinese, and Metric Units; Grains Around the World: Grain Varshney RK, Song C, Saxena RK, et al. (2013) Draft genome sequence of chickpea
(Cicer arietinum) provides a resource for trait improvement. Nature Biotechnology
Production and Consumption: Europe; Grain Production and 31: 240246.
Consumption in Oceania: Australia and Pacific Countries; Grain Yadav SS, Redden R, Chen W, and Sharma B (eds.) (2007) Chickpea Breeding and
Production and Consumption: Overview; Grain Production and Management. Wallingford: CAB International.
Consumption: South America; Grain Production and Consumption:
Africa; Production and Consumption of Grains: India; Non-food
Products from Grains: Cereal Grains as Animal Feed; Proteins: Relevant Websites
Nitrogen in Grain Production Systems; Nitrogen Metabolism; Protein
Synthesis and Deposition; The Protein Chemistry of Dicotyledonous http://www.ars.usda.gov R&D undertaken on chickpea in the USA.
http://www.cgiar.org Consultative Group on International Agricultural Research.
Grains; Scientific Organizations Related to Grains: Research http://www.icarda.org Information on kabuli chickpeas, mainly in Mediterranean-type
Organizations of the World: Asia-Pacific, CentralSouth America, and environments.
AfricaMiddle East; Research Organizations of the World: CGIAR; The http://www.icrisat.org Information on chickpeas (mainly desi) in semiarid areas.
Pea: Overview
TN Khan, University of Western Australia, Crawley, WA, Australia
A Meldrum, Pulse Australia, Perth, WA, Australia
JS Croser, University of Western Australia, Crawley, WA, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights be used as a green manure crop. Dry peas are used whole in
confectionery and snacks or milled to produce split peas for
Pea is the second most important grain legume crop in the making soups, dal (a curried soup-like preparation widely
world and is widely adapted to cooler climates. The major used in the Indian subcontinent), flour, and canned products
pea producers are Canada, parts of Europe, the United such as mushy peas. Dry peas are also used in the feed industry,
States, and Australia. particularly in the diet of pigs and poultry due to their amino
Pea is used as human food, both as a grain crop and as a acid balance.
popular vegetable. It is also used for animal feed. Pea seed is World pea production has declined by 34% since 1993 to
high in protein and a good source of vitamins such as just under 100 million tonnes (Figure 1). Competition for
riboflavin and niacin. land from cereals and oilseed crops, which usually deliver
Pea is a climbing plant with a weak stem where tendrils higher profits, is the main reason. There are over 6 million
allow it to trail if support is available. Over the last few hectares (Mha) of field pea grown in the world today (Table 1).
decades, the afila gene that modifies leaflets into tendrils This figure has been stable for the last 10 years. As of early
has been exploited to develop pea varieties with improved 2010s, Canada has the largest area under pea cultivation, fol-
standing ability. lowed in order by the Russian Federation, China, India, and
The genetics of the pea plant have been studied in great France (Table 1). Canada has maintained its pea cultivation
detail and dense linkage maps are now available. area at over 1.0 Mha since the 1990s, reaching a high of
Biotechnology techniques including genetic transforma- 1.58 Mha in 2008. Canada leads total production with more
tion, embryogenesis, and fast-track technologies are well than 3.5 million tonnes produced in 2008 and 3.3 million
established for peas. tonnes (Mt) of crop produced in 2012, followed by the Russian
Pea production and consumption are likely to remain sta- Federation with a production of 1.6 Mt. Sixty-nine percent of
ble with increases likely to be proportional to human pop- the total world production comes from the five highest field
ulation increases. pea-producing nations (Table 1).
The yield per hectare varies widely between growing regions
(Table 1). Among the largest producers, France has the highest
Learning Objective yields per hectare of over 4 t against the world average of 1.62 t,
while the leading producer Canadas average yield is 2 t.
To understand pea as a grain legume crop in human food Yields generally reflect the degree of industrialization of field
and animal feed and its significance as a legume in crop pea production and the inherent variability in the climate.
rotations. Average yields in India are below 1 t ha1. In most countries
where it is produced, pea seed is retained for domestic con-
sumption, as are other pulse crops. The United States and
Introduction Australia are the other major exporters of field pea.
14
10
0
1992 1994 1996 1998 2000 2002 2004 2006 2008 2010 2012 2014
Source: FAO Stats
Figure 1 Global dry pea area, production, and yield from 2003 to 2012. Source: FAO Statistic.
Table 1 Area, production, and yield figures for the five highest producers as compared with the world total
Aa Pb Yc A P Y A P Y A P Y A P Y A P Y
2002 0.98 1.28 1.31 0.76 1.27 1.67 0.95 1.50 1.58 0.67 0.61 0.91 0.34 1.66 4.92 6.02 9.63 1.60
2003 1.15 1.93 1.68 0.72 1.05 1.47 0.94 1.40 1.49 0.66 0.59 0.89 0.37 1.62 4.41 6.15 9.89 1.61
2004 1.24 3.10 2.49 0.72 1.24 1.72 0.88 1.06 1.21 0.71 0.73 1.02 0.36 1.68 4.71 6.34 11.74 1.85
2005 1.27 2.99 2.36 0.71 1.13 1.58 1.00 1.16 1.16 0.79 0.79 0.99 0.32 1.33 4.21 6.57 11.29 1.72
2006 1.23 2.52 2.05 0.71 1.15 1.62 0.90 1.01 1.13 0.78 0.71 0.91 0.24 1.02 4.21 6.39 9.82 1.54
2007 1.44 2.93 2.03 0.62 0.86 1.39 0.98 1.08 1.10 0.81 0.85 1.05 0.16 0.59 3.61 6.32 9.37 1.48
2008 1.58 3.57 2.26 0.64 1.26 1.98 0.93 1.10 1.19 0.68 0.75 1.10 0.10 0.45 4.50 6.11 10.07 1.65
2009 1.49 3.38 2.27 0.77 1.35 1.75 0.88 0.96 1.10 0.72 0.66 0.92 0.12 0.55 4.72 6.38 10.46 1.64
2010 1.39 3.02 2.17 0.82 1.19 1.45 0.87 0.91 1.05 0.76 0.67 0.88 0.24 1.07 4.44 6.58 10.31 1.57
2011 0.91 2.12 2.31 1.11 2.02 1.82 0.87 1.19 1.36 0.73 0.59 0.82 0.19 0.67 3.58 6.14 9.73 1.58
2012 1.31 3.34 2.14 1.16 1.66 1.43 0.93 1.11 1.20 0.74 0.63 0.85 0.14 0.57 4.05 6.33 9.86 1.56
a
Area: ha 106.
b
Production: t 106.
c
Yield: t ha1.
http://apps.fao.org (February 2015).
identified as a source of useful variation for disease and insect with red cotyledon color are also known. The testa color largely
pest resistance and drought tolerance. determines the seed appearance, which is of commercial sig-
The plant, a spreading or tendril-climbing herb, is typically nificance. Traders recognize five field pea seed types (Figure 4).
30150 cm tall (Figure 2). Papilionaceous flowers may be The round white (also known as yellow), as the name suggests,
white or purple or various shades of purple including pink. It is round with white to cream testae and yellow cotyledons. The
is predominantly self-pollinated with cross-pollination rarely dun type is variously dimpled with greenish brown (dun)
exceeding 1%. The stem is typically weak, leading to lodging as testae and yellow cotyledons. The maple type is round with a
the plant gains biomass. The compound leaves generally have brown testa that is mottled with light-colored spots giving it a
one or more leaflets modified into tendrils, although variants marbling effect and yellow cotyledons. The blue type (also
are known that have no tendrils. Of significance are the forms known as green) has a translucent testa with green cotyledons
where an entire leaf modifies to form tendrils (Figure 3) as this giving the seed a bluish hue. The marrowfat is typically a large
may help the crop to stand better by intertwining with plants. seed (>280 mg), slightly dimpled and flattened, with blue/
Greater attention to this character will be given while discuss- green testae and green cotyledons.
ing genetic improvement. White-flowered plants give rise to The diploid chromosome number is 14. Chromosomal
white/cream or translucent testae, whereas colored flowers interchanges can be common. Considerable attention has
produce variously colored testae. Cotyledon color ranges been given to the genetics of pea following the pioneering
from various shades of yellow to green, although genotypes experiments of Mendel in the nineteenth century. Several
326 THE LEGUMES AND PSEUDOCEREALS | Pea: Overview
thousand mutants have been identified and those conforming Nutritional Value, Processing, and Utilization
to Mendelian genetics were placed on a linkage map as early as
the middle of the last century. A book assembling a compre- The field pea seed is a rich source of protein, carbohydrate, and
hensive account of pea genome was published toward the end some minerals, although the nutritional content of the seed
of the last century. Since then, rapid advances have been made varies with the environment and genetic factors. By far, the
and are covered in the section Biotechnology and In vitro highest proportion of nutrient value in field pea is in the coty-
Culture Techniques for Field Pea Improvement. Updated ledons, with the embryo and seed coat contributing less than
maps, now including biochemical and molecular markers, 10% to nutritional value. The protein content is typically about
have regularly appeared in Pisum Genetics. 22% but ranges widely, depending on the genotype and growing
conditions (Tables 2 and 3). Although sulfur-containing amino
acid is low, it compares favorably with other grain legumes.
About 60% of the carbohydrate content of the seed is made up
of sucrose and oligosaccharides, starch, and crude fiber. The fat
content is low and the seeds are a good source of vitamins such
as thiamine, riboflavin, and niacin, although considerable loss
of vitamins may occur in processing.
Antinutritional factors are present in quantities lower than
in other grain legumes; the field pea has the lowest trypsin
inhibitor activity among grain legumes commonly used as
food in India. The flatulence activity resulting from oligosac-
charides of the raffinose family is also relatively low, as are
phytic acid and saponins. Lipoxygenase activity that causes
deteriorative changes during the processing and storage of
field peas is high, but these enzymes can be inactivated via
heat treatment.
The dry pea seed is processed via soaking, germination,
milling, cooking, roasting, or fermentation. In Western diets,
the seed is mainly used in soups or in mushy pea preparations.
In South Asian diets, it has varied uses: as dal (spicy soup-like
preparation in the Indian subcontinent), whole pea boiled as a
snack food, as sweet and savory snacks made from pea flour, or
as a supplement to wheat flour to make nutritious breads. In
Figure 2 P. sativum normal leaf, white-flowered type showing pod and addition, peas can be processed into protein, starch, and fiber
seed development. fractions. These products can then be used in baked goods,
Figure 3 P. sativum types: (a) semileafless, colored flowered type, and (b) normal leaf, white-flowered type.
THE LEGUMES AND PSEUDOCEREALS | Pea: Overview 327
Figure 4 The five seed types of P. sativum: top left, yellow; top right, marrowfat; center, dun; bottom left, green; and bottom right, maple.
Table 2 Nutritional content of field pea seed measured from a Table 3 Amino acid content of field pea seed measured from a
representative number of samples representative number of samples
serious abiotic constraints to field pea productivity. No true Genetic relationships among Pisum accessions have also
resistance to waterlogging or frost has been reported so far. been assessed using genomic techniques. Molecular analysis
There are reports regarding cold tolerance, but whether it of pea collections has shown that although substantial varia-
relates to avoiding damage during frost is not known. There tion is present within the cultivated Pisum gene pool, wild
are two aspects of freezing tolerance. In higher latitudes, such Pisum species offer the possibility to incorporate novel traits
as Europe, freezing tolerance is important during the vegetative that may have been inadvertently eliminated from the culti-
phase in the winter-sown crop, whereas in subtropical and vated gene pool via selection. Association mapping analysis of
Mediterranean climates, frost tolerance during the reproduc- diverse pea germplasm will result in the identification of genetic
tive stage is more important. A deeper root system has been variation related to desirable agronomic traits. The availability
reported to increase tolerance to moisture stress. However, of high-throughput omics methodologies will translate in the
osmoregulation/osmotic adjustment is a more measurable future into the development of novel, highly accurate selective
character, and it has shown good correlation with yield under breeding tools. Interspecific hybridization to transfer useful
water stress in studies done in Spain and Western Australia. traits from P. fulvum and wild-type P. sativum accessions such
Studies in Western Australia have shown that the most adapted as ssp. P. elatius to cultivated P. sativum would also benefit from
field pea variety showed one of the highest levels of osmotic techniques such as MAS and following QTLs. Targeted applica-
adjustment. tion of existing molecular techniques and the advent of new
techniques such as the analysis of gene function through func-
tional genomics will yield rapid gains in the understanding of
Biotechnology and In Vitro Culture Techniques field pea genetics.
for Field Pea Improvement
Molecular Markers and Linkage Mapping In Vitro Mediated Interspecific and Intergeneric Hybridization
Pea was the model species used by Austrian monk Gregor Men- Targeted utilization of wild relatives for adaptation to climate
del in 1866 to determine the laws of inheritance in plant genet- change will almost certainly be an urgent priority this century.
ics. Unfortunately, the large size of the pea genome, coupled Legume crops and pastures are estimated to use 3560% less
with its repetitive nature, has constrained the achievement of a fossil energy than N-fertilized cereal or grasslands. The reduced
complete genomic sequence. However, comprehensive genomic energy use was primarily attributed to reduced N fertilizer use
and postgenomic resources have been developed for peas in the legume and subsequent nonlegume crops. Interspecific
including BAC libraries, transcriptome and proteome data sets, hybridization is the crossing of two species from the same
and several types of molecular marker sets. Molecular markers genus. This allows the exploitation of useful genes from wild,
offer the potential to advance plant breeding by analysis of unimproved species for the benefit of the cultivated species.
genetic variation, unequivocal identification of genotypes and The Pisum species are diploid self-pollinators sharing a similar
true hybrids, purity testing of breeding lines, genome mapping, karyotype. P. fulvum is the only separate wild species and a
and marker-assisted selection (MAS). In field peas, classical useful source of disease and insect resistance. Fertile hybrids of
physiological, biochemical, and hybridization-based markers P. sativum P. fulvum have been produced via conventional
have been joined by polymerase chain reaction-generated unidirectional crossing with P. fulvum as the pollen donor.
markers. The polymorphic markers identified using these tech- Intergeneric crossing is another technique for improving
niques have been used to construct dense linkage maps of the genetic diversity within species. It is possible to regenerate
field pea genome, indicating the position and relative genetic plants from protoplasts in field pea via organogenesis and
distances between markers along chromosomes. somatic embryogenesis. Somatic protoplast fusion, whereby
The recent availability of full genome sequences for three genomes from different genera are combined without pollina-
other legume species has provided opportunities for genome- tion through protoplast fusion, has been attempted for inter-
wide comparison. This approach has led to the identification generic hybridization between field pea and grass pea. A low
of genes underlying agronomically important traits, some of frequency (5%) of protoplast fusion has been achieved; how-
which have been applied to MAS programs to improve Pisum ever, these hybrids have not been regenerated to whole plants.
germplasm. When markers that are closely linked to desirable
genes have been identified and validated in specific environ-
Transformation
ments, plant breeders can use them to select plants on the basis
of genotype, rather than phenotype. The application of MAS Plant genetic engineering provides an opportunity to introduce
can reduce unreliable phenotypic screening and cultivar devel- traits such as pest and disease resistance, improved protein
opment time as markers enable the selection of genotypes at quality, and herbicide tolerance from previously unavailable
the seedling stage. Molecular linkage maps and trait mapping sources. Since the first production of diploid transgenic field
are valuable tools for characterizing the genetics of disease pea plants in the early 1990s, Agrobacterium-mediated genetic
resistance, localizing resistance loci on linkage maps, and iden- transformation has become a routine procedure in a number of
tifying linked polymorphic DNA sequences that might be used laboratories worldwide. The embryonic axis of immature seeds
for MAS. This is particularly true for traits that are inherited in a or the lateral cotyledonary meristems from germinating seeds
quantitative manner, for which quantitative trait loci (QTLs) are the most responsive target explants, and regeneration is
may be identified. Despite progress in the area of marker generally via organogenesis.
discovery, complete translation to pea breeding is yet to be In addition to Agrobacterium-mediated transformation, sev-
achieved. eral other approaches have been developed to produce
330 THE LEGUMES AND PSEUDOCEREALS | Pea: Overview
transgenic field pea plants: transgenic callus has been obtained With blue peas, resistance to bleaching is an important
by electroporation of protoplasts, as has transient expression criterion. Round white types must have a smooth round sur-
following particle bombardment of meristems. Microinjection, face with white to creamy color and bright yellow cotyledons.
particle gun, agroinfection, and the pollen tube pathway are at The dun types should ideally have a mix of greenish to brown-
present less efficient or require expensive and sophisticated ish testae and fewer and shallower dimples with bright yellow
equipment. As in other species, transformation efficiency is cotyledons. Maple type, mainly used for pigeon and bird feed,
dependent on the genotype, explant type, and other physical must have white markings on a brown testa and yellow
parameters. Despite transformation being a well-established cotyledon.
technique in peas, there are currently no commercially avail- Both round white and dun types are milled, as are blue
able pea cultivars developed using this technology. types. However, currently, there are no standards for milling
recovery. Seed moisture content at sale is normally between
10% and 15%. All types of field peas are generally accepted by
Double Haploid Production the feed industry, but a preference is made for the noncolored
Doubled haploids (DHs) are plants derived from a single imma- type as they tend to have less antinutritional factors. The
ture pollen grain and doubled artificially to form homozygous Southeast Asian market for sprouting field pea prefers dun
diploids. A DH individual has two identical homologues so that types with large seed size and fewer hard seeds. Irrespective of
the amount of recombination information is equivalent to a the type, the following qualities are important for visual assess-
backcross. The advantage of using a DH population in molecular ment in the trade:
mapping is that all individuals are homozygous. Therefore, DHs Levels of admixture
may be transferred between different laboratories and environ- Levels of insect damage and presence of live or dead insects
ments for assessing the effect of the environment on gene expres- Product color
sion. Pea, along with many of the large-seeded leguminous Product size
species, has traditionally been considered recalcitrant to this Product cleanliness
technique. In 2009, the first production of confirmed haploid Product uniformity
plantlets of pea from the culture of immature pollen was
reported. Research is ongoing to develop a robust protocol that Today, almost 50% of the field pea production is traded
will be applicable across a range of genotypes. The mechanisms throughout the world compared with just 20% 15 years ago
for switching the immature pollen from normal gametic devel- and double that traded in 2002 (Figures 3 and 4), which
opment have also been recently elucidated for peas. Successful compares with temperate cereals where 5% is traded. The
production of DHs on a routine basis would reduce cultivar world field pea production has been mostly stable in the last
development time and provide excellent recombinant inbred decade, but trading across the board is showing a strong
lines for molecular mapping applications. increasing trend. The prices have been generally steady since
the early 2000s, but wide variations occur depending upon the
season, time of the year, and proximity to the market. Canada
In vitro Generation Acceleration and the Russian Federation are currently the worlds leading
exporters with Canada having dominated the export market for
In the absence of a robust, widely applicable, DH protocol, an many years. The quantity exported from France has halved
in vitro modified SSD system has been developed to accelerate the since 1999 (Figures 6 and 7). India and China are now the
achievement of homozygosity in peas. Conventional SSD con- largest importer of field pea with Bangladesh and Pakistan
sists of randomly selecting one seed per plant from an F2 popu- importing dry pea as demand requires. Germany, Spain, and
lation following a cross and advancing it through the early Belgium continue to be consistent importers among European
segregating generations, thus permitting rapid fixation of genes countries (Figures 8 and 9).
in breeding lines. As only one seed is needed to produce the next In general, the market for field pea is rather unsophisticated
generation, plants can be grown under conditions that accelerate in terms of both product definition and terminology used. For
flowering and seed set, but do not encourage high yield. Current instance, in dun-type field peas that are normally milled,
SSD methodologies enable a maximum of three generations per traders often use slight variation in testa color in an attempt
year. The in vitro SSD technique involves further shortening of to discount price that is irrelevant to the product utilization.
the generation time by culturing immature seeds developed after There are various terminologies used to describe the same
forced in vitro flowering under conditions for accelerated flower- product as in the case of white round type that is traded as
ing and seed set. This leads to rapid generation turnover by yellow pea from North America. Work continues at an inter-
reducing the length of the generation cycle, enabling up to national level to improve the marketing of pulses through
eight generations per year. development of standardized nomenclature and methods of
testing quality parameters in order to meet consumer needs.
The International Pulse Trade and Industries Confederation
Grain Quality and Marketing (CICILS/IPTIC) and various major importing organizations
are collaborating to develop standardized nomenclature and
Market requirements for grain quality relate predominantly to methods for quality evaluation that will facilitate improvement
shape, size, and color (testa and cotyledon). For all seed types, of quality assurance in the international pulse trade including
large size is desired, and in marrowfat, there may be a limit for field peas. Progress is available on the CICILS website (http://
the minimum seed size from the buyer. www.cicilsiptic.org/).
THE LEGUMES AND PSEUDOCEREALS | Pea: Overview 331
391,022
Canada
Russian Federation
835,857 672,731
United States of America
France
Australia
Figure 6 Dry pea exports by the top six exporting countries in 2002.
243,791
324,883
335,570
Canada
Russian Federation
460,797 2,854,580
United States of America
France
Australia
Figure 7 Dry pea exports by the top six exporting countries in 2011.
215,494 Spain
869,803 Belgium
214,545
Pakistan
277,199 Germany
Italy
Future Prospects yield and quality, some serious challenges need to be addressed,
which include both biotic and abiotic stresses.
The pea crop is likely to retain its importance as human food and Among biotic stresses, the black spot diseases and major
animal feed. While yield improvement per se will continue, seed insect pests continue to be major limitations to achieving high
quality will become critical as market sophistication develops. yields. Abiotic stresses such as tolerance to drought, frost, and
While pea breeding will continue to make improvements in waterlogging also need attention. Advances in new
332 THE LEGUMES AND PSEUDOCEREALS | Pea: Overview
211,784 Spain
Belgium
730,434 1,866,735
Pakistan
Germany
Italy
technologies such as those that accelerate generation turnover, Continuous improvement of doubled haploid production
DH, and molecular markers will greatly assist in addressing to enable reliable and cost-effective techniques for acceler-
future pea breeding objectives. Genetic modification will play a ating breeding.
more dramatic role in addressing some key challenges where More targeted studies in the available gene pool of P. fulvum
natural variation in pea germplasm is deficient. to find traits of interest for transfer to P. sativum.
Frost resistance/tolerance at pod development.
Challenges of possible climate change: heat and drought
Exercises for Revision resistance/tolerance.
Khan TN, Timmerman-Vaughn GM, Rubiales D, et al. (2013) Didymella pinodes and its Smykal P, Aubert G, Burstin J, et al. (2012) Pea (Pisum sativum L.) in the genomic era.
management in field pea: Challenges and opportunities. Field Crops Research Agronomy 2: 74115.
148: 6177.
Kraft JM and Pfleger FL (eds.) (2001) Compendium of Pea Diseases and Pests, 2nd
edn. St Paul, MN: The American Phytopathological Association. Relevant Websites
Makasheva RK (1986) The Pea. Rotterdam: A.A. Balkema.
Ochatt S, Pech C, Grewal R, Conreux C, Lulsdorf M, and Jacas L (2009) Abiotic stress http://www.cicilsiptic.org/ CICILS/IPTIC world trade association for pulses.
enhances androgenesis from isolated microspores of some legume species http://www.grdc.com.au/ Australian Grains Research and Development Corporation
(Fabaceae). Journal of Plant Physiology 166: 13141328. (GRDC) field pea growing guidelines including disease information.
Ribalta FM, Croser JS, Erskine W, Finnegan PM, Lulsdorf MM, and Ochatt SJ (2013) http://hermes.bionet.nsc.ru/pg/ Pisum Genetics, published periodically by Pisum
Antigibberellin-induced reduction of internode length favors in vitro flowering and Genetics Association, Department of Plant Sciences and Plant Pathology, Montana
seed-set in different pea genotypes. Biologia Plantarum 58(1): 3946. http://dx.doi. State University, Bozeman, MT 59717, USA.
org/10.1007/s10535-013-0379-0. http://www.pea-lentil.com/ USA dry pea and lentil council site with USDA policy
Ribalta FM, Croser JS, and Ochatt SJ (2012) Flow cytometry enables identification of positions and research updates.
sporophytic eliciting stress treatments in gametic cells. Journal of Plant Physiology http://www.pgro.org/ Processors and Growers Research Organisation (PGRO), United
169: 104110. Kingdom.
Schroeder HE, Schotz AH, Wardley-Richardson T, Spencer D, and Higgins TJV (1993) http://www.pulseaus.com.au/ Pulse Australia is the peak industry body representing
Transformation and regeneration of two cultivars of pea (Pisum sativum L.). Plant all sectors of the Australia Pulse Industry.
Physiology 101: 751757. http://www.pulsecanada.com/ Pulse Canada is the national industry association that
Siddique KHM, Erskine W, Hobson K, et al. (2013) Cool-season grain legume represents growers, processors and traders of pulse crops in Canada.
improvement in Australia: Use of genetic resources. Crop and Pasture Science http://www.saskpulse.com/ Saskatchewan Pulse Growers Group with agronomic
64: 347360. advice, markets and food uses.
Peanuts: Overview
RCN Rachaputi, University of Queensland, Kingaroy, QLD, Australia
G Wright, Peanut Company of Australia, Kingaroy, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights Brazil in South America. The Spaniards are believed to have
taken the crop to the western Pacific, Indonesia, and China
Background, distribution, utilization, and trade. early in the sixteenth century.
An overview of morphology, growth, and development. Arachis hypogaea is an annual member of the family Papilio-
Yield and quality constraints. naceae, which shows a unique feature of maturing their fruits
Peanut as a source of oil and quality snack food. underground (Figure 1). The plant comprises a number of
Food safety issues. subspecies Arachis fastigiata var. fastigiata (Valencia) and
Advances in breeding. var. vulgaris (Spanish), both bunch forms, and subspecies
Innovations in biotechnology. Arachis hypogaea var. hypogaea (Virginia bunch and runner).
Future prospects for increased yield and quality. Large variations in growth habit (erect vs. spreading), branch-
ing type (alternate or sequential), maturity (90160 days), and
traditional market types (Virginia, Spanish, or Valencia) exist.
Learning Objective
Leaf
Flower
Pod
Kernel in pod
Peg
Embryo
Roots
Figure 1 Diagram of a peanut plant showing its unique underground fruiting habit. Source: http://upload.wikimedia.org/wikipedia/commons/a/a4/
Arachis_hypogaea_-_K%C3%B6hler%E2%80%93s_Medizinal-Pflanzen-163.jpg.
Table 1 Annual peanut production and yields for the ten major world-producing countries during the periods
Production % Change
(million Mt) Production (million since Average yield (dry pods) Average yield (dry pods) %Change
Country 200809a Mt) 201112b 200809 (kg ha1) 200809 (kg ha1) 201112 since 2009
Trade
Brazil, which in total account for 71% of total world exports.
Peanut is generally traded in two product qualities, that is, as The peanut trade has steadily shifted from a crush for oil to a
food quality (snack food) and crush quality (cooking oil). The snack food market, which meant rising demand for higher-
crush-quality trade is generally mostly domestic to meet quality peanuts, especially with low aflatoxin contamination
demand for cooking oil. Food-quality peanut in general is (see section Aflatoxin).
international and receive a quality premium relative to crush- More than 48 peanut-growing countries have established
quality groundnuts. limits for total aflatoxins in food, but standards varied widely,
The world trade market for peanuts can be considered as a ranging from 0 to 50 parts per billion. Very strict aflatoxin
residual market, as only < 6% of the world production is regulatory limits in European markets (e.g., <4 ppb total afla-
traded internationally owing to most of the production being toxins) have impeded a number of Asian (China, Vietnam, and
utilized domestically. The major producersexporters of pea- India) and African (Nigeria and Sudan) countries in competing
nuts are the United States, Argentina, Sudan, Senegal, and in the edible peanut market.
336 THE LEGUMES AND PSEUDOCEREALS | Peanuts: Overview
Morphology and Growth Physiology grown as an intercrop with longer season annual and perennial
crops, including castor, cassava, maize, and sugarcane. Peanut
Peanut varieties grown in the world can be grouped into four can also be grown underneath perennial tree canopies such as
major botanical types, Spanish, Virginia bunch, Virginia coconut, cocoa, oil palm, and rubber, owing to its effective
runner, and Valencia, distinguishable by their branching, shade tolerance. Under these systems, peanut can provide cash
flowering, and fruiting habit. income until the plantation trees are old enough to produce
Being an indeterminate in growth, peanut supports simul- their end products.
taneous growth of vegetative and reproductive structures that
compete for nutrients and assimilates throughout most of the
crop life cycle. Flowers appear about 2535 days after planting, Biotic and Abiotic Constraints to Yield
depending on the botanical type of the variety and ambient
temperature during the crop growth, and continue to be pro- Biotic Constraints
duced until final maturity. The rate of flower initiation Insect pests
increases to a maximum and then declines as the pod load Around 360 species of arthropod pests are known to attack in
increases and the crop approaches maturity. Bursts of flowering different regions of the world. In general, the foliage pests are
are common when crop recovers from dry spells after rain of lesser importance than the soil-inhabiting pests, which can
during the vegetative phase and can lead to more synchronous cause direct economic damage to pods. A large number of
pod set and higher yields and quality given favorable growing thrips species that attack peanut and have become vectors of
conditions during the pod-filling period. Peanut has a self- some of the debilitating peanut viruses such as tomato spotted
fertilized flower that fades and withers soon after fertilization. wilt virus (TSWV). The most important soilborne pests in the
Rates of natural outcrossing are low and have been estimated developed world are the lesser cornstalk borer, Elasmopalpus
between 0.7% and 2.5%. lignosellus, and white grubs, the larvae of which attack pegs and
After 57 days of fertilization, the base of the fertilized ovary pods and cause substantial economic damage. Etiella behrii is
(gynophore or peg) begins to elongate toward the soil, pene- endemic to Australia and is known to be only Etiella species
trating the soil up to a depth of 45 cm. The ovary then begins that affects peanuts. The pod damage caused by the soil insects
to swell and turns horizontally away from the plant (Figure 1), can lead to aflatoxin contamination in kernels following infec-
which takes between 15 and 20 days from the initial flowering tion via the fungus Aspergillus flavus.
event. There may be 14 kernels per pod depending on the
botanical type. The pod volume expands as the kernels inside
Fungal pests
the pod inside fill, reaching full dimensions characteristic of the
The most important fungi causing economic damage to peanut
variety over the next few weeks. Because of the slow rate of pod
crops around the world are the leaf spots (early and late) and
addition, many pods must be added and seeds initiated to form
leaf rust. These diseases cause leaf necrosis and can lead to
a full pod load on the plant. Pod growth rates vary depending
severe yield losses if not effectively controlled by fungicides.
on temperature, water status, and varietal maturity and parti-
A number of highly efficient chemical formulations have been
tioning ability.
developed over recent years to control both diseases. Genetic
resistance is also available; however, there is a known negative
association between foliar disease resistance and yield
Farming Systems potential.
A number of soilborne diseases can cause significant eco-
Peanuts are grown as a rotation crop in a range of irrigated or nomic losses in peanuts, via seedling and plant death, as well
rainfed cropping systems around the world. In most developed as via direct pod damage due to pod rots. The most important
countries, peanut is grown as a sole crop in rotation with cereal diseases include white mold (Sclerotium rolfsii), Cylindrocladium
or sugarcane crops under rainfed or irrigated conditions, using black rot, and Sclerotinia blight. More detailed coverage of this
fully mechanized cultural practices from sowing to harvest. In topic can be found in the references suggested in the
countries such as the United States and Australia, it is not bibliography.
uncommon for one farming family to farm 500 or more hect-
ares per annum. Peanuts are generally grown in a rotation of Viruses
one peanut crop in two or three summer cereal crops in order Many viruses infect peanut, and historically, they have not
to minimize soilborne diseases and use of nitrogen fertilizers caused widespread economic losses. This situation has chan-
and allow more effective weed control. ged over the past decade, with the arrival of the potyvirus
In developing countries throughout Asia and Africa, peanut stripe virus (PStV) in Asia and the United States and
smaller-scale peanut production on 12 ha plots is more com- TSWV in the United States. A system of cultural practices have
mon, under a variety of cropping systems. been implemented to control these diseases, including plant-
In most regions in Asia, peanut is often grown as a cash crop ing date and plant and row spacing to control thrips vectors
with both kernels (as food) and haulms (as fodder) are sold in such as the western flower thrips. Some genetic tolerance has
markets. Peanuts are important component in rice-based crop- been identified for TSWV in the United States, but there is no
ping systems throughout Southeast Asia. The crop is either known genetic resistance to PStV, although a recent collabora-
planted with irrigation before a rice crop or grown on residual tive project involving Australian, Indonesian, and Chinese sci-
soil moisture after the rice harvest. In Asia, peanut is also entists has developed resistant breeding lines using molecular
THE LEGUMES AND PSEUDOCEREALS | Peanuts: Overview 337
approaches. More information on virus-impacting peanut can reproductive efficiency increasing at higher temperatures (e.g.,
be found in the website listed under bibliography. mean daily temperatures exceeding 26 C).
Soil nutrition
Abiotic Constraints The specific nutritional issues of most importance for peanut
About 80% of the worlds groundnut production comes from production include calcium (Ca) and biologically fixed nitro-
dryland regions of the semiarid tropics, which are characteri- gen (BNF).
zed by poor soil fertility and drought coupled with elevated Peanut pods absorb Ca directly from ambient soil solution.
seasonal temperatures. The Ca absorbed by the roots and transported via the transpi-
ration stream (xylem) is unavailable to the pegs and pods of
Drought the peanut plant as these subterranean organs do not transpire.
The major effect of soil and crop water deficits on peanut yield Low soil water content in the podding zone can decrease Ca
relates to reduction in canopy development and pod growth uptake and induce Ca deficiency in peanut leading to empty
ultimately affecting pod yield. Peanut has a reputation of being pods, referred to as pops. Calcium, usually applied as gypsum
a drought-tolerant crop, which is associated with a number of at during early pod set phase, has been widely practiced to
important plant traits. Significant genetic variation in these increase Ca supply to pods and yield of peanut under drought
traits exists and breeding programs are exploiting these traits conditions. Large-seeded Virginia types have been shown to be
to further enhance peanut drought resistance. Readers are more sensitive to drought-induced Ca deficiency than smaller-
referred to detailed reviews on this subject by Boote and seeded Spanish types.
Ketring, Wright and Nageswara Rao, and Nageswara Rao et al. Peanuts as other legumes have the unique ability to fix
atmospheric nitrogen by forming a symbiotic relationship
Temperature with Bradyrhizobium bacteria in the root nodules. Inoculation
Peanut is known to grow well under a wide range of tempera- of seeds with effective rhizobia has led to substantial increases
tures, with daily optima for the rate of development, growth in peanut yield in fields that have not previously grown pea-
processes, and yield being around 2530 C. High tempera- nuts. It is generally accepted that once a field has successfully
tures in excess of 35 C can reduce leaf area development, grown a peanut crop, it should not require future Rhizobium
reduce the number of pods, and result in lowered harvest applications. Estimates of fixed nitrogen as large as
indices and yields. The interactions of water deficits with tem- 300 kg N ha1 have been reported for well-watered crops,
perature cannot be ignored as high temperatures can accentu- with reports that BNF can supply over 80% of the plants
ate negative impacts of water deficit. In fact, the subterranean total N uptake. Although most of the N in the crop is removed
fruiting habit of groundnut renders it more sensitive when soil when the pods are harvested, numerous experiments have
surface temperature exceeds 40 C even under well-watered shown that significant N may be left in the soil for uptake by
conditions. The threshold temperature for growth and devel- subsequent crops. A number of studies showed that the N
opmental processes in peanut is listed in Table 2. fixation process is extremely sensitive to soil water deficits.
Photoperiod
Peanut Products, Quality, and Food Safety
While growth and yield of peanut are known to be linearly
related to total solar radiation receipt, reproductive develop- Peanut Products
ment and quality can be strongly related to photoperiod (day
Nearly two-thirds of the world production of peanut is crushed
length) in sensitive cultivars. Peanuts were previously thought
for cooking oil, making it one of the worlds leading oil crops.
to be day-neutral, with phenology mainly driven by tempera-
Peanut oil is mostly composed of triglycerides of eight fatty
ture. Controlled environment and field experiments have since
acids. Around 80% of these fatty acids are either oleic acid
determined that reproductive efficiency can be modified con-
(monounsaturated, C18:1) or linoleic acid (polyunsaturated,
siderably by varying photoperiod. Studies have found that
C18:2). Generally, these two fatty acids vary inversely. Of the
longer photoperiods reduced shelling percentage in sensitive
eight commonly detectable fatty acids in peanut oil, linoleic
cultivars. More recent data have shown that significant photo-
acid is the only one that is polyunsaturated. The second double
thermal interactions exist in peanut, with the sensitivity of
bond in a linoleic acid molecule renders it far more susceptible
to oxidation. This oxidative rancidity generates off-flavors in
Table 2 Temperature limits of various growth and developmental peanut products and hence limits the shelf life of products in
processes in groundnut
retail outlets. The monounsaturated oil type in peanuts is also
Process Tbase ( C) Topt ( C) Tmax ( C) known to be heart-healthy. It was also found that diets includ-
ing peanuts and peanut butter lowered cardiovascular disease
Germination 811.6 2936.5 4147 risk by 21%, whereas the low-fat diet decreased the risk by only
Leaf appearance 10 30 12%.
Leaf photosynthesis 2936 In the late 1980s, researchers at the University of Florida
Onset of anthesis 11 2933 38
discovered a (naturally occurring) mutant peanut plant with
Pollen viability and seed setting 2732 3743
over 75% oleic acid and linoleic acid below 5% of fatty acids,
Pod size and pod growth rate 2428
Harvest index 2732 3744 which they subsequently described as high-oleic acid germ-
plasm. This trait has recently been bred into adapted
338 THE LEGUMES AND PSEUDOCEREALS | Peanuts: Overview
commercial varieties, which are currently being released in the products. The human impact, in terms of mortality and mor-
United States and Australia. This characteristic will provide bidity, in peanut-growing countries in developing world is
even greater benefits to both product shelf life and human enormous, with estimates of up to 20 000 deaths per year
health. The increased shelf life is best illustrated by the arising from aflatoxin-related illnesses being made in some
peroxide value (PV; a measure of shelf life) data shown in Southeast Asian countries. In developed countries, consumers
Figure 2, which shows how high-oleic acid peanuts maintain are more aware of food safety issues such as aflatoxin and are
low PVs (and hence no rancid off-flavors) during an acceler- increasingly demanding peanut products meet strict regulatory
ated aging test. standards. Current strategies to reduce aflatoxin levels to below
regulatory limits (e.g., 15 ppb in retail products in the United
States and Australia) involve a post-farm gate process of selec-
Nutritional Aspects tive segregation achieved through blanching and color sorting
Peanuts are an excellent source of nutrition for both humans in shelling plants. In developing countries, aflatoxin regulatory
and animals. Tables 3 and 4 give a proximate nutritive com- limits for peanut have been stringently applied and have
position of peanut kernels and dry peanut hay, respectively. recently affected export of peanut and its products.
Clearly, they are a rich source of protein, oil, carbohydrate, and A number of potential solutions have been proposed to
minerals. Peanuts are also a good source of niacin, folic acid, minimize aflatoxin at its source, the farm. The use of agro-
phosphorous, vitamin E, and phytosterols. nomic management methods, including pre- and postharvest
strategies that minimize aflatoxin contamination by the fungi
Aspergillus flavus and Aspergillus parasiticus, has shown to be
Bioactives effective. A number of peanut varieties have been shown to
have some aflatoxin tolerance, which seems to be associated
Peanuts contain all nine amino acids essential to humans and with drought avoidance and escape mechanisms. Recent
nine nonessential amino acids. Recent studies have shown that research is also investigating the potential for using a bio-
peanuts also contain many compounds with antioxidant prop- control approach, where nontoxigenic strains of the Aspergillus
erties such as tocopherols, p-coumaric acid, and t-resveratrol, flavus fungus are applied to the soil and infect kernels and
which can be termed as antioxidants. hence competitively exclude toxigenic strains.
Figure 2 Peroxide value over time as a measure of oxidation of Source: Khan MT, Khan NA, Bezabih M, Qureshi MS, and Rahman A. (2013) The
high- and low-oleic acid peanut oils. Reproduced From OKeefe, Wiley, nutritional value of peanut hay (Arachis hypogaea L.) as an alternate forage source for
and Knauft (1993) Journal American Oil Chemists Society, 70, 489492. sheep. Tropical Animal Health and Production, 45(3), 849853.
THE LEGUMES AND PSEUDOCEREALS | Peanuts: Overview 339
Cadmium been identified for specific traits such as high oleic acid and
Cadmium (Cd) is a heavy metal, which can accumulate in nematode resistance, the marker-assisted selection has been
some foods and potentially lead to kidney damage if high successful in the development and commercial release of
levels of intake occur over prolonged periods. Recent research varieties.
has shown that peanut can accumulate high Cd levels on acidic The advent of next-generation sequencing (NGS) provides a
sandy soils, especially where there has been a history of phos- cost-effective strategy to identify molecular markers, at whole-
phatic fertilizer application, which had contained Cd as a by- genome level, associated with traits of interest and can expedite
product. A number of management practices have been the development of new peanut varieties. The size and com-
proposed to minimize Cd accumulation in peanuts, and readers plexity of the cultivated peanut genome have previously been
are referred to the following website for more information. considered to be a barrier to progress in molecular breeding
research; however, as the cost of NGS continues to decrease,
Anaphylaxis this appears to be no longer the case.
Anaphylaxis is a sudden, severe, potentially fatal, systemic aller- The International Peanut Genome Initiative (IPGI) estab-
gic reaction to peanuts that can involve various areas of the lished in 2004 and currently renamed as Advances in Arachis
body (such as the skin, respiratory tract, gastrointestinal tract, through Genomics and Biotechnology (AAGB) is aimed to
and cardiovascular system). Anaphylactic reactions can be mild facilitate the rapid discovery of progress in the development
to life-threatening and seem to have an annual incidence of of genomic resources across the international peanut commu-
about 30 per 100 000 persons. A number of the proteins nity. It is expected that a high-quality reference genome for the
responsible for peanut allergy have already been identified. cultivated peanut will be available by end of 2015.
Among major peanut allergens, Ara h 1, Ara h 2, and Ara h 3, Several research institutions have been able to transform
are storage proteins that comprise a substantial proportion of peanut with exogenous DNA and generate fertile plants from
the total protein in the seed. These proteins that are critical for transformed tissues. This technology has opened up enormous
cellular function most likely could not be eliminated without possibilities for expanding the genetic diversity of peanut.
deleterious consequences for plant growth and development. Recent applications of this technology include genetic trans-
However, breeding or bioengineering research is underway to formation for disease resistance to PStV and TSWV.
explore possibilities of substituting a nonallergenic for an aller-
genic form of proteins.
There is a general consensus that further genetic enhancement What makes peanut unique compared to other grain
legumes?
of crops including peanut for improved adaptation is best
achieved by trait-based breeding strategies. While conventional What are the major peanut-producing countries in the
world?
empirical breeding for yield improvement has been generally
successful, the rate of genetic gain has been slow and increas- What is the major food safety issue with peanut?
ingly becoming cost-prohibitive. Recent studies in Australia What parts of peanut produce aflatoxin?
lead to the development of speed breeding strategies, which What factors contribute to aflatoxin production in peanut?
can potentially half the time taken for the release of a new How can aflatoxin contamination be minimized?
variety with desirable agronomic traits. What are allergens? What implications do they have for
human health?
There is enormous genetic variability in peanut germplasm
for most traits of interest, including host-plant resistance to
disease and insects and along with important agronomic and
quality traits. This germplasm has been collected and pre- Exercises for Readers to Explore the Topic Further
served. The most significant collection is housed at the Inter-
national Crops Research Institute for the Semi-Arid Tropics
(ICRISAT) in India, where some 12 000 accessions are main-
Peanut allergens are becoming a major issue in developing
countries only. What could be the possible reasons?
tained, along with more than 70 wild Arachis species.
Allergens are some specific proteins in peanut kernel. How
many proteins are associated with allergens in peanut?
Marker-Assisted Breeding What makes molecular breeding of peanut complicated?
The biotechnology revolution occurring in a range of crops has
yet to be realized in peanut. This has been largely due to the
lack of polymorphism for allozymes or restriction fragment See also: Food Grains and the Consumer: Grains and Health;
length polymorphism (RFLP), which has restricted the use of
Food Grains: Intolerance, Allergy and Diseases: Cereal
marker technology for improved selection of quantitative traits
Allergens; The Basics: The Grain Chain: The Route from Genes to
such as yield. The peanut genome is also very large, around Grain-Based Products; The Grain Crops: An Overview; The Legumes
2800 Mbp, and is classed as a tetraploid.
and Pseudocereals: Amaranth: Overview; Beans: Overview;
Due to the low number of polymorphic markers available
Chickpea: Overview; Grain Legumes and Their Dietary Impact:
for the cultivated peanut, there has been very little success in Overview; Pea: Overview; The Oilseeds: Oilseeds: Overview;
marker-assisted breeding in peanut. Where the markers have
Soybean: Overview; Sunflower: Overview.
340 THE LEGUMES AND PSEUDOCEREALS | Peanuts: Overview
Further Reading Reddy PS (1988) Groundnut. New Delhi, India: Publications and Information Division,
Indian Council of Agricultural Research.
Ashley JM (1984) Groundnut. In: Goldsworthy PR and Fisher NM (eds.) The Physiology Revoredo CL and Fletcher SM (2002) World Peanut Market: An overview of the past 30
of Tropical Field Crops, pp. 453494. New York: Wiley. years. Research Bulletin No. 437. The Georgia Agricultural Experiment Stations
Bell MJ, Bagnall DJ, and Harch G (1991) Effects of photoperiod on reproductive College of Agricultural and Environmental Sciences, University of Georgia, Athens,
development of peanut (Arachis hypogaea L.) in a cool subtropical environment. II. GA, May 2002. (www.ces.uga.edu/ES-pubs/RB437.htm).
Temperature interactions. Australian Journal of Agricultural Research Wright GC and Nageswara Rao RC (1994) Groundnut water relations. In: Smartt JT (ed.)
42: 11511161. The Groundnut Crop, pp. 281335. London: Chapman Hall, Chapter 9.
Boote KJ and Ketring DL (1990) Peanut. In: Stewart BA and Nielsen OR (eds.) Irrigation Wright GC, Hubick KT, Farquhar GD, and Rao RCN (1993) Genetic and environmental
of Agricultural Crops. Agronomy Monograph 30, pp. 625717. Madison, WI: variation in transpiration efficiency and its correlation with carbon isotope
American Society of Agronomy. discrimination and specific leaf area in peanut. In: Ehleringer JR, Farquhar GD, and
Chu Y, Holbrook CC, Timper P, and Ozias-Akins P (2007) Development of a PCR-based Hall AE (eds.) Stable Isotopes and Plant Carbon-Water Relations, pp. 247268. San
molecular marker to select for nematode resistance in peanut. Crop Science Diego, CA: Academic Press.
47: 841847.
Chu Y, Wu CL, Holbrook CC, Tillman BL, Person G, and Ozias-Akins P (2011) Marker-
assisted selection to pyramid nematode resistance and the high oleic trait in peanut. Relevant Websites
Plant Genome 4: 110117. http://dx.doi.org/10.3835/plantgenome 2011.01.0001.
IPGI (2013) The Peanut Genome Project. In: Wilson RF (ed.) International Peanut http://agnews.tamu.edu/stories/PATH/aflares.htm Texas A. & M. University, USA.
Genome Initiative. http://www.peanutbioscience.com/homepage.html. http://www.ars.usda.gov/is/AR/archive/jan02/peanut0102.htm Aflatoxin Biocontrol.
Isleib TG, Wynne JC, and Nigam SN (1994) Groundnut breeding. In: Smartt JT (ed.) The http://www.ars.usda.gov/is/pr/1999/990712.htm Agricultural Research Service, USDA.
Groundnut Crop, pp. 552623. London: Chapman Hall Chapter 14. http://www.clw.csiro.au/publications/general2002/grain_legumes_brochure.pdf
Nageswara Rao RC, Sheshshayee MS, Nataraja Karaba N, Rohini S, Rama N, Cadmium in Grain Legumes.
Kumaraswamy S, Prasad TG, and Udaykumar M (2012) Genetic approaches to http://cropgenebank.sgrp.cgiar.org/index.php/management-mainmenu-433/stogs-
enhance adaptation of groundnut (Arachis hypogaea L.) to abiotic stresses. mainmenu-238/groundnut/guidelines/viruses Peanut Viruses.
In: Tetuja N, et al. (ed.) Improving Crop Resistance to Abiotic Stress, pp. 304360. http://www.dpi.qld.gov.au/fieldcrops/3027.html Managing Aflatoxin.
Germany: Wiley-Blackwell, Wiley-VCH Verlag GmbH & Co Chapter 14. http://www.foodallergy.org Anaphylaxis in Peanuts.
Nigam SN (1992) Groundnut A global perspective. In: Proceedings of an International http://www.peanut-institute.org/ The Peanut Institute.
Workshop, 2529 November, 1991, 548 pp. Patancheru, India: ICRISAT Centre. http://www.peanutsusa.com The American Peanut Council.
Pattee HE and Young CT (1982) Peanut Science and Technology. Yoakum, TX: http://sacs.cpes.peachnet.edu/nprl/oldver/atoxin.htm National Peanut Research
American Peanut Research and Education Association. Lab, USA.
Quinoa: Overview
SA Valencia-Chamorro, Escuela Politecnica Nacional, Quito, Ecuador
2016 Elsevier Ltd. All rights reserved.
Topic Highlights quinoa is cultivated from sea level in Chile to 4000 m in the
Altiplano of Peru and Bolivia.
Quinoa is an ancient crop from the Andean region. In the late 1970s, the main production areas of quinoa were
This grain exhibits great adaptability to extreme climate, located in the Andean region. In Peru and Bolivia, this crop was
from sea level to 4000 m, and is able to grow in poor and of great importance. It was cultivated not only for domestic
marginal soils with low annual rainfall. consumption but also for export. The cultivation of quinoa
It presents a wide genetic variability with differences in then spread from the Andean region to several countries in the
form, in grain color and size, in grain yield and also in world. In the 1980s, quinoa was grown in Canada, Denmark, the
terms of its by-products. United States, and the United Kingdom. In the late 1990s, an
Quinoa, a very nutritious grain crop, is a unique grain, American and European project on quinoa to promote the
which contains the correct balance of essential elements regional interchange of excellent genetic material among research
and vitamins, and it is gluten-free. institutes and universities from both continents was undertaken.
The main producers of quinoa are Bolivia, Peru, and Ecua- Today, after more than 20 years of work, quinoa has worldwide
dor. However, production is expanding to other continents distribution. Quinoa is being cultivated in several countries
such as Africa, Asia, Australia, and Europe. of the Andean region (Colombia, Bolivia, Ecuador, Chile, and
Quinoa production is around 80 000 t (8 107 kg) Peru), in Africa (Kenya), Asia (China, Japan, the plains of
annually. Northern India, and the Himalayas), Australia, Europe (England,
Processed foods from quinoa include a wide range of bread Denmark, France, Germany, Italy, the Netherlands, Portugal,
types, pasta, noodles, breakfast cereals, beer, porridge, etc. Spain, and Sweden), and North America (United States and
NASA has designated quinoa as a potential crop for its Canada). Quinoa production is in a process of expansion into
Controlled Ecological Life Support System (CELSS). different geographic areas of the planet due to its ability to adapt
Quinoa presents high potential to contribute to food secu- to a wide range of agroecological zones with extreme ecological
rity and nutrition for humankind in several regions conditions.
worldwide.
Classification
Learning Objectives
Quinoa belongs to the Chenopodiaceae family, genus Cheno-
To achieve understanding of the adaptability, variability, podium. Its botanical name is C. quinoa, Willd. Common names
nutritional quality, and uses of quinoa grain, the ancient used in the Andean region are quinua, kiuna, or parca in
crop from the Andean region. Ecuador, Peru, and Bolivia; supha, jopa, jupha, jiura, aara,
To be able to assess the future prospects for quinoa as a ccallapi, or vocali in Bolivia; quinhua in Chile; and suba or
major grain crop. pasca in Colombia.
The classification of quinoa was first made based on the
color of the plant and fruits. Subsequently, it was based on the
Introduction morphological types of the plant. Despite the wide variation
observed, quinoa is considered to be one single species. For
Quinoa (Chenopodium quinoa, Willd) is an indigenous crop of practical reasons, quinoa, like maize, was classified in terms of
the Andean region of South America. It is one of the oldest races.
crops of the American continent. Archaeological findings in The most extensive collection of different quinoa races
northern Chile showed that quinoa was used prior to 3000 belongs to Peru and Bolivia; each has over 2000 ecotype sam-
BCE. In Ayacucho, Peru, evidence has been found that quinoa ples. However, other collections exist in Argentina, Colombia,
was cultivated before 5000 BCE. Chile, Ecuador, England, the United States, and the former
The quinoa plant was widely cultivated in the entire Soviet Union.
Andean region, in Colombia, Ecuador, Peru, Bolivia, and Quinoa collected in Ecuador, Peru, and Bolivia has been
Chile, before the Spanish conquest. However, habits and tra- classified into 17 races; however, more races may exist. Two
ditional foods of the native people were replaced by foreign types of inflorescence are described:
crops such as wheat and barley. Therefore, quinoa was culti-
1. Glomerulates are small groups of flowers (glomeruli)
vated either in small plots in rural areas for domestic consump-
originating from tertiary axes.
tion or as borders for other crops such as potatoes or maize. For
2. Amaranthiformes have glomeruli originating mainly from
that reason, it was classified as food for poor people.
secondary axes.
Quinoa is grown in the highlands from 5 N in southern
Colombia to 30 S in northeastern Argentina. At sea level, it is According to this, the races of quinoa are classified as follows:
cultivated between 36 S and 40 S, in central Chile. Thus, first, glomerulate inflorescence (Cajamarca, Copacabana,
Cuzco, Challapata, Cochabamba, Sicuani, Junn, Ancash, clustered in different forms either amaranthiforme or glomer-
Glorieta, and Dulce) and, second, amaranthiforme inflo- ulate. The flowers are incomplete; they do not have petals.
rescence (Achacachi, Puno, Real, Potosi, Puca, Sucre, and Quinoa has both hermaphrodite flowers, located at the distal
Pichincha). end of a group, and female flowers located at the proximal end.
Quinoa grows from sea level to the Andean highlands. Thus, The fruit of quinoa is an achene. It is covered by
one of the most useful classifications is that describing five perigonium, which is the same color as the plant: white, yel-
ecotypes (Table 1): sea level, valley, subtropical, salar, and low, gray, light brown, pink, black, or red (Figure 2). It is easily
altiplanic. removed when it is dried. It produces small, circular-shaped
Quinoa presents a great genetic diversity, with differences in seeds, range 1.54 mm (1.5 10 34.0 10 3 m) diameter
color, form, and size, as well as in quality and quantity of (250500 seeds per gram).
primary elements and secondary metabolites. Quinoa germ-
plasm banks in the Andean region have been in existence since
the 1960s. Germplasm collection has continued in several
Andean countries. The main quinoa germplasm collection is
preserved in Bolivia, which conserves more than 5000 quinoa
accessions. Peru, Ecuador, and Chile hold more than 536, 642,
and 25 quinoa accessions, respectively.
Growth
Ecotypes Location altitude (m) Varieties Characteristics
Sea level South of Chile <500 Chilean varieties Unbranched, long day plants (<2 m tall),
yellow, bitter seeds
Valley Andean valley 20004000 Blanca de Junn, Rosada de Jun, Big plants (23 m tall), branched, short growth
Amarilla de Matangani, Dulce de period
Quitopamba, Dulce de Lazo
Subtropical Subtropical area of 25003000 Plants with intense green color that turn orange
Bolivia (Yungas) as they mature; small seeds, white or orange
Salar Bolivian Salares 37003800 Real Plants adapted to salty and alkalinic soils;
bitter seeds; high saponin content
Altiplanic Area around Lake 35004000 Cheweca, Kancolla, Blanca de Jul Short plants with straight stems (11.8 m tall);
Titicaca short growth period; resistant to frost
Source: Risi J and Galwey NW (1984) The Chenopodium grains of the Andes: Inca crops for modern agriculture. Advances in Applied Biology 10: 145216. Reproduced from
Encyclopedia of Food Sciences and Nutrition, second ed. (2003), p. 4896. Elsevier Ltd.
THE LEGUMES AND PSEUDOCEREALS | Quinoa: Overview 343
intensive monoculture using mechanization. Thus, some con- grains such as rice, maize, and barley. However, it represents
cerns with regard to environmental sustainability have arisen. less than half of the quantity of protein in most legumes. In
quinoa, most of the protein is located in the embryo. The
nutritional value of quinoa varies considerably among varie-
Chemical Composition and Nutritional Value ties, which may depend on soil and fertilization regime. Sev-
eral studies on the chemical composition of quinoa have been
The diet of ancient inhabitants of the Inca Empire has gener- made in the past decades. For instance, in Bolivia, variation in
ated interest due to its being extremely nutritious. nutritional values of 20 different quinoa accessions from the
Like cereals, quinoa grains contain carbohydrates, protein, fat, Banco Nacional de Germoplasma de Granos Altoandinos
minerals, fiber, and vitamins (Table 2). The chemical composi- (BNGA) was reported. The protein content was in the range
tion of quinoa depends on the variety, soil, cultural practices, and 12.4715.48%, which is low.
environment of its cultivation. Values for the nutritional charac- Albumins and globulins are the major protein fraction
teristics of 555 quinoa accessions have been reported: protein (4477% of total protein) in quinoa, which is greater than
(10.2118.39%), fat (2.0510.88%), dietary fiber (3.469.68%), the prolamins (0.57.0%). Thus, quinoa is considered to be
ash (2.125.12%), carbohydrates (52.3172.98%), and energy a gluten-free grain because it contains very little or no prola-
(1.31 1041.68 104 kJ kg 1). Quinoa accessions show wide min. Therefore, quinoa provides a nutritional food source,
variability for most of the studied parameters, which is indicative which is of particular relevance to gluten-intolerant people.
of their genetic potential. The good nutritional quality of quinoa is primarily due to
its high protein content and quality. Quinoa protein has a
good balance of essential amino acids (Table 3). It is excep-
Quinoa Grains
tionally high in lysine, an amino acid, which is generally
Protein not overly abundant in plant foods. It is also a good comple-
The protein content of quinoa grains can vary from 8% to 22%, ment for legumes, which are often low in methionine and
which on an average is higher than that of common cereals cystine.
Table 2 Chemical composition of quinoa and some cereals and legumesa (10 2
kg per kg dry wt)
Quinoa Barley Corn Rice Wheat Oat Rye Bean Lupin Soybean
Protein 16.28 11.02 10.51 7.94 17.65 18.40 11.57 26.72 40.39 39.90
Fat 7.00 1.29 5.29 2.92 2.20 7.52 1.82 0.94 10.88 21.80
Ash 2.74 1.23 1.34 1.53 2.17 1.87 1.76 4.34 3.66 5.32
Carbohydrateb 73.99 86.44 82.85 77.24 77.98 72.21 84.85 68.00 45.08 32.98
Fiber, total dietary 8.07 17.35 8.14 3.50 13.98 11.55 16.89 28.22 21.10 10.28
kJ kg1(104)c 1.54 1.47 1.53 1.55 1.34 1.63 1.41 1.39 1.55 1.87
a
Quinoa, uncooked; Barley, pearl, raw; Corn, yellow; Rice, brown, long-grain, raw; wheat, hard red spring; Beans, kidney, all types, mature seeds, raw; Lupins, mature seeds, raw;
Soybeans, mature seeds, raw.
b
Carbohydrate includes fiber.
c
kJ kg1: 4 (%protein + carbohydrates) + 9 (%fat).
Source: Adapted from US Department of Agriculture, Agricultural Research Service. 2013. USDA National Nutrient Database for Standard Reference, Release 26. Nutrient Data
Laboratory Home Page, http://www.ars.usda.gov/nutrientdata (accessed 21-02-2014).
Table 3 Essential amino acids in quinoa and other foodsa (103 kg per kg protein)
Histidine 29 11 25 21 51 30 15
Isoleucine 36 36 42 35 44 52 30
Leucine 60 70 83 67 80 95 59
Lysine 54 32 38 26 69 84 45
Methionineb 36 29 35 41 26 32 22
Phenylalaninec 61 75 89 77 15 10 38
Threonine 30 31 37 28 42 43 23
Tryptophan 12 11 13 13 12 13 6
Valine 42 49 57 44 96 63 39
a
Quinoa, uncooked; Corn, yellow; Rice, brown, long-grain, raw; Wheat, hard red spring; Beans, kidney, all types, mature seeds, raw; Milk, whole, 3.25% milk fat, with added vitamin D.
b
Methionine + cystine.
c
Phenylalanine + tyrosine.
Source: Adapted from US Department of Agriculture, Agricultural Research Service (2013). USDA National Nutrient Database for Standard Reference, Release 26. Nutrient Data
Laboratory Home Page, http://www.ars.usda.gov/nutrientdata (accessed 21-02-2014); World Health Organization (2002). Joint FAO/WHO/UNU Expert Consultation on Protein and
Amino Acid Requirements in Human Nutrition. WHO technical report series; no. 935. Geneva: WHO.
THE LEGUMES AND PSEUDOCEREALS | Quinoa: Overview 345
The nutritional evaluation of quinoa protein has been Table 4 Mineral composition (105 kg per kg dry wt) and vitamin
reported in several studies. The protein efficiency ratio of raw concentration (105 kg per kg dry wt) of quinoa and some cerealsa
debittered quinoa was 7893% of that of casein. These figures
Quinoa Wheat Rice Barley
increased when quinoa was cooked and became 102105% of
that of casein. Thus, the protein quality of quinoa matches that Minerals (105 kg per kg)
of the milk protein casein. Ca 54.2 28.7 25.7 32.3
Mg 227.2 142.1 159.5 87.9
Carbohydrates and fiber K 649.2 389.7 248.8 311.4
The major component in quinoa is carbohydrates, which varies P 526.9 380.6 371.5 245.8
from 67% to 74% of the dry matter. Starch is about 5260%. Fe 5.3 4.1 1.6 2.8
The starch is located in the perisperm of the seed; starch may be Cu 0.7 0.5 0.3 0.5
Zn 3.6 3.2 2.3 2.4
as simple units or as spherical granules. The diameter of quinoa
Vitamins (105 kg per kg)
starch granules is in the range of 0.42.0 mm (4 10 7
Thiamin (B1) 0.42 0.58 0.45 0.21
2 10 6 m). Starch granules in quinoa are smaller than those Riboflavin (B2) 0.37 0.13 0.10 0.13
reported for maize (range 123 mm (1.0 10 6 Niacin (B3) 1.75 6.55 5.68 5.12
2.3 10 5 m)) and for wheat (240 mm (2 10 6 Ascorbic acid NR 0.00 0.00 0.00
4 10 5 m)). The gelatinization temperature for different qui- -Tocopherol 2.81 1.16 1.34 0.02
noa cultivars was found to be in the range of 55.572.0 C. The -Carotene 0.01 0.01 0.00 0.01
starch amylose content is about 11%, which is lower than in
NR, not reported.
cereals, for example, rice (17%), wheat (22%), and barley a
Quinoa, uncooked; wheat, hard red spring; rice, brown, long-grain, raw; barley, pearl,
(26%). The total dietary fiber ranges from 11.0 to 13.3% in
raw.
raw and heat-treated quinoa. The crude fiber content is in the Source: Adapted from US Department of Agriculture, Agricultural Research Service
range 2.53.9%, and pentosans 2.93.6%. The high content (2013). USDA National Nutrient Database for Standard Reference, Release 26. Nutrient
of dietary fiber has many positive health effects, for example, it Data Laboratory Home Page, http://www.ars.usda.gov/nutrientdata (accessed 21-02-
can reduce the level of cholesterol in the blood and improve 2014).
digestion. For this reason, consumers in developed countries
may also have an interest in including quinoa in their diets.
Other carbohydrates are found in small amounts, such as acid, hederagenin, 30-o-methylspergulagenate, and phytolac-
monosaccharides (2%) and disaccharides (2.3%). cagenic acid are identified in sweet and bitter genotypes of
quinoa. Saponins show hemolytic activity and are in general
Fat, vitamins, and minerals toxic to cold-blooded animals, which obtain oxygen directly
Quinoa contains from 2% to 10% fat. Quinoa and soy oils from water.
exhibit similar fatty acid composition. Thus, quinoa is a rich Reduction of plasma cholesterol has been attributed to the
source of essential fatty acids such as linoleic (18:2n 6: 52%) presence of certain saponins in the diet. However, some sapo-
and linolenic (18:3n 6: 4%). The fatty acid composition is nins can form insoluble complexes with minerals, such as zinc
similar to maize germ oil. Quinoa is also a good source of and iron, which make the minerals unavailable for absorption
minerals (Table 4). It contains more calcium, magnesium, in the gut.
iron, and zinc than common cereals, and the iron content is Phytic acid (myoinositol 1,2,3,4,5,6-hexakis dihydrogen
particularly high. phosphate) is found in most cereals and legumes, fruits, and
Polishing and washing quinoa grain reduce the mineral vegetables. In quinoa seeds, phytic acid is located in the external
content to some extent, by 1215% for iron, zinc, and potas- layers as well as in the endosperm. Even small amounts
sium, 27% for copper, and 3% for magnesium. Quinoa con- (0.5 mmol g 1 (5.0 10 4 mmol kg 1)) of inositol hexapho-
tains more riboflavin (B2) and a-tocopherol than rice, barley, sphate (IP6) or pentaphosphate may reduce the solubility of
or wheat. The concentrations of the natural antioxidants g- and iron. IP6 was mainly found in varieties from Ecuador: sweet
a-tocopherol in quinoa were found to be 797.2 and 788.4 ppm INIAP-Tunkahuan (11.3 mmol g 1 (1.1 10 3 mmol kg 1)) and
(7.97 10 47.88 10 4 kg kg 1), respectively. Thus, quinoa bitter INIAP-Ingapirca (8.6 mmol g 1 (8.6 10 3 mmol kg 1)).
can be a source of vitamin E. These levels were reduced to just 0.3 mmol g 1 (3.0 -
10 4 mmol kg 1) by fermentation of germinated quinoa
Other components flour. At the same time, a five- to eightfold increase in the amount
Saponins (after removal), phytic acid, trypsin inhibitors, and of soluble iron was found.
tannins are present in quinoa in low levels. As stated, saponin, The trypsin inhibitor activity in eight varieties of quinoa
a bitter-tasting compound, is located in the outer layers of (range 1.365.04 TIU mg 1 (1.36 10 65.04 10 6 TIU -
quinoa seeds (at levels of 0.15%). It protects the seeds from kg 1)) was found to be much lower than in soybean (24.5 -
birds and insects. The amount of saponins present depends on TIU mg 1 (2.5 10 5 TIU kg 1)). Trypsin inhibitor is a
the variety. Bitter-flavor varieties contain more saponin than thermolabile protein, which is inactivated by heat treatments.
sweet varieties. Saponins are glycosidic compounds, which The total phenolic content (using the FolinCiocalteu
occur in two groups. According to the nature of the sapogenin method) of precooked quinoa seeds indicated that bitter seeds
moiety, they are conjugated with hexoses, pentoses, or uronic contained higher amount of phenolics (range 7.2310.05 mg
acids. The sapogenins are steroids (C27) or triterpenoids gallic acid equivalent per 100 g (7.2 10 51.0 10 4 kg gallic
(C30). Using gas chromatography, the sapogenins oleanolic acid equivalent per kilogram)) than sweet ones (range
346 THE LEGUMES AND PSEUDOCEREALS | Quinoa: Overview
6.059.39 mg/100 g (6.1 10 59.4 10 5 kg gallic acid equiv- Quinoa can be eaten as a rice replacement, as a hot breakfast
alent per kilogram)). Moreover, bitter quinoa precooked seeds cereal, or boiled in water to make an infant cereal (Figure 3).
exhibited a higher flavonoid content (1.041.74 mg catechin The grains can even be popped like popcorn. The seeds can be
equivalent per 100 g (1.0 10 51.7 10 5 kg catechin equiv- also sprouted. The sprouts need to turn green before they can
alent per kilogram)) than sweet seeds (0.710.91 mg/100 g be added to salads.
(7.1 10 69.1 10 6 kg/kg 1)). Thus, bitter and sweet quinoa Quinoa flour can be mixed with maize or wheat flour for
seeds varieties are a rich source of antioxidant compounds, the making bread and pasta, and it is used for making biscuits,
former being higher in antioxidants. cakes, and pastries in general.
Several levels of substitution of quinoa flour have been
reported, for instance, in bread (1013% quinoa flour),
Quinoa Leaves
noodles and pasta (3040% quinoa flour), and sweet biscuits
The leaves of quinoa are comparable to spinach as regards (60% quinoa flour). All levels produce quinoa products of
flavor. They contain (on a wet basis) carbohydrate 4.8%, pro- excellent quality. Quinoa flour can also be drum-dried and
tein 3.3%, fat 1.8%, ash 3.3%, and fiber 1.9%. The protein extruded, producing products with good physical, sensorial,
content of quinoa leaves is also similar to spinach. However, its and nutritional qualities. Solid-state fermentation of quinoa
protein contains slightly more isoleucine (5.8% per 100 g pro- with Rhizopus oligosporus Saito has been performed, to produce
tein) and valine (7.5% per 100 g protein). The amount of tempeh of good quality.
saturated fatty acids such as palmitic (16:0; 16.7%) and stearic In Bolivia, in 1975, the government adopted a resolution
(18:0; 1.3%) is higher than in the grains. Quinoa leaves are a mandating that 5% of quinoa flour must be added to all pastas,
rich source of vitamin A containing 2085 mg RE (retinol equiv- crackers, and breads.
alents) per 100 g (fresh wt.) (2.9 10 5 kg RE per kilogram Quinoa leaves, like grains, can also be cooked, made into a
(fresh wt.)) and vitamin E 2.9 mg a-TE (alpha-tocopherol) per spinach-like dish, or may be served raw in a salad.
100 g (2.9 10 8 kg a-TE (alpha-tocopherol) per kilogram). During four years, from 2002 to 2006, an Andean Quinoa
Fresh quinoa leaves contain more magnesium (83 mg per Project was undertaken in three countries, Bolivia, Peru, and
100 g fresh wt. (8.3 10 4 kg per kilogram fresh wt.)) and Colombia. Research goals included agriculture and agro-
sodium (289 mg per 100 g fresh wt. (2.9 10 3 kg per kilog- industrial activities. Several quinoa enriched quinoa products
ram fresh wt.)) than spinach leaves. Also, quinoa leaves have were developed. Additionally, more than 30 different pro-
low levels of the antinutrients nitrate and oxalate. cessed products from quinoa grains and plants have been
Sapogenins in the leaves increased as the plant matures. reported. Quinoa has jumped from a staple food to gourmet
After 120 days of from sowing, the sapogenin content in leaves kitchen food.
of sweet genotypes varied between 0.013% and 0.017% (dry Saponins from quinoa, due to their chemical qualities, have
matter) and in bitter varieties varied between 0.02% and multiple purposes. They are used in the manufacture of soap,
0.16% (dry matter). Hederagenin was the major sapogenin liquid detergents, shampoos, toothpaste, stain removers, and
present in the leaves. foam stabilizer; in beverages, especially beer; as an insecticide;
in a compound for fire extinguisher; and in photo processing.
The grains, the stems, and the leaves have been used for their
Uses medicinal properties, for cicatrizing, as an anti-inflammatory, an
analgesic, and a disinfectant. The plant-dried stalks are used as
In Chile, Ecuador, Peru, and Bolivia, the main uses of whole fuel or may be used in preparing bleach or dyes or for cardboard
quinoa seeds are in soups, salads, casseroles, and stew. Cooked and paper. The natural colorant from quinoa (betalains) was
grains swell to about two or three times their original size. traditionally used for cloth dyeing and food preparation.
Figure 3 Quinoa products. Courtesy of Lcdo. Guillermo Tapia, Cereales Andinos Ecuador.
THE LEGUMES AND PSEUDOCEREALS | Quinoa: Overview 347
Because of its excellent nutritional characteristics, NASA has What important nutritional characteristics does quinoa
included quinoa as a potential crop for its CELSS project. The have? How can these be exploited in development of new
CELSS concept utilizes plants to remove carbon dioxide from food products?
the atmosphere and generate food, oxygen, and water for the Which environmental concern is related to increased pro-
crews of long-term human space missions. duction in the Bolivian highlands?
How is quinoa being promoted all over the world? Do you
think it has potential to be major grain worldwide?
Future Perspectives
Exercises for Readers to Explore the Topic Further
Quinoa has long been recognized as nutritious food. The
importance that quinoa could play in nutritional behavior is
Quinoa is a unique food crop, which contains all the essen-
now being emphasized, not only in developing countries but tial amino acids in the correct balance. Taking this into
also in the developed world. Quinoa could be a strategic crop account, what are the agro-industrial prospects for this
used to complement the diet in rural/marginal regions where grain?
protein-energy malnutrition affects most of the population.
Production of quinoa was originally in Andean countries;
On the other hand, quinoa represents an exotic and healthy now, quinoa is cultivated in other countries. What are the
rediscovery in the developed world. Germplasm collection challenges arising from this fact?
should continue in countries of the Andean region.
The production of the grain has notably increased in the
Andean region. It is also in a process of expansion in other See also: Carbohydrates: Carbohydrate Metabolism; Cereals:
parts of the world. The potential for improving quinoa pro- Chemistry and Physicochemistry of Non-starchy Polysaccharides;
duction seems promising. Therefore, new varieties (precocity, Glycemic Index, Glycemic Load, and Their Health Benefits; Grains other
grain size and color, resistance to biotic and abiotic factors, and than Cereals: Non-starch Polysaccharides; Health Effects of b-Glucans
grain yield) and by-products should be obtained through Found in Cereals; Resistant Starch and Health; Starch: Analysis of
genetic improvement. Participatory plant breeding for quinoa Quality; Starch: Chemistry; Starch: Starch Architecture and Structure;
(including plant breeders, agronomists, communities, farmers, Starch: Synthesis; Fats: Healthy Fats and Oils; Lipid Chemistry; Food
and others) is desirable. Improvement of quinoa yields by Grains: Intolerance, Allergy and Diseases: Celiac Disease;
improving cultivation practices should be performed. Cereal Allergens; The Gluten-Free Diet; Grain Composition and
There is need for intensive cultivation of quinoa. This could Analysis: The Composition of Food Grains and Grain-Based
meet the quantity and quality needs of the food industry. Products; Proteins: Nitrogen in Grain Production Systems; Nitrogen
Traditional production of quinoa includes crop rotation. In Metabolism; Protein Synthesis and Deposition; Proteomic Analysis
the Bolivian Southern Altiplano, production is often made Relevant to Cereal Grains; The Enzymes Associated with Food Grains
without crop rotation. This could lead to very unsustainable and Their Functions in Processing; The Protein Chemistry of
production systems. Further research is needed in order to Dicotyledonous Grains; Thiolomics of the Gluten Protein Network of
identify the agroecological and social basis of quinoa sustain- Wheat Dough; The Basics: Taxonomic Classification of Grain
ability in traditional and new cultivation regions. Species; The Grain Chain: The Route from Genes to Grain-Based
Improving methods for removing saponins without signif- Products; The Grain Crops: An Overview; The Grains that Feed the
icant modification of nutritive value is encouraged. The selec- World.
tion of sweet genotypes with low saponin content in the seeds
is the main breeding goal. Further research is needed to find
markers for indirect selection of sweet genotypes.
Aggressive promotion campaigns should be carried out to Further Reading
encourage greater consumption of quinoa, as it is a very healthy
Brkic M, Garca A, 2013 Un cultivo ancestral para apuntalar el futuro. http://www.
and gluten-free grain. Quinoa has been spread outside of the alimentosargentinos.gov.ar/contenido/revista/pdfs/57/57_04_QUINOA.pdf
Andean region, owing to its great adaptability to different agro- (accessed 01.10.13.).
ecological systems. The year 2013 was declared the FAO 2011 Quinoa: An ancient crop to contribute to world food security. http://www.fao.
International Year of the Quinoa. Several international events org/fileadmin/templates/aiq2013/res/en/cultivo_quinua_en.pdf (accessed
01.02.13.).
were carried out all over the world. Quinoa should be promoted Gandarillas H (1979) Historia y distribucion geografica. In: Tapia M, Gandarillas H,
as a food of the twenty-first century, as it has a high potential to Alandia S, Cardoso A, Mujica A, Ortiz R, Otazu V, Rea J, Salas B, and Zanabria E
contribute to food security in several regions worldwide. (eds.) La Quinua y la Kaniwa. Cultivos Andinos, pp. 2144. Bogota: Centro
Internacional para el Desarrollo.
Giusti K (1970) El genero Chenopodium en la Argentina. I. Numero de Cromosomas.
Darwiniana 16: 98105.
International Year of the Quinoa. www.fao.org/quinoa-2013 (accessed 13.08.13.).
Exercises for Revision Jacobsen SE, Mujica A, and Ortiz R (2003) The global potential for quinoa and other
Andean crops. Food Rev. Int. 19: 39148.
What region is the origin of quinoa? Where is most genetic Masterbroek H, Limburg H, Gilles T, and Marvin H (2000) Occurrence of sapogenins in
leaves and seeds of quinoa (Chenopodium quinoa Willd). J. Sci. Food Agr.
variation to be found? 80: 152156.
What is the adaptability of this crop to soil and climate Mujica A, Jacobsen SE, Izquierdo J, 1998 Libro de Campo: Prueba Americana y
conditions? Could quinoa be produced near where live? Europea de Quinua (Chenopodium quinoa Willd). Universidad Nacional del
348 THE LEGUMES AND PSEUDOCEREALS | Quinoa: Overview
Altiplano, Puno, Peru; Red de Cooperacion Tecnica en Producion de Cultivos Balancing Market Potential and Sustainable Livelihoods, pp. 2574. Rome, Italy:
Alimenticios de la Oficina Regional de la FAO para America Latina y el Caribe. Bioversity International.
Mujica A, Ortiz R, Bonifacio A, et al., 2006. Proyecto Quinua: Cultivo Multiproposito Tapia M (1979) Historia y distribucion geografica. In: Tapia M, Gandarillas H, Alandia S,
para los Pases Andinos PNUD- PROY/INT/01/K01- Peru- Bolivia- Colombia. Cardoso A, Mujica A, Ortiz R, Otazu V, Rea J, Salas B, and Zanabria E (eds.)
Informe Final. La Quinua y la Kaniwa. Cultivos Andinos, pp. 1119. Bogota: Centro Internacional
Peralta E, Mazon N, Murillo A, et al., 2012. Manual Agrcola de Granos Andinos: para el Desarrollo.
Chocho, Quinoa, Amaranto y Ataco. Tercera edicion. Publicacion Miscelanea No. U.S. Department of Agriculture, Agricultural Research Service, 2013. USDA
69. Programa Nacional de Leguminosas y Granos Andinos. Estacion Experimental National Nutrient Database for Standard Reference, Release 26. Nutrient
Santa Catalina, INIAP. Data Laboratory Home Page, http://www.ars.usda.gov/nutrientdata
Repo-Carrasco R, Espinoza C, and Jacobsen S-E (2003) Nutritional value and use of the (accessed 21.02.14.).
Andean crops quinoa (Chenopodium quinoa) and Kaniwa (Chenopodium Valencia S, Svanberg U, Sandberg A-S, and Ruales J (1999) Processing of quinoa
pallidicaule). Food Rev. Int. 19: 179189. (Chenopodium quinoa Willd): Effect on the in vitro iron availability and phytate
Risi J and Galwey NW (1984) The Chenopodium grains of the Andes: Inca crops for hydrolysis. Int. J. Food Sci. Nutr. 5: 203221.
modern agriculture. Adv. Appl. Biol. 10: 145216. World Health Organization, 2002. Joint FAO/WHO/UNU expert consultation on protein
Soto JL and Carrasco E (2012) Market potential of Andean grains. In: Giuliani A, and amino acid requirements in human nutrition. WHO Technical Report Series: No.
Hintermann F, Rojas W, and Padulosi S (eds.) Biodiversity of Andean Grains: 935. WHO: Geneva.
Grains Around the World
Contents
Grain Production and Consumption: Overview
Grain Production and Consumption: Africa
Production and Consumption of Grains: India
Grain Production and Consumption: China and South-East Asia
Grain Production and Consumption: Europe
Grain Production and Consumption: Cereal Grains in North America
Grain Production and Consumption: Oilseeds in North America
Grain Production and Consumption in Oceania: Australia and Pacific Countries
Grain Production and Consumption: South America
Topic Highlights grain per hectare is close to the maximum obtainable through
photosynthesis; hence, world grain production has not
Wheat, rice, and maize contribute over 90% of global cereal matched the increase in worlds population. Low grain prices
production. at planting time, high temperatures, and water shortages are
World production of wheat, paddy rice, and maize was 652, the major factors contributing to reduced grain harvests. The
701, and 875 million tonnes (Mt) in 2012. fall in production has triggered an increase in the prices of
The major wheat-producing regions are the EU, FSU, NC wheat and maize and a corresponding increase in products
America, South America, Oceania, and Asia. (bread, breakfast cereals, pasta, and livestock products, includ-
The major rice exporters include India, Thailand, Vietnam, ing meat, milk, and eggs) derived from these grains.
the United States, and Pakistan. World production of wheat, rice (paddy), and maize in
Maize is the preferred crop in Africa and South America. 2012 was 652, 701, and 875 million tonnes (Mt), respectively
World oilseed production was 454 Mt in 2012 with soy- (Table 1). Total cereal production is shown in Figure 2 as an
bean as the predominant grain crop. average of 10-year periods beginning 1961. There was a 30%
increase in cereal production during the period between
197180 and 198190. The rate of increase in cereal produc-
Learning Objectives tion has been slowing since the days of the Green Revolution in
the 1970s. Table 1 shows world production of cereals, wheat,
Achieve understanding of the major grains that are pro- rice, maize, barley, sorghum, millet, and rye for the period
duced and consumed worldwide. 201012. Data on yields and area harvested are also listed.
The worlds wheat was produced in Asia (47%), Europe
(29%), and North and Central (NC) America (13%) in 2012.
Introduction About 90% of the worlds rice was produced in Asia. Africa and
South America contributed 4% and 3% to the worlds rice
The term grain is used to include cereals (wheat, rice, and the production, respectively. NC America, Asia, South America,
coarse grains such as maize, barley, sorghum, oats, and rye), and Europe contributed 36, 33, 13, and 11%, respectively, of
oilseeds, and pulses. World grain production, particularly in the worlds maize production. Wheat, rice, and maize contrib-
wheat and rice, increased substantially between 1961 and 1980 uted to over 90% of global cereal production. World exports
due to the increase in yields per hectare or the Green and imports of cereals by region in 2009 and 2010 are given in
Revolution (Figure 1). Since then, per capita (per person) Figures 3(a) and 4(a), respectively. Pulses (including dry
grain production has not increased much as most of the easily beans, dry peas, chick pea, and pigeon pea) were 3% of the
realizable benefits of plant breeding, fertilizer, machinery, total cereal production. Among oilseeds, soybean was 10% of
and irrigation have already been achieved. Production of world cereal production.
2800
2600
2400
2200
2000
Production (mt)
1800
1600 Cereals, total (Mt)
1400 Rice, paddy (Mt)
1200
Wheat (Mt)
1000
800 Maize (Mt)
600
400
200
1960
1961
1962
1963
1964
1965
1966
1967
1968
1969
1970
1971
1972
1973
1974
1975
1976
1977
1978
1979
1980
1981
1982
1983
1984
1985
1986
1987
1988
1989
1990
1991
1992
1993
1994
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
2006
2007
2008
2009
2010
2011
2012
Year
Figure 1 World grain production for the period 19612012. FAOSTAT Database 2013.
Area harvested (Mha) Yield (kg ha 1) Production (Mt) Data on world wheat production, area harvested, and yield for
the period 201012 are given in Table 1. World wheat produc-
Cereals, total
tion decreased by 49 Mt in 2012 from the previous year
2010 693 3571 2547
2011 707 3660 2589 (Table 1). The area harvested declined by 4 million hectares
2012 703 3622 2474 (Mha) and yields were 60 kg ha 1 less than in 2011. The
Wheat average global wheat production increased tremendously dur-
2010 217 3003 675 ing the period 196080 largely due to an increase in yield. In
2011 221 3175 701 developed countries, wheat yields tripled between 1950 and
2012 217 3115 652 1996 due to new wheat varieties and improved technologies
Rice, paddy including methods of sowing, irrigation, fertilizer application,
2010 162 4336 718 moisture retention, and integrated pest management. Potential
2011 163 4429 723
yields are still to be realized in African and Asian developing
2012 163 4395 701
countries due to several constraints including inadequate
Maize
2010 164 5187 850 water, infertile soils, poor weather, and lack of inputs. The
2011 172 5154 885 major wheat-producing countries during 2012 by regions are
2012 177 4944 875 EU (France, Germany, and the United Kingdom), FSU (the
Barley Russian Federation and Ukraine), NC America (Canada),
2010 47 2606 124 South America (Argentina), Oceania (Australia), and Asia
2011 48 2747 133 (China, India, Pakistan, Turkey, Iran, and Kazakhstan).
2012 49 2684 132 Bread wheat is planted on 93% of world wheat-growing
Sorghum area while durum and soft wheat occupy the remainder. Spring
2010 41 1439 59
types are sown on two-thirds of the land devoted to wheat in
2011 42 1384 59
developing countries.
2012 38 1535 58
Millet The five major wheat exporters are France (EU), the United
2010 36 905 32 States and Canada (NC America), the Russian Federation
2011 34 803 27 (FSU), and Australia (Oceania) (Figure 3(b)). India and
2012 31 820 26 Pakistan produce enough wheat to meet their country needs.
Oats Developing countries in Africa, Asia, and South America
2010 9 2163 20 accounted for at least 50% of world wheat imports in 2009
2011 10 2348 23 and 2010 (Figure 4).
2012 10 2179 26 Figure 5 shows consumption of wheat in the EU, FSU, NC
Rye
America, South America, NE Asia, FE Asia, and Africa in 2009.
2010 5 2374 12
Most wheat is used for human consumption with a limited
2011 5 2565 13
2012 6 2617 15 amount going into livestock feed. World wheat production and
consumption are projected to increase in 2013 and 2014.
Source: FAOSTAT Database 2013. Exports to China are expected to increase in 2013 due to a
GRAINS AROUND THE WORLD | Grain Production and Consumption: Overview 351
700
600
400
300
100
0
1961-70 1971-80 1981-90 1991-2000 2001-10
700
600
500
400
300
200
100
0
1961-70 1971-80 1981-90 1991-2000 2001-10
700
600
500
400
300
200
100
0
1961-70 1971-80 1981-90 1991-2000 2001-10
800
700
600
500
400
300
200
100
0
1961-70 1971-80 1981-90 1991-2000 2001-10
Figure 2 World cereal production, averages for 10-year periods. FAOSTAT Database 2013.
poor quality domestic crop. Wheat dry milling produces flour, bran, a by-product of milling, is used in high-fiber foods and as
semolina, bran, and germ. The primary products of milling are an animal feed ingredient. Wheat is also used as a valuable feed
used largely in baking and extrusion to produce breads and ingredient for milk and beef livestock as it is more nutritious
pastas. Wheat germ oil and meal are valuable coproducts. The than maize, sorghum, and barley.
Other
ME Asia
0% Africa
6% Africa S America Other
ME Asia 1%
FE Asia 1% 10% 1%
FE Asia 7%
6%
6%
S America
12%
EU
25% NC America EU
31% 26% NC America
33%
FSU FSU
15% 8%
Oceania Oceania
(a) 6% 6%
S America
ME Asia Other S America Other
ME Asia 5%
3% 1% 5% 1%
5%
Africa
0% Africa
FE Asia 0%
0% FE Asia
EU
0%
32% EU
NC America
35% NC America
29%
32%
FSU FSU
20% Oceania 11%
(b) 10% Oceania
11%
FE Asia FE Asia
53% 51%
(c)
ME Asia Africa
Other 2% ME Asia Other Africa
3%
FE Asia 3% 2% 2% 2%
2%
FE Asia
1%
EU
EU S America FSU 15%
16% 18% 3% S America
27%
Oceania
FSU 0%
8%
NC America NC America
Oceania
48% 48%
0%
(d)
Figure 3 (a) World exports of cereals by region for 2009 (330 Mt) and 2010 (334 Mt); regional totals exclude processed secondary products.
(b) World exports of wheat by region for 2009 (146 Mt) and 2010 (145 Mt); regional totals exclude processed secondary products. (c) World exports of
milled rice by region for 2009 (30 Mt) and 2010 (33 Mt); regional totals exclude processed secondary products. (d) World exports of maize (corn)
by region for 2009 (101 Mt) and 2010 (108 Mt); regional totals exclude processed secondary products. FAOSTAT Database 2013.
GRAINS AROUND THE WORLD | Grain Production and Consumption: Overview 353
Other
Other 3%
4%
Africa Africa
19% ME Asia 20%
ME Asia
18%
20%
S America S America
7% 8%
NC America
NC America FE Asia
FE Asia 8%
8% 23%
21%
Oceanla
EU Oceanla EU
FSU 0%
21% 0% 20%
FSU 0%
(a) 0%
Other
Other
3%
3%
Africa
ME Asia Africa
ME Asia 24%
17% 26%
23%
S America FE Asia
FE Asia 7% 17%
15% S America
NC America 9%
EU 5% EU
23% Oceanla NC America
23%
0% 5%
FSU FSU Oceanla
(b) 0% 0% 0%
Other
Other 4%
4%
Africa Africa
ME Asia 31% ME Asia 29%
23% 24%
S America
FE Asia 4%
S America FE Asia
18% NC America
EU 4% 21% EU
10% NC America 9% 6%
7% Oceanla
FSU Oceanla FSU
(c) 1% 2% 1% 2%
Other Other
2% 3%
Africa Africa
ME Asia ME Asia 13%
14% S America
12% S America 13% 8%
9%
NC America NC America
12% 12%
FE Asia
33% FE Asia
35% Oceanla
EU 0%
EU Oceanla FSU
16%
18% 0% 0%
FSU
(d) 0%
Figure 4 (a) World imports of cereals by region for 2009 (328 Mt) and 2010 (335 Mt); regional totals exclude processed secondary products.
(b) World imports of wheat by region for 2009 (147 Mt) and 2010 (144 Mt); regional totals exclude processed secondary products. (c) World imports of
milled rice by region for 2009 (30 Mt) and 2010 (31 Mt); regional totals exclude processed secondary products. (d) World imports of maize by
region for 2009 (100 Mt) and 2010 (107 Mt); regional totals exclude processed secondary products. FAOSTAT Database 2013.
354 GRAINS AROUND THE WORLD | Grain Production and Consumption: Overview
100%
80%
60%
40%
20%
0%
EU
ica
ica
ia
ia
ca
ld
FS
As
As
or
ri
er
er
Af
W
Am
Am
FE
M
NC
S
(a)
100%
80%
60%
40%
20%
0%
EU
ica
ica
ia
ia
ld
ric
FS
As
As
or
er
er
Af
W
Am
Am
FE
M
NC
(b)
100%
80%
60%
40%
20%
0%
EU
ica
ica
ia
ia
ld
ric
FS
As
As
or
er
er
Af
W
Am
Am
FE
M
NC
(c)
Rice
doubled from 216 to 448 Mt during the period 196183. The
World production of paddy rice (unmilled or rough rice) was increase in average rice production was 15% between the
701 Mt in 2012, a decrease of 22 Mt from the previous year. 10-year periods 198190 and 19912000 and 19912000
The area harvested remained the same but yields decreased by and 200010 compared with 30% for the periods 197180
34 kg ha 1 during the same period. World rice production and 198190 (Figure 2).
GRAINS AROUND THE WORLD | Grain Production and Consumption: Overview 355
Most of the rice is grown in developing countries. In 2012, exporters are the United States, Argentina, Brazil, France, Hun-
Asia was responsible for 93% of the world rice production, gary, and Ukraine. The United States accounts for almost half
51% of the total crop being produced by China and India. of the world maize exports. China supplies grain to its neigh-
Only small amounts (< 10%) of world rice produced are boring countries. Japan and the Republic of Korea are the
traded internationally. World exports of rice by region for largest importers, each importing 16 and 9 Mt, respectively,
2009 and 2010 are given in Figure 3(c). The major rice in 2010.
exporters include India, Thailand, Vietnam, the United States, Globally, the total domestic supply of maize was 799 Mt in
and Pakistan. Many countries in NC and South America, 2009. The primary product of maize milling is the meal used
Europe, and Africa are net importers (Figure 4(c)). Total rice for feed and food purposes. At least 55% of world maize
consumption (milled equivalent) for 2009 in Mt was 3 production is used to feed livestock and 14% is used for
(EU15), 1 (FSU), 5 (NC America), 15 (South America), 130 human food. Other uses include industrial processing and
(ME Asia), 245 (FE Asia), and 21 (Africa), giving a total of 423 seed. Secondary and derived products are obtained through
(world). Global rice trade is predicted to remain the same in dry-milling and wet-milling processes. Global maize trade is
2013 and 2014 as there are limited opportunities for expan- forecast to reach a 6-year high in 2013 as the demand in China
sion. Both production and consumption are expected to is increasing.
increase slightly worldwide.
Barley
Coarse Grains
World barley production was 132 Mt in 2012, down by 1 Mt
World production of coarse grains (including maize (corn), from 2011 (Table 1). The area planted has steadily increased by
barley, sorghum, oats, rye, millet, triticale, buckwheat, and 1 Mha from 2010 to 2012 but yields declined by 63 kg ha 1
quinoa) was 1153 Mt in 2012 versus 1121 Mt in 2010. Among from 2011 to 2012. In 2012, the major producers by region
the coarse grains, maize, barley, and sorghum are the most included the EU (Germany 10, France 11, and Spain 6), FSU
important in terms of total cereal production. World produc- (Russian Federation 14), NC America (Canada 8 and the United
tion of maize, barley, and sorghum was 875, 132, and 58 Mt, States 5), Asia (Turkey 7), and Oceania (Australia 8). The EU is a
respectively, in 2012 (Table 1). Production of other coarse major exporter of barley. The Russian Federation and Ukraine
grains was below 30 Mt each. A record of 1295 Mt, an increase have become competitors with the EU in export markets.
of 11.4% from 2012 in global production of coarse grain in Exports from the EU also face competition from Australia
2013, has been attributed to a strong recovery in maize pro- and Canada especially for supply of malting barley. ME Asia
duction in the United States following the 2012 drought- and FE Asia imported 10 and 4 Mt, respectively, of the total 25
reduced level. Leading producers in coarse grain production Mt traded in 2010. Saudi Arabia, China, and Japan were the
include the United States, China mainland, EU, Brazil, India, major Asian importers along with European countries Germany,
Argentina, the Russian Federation, Ukraine, Mexico, Canada, Belgium, and the Netherlands. Global domestic supply of barley
Nigeria, Indonesia, Ethiopia, Turkey, and South Africa. Major totaled to 140 Mt in 2009. Feed use accounted for 65% of barley
exporters of coarse grains are Argentina, Australia, Brazil, consumption while food use remained very low (4.5%).
Canada, EU, Russian Federation, Ukraine and the United
States.
Sorghum
Maize The area under production for sorghum was 38 Mha in 2012
compared with 42 Mha in 2011 (Table 1). Global yield
Maize composes 35% of total cereal production in the world. increased by 151 kg ha 1 from 2011 to 1535 kg ha 1 in
Maize is the preferred crop in Africa and South America. In 2012. World sorghum production decreased 1 Mt from 2011
Asia, it occupies the third position after rice and wheat. World to 58 Mt in 2012. Nigeria, Mexico, the United States, India,
maize production fell by 10 Mt from 885 Mt in 2011 (Table 1). Argentina, China, and Ethiopia are the leading producers. Most
The area harvested in 2012 was 177 Mha. Yields have remained sorghum is produced by small-scale and subsistence farmers in
relatively low in developing countries (1980 kg ha 1) com- semitropical regions of Africa and Asia and by other farmers in
pared with the global average (4944 kg ha 1) due to environ- the United States and Latin America. Africa produced 40% of
mental, technological, and socioeconomic factors. Developed the world total in 2012 on 23 Mha, more than half the world
countries make use of adequate inputs and a well-mechanized total area. India has the largest area harvested for sorghum,
system for maize production. In 1961, world maize production although there has been a significant reduction from 16 Mha in
was 205 Mt. The figure rose to 615 Mt in 1998. After the 1989 to 6.3 Mha in 2012. Production decreased in China from
1998 peak in world maize production, rising production costs a peak of 6 Mt in 1994 to 2 Mt in 2012 due to a decline in area
and shortage of foreign exchange in many developing countries harvested. The United Arab Emirates had the largest average
have led to diminished production and, consequently, maize yield in 2012 at 84 667 kg ha 1. World trade of sorghum was
trade. There was a 50% increase in average global maize pro- 13 Mt in 2010 with the United States and Argentina supplying
duction between the periods 196170 and 197180 (Figure 2). 3.9 and 1.7 Mt to the export market, respectively. In 2010,
World exports and imports of maize by region for 2009 and Mexico and Japan imported 2.3 and 1.6 Mt, respectively.
2010 are given in Figures 3 and 4, respectively. The main maize Mexico, the most important producer of sorghum at 7 Mt in
356 GRAINS AROUND THE WORLD | Grain Production and Consumption: Overview
2012, is also the largest importer. Chile, Ethiopia, the Republic respectively, in 2012. These grains composed 1% of total
of the Sudan, and Spain are also main importers. Sorghum is cereal production. Germany, Poland, the Russian Federation,
also an important commercial and export crop for Australia and Belarus are the major producers of rye. World buckwheat
and Argentina. The crop is utilized for feed (43%), food (44%), production increased between 2010 and 2011 as area har-
and other uses (13%). Mexico, the United States, Australia, vested and yields grew slightly. Decreases in area harvested
Brazil, Japan, and Nigeria are principal feed users. resulted in a decline in global canary seed production between
2010 and 2011. Worldwide production of both canary seed
and buckwheat remained steady between 2011 and 2012.
Millets
World quinoa production has steadily increased while global
triticale and fonio production has remained fairly constant
World production of all millets was 26 Mt in 2012 (Table 1),
from 2010 to 2012. Canada contributed 62% of world
46% of which was produced in Africa. West Africa (Nigeria,
canary seed production in 2012. Canary seed is currently
Niger, Burkina Faso, Chad, Mali, and Senegal) produces 75%
used almost exclusively as birdseed. Rye, triticale, buckwheat,
of the millet output in Africa. Asia and Africa account for
fonio, and quinoa are used for human food mostly as ingredi-
96% of the global output of millet. India, China, Niger,
ents in baked products. It is projected that the demand for
Burkina Faso, and Nigeria are the leading producers of millet.
these grains will grow as whole grains continue to be promoted
Small-scale farmers produce almost all millet for household
as healthy foods.
consumption and localized trade. Very little millet (0.9 Mt or
3.5% of world millet production) is traded internationally.
India, the United States, China, Ukraine, and Austria are the
major exporters of millet. The European community accounts Pulses
for more than 23% of the global imports. Millet is primarily
dry-milled to produce cracked grain, grits, meal, and flour from Table 2 gives data on world pulse production for the period
which a number of secondary and derived products are made. 201012. Among pulses, beans were produced at 23.2 Mt
followed by chickpeas (10.9 Mt) and peas (10.2 Mt) in
2010. Asia contributes 45% of world pulse production.
Oats
Pulse production quadrupled in NC America and Africa since
1961. In the EU, pulse production peaked in 1990 at 7.4 Mt
World oat production in 2012 was 21 Mt, down by 2 Mt from
and steadily declined to 3.2 Mt in 2012. India, Brazil, and
2011 due to a decrease in crop yields (Table 1). The total area
Myanmar are the leading producers of dry beans. The major
harvested was 10 Mha with an average yield of 2179 kg ha 1.
chickpea producer is India followed by Australia, Turkey,
World oat production for 2012 by region, in Mt, was 7.9 (EU),
Myanmar, and Ethiopia. Canada leads in both production
4.7 (FSU), 3.7 (NC America), 1.3 (Oceania), 0.6 (FE Asia), and
and export of dry peas in the world. Russia, France, China,
0.3 (ME Asia). Russia and Canada are the leading producers
and India are also significant producers. World lentil produc-
followed by the United States and Australia. Canada is the
tion is 4.6 Mt with Canada (1.9 Mt) and India (0.9 Mt) as
worlds largest exporter with the United States accounting for
leading producers. India is a major importer of dry peas and
over 90% of Canadas oat exports. The next largest export
dry beans. Over 70% of pulse crops are consumed as human
regions are the EU (mainly Finland and Sweden) and Australia.
food globally. Peas are used for both livestock feed and human
Russia generally consumes its feed quality oats domestically
food. The proportion for the latter use is higher in Asia and
even though it is the worlds largest single producer of oats. The
Latin America than in Europe.
United States dominates global oat imports, accounting for
80%, followed by Germany, Mexico, and Japan. Oats is used
primarily for animal feed (71%) and human food (16%). High
levels of soluble fiber in oats have contributed to its promotion Oilseeds
as a healthy food ingredient.
Major oilseeds produced include soybean, rapeseed, cotton-
seed, groundnuts, sunflower seed, palm kernels, and copra.
Rye, Triticale, Buckwheat, Fonio, Quinoa, World oilseed production was 454 Mt in 2012 with soybean,
and Canary Seed rapeseed/canola, cottonseed, groundnuts (unshelled), sun-
flower seed, and palm kernels making up 53, 13, 10, 8, 8.6,
World production of rye, triticale, buckwheat, fonio, quinoa, and 3%, respectively, of the total production. World oilseed
and canary seed was 13, 14, 2.3, 0.6, 0.08, and 0.2 Mt, production has risen tremendously since the 1970s due to the
Table 2 World pulse and oilseed production (Mt) for the period 201012
Year Total pulses Dry beans Dry peas Chickpeas Cow peas Pigeon peas Soybean Rapeseed
2010 70 23 10 11 7 4 265 60
2011 68 23 10 12 5 4 262 63
2012 70 23 10 11 6 4 253 65
300
250
Production (Mt)
200
150
100
50
0
1960 1970 1980 1990 2000 2010
Year
Figure 6 World soybean production for the period 19612012. FAOSTAT Database 2013.
358 GRAINS AROUND THE WORLD | Grain Production and Consumption: Overview
1200
1000
Population (millions)
800 Total
600
Rural
400
Urban
200
0
1965 1970 1975 1980 1985 1990 1995 2000 2005 2010 2015
Year
Figure 1 Africas population growth from 1965 to 2015. Data from FAOSTAT: http://faostat.fao.org. Population (accessed 21.04.14.).
Cereals
Cereals (total) 172 184 106 955 1.61 Nigeria, Egypt, Ethiopia, South Africa
Barley 6023 (3.5)a 4997 1.21 Ethiopia, Algeria, Morocco
Fonio 587 (0.3) 566 1.04 Guinea, Nigeria
Maize 69 637 (40.4) 33 710 2.08 South Africa, Nigeria, Egypt, Ethiopia
Millet (mainly pearl millet) 16 083 (9.3) 19 978 0.81 Nigeria, Niger, Mali
Oats 276 (0.2) 199 1.39 Algeria, South Africa, Morocco
Rice (paddy) 26 824 (15.6) 10 538 2.54 Egypt, Nigeria, and Madagascar
Rye 94 (<0.1) 52 1.80 Egypt
Sorghum 23 313 (13.5) 23 169 1.01 Nigeria, Ethiopia, Burkina Faso, and Sudan
(former)
Triticale 26 (<0.1) 13 2.01 Tunisia
Wheat 24 711 (14.4) 10 217 2.42 Egypt, Morocco, Ethiopia, and Algeria
Pseudocereals
Buckwheat 10 11 0.97 Tanzania
Pulses (grain legumes)
Pulses (total) 15 865 24 447 0.65 Ethiopia, Nigeria, Tanzania, and Niger
Bambara bean 159 (1.0)b 233 0.68 Burkina Faso, Niger, Cameroon, and
Mauritania
Dry beans (common beans) 5070 (32.0) 7829 0.65 Tanzania, Kenya, Ethiopia, and Rwanda
Broad bean 1505 (9.5) 670 1.55 Ethiopia
Chickpea 697 (4.4) 595 1.17 Ethiopia
Cowpea 5403 (34.1) 11 043 0.49 Nigeria, Niger
Lentil 187 (1.2) 188 0.99 Ethiopia
Peas (dry) 596 (3.8) 715 0.83 Ethiopia and Tanzania
Pigeon pea 632 (4.0) 754 0.84 Malawi and Tanzania
Pulses (other) 1262 (8.0) 1890 0.67 Mozambique, Sudan (former)
Oil-rich legumes
Peanuts (groundnuts) 10 942 11 823 0.93 Nigeria, Sudan (former), and Tanzania
(in shell)
Soybean 1986 1638 1.21 South Africa and Nigeria
Oilseeds
Rapeseed 201 113 1.79 South Africa, Ethiopia, and Algeria
Sunflower seed 2224 1780 1.24 Tanzania and South Africa
a
Percentage of cereal production.
b
Percentage of pulse production.
Data from FAOSTAT http://faostat.fao.org. Production (accessed 21.04.14.).
GRAINS AROUND THE WORLD | Grain Production and Consumption: Africa 361
Production takes place throughout Africa, with 47 countries with an 73% increase over the entire period. In contrast,
recorded as significant producers. However, Nigeria produces yield initially increased over the first 10 years and then
by far the largest quantity of peanuts, 3.1 Mt. Significant remained static until the mid-1990s and has subsequently
quantities of oil-bearing grains other than peanuts are also increased substantially in recent years. Over the whole period,
produced in Africa. The most important are sunflower and yields increased by 83%. Increased area of harvest and
soybean, with a total production of 2.2 and 2.0 Mt, respec- increased yields have resulted in a continuous increase in cereal
tively. There is also some production of rapeseed (canola). production, which has accelerated since the mid-1990s. Over-
South Africa is the major producer of both soya and canola. all, cereal production increased 3.2-fold, keeping pace with
but not exceeding the increase in population 3.1-fold. As a
consequence, Africas cereal imports have grown at a rapid but
Grain Agriculture linear rate over the period, increasing from 11% to 34% of
cereal production. Imports of maize now represent some 26%
Figure 3 shows the trends in cereal production and imports for of local production (Table 2).
Africa over a 45-year period since the mid-1960s when most The need to import an increasing proportion of cereal
African countries gained independence, to 2010. Over the grains is related to the fact that much of Africas agriculture is
period, the area harvested remained more or less constant up still subsistence farming. However, small-scale commercial and
until the about 1980 and then steadily but gradually increased, large-scale mechanized farming are widespread and increasing.
180 1.80
160 1.60
Production/area harvested/imports (Mt)
140 1.40
120 1.20
Yield (tonnes/Ha)
100 Yield 1.00
80 Area 0.80
Production harvested
60 0.60
40 0.40
Imports
20 0.20
0 0.00
1965 1970 1975 1980 1985 1990 1995 2000 2005 2010
Year
Figure 3 Africas cereal production and imports from 1965 to 2010 (data from FAOSTAT5). Data from FAOSTAT. Production: Crops http://faostat.fao.
org/ (accessed 21.04.14.).
Table 2 Africas domestic grain-based food and beverage supply (2009) (t 103)
Cereals (total) 146 217 55 987 38.3 4065 2.8 1163 199 303 136.3
Maize 57 702 14 995 26.0 2132 3.7 3498 67 067 116.2
Wheat 26 074 30 288 116.2 897 3.4 126 55 339 212.2
Barley 5827 1744 29.9 29 0.5 306 7237 124.2
Lager and stout beers 9758 510 5.2 188 1.9 199 9882 101.3
Pulses (total) 12 086 1262 10.4 577 4.8 322 12 803 105.9
Peanuts (groundnuts) 9604 227 2.4 148 1.5 16 9670 100.7
(in shell)
Soybeans 1612 1133 70.3 192 11.9 170 2722 168.9
Sunflower seed 1689 175 10.4 14 0.8 116 1734 102.7
Groundnut oil 1374 9 0.7 48 3.5 16 1319 96.0
Soybean oil 300 1449 483.0 127 42.3 87 1535 511.7
Sunflower seed oil 597 596 99.8 90 15.1 132 971 162.6
Figure 5 Selection of African grain foods and beverages. (a) Kisa flatbread (sorghum) Sudan, (b) to porridge (maize) with chickpea relish and
goat meat Burkina Faso, (c) Ugali porridge (sorghum) Kenya, (d) Baladi bread (wheat) Sudan, (e) Fonio couscous and peanut sauce (Burkina Faso),
(f) Mealie bread (maize and wheat) South Africa, (g) Samp and beans (maize and dry beans) southern Africa, (h) Opaque sorghum beer
(maize and sorghum) southern Africa, (i) Mageu fermented nonalcoholic beverage (sorghum and maize) Zimbabwe and South Africa.
dry roasted grain milled into powder, spiced, and made production may seem surprising. Part of the explanation is that
into a sauce, for example, shiro, pea, broad bean, or chick- a substantial proportion of these beers are now brewed using
pea (Ethiopia). locally grown sorghum as either the sole or major ingredient,
especially in West and East Africa.
However, food consumption patterns in Africa are changing
rapidly as a result of urbanization. The so-called nutrition
transition is resulting in a dramatic increase in consumption
of Western-style convenience foods, including bread, potato Developments and Challenges in Grain Agriculture
chips (French fries), fried chicken, and beef burgers, and bev- and Grain Handling
erages, in particular carbonated soft drinks and lager beer. For
example, with regard to convenience foods, the largest South The high dependence on imported grains has a very adverse
African fast-food franchise company opened 18 restaurants in effect on the fragile economies of many African countries. To
other African countries in the last quarter of 2012. In the case prevent increasing poverty and malnutrition resulting from
of lager beer, the figures are if anything even more dramatic. rapid population growth and urbanization, it is essential that
From 2007 to 2012, the largest brewing company in Africa the effectiveness of African grain agriculture is improved. How-
recorded an average annual 12% increase in beer volume. ever, the improvement of agriculture in Africa faces a number
These changing food consumption patterns are severely of severe challenges. These include the acute need to introduce
impacting on the grain food supply situation in Africa. With modern agricultural production technologies and chronic
regard to cereals, the continent is importing as much wheat as environmental problems such as decreasing soil fertility and
is produced locally (Table 2). In the case of barley, imports desertification, resulting from farming more marginal land.
amount to 30% local production. The situation with regard There are also sociopolitical issues. Governmental policies are
to soybeans, the most important protein source in formulated often not conducive to local agricultural development. As of
animal feeds, is worse, with domestic supply being 170% of 2013, only eight countries in Africa had met the UN Millen-
production. With regard to seed oils, which are majorly used to nium Development Goal of halving the proportion of hungry
manufacture plant-based fats like margarine, the situation is people. Persistent problems include keeping food prices artifi-
even more dire. Africa is importing nearly five times the quan- cially low to subsidize the urban consumer, but at the cost of
tity of soy oil it produces and as much sunflower seed oil as it the farmers livelihood, and poorly planned and executed land
produces. Taking the previously mentioned into account, the reforms that disrupt large-scale commercial farming
fact that Africa is essentially self-sufficient in lager and beer operations.
GRAINS AROUND THE WORLD | Grain Production and Consumption: Africa 365
Explain what is the impact of rapid population growth and Popkin BM (2003) The nutrition transition in the developing world. Development Policy
urbanization on food security in Africa. Review 21: 581597.
Explain the concept of biofortification and its benefits. Rohrbach DD (2003) Improving the commercial viability of sorghum and pearl millet in
Africa. Afripro Paper 22. http://www.afripro.org.uk (accessed 21.04.14).
Saltzman A, Birol E, Bouis HE, et al. (2013) Biofortification: Progress toward a more
nourishing future. Global Food Security 2: 917.
Taylor JRN, Dlamini BC, and Kruger J (2013) 125th Anniversary Review: The science of
Exercises for Readers to Explore the Topic Further the tropical cereals sorghum, maize and rice in relation to lager beer brewing.
Journal of the Institute of Brewing 119: 114.
Improving smallholder agriculture productivity in Africa Van Wyk B-E and Gericke N (2000) Peoples Plants: A Guide to Useful Plants of
using appropriate technologies. Southern Africa. Pretoria: Briza.
Topic Highlights of Indian agriculture can be divided into six phases viz. pre-
green revolution period (196061 to 196869), early green
Agriculture accounts for considerable importance in Indias revolution period (196869 to 197576), period of wider
economic development, as it provides food for more than technology dissemination (197576 to 198889), period of
1.2 billion people and total employment to about 54.6% of diversification (198889 to 199596); postreform period
the population. (199596 to 200405), and period of recovery (200405 to
India holds the second-largest agricultural land (179.9 201011). The green revolution period was the golden period
million hectares) in the world. for Indian agriculture, as it witnessed tremendous growth in
Food grain production covers the dominant part of the agricultural production. During the period of wider dissemi-
cropped area (65%) in Indian agriculture. nation of technology, the spread of green revolution technol-
India is the worlds largest producer of millets and second- ogies pushed to maintain the increasing rhythm of growth
largest producer of wheat, rice, and pulses. across various regions realized during the previous period.
The highest food grain production of 259.32 million However, a visible deceleration of growth was experienced
tonnes was recorded in 201112. during the postreform period, as both public and private
The cereals other than rice and wheat contributed 1 kg per investments suffered a setback, which resulted in the sluggish
person per month in rural India and 0.8 kg per person per performance of the sector. Moreover, yield levels of most of the
month in urban India in 201112. crops suffered a deceleration because use of primary resources
The consumption of all food grains per person in the rural in the sector also slowed down. The descending trend of
areas is projected to decline from 15.3 kg per month in growth continued up to 200405, after which a sharp return
2000 to 13.8 kg per month by 2050 and only slightly was attained that can be attributed to a conscious increment in
decrease from 11.8 to 11.6 kg per month in the urban areas. public and private ventures and substantial improvement in
The nongrain food crops and animal products contributed trade in favor of the agricultural sector.
27% and 8%, respectively, of the total caloric requirement.
Meat consumption in India is very low due to cultural and
religious reasons, and most of the calories from animal
products are obtained from milk and milk products. Food Grain Production
300
200
150
100
50
0
2010-11 2011-12 2012-13
Year
Figure 1 Production of food grains in India.
40
33.8
35
Area (percentage)
30
25
20
14.8
15 12.7
9.7 8.7
10 7.8
3.5 2.8
5 1.8 1.5 1.5 1.4
0
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at
sh
sh
ab
na
s
ha
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ha
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es
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ar
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en
th
Bi
ad
st
uj
as
Pu
ar
ra
ra
O
ra
tB
ja
G
Pr
ar
H
lP
rP
Ra
tta
ah
es
a
ha
tta
U
hy
ac
U
ad
im
M
States
Figure 2 Share of states in India in terms of area under cultivation of wheat.
such as Iran, Saudi Arabia, Indonesia, UAE, and Bangladesh Uttaranchal, Himachal Pradesh, and Jammu and Kashmir
mainly imported cereals from India during 201314. (Figure 3). These states contribute about 99.5% of total
wheat production in the country. The remaining states, namely
Jharkhand, Assam, Chhattisgarh, Delhi, and other
Wheat northeastern states, contribute only about 0.5% of the total
The area and production under wheat has increased from wheat production in the country. Among food grains, wheat in
27.99 million hectares in 200607 to 29.86 million hectares India stands next to rice both in area and production.
in 201112 and from 75.81 million tonnes in 200607 to an
all-time record high of 94.88 million tonnes in 201112,
Rice
respectively. Figure 2 shows the percent share of states in
India in terms of area. Uttar Pradesh, Madhya Pradesh, Punjab, India is the second-largest producer and consumer of rice in
Rajasthan, and Haryana are the leading states in terms of area the world and accounts for 22.3% of global production. Sev-
under wheat cultivation. Uttar Pradesh accounts for the maxi- eral programs run by the government such as National Food
mum area under cultivation and production, whereas Punjab Security Mission (NFSM) launched during 200708 and Bring-
and Haryana are yielding higher productivity with lesser land- ing Green Revolution in Eastern India (BGREI) during
holding under wheat cultivation. 201011 increased production and productivity of rice from
The increase in production of wheat has been observed in 96.7 million tonnes and 2202 kg ha 1 in 200708 to 105.31
the states of Haryana, Punjab, Madhya Pradesh, and Uttar million tonnes and 2393 kg ha 1, respectively, in 201112.
Pradesh. The normal national productivity is about 2.9 The NFSM is presently under implementation in 482 dis-
tonnes per hectare. The major wheat-producing states are tricts of 19 states of the country with a view to enhance the
Uttar Pradesh, Punjab, Haryana, Madhya Pradesh, Rajasthan, production of rice, wheat, and pulses through area expansion
Bihar, Maharashtra, Gujarat, Karnataka, West Bengal, and productivity enhancement, restoring soil fertility and
GRAINS AROUND THE WORLD | Production and Consumption of Grains: India 369
40
33.7
35
Production (percentage)
30
25 21.2
20
15 12.9
9.6 8.9
10 6.2
5 1.7 1.6 1.2 1 0.8 1.2
0
ab
na
ra
al
sh
rs
at
ga
ha
es
es
ha
ch
e
ht
ar
de
ya
nj
en
th
Bi
ad
ad
st
as
uj
an
Pu
ar
ra
O
tB
ja
G
Pr
Pr
ar
r
H
lP
Ra
tta
ah
es
r
ha
tta
U
hy
ac
U
ad
im
M
H
States
Figure 3 Share of states in India for production of wheat.
25
Production (million tonnes)
20
15
10
5
4
-0
-0
-0
-0
-0
-1
-1
-1
-1
-1
04
05
06
07
08
09
10
11
12
13
20
20
20
20
20
20
20
20
20
20
Year
Figure 4 Status of maize production in India.
productivity, creating employment opportunities, and enhanc- improving, and its cultivation has spread to other parts of
ing farm level economy to restore confidence of farmers. The Uttar Pradesh and Madhya Pradesh.
basic strategy of the mission is to promote and extend
improved technologies that is, seed, micronutrients, soil
Maize
amendments, integrated pest management, farm machinery,
and resource conservation technologies, along with capacity After rice and wheat, maize is emerging as the third-most-
building of farmers with effective monitoring and better man- important crop in India. Its importance lies in the fact that it
agement in the high-potential districts to bridge the yield gaps. is used for human food as well as animal feed, and it is also
BGREI aims to increase rice productivity by intensive culti- widely used for the starch industry, oil production, baby corns,
vation through promotion of recommended agriculture tech- and so on. However, despite the production strength, Indian
nology and a package of practices by addressing the underlying corn yields are significantly below the yields in major corn-
constraints of different agroclimatic subregions. It is under producing countries. There is immense scope for an increment
operation in seven states including Uttar Pradesh, Jharkhand, in Indias corn production by increasing the area under
Bihar, West Bengal, Assam, Orissa, and Chhattisgarh. Rice hybrids, adoption of better genetics, and improved agronomic
contributes to more than 40% of total food grain production practices. Maize is primarily a kharif crop (summer seasons
and is cultivated throughout the country. Rice production in crop) with 85% of the area under cultivation and accounting
India shows a steady upward trend, but it is subjected to wide for 9% of total food grain production. In India the CAGR
year-to-year fluctuations compared to wheat, as a significant (cumulative annual growth rate) of maize production has been
portion of the crop is not irrigated. Indian Basmati rice is 5.5% over the last 10 years from 14 million tonnes in 200405
traditionally grown in Punjab, Haryana, and western Uttar to 23 million tonnes in 201314 (Figure 4). There was a
Pradesh. With the introduction of high-yielding PUSA 1121 decline in the production of maize during 200910 due to
variety, Indias long-grain basmati rice production has been drought conditions in India.
370 GRAINS AROUND THE WORLD | Production and Consumption of Grains: India
6 1.5
5
4
-0
-0
-0
-0
-0
-1
-1
-1
-1
-1
04
05
06
07
08
09
10
11
12
13
20
20
20
20
20
20
20
20
20
20
Years
Figure 5 Yield and area under cultivation of maize.
3000
2500
2000
1500
1000
500
0
2007-08 2008-09 2009-10 2010-11 2011-12 2012-13
Year
Figure 6 Area, production, and yield of barley in India.
In Figure 5 area under maize cultivation, yield, and CAGR of habits and lifestyle has reduced the demand for barley for
area are mentioned. During the last decade (200405 to human consumption, thereby causing the area in cultivation
201314), CAGR of the area under maize cultivation has of barley to plummet during last few decades. Barley is utilized
increased 2.5%, and cultivated area increased from 7.5 million for beer, food processing, and feed industries apart from direct
hectares in 200405 to 9.4 million hectares in 201314. Factors human consumption in India. Recently, a rise in demand for
such as adaptability to diverse agroclimatic conditions, lower beer in urban India has directly increased the demand for
labor costs, and lowering of the water table in the rice belt of barley malt from Indian beer manufacturing industries.
India have contributed to the increase in acreage of maize. The country is self-sufficient in barley production, although
Productivity of maize has accelerated at a CAGR of 2.9% from production is low, besides exporting 0.10.2 million tonnes of
1.9 million tonnes per hectare in 200405 to 2.5 million tonnes barley annually. In recent years, Indias annual production has
per hectare in 201314. Yield of maize increased by 20% due to been stagnant at 1.21.5 million tonnes, with production in
introduction of single-cross hybrid seeds coupled with adequate 201213 at 1.74 million tonnes against 1.61 million tonnes
rainfall in 200708. In India 85% of maize production and 80% reported last year. Similarly, the area under cultivation has also
of the area under cultivation come from nine states viz. remained stable at 600700 ha, with a per-hectare yield of
Karnataka, Andhra Pradesh, Tamil Nadu, Rajasthan, Maharash- 2508 kg (Figure 6).
tra, Bihar, Uttar Pradesh, Madhya Pradesh, and Gujarat. In India, major barley-growing states are Uttar Pradesh,
Rajasthan, and Madhya Pradesh, with a contribution of 34%,
30%, and 12%, respectively, in total acreage. Compositely,
these states hold about 80% of the whole acreage (Figure 7).
Barley
Rajasthan ranks second in terms of acreage under barley
Barley is the fourth-most-important and most adapted crop of and holds first place in terms of production due to a higher
all cereals. It is used for both feed and food. A shift in the food yield level. Rajasthan accounts for 40% of total production of
GRAINS AROUND THE WORLD | Production and Consumption of Grains: India 371
0
1961-71 1971-81 1981-91 1991-01 2001-11
Years
Figure 9 Growth rate of population and food grain production in India.
consumption to nongrain food and animal product consump- Exercises for Readers to Explore the Topic Further
tion. Rath suggested that the per capita grain consumption will
further decrease due to the reduction in physical labor require- Why are Indias food grain production and productivity still
ment in rural areas due to the adoption of more mechanized lagging behind?
and modernized agriculture. Furthermore, improving medical What would be the implications of economic development
facilities and infrastructure also contribute to the reduction in and urbanization on the food grain consumption patterns
energy requirements and thus less cereal consumption. It has of the Indian population?
been suggested that a reduction in the intake of food grains on What would be the effects of switching consumption pat-
this account should not be taken as deterioration in human terns on the future production estimates of grains?
welfare.
Sustained economic growth and changing lifestyles are
accounting for a significant shift in Indian food culture that is See also: Food Grains and the Consumer: Consumer Trends in
inclined toward high-value horticultural and animal products Grain Consumption; Cultural Differences in Processing and
and away from staple food grains. Rising per capita income and Consumption; Grains Around the World: Grain Production and
urbanization are leading to a dietary transition primarily from Consumption: Overview; The Basics: The Grain Chain: The Route
cereals with low-price calorie foods (foods such as wheat and from Genes to Grain-Based Products; The Grain Crops: An Overview;
rice with low price and rich in carbohydrates) toward high- The Cereal Grains: An Overview of the Family of Cereal Grains
price calorie foods like livestock products and processed or Prominent in World Agriculture; Barley: An Overview of a Versatile
ready-to-eat products. The transitions are also attributed to Cereal Grain with Many Food and Feed Uses; Maize: Overview; Rice:
farming patterns, as farmers are adopting new technologies Overview; Wheat: An Overview of the Grain That Provides Our Daily
that have resulted in an increase in farm productivity. Bread.
In accordance with Engels law, as the earnings of people
rise, the cheaper source of energy for living, which comes from
the staple food consumption, is likely to drop. A decline in
cereal consumption is substituted with the increased consump- Further Reading
tion of high-value commodities. Over time the shifting of
Amarasinghe UA, Shah T, and Singh OP (2007) Changing consumption patterns:
cereal consumption pattern will be prominent across all house- Implications on food and water demand in India. Colombo, Sri Lanka: International
holds in different expenditure groups. The devolution in cereal Water Management Institute, 43 p. (IWMI Research Report 119).
consumption can also be credited to the diversification of food Annual Report, 201112. Department of Agricultural Research (DARE) & Education/
production, easy access of high-value commodities, and Indian Council of Agricultural Research (ICAR).
Bansil PS (1999) Demand for foodgrains by 2020 AD. New Delhi: Observer Research
switching tastes and preferences. This agrees with the argu-
Foundation.
ments by Kumar and Mathur that the demand for food is not Directorate of Economics and Statistics, 1213. Department of Agriculture and
only influenced by the income of a person but also by differ- Cooperation, Ministry of Agriculture, Government of India, New Delhi.
ences in urban and rural lifestyles, the development of more Kumar P and Mathur VC (1996) Structural changes in demand for food in India. Indian
advanced marketing systems, and occupational changes are Journal of Agricultural Economics 51(4): 664673.
Kumar P and Mathur VC (1997) Agriculture in future: Demand and supply prospective.
closely linked with increasing per capita income. Tastes and In: Desai BM (ed.) Agricultural Development Paradigm for the Ninth Plan Under New
preferences are also moving toward high value commodities. Economic Development. New Delhi, India: Oxford and IBH Publishing.
With the availability of cash money and high-value commod- Mittal S (2007) What affect changes in cereal consumption? Economic and Political
ities in the food basket, the decline in calories by reduced cereal Weekly: 444447, February.
National Sample Survey (NSS)-68th round July 2011-June 2012. Household
consumption is partly compensated for by intake of highly
consumption of various goods and services in India, Government of India.
nutritive and vitamin-rich food products. Some other studies Radhakrishna R and Ravi C (1990) Food Demand Projections for India. Hyderabad:
have also attributed the change in per capita consumption of Centre for Economics and Social Studies.
cereals to economic growth, rise in per capita income, urban- Radhakrishna R and Ravi C (1992) Effects of growth, relative price and performances of
ization and preferences, market integration, and many more as food and nutrition. Indian Economic Review 27: 303323.
Rao CHH (2000) Declining demand for foodgrains in rural India: Census and
the dominating factors. Implications. Economic and Political Weekly, January 22.
Rath N (2003) Linking of rivers: Some elementary arithmetic. Economic and Political
Weekly: 30323033, July.
Exercise for Revision Singh SK (2014) Grain and feed annual, India. Global Agricultural Information Network
(GAIN) report-IN4005.
The Economic Survey 201415, Agricultural and Processed Food Products Export
How has Indian agriculture been successful in changing its Development Authority (APEDA).
face postindependence? The Tribune, Chandigarh, February 21, 2014. Sale of beer on the rise, scientists for
In what ways has the Indian government contributed to the contract farming of barley.
growth of agriculture in India?
What factors govern the changing consumption patterns of
food grains in India, and how is it changing? Relevant Websites
What is the total production of different grains in India, and
which are the major grain-producing states? faostat.fao.org FAOSTAT, data.
Grain Production and Consumption: China and South-East Asia
H Corke and YZ Cai, The University of Hong Kong, Hong Kong, Peoples Republic of China
2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition volume 2, pp 7786, 2004, Elsevier Ltd.
to seek lives with less physical hardship and better incomes populous nation in the not too distant future. The limited land
elsewhere. supply (an index of population density) in China is going to be
In a 1996 interview, K. Lampe, then Director General of matched across Asia as a whole as the population of the rest of
IRRI, summarized the problems of developing tropical Asian Asia continues to increase (Figure 1). Although the population
economies such as Philippines. density of India is greater than China, the supply of arable land is
also greater (Figure 2). Widely quoted figures are that China has
22% of the worlds population but only 7% of the worlds arable
In 30 years from now we will have for example in the Philippines land. This creates a stark problem how to feed 5 billion
120 million people, 60 million more than today. How to feed them, Asians in 25 years time without increased supply of high-quality
nobody seems to care. Over the last 30 years rice has had to increase land (indeed, with high-quality land continuing to be lost to
to such an extent that 600 million people can eat rice today who
otherwise would not be able to. Another 30 years and we are
urbanization and industrialization). Six Asian countries had
predicting a world population of more than 8 billion people. More populations of 100 million or more in 2001 (Figure 2). Their
than half of them will be rice eaters. A prediction from United agricultural populations accounted for 4366% of their total
Nations data is that about 400 million people will move over the population, except for Japan, which had only 3.6% of its people
next 10 years from rural areas in Asia to the big cities. What will
engaged in agriculture. Japan, as a highly advanced economy
happen is that the belt of poverty around these mega-urban con-
glomerations will become most probably unbearable. And what we which is already committed to food import rather than food self-
will face is most probably social unrest of unprecedented magni-
tude. Our task is to grow more rice on less land with less fertilizer,
less pesticides, and less labor, because producing rice is one of the China
most tedious tasks for a farmer.
India
Kazakhstan
Saudi Arabia
Indonesia
Iran
Production Statistics Mongolia
Pakistan
Using the Food and Agriculture Organization of the United Turkey
Nations (FAO) definition of Asia, the population of Asia has Myanmar
Others
more than doubled in the past 40 years (Figure 1), and is
dominated by its two most populous countries (China and 0 20 40 60 80 100
India). The population of China is more or less under control, Land area (10 Mha)
such that India is likely to overtake it as the worlds most
India
China
Turkey
6000
Kazakhstan
5000 Pakistan
Population (million)
Asia Indonesia
4000
Thailand
3000 Iran
Myanmar
2000 Bangladesh
China
1000 Others
India
0 40 80 120 160 200
0
Arable land (Mha)
1950 1965 1980 1995 2010 2025 2040 2055
Years China
1.9 India
Indonesia
Land area (ha per capita)
1.7
Pakistan
1.5 Aisa Bangladesh
1.3 Japan
1.1 Vietnam
0.9 Philippines
0.7 China Iran total pop. (M)
0.5 Turkey
agric. pop. (M)
India Others
0.3
0.1 0 200 400 600 800 1000 1200 1400
1950 1965 1980 1995 2010 2025 2040 2055 Population (million)
Years
Figure 2 Land area, arable land, and total and agricultural populations
Figure 1 Estimated population (19502001) and land area per capita of top ten population countries in Asia in 2001. (Data from FAOSTAT
(19612000) of all Asia, China, and India, with projections till 2050. Database).
376 GRAINS AROUND THE WORLD | Grain Production and Consumption: China and South-East Asia
sufficiency, has substantially different problems from other predominantly in China, is substantial. China ranks second
Asian countries. It focuses on managing the quality and suitabil- to the US in maize production. It is interesting that the center
ity of its imports, and on maintaining a domestic agricultural of origin of maize is in America (Mexico) but it has become
sector (e.g., a meat industry using imported grain), which com- globally widespread and successful, including in Asia. How-
petes poorly on price but which is politically and socially ever, soybean, a crop originating in Asia, is more commercially
important. significant in the US than in Asia. Sorghum and millets are
China produces 41% of Asian grain on 26% of the area significant crops in arid parts of Asia, typically areas of China
under cultivation; India produces 22% on 30% of the area and India too dry for maize production. Many local traditional
(Figure 3). This reflects a wider adoption of intensive cultural uses of these cereals have developed, for example typical use of
practices and some favorable environmental factors in China. sorghum in China is in distilled alcoholic beverage production.
Indonesia, another highly populous country, also has greater Buckwheat also has a significance beyond its relatively small
production per unit area than the average for Asia, whereas production, with uses in traditional soba (Japanese buck-
Thailand shows below-average production. These figures can wheat noodles) and some Chinese traditional foods such as
be used to indicate possible areas for greater technology adop- noodles, vermicelli, and fermented vinegars. Barley is a sub-
tion and hence increased production using existing land under stantial crop, fourth after maize, partly reflecting Chinas rise to
cultivation. rank first in the world for beer production.
How is Asias one billion tonnes (Gt) annual grain produc- Asias grain imports in 2001 were worth about US$ 25
tion distributed across crops? Asia is well-known for rice pro- billion, and exports about $8 billion (Table 2). Imports were
duction, and for the widespread dietary cultures based on rice dominated by wheat, maize, and soybean, and exports by rice.
consumption. Indeed, more than half of Asian grain produc- Industrial, processed food, and feed uses predominate for
tion is rice (Table 1, Figure 4) with 523 million tonnes (Mt) in maize and soybean. The unique properties of wheat flour
2002. Other than for India, Asia is less well known as a wheat make it impossible to substitute with any other grain. Strong
producer, yet more than 44% (Table 2) of global wheat pro- demand for wheat supplies exists for traditional uses (e.g.,
duction is from Asia, and China is by far the largest wheat noodles and steamed bread in northern China and other tem-
producer in the world. Maize production at 166 Mt, again perate Asian countries). Economic development also tends to
India 29.7%
China 26.4%
Indonesia 4.9%
Turkey 4.6%
Pakistan 3.9%
Bangladesh 3.9%
Thailand 3.7%
Vietnam 2.7%
Myanmar 2.2%
Philippines 2.1%
Others 14.1%
0 20 40 60 80 100
Harvested area (Mha)
China 40.8%
India 21.7%
Indonesia 6.2%
Bangladesh 4.0%
Vietnam 3.7%
Turkey 3.2%
Thailand 3.1%
Pakistan 2.7%
Myanmar 2.3%
Philippines 1.8%
Others 10.5%
0 90 180 270 360 450
Grain production (Mt)
Figure 3 Harvested area and grain production of top ten grain-producing countries in Asia in 2002. Data from FAOSTAT Database. Percentage (%) is
based on total harvested area or total grain production in Asia.
GRAINS AROUND THE WORLD | Grain Production and Consumption: China and South-East Asia 377
Table 1 Area, production, and yield of grains (cereals, soybeans, and pulses) in Asia (2002)a
Crop Mha % of Asia total Mt % of Asia total % of world total Yield (kg ha 1) Countries (number)
250
450 China
China Bangladesh
Bangladesh
400 India Indonesia
India
Total grain production (t)
300 150
250
200 100
150
100 50
50
0 0
1960 1965 1970 1975 1980 1985 1990 1995 2000 2005 1960 1965 1970 1975 1980 1985 1990 1995 2000 2005
Years Years
140
120 China India China India
Iran Kazakhstan 120 Indonesia Philippines
100 Pakistan Turkey Thailand Turkey
100
Wheat production (t)
80
80
60 60
40 40
20 20
0 0
1960 1965 1970 1975 1980 1985 1990 1995 2000 2005 1960 1965 1970 1975 1980 1985 1990 1995 2000 2005
Years Years
Figure 4 Trends of yield production in major Asian countries producing principal cereal crops from 1961 to 2002. (Data from FAOSTAT Database).
increase demand for wheat products (e.g., baked bread) at the (far past the rapid growth stage) and therefore stable dietary
expense of rice consumption. Thus, many tropical Asian coun- patterns. China was a major importer but also a significant
tries that do not produce any wheat require substantial imports exporter (in 2001). But its importexport pattern is highly
(e.g., Philippines, Indonesia, Malaysia). Country data (Table 3) variable, as slight percentage changes in its large production
show that Japan is the largest net importer of grain in Asia. Its base, will cause dramatic changes in its import needs and
demand is likely to be stable, due to constant (or slightly export capacity (Figure 7). India was a net exporter in 2001,
declining) population and its advanced economic status and should be able to maintain this capability in the short
378 GRAINS AROUND THE WORLD | Grain Production and Consumption: China and South-East Asia
Table 2 Quantity and value of grain imports and exports to/from Asia in 2001
Crop Imports qty (Mt) Imports value (1000 US$) Exports qty (Mt) Exports value (1000 US$)
Cereals 106 478 996 16 732 160 38 394 615 7 057 804
Maize 40 913 789 5 076 869 7 013 319 759 271
Wheat 40 238 186 6 760 921 7 906 977 896 782
Barley 10 117 996 1 490 596 436 870 40 048
Rice 9 426 314 2 623 935 19 798 418 4 902 134
Sorghum 2 015 717 233 845 20 587 3061
Rye 381 226 46 196 15 983 2065
Oats 112 604 20 100 5597 701
Buckwheat 104 668 25 227 104 491 20 365
Millet 65 854 15 188 45 735 9099
Triticale 251 74 25 528 5653
Others 3 102 391 439 209 3 021 110 418 625
Soybean 28 045 849 6 025 734 366 951 109 096
Pulses 4 463 201 1 468 603 2 636 496 1 065 219
Table 3 Top ten Asian countries of grain (cereal and soybean) imports and exports in 2001
Top ten import Percentage Top ten export Percentage Top ten net import Percentage
countries Mt (%) countries Mt (%) countries Mt (%)
Asia, total 134.5 100.0 Asia, total 38.8 100.0 Asia, total 95.8 100.0
1. Japan 31.1 23.1 1. China 9.3 23.9 1. Japan 30.1 31.4
2. China 26.3 19.6 2. Thailand 8.2 21.2 2. China 17.1 17.8
3. Korea 13.7 10.2 3. India 5.4 14.0 3. Korea 13.7 14.3
4. Iran 10.4 7.7 4. Vietnam 3.8 9.8 4. Iran 10.4 10.8
5. Indonesia 5.9 4.4 5. Kazakhstan 3.6 9.2 5. Indonesia 5.8 6.1
6. Saudi Arabia 5.1 3.8 6. Pakistan 3.4 8.6 6. Saudi Arabia 5.1 5.4
7. Malaysia 4.4 3.3 7. Turkey 1.6 4.0 7. Malaysia 4.3 4.4
8. Philippines 4.2 3.1 8. Myanmar 1.0 2.7 8. Philippines 4.2 4.4
9. Iraq 4.0 2.9 9. Japan 1.0 2.6 9. Iraq 3.9 4.1
10. Israel 3.6 2.6 10. United Arab 0.5 1.3 10. Israel 3.6 3.7
Emirates
Others 25.8 19.2 Others 1.1 2.8 Others 2.3 2.4
W y
M t
ze
So ley
w t
at
Tr s
O le
s
ille
d
at
er
Ry
ce ot
ca
he
hu
he
ai
ad
th
M
Ba
Ri s, t
iti
rg
Bu
years (Figures 4 and 5), with substantial drops in production Cereal crops in Asia
of cereal grains other than the big three, namely, rice, Figure 5 Change in Asian grain production and harvested area of
wheat, and maize. In fact, between 1992 and 2002 total pro- total cereals by crop over the past decade (19922002). (Data from
duction (output) increased only 6%, whereas area harvested FAOSTAT Database).
GRAINS AROUND THE WORLD | Grain Production and Consumption: China and South-East Asia 379
decreased 8%. Although this reflects a healthy improvement responsible for much of development of these rice varieties,
in yield per unit area, it underscores worrying trends. One is summarized their biological features:
the diversion of agricultural land to noncereal uses, offering
higher returns to farmers, and another is the steady loss of reduction in plant height (improved harvest index) and
high-quality agricultural land. In many places, such as in parts higher biomass,
of China, production of grain on small family-based farm photoperiod insensitivity (can grow any time of year) and
units is no longer economically attractive. Other agricultural short growth duration (110 days),
activities vegetables, meat production such as ducks, or even increased yield stability through disease and insect resis-
flower production may be preferable. In some cases, land may tance (biotic stresses),
be left idle while the landholder pursues nonagricultural tolerance to adverse soil conditions (abiotic stresses),
business activities. These shifts are often beneficial in raising eating quality, and
the living standards of rural families. But they also show that comprehensive agronomic knowledge on crop manage-
more or less open markets, keeping grain prices near world ment, e.g., nutrient requirements, water management,
levels, are certainly incompatible with full self-sufficiency in mechanization techniques, and equipment.
grain. These technical inputs alone were not enough. As with wheat
in India, production conditions had to be improved. The main
factors associated with this are:
The Green Revolution
Increased provision of irrigation facilities,
India has all the food it needs, but half of it is currently being
Availability of inorganic fertilizers, and
eaten by rats, according to a western aid official quoted in a
Supportive government policies
Financial Times report of December 2002. In the three years up
to 2002, Indias reserve of rice and wheat increased from 20 to The Green Revolution is a high-input system. There is no free
60 Mt, relative to an annual production of 220 Mt. Half of lunch or nonenergy intensive way to produce such dramatic
Indian children are underweight, and many are malnourished. increases in production. Central to this is farmer knowledge
India has seemingly solved one problem the technological and participation in a system with higher risks (e.g., use of
problem for which the solution began with the Green credit to finance the required inputs) but better rewards
Revolution only to be left with a perhaps more intractable (higher economic returns).
problem, that of creating an economic system that distributes
food equitably and without undue wastage.
Development of dwarf wheat that could sustain high fertilizer The Crisis
inputs and thus produce high grain yields without lodging began Grain Supply in China
at International Center for the Improvement of Maize and Wheat
(CIMMYT) in Mexico in the early 1960s. At that time, Indias Chinese tradition and the political leadership have a clear
wheat production was low ( 10 Mt per year) and static, showing understanding of the importance of grain production, from
only random seasonal fluctuations from year to year. Like most the proverb Of all things food is the foremost necessity of
low-income, high-population countries, population growth was the people to the comment by Deng Xiaoping Without agri-
much more rapid than growth in food production, and a social culture, there is no stability, without grain, there is chaos. This
catastrophe seemed imminent. The dwarf wheat from Mexico has an impact even in relation to Hong Kong: As long as the
was introduced to India, through a system of demonstration overall political situation in China is stable and the economy
fields and a supporting package of technology that emphasized develops further, we will stand by our agreements and princi-
to farmers the uniqueness of the new varieties, and the necessity ples agreed with Britain Qiao Shi (quoted 8 January 1995).
of growing them with the high inputs of water, fertilizer, and This was in relation to the agreed transfer of sovereignty of
other agricultural chemicals. From the mid-1960s, wheat pro- Hong Kong from the United Kingdom to China from 1997,
duction in India began to increase rapidly, outstripping growth where substantial autonomy would be granted to Hong Kong
in population and leading to the present day problem of (as a Special Administrative Region) for a period of 50 years.
distribution of excess production. Over a 20-year period Pragmatically speaking, if a government cannot feed its popu-
(196180), overall food production increased 3.6% per year in lation, it cannot maintain the credibility to stay in power and
Asia as a whole, from 1981 to 2000 (sometimes referred to as carry out its normal obligations. This issue is all the more
Late Green Revolution), it increased 2.1% per year. Thus, we pressing in China with its large population, limited area of
should consider the adoption of high-yielding, high-input wheat arable land, and recent memory of the disastrous famine of
varieties as the beginning of an agricultural transformation, and the early 1960s, which caused the deaths of over 30 million
not as an abrupt one-time revolution. Present-day technologies people.
may be seen as contributors to the same continuum of progress.
If production increases in the range of 23% a year, clearly
Money, Meat, and Grain
limiting the population growth to less than this is the key to
increased production per person. The book Who Will Feed China? by US environmentalist L.
Shortly after the dwarf wheat varieties came into use, a R. Brown, (1995), was perhaps the most influential early study
parallel development started with dwarf rice varieties, distrib- in drawing the attention of western policy-makers to a fairly
uted from IRRI Philippines. G. S. Khush, the IRRI plant breeder obvious problem, i.e., grain trade and capacity for global
380 GRAINS AROUND THE WORLD | Grain Production and Consumption: China and South-East Asia
Meat (kg)
many effects on agriculture: 20
0
1960 1965 1970 1975 1980 1985 1990 1995 2000 2005
The Future
(b) Years Asia is the key to world prosperity, food security, and stability;
Figure 7 Changes in grain (cereals and soybean) imports and exports or the alternative of massive regional famines. Large increases
in Asia (a) and China (b) from 1961 to 2001. in grain prices will not much affect consumers in the industri-
alized west. Like oil-price fluctuations, these will be absorbed
into the economy. Grain raw material prices are a small per-
needs, urban consumer needs, and development of grain- centage of most consumers monthly food costs in these coun-
processing facilities in producing-yet-importing markets tries. Most Asian countries will be able to absorb any such
(China) and exporting but price-distorting markets (the US). structural increases in grain prices albeit with some economic
Recent Chinese restrictions on soybean imports have increased pain during the readjustment. Most of the poorer Asian coun-
domestic prices, benefiting farmers in Heilongjiang, the main tries are more or less self-sufficient in grain, or with increasing
soybean-producing province. Limited supplies of soy meal prices will have the motivation and capability to become self-
have increased prices of pork, and soybean oil has also become sufficient. Rapidly industrializing and developing economies
more expensive, affecting urban shoppers across the country. such as China will combine continued substantial domestic
Rising meat consumption in China has led to development of a grain production with the capacity to import additional needs
soybean feed-processing industry employing 250 000 people. through the strength of their export-focused manufacturing
Restricted imports, for whatever reason, affect their liveli- industries. Perhaps long-term grain price increases will enable
hoods. Chinas estimated consumption of soybean is 33 Mt, major producers (in North America and Europe) to remove
which is projected to rise to 50 Mt within 5 years; in the face of farm subsidies and move to more effective free-market produc-
current production of only 17 Mt. Expectations of free trade tion. An additional 100 Mt, for example, of grain entering
under World Trade Organization (WTO) are commonly under- world trade (at far higher prices than today) will be an attrac-
mined by nontariff barriers. Clearly Chinese imports of soy- tive prospect to marketing boards and farmers organizations
bean from the US should continue and a mutually in the US, Canada, and Australia. But these major exporting
beneficial trade will result. The reasons for the restrictions countries have to look carefully at the environmental costs of
(safety standards for genetically modified soybeans, and phy- meeting such increased demand. The food supply crisis,
tosanitary regulations concerning phytophthora) obviously although arising from western production constraints and
have a strong policy overtone. However, they may be placed Asian demands, will have its most serious consequences in
in context of the widespread demands from developing econ- countries, many in Africa, which have limited production,
omy nations for US farm subsidies and tariffs to be reduced in rapidly rising populations, and few competitive exports.
the same spirit of free trade. Many of these are the same countries that suffered most in
382 GRAINS AROUND THE WORLD | Grain Production and Consumption: China and South-East Asia
the oil price crises of the past few decades. With grain more Australia and Pacific Countries; Grain Production and Consumption:
than with oil, the impact of the crisis will be more immediate Overview; Grain Production and Consumption: South America.
and drastic on the health and survival of the people of these
countries.
Further Reading
Exercises for Revision Brown L (1995) Who Will Feed China?, p. 163. New York: W.W. Norton and Company.
Cassman KG and Pingali PL (1995) Extrapolating trends from long-term experiments to
Can and should China feed itself? farmers fields: the case of irrigated rice systems in Asia. In: Barnett V, Payne R, and
Was the Green Revolution really a success? Argue the pros Steiner R (eds.) Agricultural Sustainability: Economic, Environmental and Statistical
Considerations, pp. 6384. London: Wiley.
and cons, focusing on the effect on rural populations in Garnaut R and Ma G (1992) Grain in China, p. 151. Canberra: East Asia Analytical Unit,
India. Department of Foreign Affairs and Trade, Commonwealth of Australia.
How much grain is required to produce one kg of each Kaosa-ard MS and Rerkasem B (2000) The Growth and Sustainability of Agriculture in
major meat product? Asia, p. 303. Oxford: Oxford University Press.
Khush GS (1999) Green Revolution: preparing for the 21st century. Genome
42: 646655.
Exercises for Readers to Explore the Topic Further Khush GS (2001) Green revolution: the way forward. Nature Reviews: Genetics
2: 815822.
Wittwer S, Yu Y, Sun H, and Wang L (1987) Feeding a Billion, p. 462. East Lansing,
How can new storage and supply chain technologies reduce Michigan: Michigan State University Press.
grain losses from farm to consumer?
What are the prospects for a new Green Revolution based
on genetic engineering? Relevant Websites
Why is Vietnam a major rice exporter, yet Philippines strug-
gles to feed itself? apps.fao.org http://apps.fao.org The FAO database on agricultural production and
trade statistics is a comprehensive resource and invaluable to researchers working
in this area. It has been extensively used in preparation of this article.
irri.org http://www.irri.org Website of the International Rice Research Institute,
See also: Grains Around the World: Grain Production and Philippines. Information and links on sustainable rice production in Asia.
Consumption: Cereal Grains in North America; Grain Production and worldwatch.org http://www.worldwatch.org Website of Worldwatch Institute, for
Consumption: Europe; Grain Production and Consumption in Oceania: independent research for an enviornmentally sustainable and socially just society.
Grain Production and Consumption: Europe
MG Lindhauer, Max Rubner-Institut (MRI), Detmold, Germany
2016 Elsevier Ltd. All rights reserved.
Topic Highlights Consequently, data have been prepared and presented with
respect to this given political situation but concentrating more
Europe is one of the most important cereal producers of the on the main players to ensure clarity.
world.
Within Europe, the European Union is the dominant cereal
producer.
Soft (bread) wheat and maize, and to a lower extent barley,
Grain Production
rye, oats, and triticale, are the most important grain cereals.
In some parts of Europe, rye is the basis for bread making. Cereals
Other cereals are rice, sorghum, millet, and buckwheat
In Europe, cereals are by far the most important agricultural
(pseudocereal).
crops (Tables 14) and among these, wheat is the principal
The per capita consumption of cereals varies significantly
crop (Table 1). Wheat accounts for nearly half of the total
between European countries.
quantity harvested in 2012. Grain maize and barley and, to a
Cereal consumption and preference for individual cereals
distinctly less degree, oats, rye, and triticale are also of signifi-
are very much dependent on cultural traditions in diet.
cance. Triticale, the result of crossbreeding wheat and rye, is in
Europe is an important producer of pulses (beans, broad
demand in those cultivation areas where normally rye is grown
beans, peas, and lentils) and to a lower extent of soybeans.
and where farmers are looking for alternatives.
Rapeseed and sunflower are important oilseeds.
All other cereals are of minor importance. The pseudocereal
The majority of grains harvested are used for animal feed.
buckwheat a plant belonging botanically to the Polygonaceae
family was grown throughout Europe until the early 1900s. It
had mostly been grown in areas with poor soils and served for
Learning Objective the production of grits and for specific traditional dishes such
as pancakes and crepes. It has survived as a kind of specialty
To get an overview of the agronomically important grain crop, for example, in Northern Germany, Russia, Austria, and
species grown in Europe and of their importance in human individual Balkan states. In very recent years, there has been a
consumption and other utilization. certain rekindling of interest in buckwheat as a functional food
in modern nutrition systems because of constituents such as
dietary fibers that have been scientifically proved to improve
health and avoid diseases in human (animal) metabolism.
Introduction Rice is produced only in those parts of Europe that have a
suitable climate, for example, Spain, southern France, Italy, or
In Europe, grain crop production has a long tradition, and Southern Russia.
grains of different plant origins are substantial constituents of On the European continent, the EU, with its 28 member
the human diet and of animal feed since ancient times. countries, is the most important cereal producer accounting for
For the cultivation of grain crops, of cereals especially, almost far more than half of the production of all the cereals,
Europe has excellent prerequisites because of its climatic and wheat, barley, maize, and rice (Table 2). Most of the triticale is
pedological conditions and its overall efficient agricultural grown in the EU.
systems. Among EU member countries, France is the main cereal
Preference for specific grain crops varies very much among producer, followed by Germany, Poland, the United Kingdom,
European countries and larger geographic regions owing to the Spain, and Italy (Table 3). Variance of data between individual
wide cultural multiplicity and ethnic diversity in nutritional harvest years is due to specific climatic conditions as, for
habits. example, in 2010 and 2011 where drought periods during
Rye may serve as an example. It is a cereal very specific to the growth period in major production countries (e.g., France
Europe and hardly grown not only anywhere else in the world and Germany) resulted in significantly lower cereal yield. Also,
but also within Europe; it is projected as a bread-making cereal it has to be taken into consideration that data generation may
and animal feed. It is grown almost exclusively in central, differ between individual member states, making comparison
northern, and northeastern countries of Europe. of data from different sources sometimes difficult.
Another point of consideration while recording statistical Growing of cereals is a long-standing tradition in the Rus-
data is that because the European Union (EU) is politically very sian Federation and in Ukraine, though both countries are yet
diverse, it dominates all the markets including the agricultural to utilize their full agronomic capacity (Table 4).
one. This is the case, although the EU as yet does not include Also, when studying production data for single cereals of
countries like Russia and other countries of the former Soviet Russia and Ukraine (Table 4), the role of wheat as the most
Union. preferred cereal is underlined. Barley is in the second position
Table 1 Cereala production (Mt, millions of tonnes) in Europe in the Table 3 Total cereal production (1000 tonnes) in EU member
year 2013 countries in selected years
in Russia, and in the Ukraine, it is maize. Rye has a certain Wheat 37.72 15.76
importance in Russia but nearly none in the Ukraine. Maize 8.21 20.96
Yields for cereals as a whole and for individual cereal spe- Rye 2.13 0.01
cies reveal a higher cultivation efficiency within the EU Oats 4.03 0.01
(Table 5) when compared with respective data calculated on Triticale 0.46
the whole-Europe basis (Table 6) or with data from Russia or Barley 13.95 6.94
Ukraine (Table 7). The reasons for the more efficient agricul- Sorghum 0.05 <0.001
Millet 0.33 0.001
tural system in most of the European member countries are
Rice, paddy 1.05 0.002
multifold. Without a doubt, in the majority of the countries, Buckwheat 0.80 0.002
farmers are well educated and experienced and can rely on Cereals, total 68.77 45.67
well-established governmental and private consulting systems.
They have unlimited access to production means like fertil- Source: FAOSTAT: FAO Statistical Division (2014).
izers, pesticides, high-value varieties, and modern technical
equipment.
After revision of the EU common agricultural market policy wheat, maize, and barley. But due to an increasing demand
in 2007 and thus unlike as in former decades, grain interven- with coinciding high prices achievable on the world market,
tion, that is, buying for a fixed threshold price and storing cereal intervention has nearly lost its importance.
surpluses from the market, has practically come to an end. It In large parts of the EU, the climate and soil are very
has been restricted to the cereal species soft wheat, durum favorable for cereal cultivation. In the main cereal production
GRAINS AROUND THE WORLD | Grain Production and Consumption: Europe 385
Table 5 Cultivation area (1000 ha) and average yield (tonnes ha1) countries in Western and Central Europe, precipitation is
a
of cereals in the European Union (EU-28) in the year 2013 rather high and equally distributed throughout the year. Tem-
peratures are moderate, being, on average, not too low during
Species Cultivation area Yield
winter for the growth of winter-type cereals (winter wheat,
Wheat 25 683 5.6 winter rye, etc.). These cereals have a prolonged vegetation
Barley 12 339 4.8 period and are thus higher in yield.
Rye 2583 4.0 In recent years, however, there seems to be a tendency
Oats 2645 3.2 towards more frequent unusual weather events like drought
Maize 9685 6.7 and very high temperature in spring and during the main
Triticale 2676 4.3 growth period, respectively, and more frequent heavy thunder-
Sorghum 137 5.0 storms with huge amounts of rainfall that may be taken as
Millet 61 1.9
indications of climate change, the consequences of which for
Rice, paddy 430 6.5
cereal production in quantity and processing quality are hard
Buckwheat 159 1.8
Mixed grain 1201 3.0 to assess at present.
Cereals, nesb 119 4.9 Also, among EU member countries, yields of different
Total cereals 57 721 5.3 cereals are varying significantly. Data for bread wheat yield in
2013 (data not shown) may serve as an example. The average
a
Including the pseudocereal buckwheat. yield ranged from 8.6 tonnes ha1 in Belgium to
b
Including inter alia amaranth and quinoa species. 1.9 tonnes ha1 in Portugal. In countries like Belgium/Lux.,
Source: FAOSTAT: FAO Statistical Division (2014).
Denmark, France, Germany, Ireland, the Netherlands, and
the United Kingdom, yields much higher than the EU average
Table 6 Cultivation area (1000 ha) and average yield (tonnes ha1)
a are standard. To a limited degree, there is also a variation in
of cereals in Europe in the year 2013
yield within these countries. These phenomena are the result of
Species Cultivation area Yield differences in climate, soil conditions, utilization of produc-
tion means, and a question of the choice of varieties among
Wheat 57 582 3.9 which there is a broad range in genetically determined yield-
Barley 24 569 3.5 forming potential.
Rye 4978 3.0
Oats 6143 2.4
Maize 18 874 6.2 Wheat
Triticale 3418 4.0
Sorghum 328 3.5 Additional detailed information on wheat has been provided
Millet 495 1.3 because of its overall predominance in production and con-
Rice, paddy 648 6.0 sumption. Data presented in Tables 1, 2, and 4 refer to wheat
Buckwheat 1265 1.1 as a single entity. These statistical data do not distinguish
Mixed grain 1208 3.0 between different wheat species. As mentioned in the preceding
Cereals, nesb 183 3.6 text, the so-called soft aestivum wheat (botanical name Triticum
Total cereals 119 791 4.0 aestivum) is the most important wheat species in Europe and
a
Including the pseudocereal buckwheat. worldwide. Two types of T. aestivum wheat are cultivated: winter
b
Including inter alia amaranth and quinoa species. wheat and spring wheat, where winter wheat is by far the most
Source: FAOSTAT: FAO Statistical Division (2014). preferred due to its higher yield performance. Spring-type wheat
are chosen when they specifically fit into the crop rotation
Table 7 Area harvested (1000 ha) and average yield (tonnes ha1) of system or in those countries (e.g., Scandinavia) where the cli-
cereals and buckwheat in the Russian Federation and in Ukraine in the mate in winter and the growth period are too uncomfortable for
year 2012 winter-type wheat. In the main wheat-producing countries, the
contribution of spring wheat to the total wheat harvest is dis-
The Russian Federation Ukraine tinctly less than 1%. T. aestivum wheat are covering the wide
range of feed and food applications, whereas the species T.
Area Yield Area Yield
durum, known as durum or hard wheat, is specifically grown
Wheat 21 278 1.77 5630 2.80 for pasta production. (This does not mean that pasta production
Barley 7641 1.83 3293 2.11 would not be possible from soft wheat, too!)
Rye 1423 1.50 0.3 2.27 Durum wheat is more adapted to warmer climatic condi-
Oats 2853 0.14 0.3 2.09 tions. Consequently, it is preferably grown in countries with
Maize 1938 4.24 4372 4.79 Mediterranean climates, namely, in Italy, France, and Spain
Triticale 0.2 2.08 (Table 8). Since a couple of years, however, durum wheat
Sorghum 0.04 1.04 0.06 2.23 varieties more adapted to moderate climates are successfully
Millet 0.03 1.02 0.02 1.03
grown in smaller quantities in the Czech Republic, in
Rice, paddy) 0.02 0.05 0.03 6.19
Buckwheat 1030 0.77 0.03 0.87 Germany, and in Austria. In regions with suitable growth con-
Cereals, total 36 988 1.86 14 412 3.17 ditions within these countries, durum wheat means a produc-
tion alternative for farmers and a local supply of durum mills
Source: FAOSTAT: FAO Statistical Division (2014). with produce.
386 GRAINS AROUND THE WORLD | Grain Production and Consumption: Europe
Table 8 Cultivation area, yield, and production of durum wheat in the Table 9 Production of selected cereals (1000 tonnes) in Europe in
European Union (EU-28) in the year 2013 2012
far as human consumption is concerned and on recipes of Both plants are of tremendous significance, whereas other
optimized feedstuff mixtures in modern animal production. species like linseed, for example, are specialty crops in specific
At this point, it should be mentioned that there is a strong countries. Basically, for oilseeds, too, it has to be stated that
tendency not only in Europe, namely, in Russia and Ukraine, they are of lesser importance, although they may contribute to
but also in Mediterranean countries to grow soybeans. Even in a large extent to agricultural production as in France, Germany,
countries like Austria or Germany where the climate was con- or the United Kingdom, for instance.
sidered so far to be too unfavorable for this species, attempts
are made to establish soybean production. Making use of
respective progress in breeding the underlying idea is to have Consumption and Utilization
a further feedstuff raw material from home production. The Consumption and utilization of grain seeds in Europe are
majority of the huge amounts of soybeans used, however, will greatly influenced by factors as varied as climate, soil condi-
remain to be imported from overseas. tions, preference for specific crops based on tradition and
Soybean may be used not only as protein source for food ethnic peculiarities, economics, and governmental regulations.
and feedstuff but also as oil production raw material. The These factors are further compounded by the diversity in
statistics, however, reveal rapeseed and sunflower as oilseeds regions and countries found in the European continent at
of importance in European agriculture (Tables 12 and 13). large. As it is impossible to discuss every grain species in detail
or take into consideration the entire range of diversities, an
attempt will be made to focus on major trends and to outline
Table 10 Production of pulses and soybeans (1000 tonnes) in total the main fields.
in Europe, the European Union (EU-28), Russia, and Ukraine in the year Nearly all grain seeds or components of them serve as food
2012 and feed raw materials, and these are by far the most important
fields of utilization. Furthermore, these crops or components
Europe EU-28 Russia Ukraine
derived thereof may be used as basic materials for chemical or
Total pulses 6485 3173 2220 473 technical purposes in the nonfood or feed industry. For exam-
Beans, dry 488 139 7 <1 ple, starch is applied for the synthesis of adhesives.
Chickpeas 89 36 <1
Cowpeas 23 <1
Lentils 91 62 28 <1 Consumption of Cereals
Peas, dry 3420 1272 1160 <1
Pulses, nesa 1186 850 321 <1 Statistically, the EU as a whole is a self-supplier of cereals.
Soybeans 5515 863 1806 2410 Individual countries, however, especially the main grain-
producing ones like France (200%), Germany ( 140%),
a
Including inter alia: hyacinth bean (Dolichos ssp.), jack or sword bean (Canavalia and the United Kingdom (125%), produce much more
spp.), and winged bean (Psophocarpus tetragonolobus). cereal in total than can be consumed by the national popula-
Source: FAOSTAT: FAO Statistical Division (2014). tion. However, there are also countries within the EU like, for
Table 11 Production of pulses and soybeans (1000 tonnes) in selected European countries in 2013
example, Italy (40%) or Spain ( 70%) that lack sufficient of cereals, however, being much dependent on yearly fluctua-
production. This means a compensation of surpluses and tions in total production quantities.
demands, respectively, not only within the EU but also from The majority of the yearly cereal harvest is used for feed-
outside. As a whole, the EU is one of the big players in world- stuff. Germany may serve as an example: Out of c.26 million
wide cereal trade; this is specifically valid for major grain- tonnes of wheat produced per year round, about 7 million
producing member states like France or Germany where there tonnes is milled for human consumption. Food use is domi-
is a strong pressure for export outside the EU or for alternative nated by bread production and the production of other baked
utilization of cereals. The latter aspect, however, is diversely goods like pastries and cakes. Breakfast cereals also contribute
discussed and may steadily lose importance with respect to to the total consumption of cereals, though in comparison
world population growth and therefore increased demand for with bread making, this segment is small, with a permanently
cereals for human consumption. In more recent years, Russia increasing tendency. The significance of grain food use is dem-
and Ukraine have also become significant cereal exporters onstrated in Table 14 as consumption of cereals per capita and
(especially wheat) on the world market, the available amounts year in selected EU member states. Consumption differs a lot
between individual countries, with, for example, Greece having
extraordinarily high consumption rates. In the large majority
Table 12 Total production of pulses (1000 tonnes) and oil crops
of the European countries, the production of bread or compa-
(1000 tonnes) in Europe in the year 2013
rable foodstuff means production from wheat. This is also true
Pulses Oil crops for central, eastern, and northeastern countries, but here in
addition, rye is also used as a raw material for bread making.
Total Europe 6212 27 240 The extent to which rye is used is quite different between
EU-28 3165 15 137 individual countries. For example, in Germany, 850 000
Russia 2084 5294 tonnes of rye is milled for bread production; however, this
Ukraine 372 5948
tendency is slowly decreasing.
Source: FAOSTAT (2012). Bread making does not only mean utilization of pure rye
flour. Most breads offered on the market are made of mixtures
of rye and wheat flour. In relation to wheat, rye has higher
Table 13 Production (1000 tonnes) of rapeseed and sunflower seed proportions of dietary fiber. Actually, this health benefit is used
in the European Union in the year 2012 in arguing in favor of increased rye consumption again. Health
aspects are also the reason of increased interest in the so-called
Rapeseed Sunflower seed
old wheat species as spelt, emmer, or einkorn.
Austria 149 53 With steadily increasing amounts, wheat and maize serve as
Belgium 48 raw materials for starch production (Table 15). The advantage
Bulgaria 271 1388 of these raw materials is based on marketable by-products like
Croatia 26 90 germ for oil extraction or protein for feed (maize) or gluten for
Cyprus 0 food use (and feed) from wheat. The so-called vital gluten from
The Czech Republic 1109 57 wheat starch plants is used in a wide range of feed and food
Denmark 485 0 applications, as a flour improver in the bread industry, for
Estonia 163
example. The main product of the starch industry, starch, is
France 5483 1575
Finland 73 0 mainly used in food applications (Table 16).
Germany 4821 63
Greece 1 160 Table 14 Per capita consumption of cereals (kg year1) in selected
Hungary 401 1302 European countries in the economic year 2011/2012
Italy 25 186
Ireland 0 Cereals, Soft durum
Latvia 304 0 total wheat Other cereals
Lithuania 633 0
Luxemburg 15 0 Austria 90.9 64.4 26.5
Malta 0 Estonia 62.3 38.0 24.3
The Netherlands 7 0 Germany 95.0 69.9 25.0
Poland 1862 Greece 313.0 308.1 4.8
Portugal 10 Hungary 106.6 89.9 16.8
Romania 159 1389 Latvia 112.0 82.9 29.0
Slovenia 17 0 Lithuania 92.2 74.4 17.8
Slovakia 213 197 Luxemburg 60.0 57.3 2.7
Spain 52 0 The Netherlands 109.9 99.3 10.6
Sweden 323 0 Romania 159.2 120.3 38.9
The United Kingdom 2564 0 The United 80.8 76.9 3.9
EU-28 Kingdoma
a
, Data not available. Data of 200910.
Source: Federal Ministry of Food and Agriculture (ed.) (2013) Statistisches Jahrbuch Source: Federal Ministry of Food and Agriculture (ed.) (2013) Statistisches Jahrbuch
uber Ernahrung, Landwirtschaft und Forsten der Bundesrepublik Deutschland, 57. Jhrg. uber Ernahrung, Landwirtschaft und Forsten der Bundesrepublik Deutschland, 57. Jhrg.
Landwirtschaftsverlag GmbH Munster-Hiltrup. Landwirtschaftsverlag GmbH Munster-Hiltrup.
GRAINS AROUND THE WORLD | Grain Production and Consumption: Europe 389
Table 15 Total raw material processed, starch production, and the In European tradition, maize as such is of little interest for
respective contribution of specific plant groups in Europe in selected food use, though actual development reveals the so-called
years Mexican food to become a segment of diet, at least in some
European countries.
Year 2001 2005 2008 2010
Minor quantities of grain maize are used for grits produc-
Total raw material (Mt) 21.2 22.6 21.6 22.0 tion, as raw material for snacks, while the great majority of
Potato (%) 41 39 35 32 grain maize is used in feed.
Maize (%) 31 31 33 34 Because of their b-glucan content with health benefits, oats
Wheat (%) 28 30 32 34 are processed to a certain amount to produce special food like
Starch produced (Mt) 9.0 9.6 9.4 9.9 flakes and extruded products. But the majority of the produc-
Potato (%) 20 18 16 14 tion is used for feed. This is also valid for barley, though a
Maize (%) 46 46 47 48
notable quantity of barley is used for beer brewing. Barley is
Wheat (%) 34 36 37 38
also rich in b-glucan, a reason why it should be treated as
Mt, millions of tonnes cereal, the potential of which for food production is
Source: German Starch Industry Association (2012). underestimated.
Table 17 Food supply quantity (1000 tonnes) and consumption per capita (kg) of pulses and soybeans in selected European countries in the year 2011
1000 tonnes kg 1000 tonnes kg 1000 tonnes kg 1000 tonnes kg 1000 tonnes kg
Albania 15.00 4.8 0.00 0.0 0.91 0.3 16.00 5.1 0.04 0.0
Austria 1.60 0.2 3.99 0.5 1.27 0.2 7.00 0.8 14.39 1.7
Belgium 8.23 0.8 13.00 1.2 2.06 0.2 23.00 2.1 0.65 0.1
Bulgaria 12.00 1.6 1.00 0.1 3.46 0.5 16.00 2.2 0.31 0.0
The Czech Republic 1.85 0.2 20.00 1.9 6.42 0.6 28.00 2.6 6.30 0.6
Denmark 0.74 0.1 4.00 0.7 1.00 0.2 6.00 1.1 0.35 0.1
Finland 0.12 0.0 6.00 1.1 0.23 0.0 6.00 1.1 0.28 0.1
France 52.70 0.8 30.18 0.5 44.04 0.7 127.00 2.0 3.53 0.1
Germany 8.00 0.1 59.18 0.7 19.00 0.2 86.00 1.0 52.33 0.6
Greece 30.83 2.8 1.53 0.1 20.50 1.8 53.00 4.8 0.16 0.0
Hungary 7.83 0.8 24.39 2.4 5.63 0.3 35.00 3.5 0.05 0.0
Italy 115.07 1.9 45.76 0.8 130.43 2.2 291.00 4.8 0.63 0.0
Poland 36.67 1.0 40.51 1.1 2.99 0.1 81.00 2.1 0.27 0.0
Portugal 23.03 2.2 3.39 0.3 10.36 1.0 36.00 3.4 0.14 0.0
Serbia 45.62 4.8 42.64 4.4 0.59 0.1 90.00 9.4 0.04 0.0
Spain 46.12 1.0 215.89 4.6 141.44 3.0 403.00 8.7 0.69 0.0
Sweden 2.56 0.3 12.00 1.3 2.83 0.3 18.00 1.9 1.23 0.1
Switzerland 2.00 0.3 10.54 1.3 2.28 0.3 15.00 1.9 1.03 0.1
The United Kingdom 10.00 0.2 118.95 1.9 21.65 0.4 151.00 2.4 5.42 0.1
Russia 260.00 1.8 1.14 0.0 261.00 1.8 5.40 0.0
Ukraine 69.82 1.5 1.09 0.0 71.00 1.6 0.61 0.0
Exercises for Revision See also: Food Grains and the Consumer: Consumer Trends in
Grain Consumption; Genetically Modified Grains and the Consumer;
Which are the main cereals produced in Europe as a whole Grains and Health: Misinformation and Misconceptions; Labelling of
and in the EU? Grain-Based Foods; Food Grains and Well-being: Functional
What are the main cereal-producing countries in Europe? Foods: Overview; Grains Around the World: Grain Production and
What cereal species is a specialty in Europe and hardly Consumption: Overview; Non-food Products from Grains: Cereal
grown elsewhere in the world? Grains as Animal Feed; Utilization of the Whole Cereal Plant to
What are the main pulses grown in Europe as a whole and Maximize its Value; Processing of Grains: Evaluation of Wheat-
in the EU? Grain Quality Attributes; Grain Quality Attributes for Cereals Other than
What are the main oilseeds produced in Europe? Wheat; Starch: Uses of Native Starch; The Basics: The Grain Crops:
What are the cereal species preferably used for human An Overview; The Cereal Grains: An Overview of the Family of Cereal
consumption in Europe? Grains Prominent in World Agriculture; Durum Wheat: Overview; Rye
What is the significance of soft wheat in bread making in Grain: Its Genetics, Production, and Utilization; Wheat: An Overview of
Europe? the Grain That Provides Our Daily Bread; The Legumes and
Are there differences in the significance for bread making Pseudocereals: Grain Legumes and Their Dietary Impact: Overview;
between individual European countries? Pseudocereals: Overview; The Oilseeds: Oilseeds: Overview;
Are there differences among countries in the preference for Soybean: Overview; Wheat Processing: Gluten and Modified Gluten;
the use of pulses? Wheat-Based Foods: Cereals: Overview of Uses: Accent on Wheat
Grain.
Topic Highlights Maize is the most commonly grown crop on the continent,
accounting for almost half of the area in cereals (Figure 2)
North Americas relatively low population density and followed by wheat with about one-third. Less than 2% of
abundant arable land contribute to its position as a major North American cereal area is in rice production compared to
grain producer and exporter. almost 25% for the world.
The US dominates global maize production, widely uses Between 2002 and 2013, the area in North American cereal
GM maize, and in recent years has further increased pro- production increased by about 6.6 Mha (8.5%). Most of the
duction to meet demand for bioethanol. increase (5.7 Mha) occurred in the United States, where the
Maize is the most important cereal crop in Mexico (70% of area in maize (corn) increased from 28 Mha to over 35 Mha.
production area), and wheat is in Canada (60% of produc- The net increase in US cereal area was partially a result of a
tion area). 3 Mha reduction in the Conservation Reserve Program (USDA
North America produces only 2% of the worlds rice, but is Farm Service Agency) and reductions in other crops, primarily
still a significant exporter. hay (down 2.5 Mha) and cotton (down 1.4 Mha) (USDA,
QuickStats). During the same period, the area in oilseeds (soy-
beans, rapeseed, sunflower, and linseed) in North America, the
main alternative to cereal crops, also increased by about 6 Mha
Learning Objective from 36 Mha in 2002 to 42 Mha in 2013. Most of the net
increase in oilseed area occurred in Canada where the area in
To provide perspective on the role of North America in rapeseed (canola) production more than doubled from about
global production and trade for the covered grains. 3.6 Mha in 2002 to over 8 Mha in 2013.
Figure 3 shows total cereal production in North America
between 2002 and 2013. During that period, production
increased from 361 million tonnes (mt) to 535 mt, an increase
Introduction of over 48%. Production increased in all three countries: by 47%
in the United States, by 86% in Canada, and by 15% in Mexico.
According to worldstat.info, the North American continent On average, the United States accounted for 82% of North
(herein defined as comprising the United States, Canada, and American cereal production over the 12-year period, with
Mexico) has 17% of the worlds arable land and 6.6% of the Canada contributing 11% and Mexico 7%.
worlds population. Given the relative abundance of arable land
(0.52 ha per person vs. world average of 0.2 ha per person), the
continent is a major producer and exporter of several grain crops
including barley, maize, oats, rice, sorghum, and wheat. Maize (Corn)
This article updates an earlier Encyclopedia of Grain Science
chapter on the North American grain sector (Lukow and Table 1 provides data on North American and world maize
McVetty) with data covering the 12-year period from 2002 to production from 2002 to 2013. Maize is the most important
2013. Like the earlier chapter, which covered the period from cereal grain in North America accounting on average for 49%
1992 to 2001, we provide an overview of production, consump- of harvested cereal acres and 71% of cereal tonnage produced.
tion, and trade for the major North American grains. Unless Maize is also the worlds top cereal crop with over 1 billion
otherwise noted, all data are drawn from the Food and Agricul- tonnes produced in 2013.
ture Organization (FAO) FAOSTAT database. We begin with a On average, North America produced almost 42% of the
brief overview of grain production on the continent and then worlds maize between 2002 and 2013, on 25% of the worlds
provide additional detail for each of the major cereal grains. harvested maize area. Yields in North America averaged
Figure 1 shows the total hectares of cereal grains (barley, 8.2 mt ha1 compared to the overall world average (including
maize, oats, rice, sorghum, and wheat) harvested annually in North America) of 4.9 mt ha1. The United States is the worlds
the United States, Canada, and Mexico between the years 2002 leading producer, accounting for 90% of North American pro-
and 2013. During the 12-year period between 2002 and 2013, duction, with Canada contributing 3% and Mexico 7%. US
an average of 82.2 Mha of cereals was harvested in North yields averaged 9.3 mt ha1 over the 12-year period, compared
America, accounting for almost 13% of the worlds harvested to 8.6 mt ha1 in Canada and 3.0 mt ha1 in Mexico. Drought
area (640 Mha). With higher yields, however, North America in the US Midwest during the 2012 growing season resulted in
accounted for 20% of world cereal production during the same yields declining to 7.7 mt ha1, the lowest average yield during
period. Within North America, the United States, with an the 12-year period. Average yields trended upward in each
average harvested area of over 57 Mha, accounted for about country during the 12-year period, by 0.03 mt ha1 per year
70% of the total harvested area, with Canada having 18% and in the United States, by 0.18 mt ha1 per year in Canada, and
Mexico 12%. by 0.04 mt ha1 per year in Mexico.
40% 36%
34%
30%
25% 25%
20%
10% 8%
6% 5% 7%
2% 2% 2%
0%
Barley Maize Oat Rice Sorghum Wheat
Figure 2 Cereal area distribution: North America versus world.
Cereal production
600 000 000
0
2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013
Since the introduction of genetically modified (GM) crops (GMO Compass). GM maize is not grown commercially in
in the late 1990s, both the United States and Canada have Mexico and its possible introduction is the subject of ongoing
experienced widespread adoption of GM maize. In 2013, GM legal challenges (Nature.com).
varieties of maize accounted for 90% of the planted area in the The area of maize harvested in the United States increased
United States (USDA, ERS) and 98% of the area in Canada by over 25% between 2002 and 2013, from 28 Mha to over
GRAINS AROUND THE WORLD | Grain Production and Consumption: Cereal Grains in North America 393
Table 1 Maize production: Area harvested (000 ha) and production (000 mt)
Year Area Prod. Area Prod. Area Prod. Area Prod. Area Prod.
2002 28 057 227 765 1283 8999 7120 19 298 36 460 256 062 137 609 604 872
2003 28 710 256 227 1226 9587 7521 20 701 37 457 286 516 144 701 645 165
2004 29 798 299 874 1072 8837 7688 21 670 38 558 330 381 147 454 728 971
2005 30 399 282 261 1085 9332 6606 19 339 38 089 310 932 148 035 713 682
2006 28 587 267 501 1061 8990 7295 21 893 36 942 298 384 146 741 706 847
2007 35 014 331 175 1369 11 649 7333 23 513 43 716 366 337 158 390 790 115
2008 31 797 307 142 1169 10 592 7354 24 320 40 319 342 054 162 689 830 611
2009 32 169 332 549 1142 9561 6223 20 143 39 534 362 253 158 743 820 203
2010 32 960 316 165 1203 11 715 7148 23 302 41 311 351 181 164 030 851 271
2011 33 990 313 949 1202 10 689 6069 17 635 41 260 342 273 172 257 887 855
2012 35 359 273 820 1418 13 060 6924 22 069 43 701 308 949 178 552 872 792
2013 35 478 353 699 1480 14 194 7096 22 664 44 054 390 557 184 192 1 016 736
Avg. 31 860 296 844 1266 10 600 7031 21 379 40 117 328 823 158 616 789 093
35 Mha, with its share of US cereal area increasing from 52% to to livestock feed, processing is the amount used for manufacture
over 60%. Higher yields, in conjunction with the increase in of processed food items (e.g., high-fructose corn syrup), food
area, resulted in a 55% boost in production over the same comprises amounts not detailed otherwise in the balance sheet
period. The acreage increase can be attributed largely to that are available for human consumption, while other uses
increased demand for ethanol production spurred by the US comprise quantities used for the manufacture of nonfood
renewable fuels mandate introduced in the 2005 Energy Policy products (e.g., ethanol).
Act (US Environmental Protection Agency) and the attendant In the United States and Canada, maize is grown primarily
increase in maize price. US maize production is concentrated for livestock feed and ethanol with relatively small amounts
in the Midwest corn belt states, with Iowa and Illinois typi- used for human food consumption. In Mexico, however, white
cally accounting for around one-third of the US crop. The maize targeted toward human consumption (e.g., tortillas)
recent expansion has boosted production in other states such accounts for two-thirds of production (USDA, FAS). Over the
as Kansas where the area under maize increased from 1 million 10-year period from 2002 to 2011, food use accounted for 63%
to 1.6 million hectares between 2002 and 2013 (USDA, Quick of maize produced in Mexico, compared to 6% in Canada and
Stats). Approximately 15% of US corn harvested area is irri- only 1.3% in the United States. Mexico has traditionally been a
gated (USDA, Census of Agriculture). major importer of feed corn for its domestic livestock sector,
Maize is by far the most important cereal crop in Mexico, particularly the poultry sector, with imports coming primarily
accounting for over 70% of the area under cereals, with pro- from the United States. Over the 10-year period from 2002 to
duction occurring throughout the country. The total area in 2011, average imports of 7.3 mmt were equal to 34% of
maize was stable during the period under review, with almost average domestic production. In 2002, Mexico imported
identical areas harvested in 2002 and 2013. Higher yields 5.2 mmt and ranked as the worlds third largest importer
resulted in a 17% increase in production between 2002 and after Japan and South Korea. In 2011, imports had doubled
2013 but productivity gains have been limited by structural to 11.2 mmt, with Mexico as the #2 ranked import market after
problems. More than 85% of Mexicos 2 million corn growers Japan.
have fewer than 5 ha and have limited access to improved Between 2002 and 2011, Canada was a net maize importer,
genetics, fertilizer, etc. (USDA, FAS). In Canada, maize is a but as maize production increased, the country gradually
relatively minor crop, accounting for an average of only 8.2% shifted from being a relatively small net importer to a relatively
of the area in cereals, with production concentrated in Ontario small net exporter. In 2002 and 2003, Canadian net imports
and Quebec (Statistics Canada). The area in maize increased amounted to 3.6 mmt and 1.7 mmt, respectively, while in
slightly from 1.28 Mha in 2002 to 1.48 Mha in 2013, but 2011, net imports had fallen to 0.4 mmt. For 2012 and 2013,
higher yields contributed to an overall increase in production Canada was estimated to have net maize exports of between 1
of about 58%. and 1.25 mmt.
In the United States, the growth of the ethanol sector during
the past decade consumed an increasing share of the US maize
Consumption and Trade crop. Of the US maize crops produced in 2002 and 2003, an
Table 2 provides 10-year average balance sheet data on pro- average of 27 mmt (11% of total usage) was used for ethanol
duction, consumption, and trade for maize in North America production. Of the 2010 and 2011 crops, the average amount
for the period 2002 to 2011. In the balance sheet, domestic used for ethanol was 127 mmt, equivalent to 40% of total use
supply is calculated as production imports stock change- (USDA World Agricultural Outlook Board). The United States
imports. Thus, a negative stock change corresponds to a reduc- has long been the worlds top-ranked maize exporter and,
tion in domestic supply. Because total supply accounts for despite increasing ethanol production, has maintained the
stock change, the total use number corresponds to total supply. top export ranking due to increased domestic production.
Using FAO definitions for the various use categories, feed refers Throughout the 10-year period, the United States, on average,
394 GRAINS AROUND THE WORLD | Grain Production and Consumption: Cereal Grains in North America
Production 293 460 800 9 995 000 21 181 430 324 637 230
Imports 566 349 2 562 245 7 283 827 10 412 421
Stock change 2 678 290 (94 500) 280 889 2 864 679
Exports 50 552 700 641 963 308 182 51 502 844
Domestic supply 246 152 739 11 820 778 28 437 950 286 411 467
Use
Feed 138 788 400 9 357 673 10 081 009 158 227 082
Seed 555 280 12 840 396 066 964 185
Waste 371 117 3 208 635 3 579 752
Processing 22 336 530 1 213 531 1 409 676 24 959 737
Food 3 892 730 647 500 13 296 600 17 836 830
Other uses 80 579 890 218 127 45 968 80 843 984
Total use 246 152 830 11 820 788 28 437 953 286 411 570
Table 3 Wheat production: Area harvested (000 ha) and production (000 mt)
Year Area Prod. Area Prod. Area Prod. Area Prod. Area Prod.
2002 18 545 43 704 8711 15 961 635 3236 27 890 62 901 213 817 574 743
2003 21 474 63 803 10 215 23 049 605 2716 32 293 89 568 207 700 560 129
2004 20 222 58 697 9389 24 796 517 2321 30 128 85 814 216 569 632 144
2005 20 277 57 242 9404 25 748 635 3015 30 315 86 005 219 573 626 739
2006 18 940 49 216 9682 25 265 646 3378 29 268 77 860 211 195 602 332
2007 20 639 55 820 8636 20 054 692 3515 29 967 79 390 216 710 612 874
2008 22 541 68 016 10 032 28 611 802 4019 33 374 100 647 222 279 683 207
2009 20 191 60 366 9638 26 848 828 4116 30 658 91 329 224 634 686 720
2010 19 271 60 062 8269 23 167 679 3677 28 218 86 906 216 965 649 325
2011 18 496 54 413 8544 25 261 662 3628 27 702 83 302 220 196 699 389
2012 19 798 61 677 9497 27 205 579 3274 29 874 92 157 217 320 671 497
2013 18 274 57 967 10 442 37 530 634 3357 29 350 98 854 218 461 713 183
Avg. 19 889 57 582 9371 25 291 659 3354 29 920 86 228 217 118 642 690
exported around 17% of domestic maize production. During in 2013 representing a decline of over 25% on the 40.1 Mha
the 2002/03 crop marketing year, the United States exported harvested in 1992. Between those years, the US area in wheat
40.3 mmt compared to total world exports of 78.2 mmt, a declined by about 7 Mha, and the Canadian area by about
market share of 51.5%. The second ranked exporter was 3 Mha.
Argentina with 11.2 mmt. During the 2011/12 crop year, US Wheat is the primary cereal crop in Canada, occupying over
exports of 39.2 mmt accounted for a 50.2% share of the 60% of the area in cereals and over 20% of the total arable
77.8 mmt export market. The second ranked exporter was land. Most of Canadas wheat production is in the Prairie
Brazil with 24.3 mmt, followed by Argentina with 17.1 mmt. Provinces comprising Alberta, Saskatchewan, and Manitoba.
Canada Western Red Spring (CWRS) is the most commonly
produced class accounting for about 60% of the wheat area,
Wheat followed by Canada Western Amber Durum (CWAD) with
around 15%. The United States harvested an average of just
Following maize, wheat is the second most commonly grown under 20 Mha of wheat during the 12-year period, equivalent
cereal in North America, accounting for about one-third of the to about one-third of the countrys cereal acreage and slightly
area in cereals. Worldwide, average annual wheat production more than 10% of the arable land. In 2013, about one-third of
between 2002 and 2013 was 640 mmt, with North America US wheat was hard red winter (HRW), grown primarily in the
accounting for around 13%. Central and Southern Great Plains. Hard red spring (HRS),
Table 3 provides data on North American and world wheat grown in the Northern Great Plains, and soft red winter,
production from 2002 to 2013. grown in the eastern and southeastern states, each accounted
Between 2002 and 2013, the area harvested in North for about a quarter of production. White wheat accounted for
America remained relatively stable at about 30 Mha, peaking around 12% of production, and durum around 3%. Wheat is a
at 33.3 Mha in 2008 with production peaking at over 100 mmt relatively minor crop in Mexico, occupying only about 7% of
in the same year. Since the early 1990s, however, the area in the area in cereals and <3% of arable land, with durum being
wheat has dropped significantly with the 29.5 Mha harvested the predominant class (USDA, FAS).
GRAINS AROUND THE WORLD | Grain Production and Consumption: Cereal Grains in North America 395
In 2013, the United States was the worlds fourth largest average exports of over 30 mmt representing more than half
producer (after the EU, China, and India) and, with annual of domestic production, the United States is the worlds lead-
production of around 57 mmt, generally accounts for about ing wheat exporter, accounting in recent years for a 20% share
two-thirds of North American production. Of the remainder, of the worlds trade.
Canada, with an average of about 25 mmt, accounts for about Most of the domestic wheat supply is for human food
30%, with only 4% produced in Mexico. A bumper crop in consumption, with only 13% used for animal feed. Canada is
Western Canada in 2013 saw the Canadian share of North also a major wheat exporter with average exports of 16.7 mmt
American production rise to 38%. representing over 70% of production. Canada is typically one
North American wheat yields averaged 2.8 mt ha1 over the of the top five wheat exporters with the other major exporters
12-year period, slightly below the world average of being the EU, Australia, and Russia. In contrast to the United
2.9 mt ha1. Within North America, average yields were high- States, almost half of Canadas domestic supply is used for
est in Mexico at 5.1 mt ha1 compared to 2.9 mt ha1 in the animal feed. Mexico is a net importer of wheat with average
United States and 2.7 mt ha1 in Canada. Higher yields in imports exceeding domestic production. Imports doubled
Mexico are due to the fact that much of Mexican wheat pro- from an average of 1.8 mmt per year in the period from 1991
duction is done under irrigation. Average yields trended to 2000 to an average of 3.6 mmt in period from 2002 to 2011.
upward in each country during the 12-year period, by Approximately two-thirds of Mexicos domestic wheat supply
0.05 mt ha1 per year in the United States, by 0.1 mt ha1 is for food use.
per year in Canada, and by 0.07 mt ha1 per year in Mexico.
Sorghum
Consumption and Trade
Table 4 summarizes wheat production, consumption, and Grain sorghum is grown in Mexico and the United States, but
trade in North America for the period 2002 to 2011. With because maturity requires higher temperatures, there is no
Table 5 Sorghum production: Area harvested (000 ha) and production (000 mt)
Year Area Prod. Area Prod. Area Prod. Area Prod. Area Prod.
2002 2883 9163 0 0 1744 5206 4627 14 369 41 316 53 496
2003 3156 10 446 0 0 1973 6462 5128 16 908 44 609 59 037
2004 2637 11 523 0 0 1833 7004 4470 18 528 40 832 58 117
2005 2321 9981 0 0 1599 5524 3921 15 505 46 362 59 633
2006 1999 7032 0 0 1600 5519 3600 12 550 43 366 57 772
2007 2748 12 636 0 0 1775 6203 4523 18 839 44 818 62 810
2008 2943 11 998 0 0 1838 6611 4781 18 609 45 382 66 524
2009 2234 9728 0 0 1691 6108 3924 15 836 40 945 56 938
2010 1948 8779 0 0 1768 6940 3716 15 720 41 583 60 056
2011 1590 5447 0 0 1728 6429 3318 11 876 42 312 58 243
2012 2005 6272 0 0 1820 6970 3825 13 242 38 158 57 030
2013 2630 9882 0 0 1689 6308 4319 16 190 42 120 61 385
Avg. 2425 9407 0 0 1755 6274 4179 15 681 42 650 59 253
396 GRAINS AROUND THE WORLD | Grain Production and Consumption: Cereal Grains in North America
commercial grain sorghum production in Canada (AgAnnex). sorghum exporter. Over the 10-year period summarized in
Some sorghum is harvested for silage, but the area is small, Table 6, US exports accounted for 76% of total world trade,
typically around 5% of the US area. Table 5 provides sorghum but its share declined throughout the period as domestic use,
production data for the United States and Mexico. including some for ethanol production, increased and exports
Between 2002 and 2013, sorghum occupied about 5% of from Argentina, the second ranked exporting country, grew
North Americas area in cereals, far less than the shares taken by from < 0.5 mmt in 2002 to over 1.8 mmt in 2011. Most of
maize (49%) and wheat (36%). It is, however, the second most the US domestic supply is used for animal feed.
commonly grown cereal crop in Mexico, with 1.7 Mha Despite being a major producer, Mexico is also the worlds
accounting for about 18% of harvested cereal acreage. As the leading sorghum importer, accounting in some years for more
US area in maize has expanded, the area in sorghum has fallen, than half of world imports. Until 2013, when sorghum was
and the 2.6 Mha harvested in 2013 represented a 46% decline imported from Argentina, imports were sourced exclusively
on the 4.8 Mha harvested in 1992. North Americas area in from the United States. Mexican sorghum use is for animal
sorghum averaged 4.1 Mha between 2002 and 2013, just under feed, with the poultry industry being the major consumer
10% of the world total. With higher yields, averaging (USDA, FAS). Canadas domestic supply, which is also used
3.9 mt ha1 in the United States and 3.6 mt ha1 in Mexico exclusively for animal feed, is imported from the United States.
compared to a world average of 1.4 mt ha1, the continent
accounted for more than a quarter of world production. Inter-
estingly, US sorghum yields declined from an average of Barley
4.1 mt ha1 in the 10-year period from 1992 to 2001 to an
average of 3.9 mt ha1 between 2002 and 2013. During the Similar to wheat, barley has both spring and winter growth
same periods, Mexican average yields rose from 3.2 to forms, but North American production is predominantly of the
3.6 mt ha1. spring form (Lukow and McVetty). North America produces
In most years, the United States and Mexico are ranked about 11% of the worlds barley supply, and the crop occupies
among the top four sorghum-producing nations along with about 6% of the North American area in cereals. Table 7 pro-
Nigeria and India, with each typically producing between 10% vides data on barley area harvested and production.
and 15% of the worlds total sorghum crop. In 2013, the Between 2002 and 2013, the EU was the worlds largest
United States was the top-ranked producer with 9.8 mmt repre- barley producer, accounting for about 40% of production,
senting 16% of world production, while Mexico ranked third with North America contributing about 11%. Other leading
with 6.3 mmt. In the United States, the largest sorghum- producers included Russia, Ukraine, and Australia. Canada is
producing states are Kansas and Texas, which jointly account the largest producer in North America, supplying about two-
for almost 80% of production (USDA, Crop Production). In thirds of the continents barley, equivalent to about 7% of
Mexico, sorghum is produced throughout the country and world production. Canada is typically among the top five
approximately one-third of the planted area is irrigated producing countries. After wheat, barley is the second most
(USDA, FAS). commonly grown cereal in Canada, occupying about 20% of
the cereal area. Canadian acreage declined by 20% during the
12-year period described in Table 7, and the 2013 harvested
area of 2.6 Mha was 30% below the 1992 figure of 3.8 Mha.
Consumption and Trade
Barley is a relatively minor crop in both the United States
Table 6 summarizes sorghum production, consumption, and and Mexico, occupying between 2% and 3% of the cereal area.
trade in North America for the period 2002 to 2011. With In the United States, barley is grown primarily in northern
exports averaging 4.6 mmt per year comprising almost half of states, with Idaho, Montana, North Dakota, and Washington
domestic production, the United States is the worlds leading combining to account for almost 80% of the US harvested area.
Table 7 Barley production: Area harvested (000 ha) and production (000 mt)
Year Area Prod. Area Prod. Area Prod. Area Prod. Area Prod.
2002 1669 4940 3342 7468 282 737 5293 13 145 55 270 136 724
2003 1913 6059 4397 12 164 365 1085 6675 19 308 57 736 142 565
2004 1627 6091 3841 12 557 326 932 5794 19 579 57 567 153 905
2005 1323 4613 3634 11 678 306 761 5263 17 052 55 357 138 680
2006 1194 3923 3223 9573 315 869 4733 14 365 56 386 139 522
2007 1417 4575 3998 10 984 286 653 5701 16 211 55 860 134 175
2008 1529 5230 3502 11 781 311 781 5342 17 792 55 533 154 788
2009 1260 4949 2918 9517 239 519 4416 14 985 53 956 151 782
2010 998 3925 2387 7605 268 672 3652 12 203 47 368 123 683
2011 906 3392 2365 7756 218 487 3489 11 635 48 488 132 951
2012 1313 4796 2060 8012 328 1032 3701 13 840 49 573 133 507
2013 1214 4683 2652 10 237 297 594 4163 15 514 49 781 144 755
Avg. 1364 4765 3193 9944 295 760 4852 15 469 53 573 140 586
As in Canada, the US area in barley has also declined from food and seed. Usage for animal feed has declined in step with
2.9 Mha in 1992, to 1.7 Mha in 2002, and down to 1.3 Mha in reduced production, while the amounts used for processing
2013, a reduction of over 55% in 22 years. The decline can have been fairly steady. North America is a net exporter of
largely be attributed to the expansion of corn production. In barley, with exports accounting for around 12% of average
Mexico, the area in barley has been stable, averaging annual production.
295 000 ha. Average yields are highest in the United States, at Canada, with exports averaging about 2.2 mmt, is typically
3.5 mt ha1, and lowest in Mexico at 2.5 mt ha1, while Cana- one of the top five exporting countries. Of its remaining
dian yields averaged 3.2 mt ha1 over the 12-year period. Aver- domestic supply, almost 90% is used for animal feed. In the
age yields increased slightly in the United States and Canada, United States, however, only 30% of the available supply goes
by 0.06 and 0.1 mt ha1 per year, respectively, between 2002 to animal feed, with the majority of the crop going to the
and 2013, but showed no growth in Mexico. Overall produc- malting sector. In 2011, for example, malting varieties
tion of barley in North America is trending downward, how- accounted for 75% of the acreage planted in Idaho (USDA,
ever. In 2012 and 2013, average production was 14.7 mmt NASS, Idaho Barley Varieties) and over 90% of the acreage in
compared to an average of 21.8 mmt in 1992 and 1993. North Dakota (North Dakota Barley Council).
Table 9 Oat production: Area harvested (000 ha) and production (000 mt)
Year Area Prod. Area Prod. Area Prod. Area Prod. Area Prod.
2002 833 1684 1379 2911 42 64 2254 4658 12 436 25 424
2003 898 2096 1415 3377 46 94 2360 5567 12 128 26 541
2004 723 1679 1234 3467 67 103 2024 5249 11 663 26 036
2005 738 1667 1271 3283 77 127 2086 5077 11 250 23 697
2006 633 1357 1537 3852 76 152 2246 5362 11 661 22 612
2007 609 1313 1816 4696 72 125 2497 6134 11 990 25 806
2008 567 1294 1448 4273 100 148 2115 5714 11 279 25 863
2009 558 1351 980 2906 72 130 1610 4388 10 195 23 345
2010 511 1178 906 2480 67 111 1484 3769 9096 19 723
2011 380 779 1030 2997 52 51 1461 3826 9638 22 316
2012 423 987 985 2812 47 84 1455 3884 9580 21 312
2013 417 1016 1107 3888 49 91 1574 4995 9759 23 821
Avg. 607 1367 1259 3412 64 107 1930 4885 10 890 23 875
the area, the United States with 31%, and Mexico < 4%. Table 9 Saskatchewan, and Manitoba). Yields in the United States aver-
provides data on oat production between 2002 and 2013. aged 2.3 mt ha1 between 2002 and 2013, compared to
Worldwide, oats accounted for only 1.7% of the area in cereals, 2.7 mt ha1 in Canada and 1.7 mt ha1 in Mexico. Yields in
and between 2002 and 2013, the harvested area declined by the United States and Canada evidenced a slight upward trend
more than 20%. North America accounts for about 20% of the over the period, but as was the case with barley, there was no
worlds production. similar upward trend in Mexican yields. The area in oats in
The EU and Russia are the leading oat producers, account- Mexico ranged from 42 000 ha in 2002 to 100 000 ha in 2008,
ing for roughly 35% and 20% of production, respectively. with most production in the northwest of the country.
Canada is the third ranked producer with around 15% of
world production (Agriculture and Agri-Food Canada). In
Consumption and Trade
Canada, oats occupy about 8% of the area in cereals, a share
similar to that of maize. In the United States and Mexico, its Table 10 summarizes production, consumption, and trade in
share is much lower at 1.1% and 0.6%, respectively. oats in North America between 2002 and 2011. As with barley,
Mirroring the trend in world acreage, between 2002 and oats are used primarily for animal feed. Of the average domes-
2013, the area in oats declined in both the United States and tic supply of 5 mmt, around 3.5 mmt or 70% was used for
Canada. The decline has been sharpest in the United States animal feed, with most of the remainder used for human
where the 416 Tha harvested in 2013 represented half the area consumption. The amount of oats used for human food has
harvested in 2002 and less than a quarter of the 1.8 Mha remained relatively constant averaging 1.3 mmt per year in
harvested in 1992. In the United States, oats are grown primar- the period between 1992 and 2001 and 1.25 mmt between
ily in the North Central states, with Minnesota, North and 2002 and 2011. As production declined, the amount used for
South Dakota, Wisconsin, and Iowa jointly accounting for animal feed also declined, averaging 3.5 mmt per year between
more than half of the US harvested area. In Canada, most of 2002 and 2011, compared to 4.9 mmt between 1992
the production is in the Prairie Provinces (Alberta, and 2001.
GRAINS AROUND THE WORLD | Grain Production and Consumption: Cereal Grains in North America 399
Trade in oats is fairly limited with exports from Canada to The US area in rice declined by over 20% between 2002 and
the United States accounting for between 55% and 65% of 2013, from 1.3 Mha to just under 1.0 Mha. An average yield
annual world trade, making Canada the worlds largest increase of 0.08 mt ha1 per year tempered the decline in
exporter and the United States the largest importer. Canadas production to about 10% over the 12-year period. The leading
exports averaged 1.7 mmt, equivalent to half of domestic pro- rice-growing state is Arkansas with about 44% of the total US
duction, while US imports exceeded its own domestic produc- area, followed by California with 22%. Rice is also grown in
tion. Mexico also imports more oats than it produces Louisiana, Mississippi, Missouri, and Texas. Rice is classified as
domestically with imports coming primarily from Canada. either long grain, medium grain, or short grain depending on
the shape and length of the grain. US production is predomi-
nantly long grain, which is grown mostly in Arkansas and the
Rice other southern states and accounts for over 70% of US produc-
tion. Medium and short grain types are grown almost exclu-
Data on North American and world rice production (paddy sively in California, although Arkansas does have some
equivalents) from 2002 to 2013 are provided in Table 11. The medium grain production.
United States accounts for over 95% of North American rice
acreage and production. Mexico harvests about 50 000 ha
Consumption and Trade
annually accounting for < 0.5% of its cereal area. Because the
climate is unsuitable, no rice is grown in Canada. Table 12 provides data on rice production, consumption, and
Combined, the United States and Mexico account for < 2% trade (in paddy equivalent tonnes, not milled) in North America
of world production. Global production is concentrated in between 2002 and 2011. Although the United States is relatively
Asia, with China and India combining for over half the world small in terms of global production, it is a major exporter. On
total. Another 20% of production comes from Thailand, average, over the 10-year period considered in Table 12, the
Vietnam, Indonesia, and the Philippines. United States exported over half of its domestic production and,
Table 11 Rice production: Area harvested (000 ha) and production (000 mt)
Year Area Prod. Area Prod. Area Prod. Area Prod. Area Prod.
2002 1298 9569 0 0 50 227 1348 9796 147 628 571 388
2003 1213 9067 0 0 60 273 1273 9340 148 509 587 031
2004 1346 10 540 0 0 62 279 1408 10 818 150 554 607 986
2005 1361 10 108 0 0 57 291 1419 10 399 154 989 634 444
2006 1142 8826 0 0 70 337 1212 9163 155 581 641 207
2007 1112 8999 0 0 71 295 1183 9293 155 040 656 978
2008 1204 9241 0 0 50 224 1255 9466 159 993 688 415
2009 1256 9972 0 0 54 263 1310 10 235 158 130 686 958
2010 1463 11 027 0 0 42 217 1505 11 244 161 189 701 999
2011 1059 8389 0 0 34 173 1093 8562 162 800 726 122
2012 1084 9051 0 0 32 179 1116 9230 162 317 738 188
2013 1000 8613 0 0 33 180 1033 8793 164 722 745 710
Avg. 1211 9450 0 0 51 245 1263 9695 156 788 665 535
with an 11% share of world exports, was the fourth largest of maize for as many years as possible. Subtract the US and
exporter following Thailand, Vietnam, and India and ranking Canadian data from the world data to facilitate comparison
marginally ahead of Pakistan. Mexico is the largest export market between the United States, Canada, and the rest of the world
for US rice with other important markets including Japan, Turkey, (RoW). Divide production by area harvested to obtain aver-
and a number of Central American and Caribbean countries. age yield, and plot the average yield over time for the United
The United States also imports rice, but imports are mostly States, Canada, and the RoW. Comment on the trends.
aromatic varieties that are not grown domestically, such as
jasmine from Thailand and basmati from India and Pakistan
(USDA, ERS, Rice). Canadas domestic supply is all imported
with about half coming from the United States. Mexico is also
reliant on imports, which account for close to 80% of its
Further Reading
supply. Those imports come primarily from the United States, Lukow OM and McVetty PBE. Grain production and consumption: Cereal grains in
but Mexico also imports from Uruguay and Pakistan (USDA, North America. Encyclopedia of Food Grains, 1st edn.
FAS, Mexico Grain and Feed Annual). In North America, rice is United States Department of Agriculture (USDA), Foreign Agricultural Service (FAS).
used predominantly for human food with some use in food (2013) Global Agricultural Information Network (GAIN) Report Number MX3024.
Grain and Feed Annual, Mexico. http://gain.fas.usda.gov/Recent%20GAIN%
manufacturing (rice starch, beer, or alcoholic beverages). 20Publications/Grain%20and%20Feed%20Annual_Mexico%20City_Mexico_3-
15-2013.pdf.
Topic Highlights each North American countries and on the continent since
1960s. The last 10-year mean disposition and consumption of
The major oilseeds grown in North America are canola, each of these major oilseeds in each of the countries and on the
soybean, linseed/flax, and sunflower. continent will also be reviewed. All comments regarding crop
Whereas sunflower is native to North America, the other disposition and consumption refer to averages of the 2000s,
species have been introduced. unless specifically stated otherwise. Area and production data
Statistics are provided for disposition and consumption of for all crops relate to crops harvested for dry grain only. For
these oilseeds as 10-year means, as well as historical and disposition and consumption of commodities, the definitions
recent annual figures for acreage and production for Can- of the listed elements are taken primarily from the Food and
ada, the United States, and Mexico. Agriculture Organization (FAO) of the United Nations. Values
Canola (Brassica napus) has increased in area because it is given for production relate to the total domestic production.
adapted to cool Canadas climate and has been bred to have Imports cover all movements into the country of the commod-
low levels of erucic acid in seed oil and glucosinolates in ity. Stock variation is the changes in stocks at all levels between
seed meal. the production and the retail levels. A negative sign indicates
Canada is the major producer of canola with much of it increases in stocks (add to stock) and no sign means net
being exported. decreases (from stock). Exports cover all movements by the
The United States is the major producer of soybeans, with crop commodity out of the country. Domestic supply is defined
much of it being used domestically. as (production imports) (exports changes in stocks). Feed
Linseed and sunflower are relatively minor crops in North is the amount of the commodity in question fed to livestock,
America. whether domestically produced or imported. Seed comprises
the amount of commodity used for reproduction. Waste
includes the amount of crop commodity lost in processing,
Learning Objectives storage, and transportation. Processing is the amount of com-
modity used for the manufacture of processed products that
cannot be converted back to their originating primary commod-
To achieve understanding of the origins and production of
the major oilseed species in Canada, the United States, and ities. Food available for human consumption comprises the
Mexico. amounts of the commodity not detailed otherwise in the bal-
ance sheet. Other uses comprise quantities of the commodity
To understand the uses of these species and the extent of
their involvement in world trade. used for manufacture of nonfood products.
few oil crops that can be grown in the temperate regions of the Table 1 Area harvested (ha) for major oilseeds grown in Canada
world. Among these, B. napus is predominantly grown in North
American countries; however, B. rapa and B. juncea can also be Year Canola Soybean Linseed Sunflower
found on limited areas (<1%). B. rapa is more suited to short 196367 473 395 104 650 721 569 23 253
growing season areas, while B. juncea is more suitable to the 196872 1 269 886 139 534 830 993 49 768
hotter and drier regions. B. napus exists in both spring and 197377 1 311 010 178 307 532 374 34 845
winter growth habit forms, but it is the spring form that pre- 197882 2 296 424 296 980 621 610 116 520
dominates in North American production; winter form is 198387 2 695 840 403 880 647 360 46 660
grown on very limited areas in the United States. This oilseed 198892 3 060 920 555 400 508 800 58 360
crop was first introduced in Canada and grown at a commer- 199397 4 692 620 855 900 679 520 58 060
cial scale during the Second World War. 19982002 4 653 100 1 027 500 690 180 75 580
2003 4 689 200 1 048 600 728 400 105 600
Traditional rapeseed and mustard seeds contain a high level
2004 4 867 000 1 173 500 518 000 55 400
of erucic fatty acid in oil (>40% of the total fatty acids) and a
2005 5 175 400 1 165 200 732 600 70 800
high level of glucosinolates in seed meal (>100 mmol g1). The 2006 5 238 200 1 201 200 785 200 76 900
erucic acid is undesired in edible oil, while the seed meal 2007 6 327 600 1 171 500 524 000 78 900
containing a high level of glucosinolate is undesirable for use 2008 6 494 400 1 195 400 625 200 68 800
of this meal as protein source in animal feed. The oil and meal 2009 6 513 300 1 383 300 623 300 63 500
quality of these oilseed crops has been dramatically improved 2010 6 848 300 1 476 800 353 300 51 400
(virtually eliminated erucic acid in oil and reduced glucosino- 2011 7 471 300 1 542 400 273 200 13 800
late content to <30 mmol g1 seed meal) in 1960s1970s 2012 8 379 900 1 668 400 360 000 40 000
through research, and the name canola for this quality- 200312 6 200 460 1 302 630 552 320 62 510
improved-type Brassica oilseed crops has been adopted in Italicized 5- or 10-year means.
most parts of the world. The term double-low rapeseed or Source: Food and Agriculture Organization of the United Nations (FAO).
rapeseed or oilseed rape is still being used for this quality-
improved type in some countries as well as in some articles.
Most of the world production of Brassica oilseeds including the Table 2 Area harvested (ha) for major oilseeds grown in the
production acreages in North American countries has trans- United States
ferred to canola quality type; therefore, the word canola will
be used in this article. Year Canola Soybean Linseed Sunflower seed
Currently, most of the canola grown in North American
196367 0 13 775 857 1 078 310 37 669
countries is hybrid cultivars. About half of the canola crop is
196872 0 17 268 909 827 732 146 238
Roundup (glyphosate) herbicide-tolerant and half Liberty
197377 0 21 679 194 571 738 472 853
(glufosinate) herbicide-tolerant; Clearfield (imidazolinone) 197882 0 27 309 878 286 500 1 653 180
herbicide-tolerant crops cover a small acreage (<5%). 198387 3060 24 741 123 230 500 1 078 720
198892 38 960 23 432 403 93 014 831 596
199397 163 604 25 261 600 60 464 1 183 810
Area Harvested
19982002 515 835 29 199 730 202 993 1 157 571
Canada has harvested canola on an average from about 6.2 2003 432 695 29 330 310 237 958 889 104
million hectares (Mha) (14.4% of Canadian arable land) dur- 2004 338 240 29 930 060 206 797 692 425
ing the period of 200312 (Table 1). Since 1980s, canola 2005 451 634 28 834 570 386 480 1 056 240
2006 413 593 30 190 680 310 397 716 300
production area in Canada has increased more than twofold
2007 468 070 25 959 245 141 237 814 236
with a sharp increase in the recent years (>8 Mha in 2012)
2008 400 160 30 222 654 137 595 969 640
(Table 1). The vast majority of canola production area in 2009 329 780 30 906 980 127 070 790 560
Canada is in the Prairie Provinces Alberta, Saskatchewan, and 2010 579 880 31 003 300 169 160 758 310
Manitoba, while small production areas can be found in 2011 422 620 29 856 410 70 010 589 960
Ontario and British Columbia. 2012 700 560 30 798 530 135 980 745 030
The United States started to grow canola from the late 200312 453 723 29 703 274 192 268 802 181
1980s. This country on an average has harvested canola from
Italicized 5- or 10-year means.
about 453 725 ha (0.3% of American arable land) annually
Source: Food and Agriculture Organization of the United Nations (FAO).
during the period of 200312 (Table 2); this is more than a
twofold increase since the 1990s. The vast majority of canola
production area in the United States is the northern state of land) annually in the last 10 years (Table 3). The area of canola
North Dakota where spring canola is grown. Small production harvested in Mexico has varied widely during the years. Canola
areas can also be found in Minnesota, Montana, Idaho, Ore- production area in Mexico has reduced to less than half in the
gon, Washington, and Oklahoma. Recently, winter canola has recent years when compared with 1970s (Table 3). This varia-
captured about 15% of the canola production area in the U- tion in harvested area is typical for any small-volume crop. The
nited States, primarily grown in Oklahoma and small areas in vast majority of canola production area in Mexico is in the
the Pacific Northwest region. central area of the country.
Mexico is a very small canola producer, harvesting on an For the entire North American continent, canola was har-
average from only about 2000 ha (0.01% of Mexican arable vested from an average 6.7 Mha (2.9% of North American
GRAINS AROUND THE WORLD | Grain Production and Consumption: Oilseeds in North America 403
Table 3 Area harvested (ha) for major oilseeds grown in Mexico Table 5 Production (tonnes) for major oilseeds grown in Canada
Year Canola Soybean Linseed Sunflower Year Canola Soybean Linseed Sunflower
196367 5800 41 925 17 623 0 196367 429 557 202 003 518 155 15 310
196872 6400 151 696 17 340 13 415 196872 1 257 833 278 264 689 302 41 111
197377 5800 288 624 10 262 3354 197377 1 403 862 374 759 443 500 36 893
197882 3600 297 419 4795 9905 197882 2 697 258 663 476 609 519 142 980
198387 807 421 432 683 9545 198387 3 430 320 978 700 757 940 57 440
198892 177 315 880 85 4098 198892 3 758 120 1 309 720 546 360 85 100
199397 1025 166 454 27 1295 199397 6 120 800 2 260 540 887 780 76 340
19982002 3200 75 084 367 470 19982002 6 636 900 2 438 280 820 740 122 840
2003 1000 67 880 1350 124 2003 6 771 200 2 273 300 754 400 142 300
2004 1000 88 840 2900 90 2004 7 673 600 3 043 900 516 900 52 200
2005 4000 96 266 1400 22 2005 9 483 300 3 155 600 990 600 84 400
2006 5000 54 211 8500 175 2006 9 000 300 3 465 500 988 800 157 300
2007 3000 62 580 10 300 3 2007 9 601 100 2 695 700 633 500 124 800
2008 2000 75 767 7300 4 2008 12 642 900 3 335 900 861 100 112 200
2009 2181 64 740 2600 216 2009 12 889 200 3 506 800 930 100 101 900
2010 0 153 473 3600 1878 2010 12 773 300 4 345 300 423 000 67 600
2011 185 155 513 2600 1850 2011 14 164 500 4 246 300 368 300 19 800
2012 10 142 329 2500 1280 2012 15 409 500 4 870 160 489 000 73 600
200312 1838 96 160 4305 564 200312 11 040 890 3 493 846 695 570 93 610
Table 4 Area harvested (ha) for major oilseeds grown in North Production
America
Canada has produced on an average about 11 Mt canola annu-
Year Canola Soybean Linseed Sunflower ally during the period of 200312 (Table 5). Canola produc-
tion in Canada has varied by a factor of about 2 during the
196367 479 195 13 922 433 1 817 502 60 922 2000s (Table 5). This variation resulted from the increase in
196872 1 276 286 17 560 138 1 676 065 209 421 harvested area as shown in Table 1 and about a 25% increase
197377 1 316 810 22 146 124 1 114 375 511 051
in yield with the introduction of hybrid cultivars. The average
197882 2 300 024 27 904 277 912 904 1 779 605
yield of canola in Canada has reached to 1.78 tonnes ha1 in
198387 2 699 707 25 566 435 878 543 1 134 925
198892 3 100 057 24 303 683 601 899 894 054 the recent years as compared to 1.35 tonnes ha1 in 1990s.
199397 4 857 249 26 283 954 740 011 1 243 165 The United States has produced on an average
19982002 5 172 135 30 302 314 893 539 1 233 620 754 943 tonnes of canola annually during the period of
2003 5 122 895 30 446 790 967 708 994 828 200312 (Table 6). Canola production in this country has
2004 5 206 240 31 192 400 727 697 747 915 increased by about 1.5-fold in recent years when compared
2005 5 631 034 30 096 036 1 120 480 1 127 062 with production in the 1990s (Table 6). The average yield of
2006 5 656 793 31 446 091 1 104 097 793 375 canola in the United States has reached 1.66 tonnes ha1, sim-
2007 6 798 670 27 193 325 675 537 893 139 ilar to the Canadian average yield.
2008 6 896 560 31 493 821 770 095 1 038 444
Mexico has produced on an average 2746 tonnes canola
2009 6 845 261 32 355 020 752 970 854 276
annually during the period of 200312 (Table 7). The produc-
2010 7 428 180 32 633 573 526 060 811 588
2011 7 894 105 31 554 323 345 810 605 610 tion of this crop in Mexico has reduced to almost half in the
2012 9 080 470 32 609 259 498 480 786 310 2000s when compared with 1970s (Table 7). The average yield
200312 6 656 021 31 102 064 748 893 865 255 of canola in Mexico has been 1.49 tonnes ha1, slightly lower
than Canada and the United States.
Italicized 5- or 10-year means. For the entire North American continent, average annual
Source: Food and Agriculture Organization of the United Nations (FAO). canola production was 11.8 Mt during the period of 200312
(Table 8); this is about 22% of the worlds production of
Brassica oilseeds. In North America, there has been a sharp
arable land) annually during the period of 200312 (Table 4); increase in production in the last 10-year period mostly
this production area constitutes about 22% of the worlds reflecting Canadian production. Canada has produced
Brassica oilseeds crop acreage. The canola production area in 93.6%, the United States 6.4%, and Mexico <0.3% of the
this continent has doubled in the recent years when compared North American canola during the period of 200312. In
with the production area in the 1990s (Table 4). Of the total terms of global production, North America has produced
North American canola harvested area during the period of about 22% of the worlds Brassica oilseeds during the period
200312, Canadas share was 93.1%, the United States 6.8%, of 200312. The share of world canola for Canada, the United
and Mexico <0.5%. States, and Mexico in the last 5 year average is 22.0%, 1.4%,
404 GRAINS AROUND THE WORLD | Grain Production and Consumption: Oilseeds in North America
Table 6 Production (tonnes) for major oilseeds grown in the United Table 8 Production (tonnes) for major oilseeds grown in North
States America
Year Canola Soybean Linseed Sunflower Year Canola Soybean Linseed Sunflower
196367 0 22 592 554 682 119 40 319 196367 433 357 22 874 598 1 215 535 55 629
196872 0 31 646 150 626 942 155 861 196872 1 263 433 32 206 247 1 339 214 205 349
197377 0 40 105 814 344 966 596 032 197377 1 409 662 40 979 377 804 955 635 980
197882 0 55 071 062 233 298 2 256 880 197882 2 700 858 56 278 439 848 497 2 407 590
198387 2450 51 579 882 209 340 1 395 300 198387 3 433 783 53 326 233 967 827 1 461 224
198892 57 918 52 119 203 81 928 1 089 000 198892 3 816 190 54 051 454 628 429 1 176 857
199397 242 190 63 292 800 64 184 1 692 246 199397 6 364 140 65 843 445 952 010 1 769 907
19982002 769 017 75 110 850 247 081 1 726 263 19982002 7 410 317 77 667 860 1 068 078 1 849 665
2003 686 462 66 781 392 267 117 1 208 930 2003 7 458 662 69 180 698 1 022 477 1 351 385
2004 612 533 85 013 945 263 358 929 690 2004 8 288 133 88 191 192 782 258 982 122
2005 718 484 83 504 928 500 273 1 822 151 2005 10 206 784 86 847 763 1 491 833 1 906 582
2006 632 948 86 998 898 279 894 972 330 2006 9 641 248 90 545 511 1 274 694 1 129 808
2007 649 519 72 857 739 149 764 1 301 298 2007 10 254 619 75 641 810 790 464 1 426 105
2008 655 610 80 748 700 145 192 1 552 570 2008 13 302 510 84 237 622 1 011 392 1 664 775
2009 669 340 91 417 300 188 550 1 377 320 2009 13 561 821 95 045 000 1 120 450 1 479 552
2010 1 113 390 90 605 460 230 030 1 240 830 2010 13 886 690 95 118 425 655 530 1 312 227
2011 698 910 84 191 930 70 890 924 550 2011 14 863 440 88 643 464 440 990 947 176
2012 1 112 230 82 054 800 146 360 1 263 570 2012 16 521 880 87 172 460 637 060 1 338 478
200312 754 943 82 417 509 224 143 1 259 324 200312 11 798 579 86 062 395 922 715 1 353 821
Table 7 Production (tonnes) for major oilseeds grown in Mexico Table 9 Canada 10-year averages for disposition and consumption
(tonnes) 200009
Year Canola Soybean Linseed Sunflower
Crop Canola Soybean Linseed Sunflower
196367 3800 80 042 15 261 0
196872 5600 281 833 22 970 8376 Production 8 609 150 2 815 060 695 570 115 560
197377 5800 498 804 16 488 3056 Imports 200 832 479 988 13 000 21 823
197882 3600 543 900 5680 7730 Stock change 84 000 4000 1850
198387 1013 767 651 547 8484 Exports 4 799 534 1 242 257 530 570 82 127
198892 152 622 531 141 2757 Domestic supply 3 926 448 2 056 791 178 000 57 106
199397 1150 290 105 46 1321 Feed 288 376 336 197 5142
19982002 4400 118 730 258 562 Seed 45 593 122 440 30 260 954
2003 1000 126 006 960 155 Waste 266 662 27 000
2004 2000 133 347 2000 232 Processing 3 280 738 1 549 250 51 010
2005 5000 187 235 960 31 Food 45 080 21 804 8900
2006 8000 81 113 6000 178 Other uses 131 720
2007 4000 88 371 7200 7
2008 4000 153 022 5100 5 Statistics Canada, 2014.
2009 3281 120 900 1800 332 Source: Food and Agriculture Organization of the United Nations (FAO).
2010 0 167 665 2500 3797
2011 30 205 234 1800 2826
2012 150 247 500 1700 1308 canola used domestically. About 85% of the canola used domes-
200312 2746 151 039 3002 887 tically in Canada was used for processing and food manufacture,
7% for feed, and small amount for planting seed (Table 9).
Italicized 5- or 10-year means.
Source: Food and Agriculture Organization of the United Nations (FAO). Of the average annual 737 320 tonnes canola produced in
the United States and the 536 768 tonnes imported each year
during 2000s, 965 542 tonnes was used domestically and
and 0.003%, respectively. Currently, Canada ranks in the top
309 171 tonnes was exported (Table 10). Of the total domestic
five countries of the world for canola production.
supply in the United States, almost the whole amount was used
for processing and food manufacture (Table 10) and a small
amount for planting seed.
Disposition and Consumption
Mexicos domestic supply of canola is heavily dependent on
During 200009, of the average annual 8.6 Mt canola produced imports. During 200009, the country imported on an average
in Canada and the 200 832 tonnes imported, 3.9 Mt was used 1.1 Mt annually while producing only 5017 tonnes per year
domestically and 4.8 Mt was exported (Table 9). Thus, (Table 11). About 99% of the Mexicos domestic supply was
Canadian production was more than double the quantity of used for processing and food manufacture (Table 11).
GRAINS AROUND THE WORLD | Grain Production and Consumption: Oilseeds in North America 405
Production was used domestically and 871 tonnes was exported during the
period of 200009 (Table 11). Approximately 55% of the
Canada has produced on an average 3.5 Mt of soybean annu-
soybean used domestically in Mexico was used for food pro-
ally during the period of 200312 (Table 5). The production of
cessing (Table 11). Another 20% of the soybean domestic
soybean in Canada has varied by a factor of almost 3 during the
supply was used as animal feed and 20% was used in other
1990s and 2000s (Table 5), paralleling the variation in har-
uses. Minor amounts were used for planting seed and food
vested area for soybean in Canada shown in Table 1. The
(<3%).
average yield of soybean in Canada has been 2.8 tonnes ha1.
Of the average annual 82.5 Mt of soybean produced in
The United States has produced on an average 82.4 Mt of
North America and the 4.5 Mt imported, 57.4 Mt was used
soybean annually during the period of 200312 (Table 6).
domestically, and 31.1 Mt was exported during the period of
Soybean production has increased by more than 40% in the
200009 (Table 12). Over 60% of the soybean used domesti-
United States during the 1990s, from 52.1 Mt at the beginning
cally in North America was used for food processing (Table 12).
of the 1990s to 90.6 Mt at the end of the 2000s (Table 6). The
Small amounts were used for feed, seed, food, and other uses
average yield of soybean in the United States has been
(<8%).
2.9 tonnes ha1, similar to the Canadian average yield.
Mexico has produced on an average 151 000 tonnes of soy-
bean annually during the period of 200312 (Table 7). The
Linseed (Flax)
production of soybean in Mexico has varied by a factor of
8 during the 1990s, from a low of 81 000 tonnes early 1960s Introduction
to a high of 622 000 tonnes in early 1990s (Table 7). This
Linseed (Linum usitatissimum), also known as flax in North
variation in production is typical for any medium volume
America, is one of the earliest cultivated field crops, initially
crop. The average yield of soybean in Mexico during the
grown for its fiber. In North America, in the last two centuries,
1990s has been 1.7 tonnes ha1, lower than the soybean yields
linseed has been grown primarily for its oil. It is a spring
seen for Canada and the United States. Soybeans are grown in
annual, adapted to a wide range of soil and climatic conditions
Mexico under water-limiting rain-fed conditions.
in the cool temperate zones of the northern hemisphere. Lin-
For the entire North American continent, average annual
seed in North America is used primarily to produce industrial
soybean production during the period of 200312 was
oil and animal feed meal. Linseed oil is a drying oil used in
86.1 Mt (Table 8). There has been a year-to-year variation in
paints and varnishes and for the manufacture of linoleum
production, with a trend toward increased production, not
flooring. A trend toward bioproducts has increased this market
only mostly in the United States in recent years but also in
in the 2000s. Because linseed oil contains alpha-linolenic acid
Canada. In terms of North American production proportions,
(ALA), an essential omega-3 fatty acid, it has a variety of uses.
Canada has produced an average 4.6%, the United States
Whole linseed is fed to poultry to produce omega-3 fatty acid-
94.9%, and Mexico <0.5% of the soybean produced in
enriched eggs. Linseed meal is a component of rations fed to
North America. In terms of global soybean production,
pets, poultry, and animal. Small quantities of linseed are also
North America has produced on an average 33.0% of the
used in food products (breads and breakfast cereals), but a
soybean produced in the world. The share of world soybean
significant number of products are being developed using
production for Canada, the United States, and Mexico is
linseed, in response to the variety of health benefits, including
1.0%, 32.5%, and <0.2%, respectively. The United States
reducing heart disease and cancer risk. Additional products
is the top soybean producer of the world; however, it falls
have been created based on the health and nutrition claims
behind Brazil and Argentina, which represent almost a half
of linseed oil, lignans, and antioxidant flavonoids. Small quan-
(49%) of world production combined.
tities of linseed are used for (re)planting seed.
For the entire North American continent, linseed was har- Sunflower
vested from an average 748 000 ha (0.32% of North American
arable land) annually between 2003 and 2012 (Table 4). The Introduction
area harvested has varied widely during the 2000s (Table 4). Sunflower (Helianthus annuus) was domesticated as a food crop
Canadas average linseed harvested area was 73.7%, United in North America, perhaps as early as 3000 BC. This was
States 25.6%, and Mexico 0.6% of the total North American introduced to Europe in the 1600s and was successfully devel-
linseed harvested area. oped as an oil crop in Russia in the early 1800s. Russian plant
breeders were able to increase the oil content in sunflower seed
Production from less than 30% to over 50% the major factor permitting
the development of sunflower as oil crop for the temperate
Canada has produced on an average 695 570 tonnes of lin- areas of the world.
seed annually between 2003 and 2012 (Table 5). The pro- Sunflower is used primarily for edible vegetable oil and the
duction of linseed in Canada has varied widely during the seed meal is used for animal feed (Table 10). Moderate quan-
2000s, from a low of 368 000 tonnes to a high of 0.99 million tities of sunflower are directly used for animal feed and food
tonnes (Mt) (Table 5), paralleling the widely varying produc- manufacture (confectionary and bakery products) (Table 10).
tion area for linseed harvested in Canada as shown in Table 1. Small amounts are used for planting seed (<5%).
The average yield of linseed in Canada between 2003 and
2012 was 1.26 tonnes ha1.
Area Harvested
The United States has produced on an average
224 000 tonnes of linseed annually (Table 6). The production Canada has been growing sunflower commercially since the
of linseed in the United States has varied widely during the early 1940s; however, the production area remains small. Dur-
2000s, from a low of 71 000 tonnes to a high of 500 000 tonnes ing the period of 200312, Canada on an average has har-
(Table 6), paralleling the widely varying production area for vested sunflower from about 62 500 ha (about 0.1% of
linseed harvested in the United States shown in Table 2. The Canadian arable land) annually (Table 1). The area of sun-
average yield of linseed in the United States has been flower harvested in Canada varied greatly during the years
1.02 tonnes ha1, slightly lower than the Canadian average since the 1960s from a low of <15 000 ha to a high of
yield. >100 000 ha (Table 1), which is typical for a small-volume
For the entire North American continent, average annual crop. The vast majority of sunflower production area in
linseed production was 923 000 tonnes annually between 2003 Canada is in the Prairie Province of Manitoba.
and 2012 (Table 8). In terms of North American production The United States on an average has harvested sunflower
proportions, Canada has produced an average of 75%, United from about 0.8 Mha (0.5% of American arable land) annually
States 24%, and Mexico 0.3% of the linseed produced in North during the period of 200312 (Table 2). This is about a 15-fold
America. In terms of global linseed production, North America increase in the production area when compared with the pro-
has produced on an average 43.6% of all the linseed produced in duction area in 1960s; this is, however, about a 30% reduction
the world. The share of world linseed production for Canada and when compared with the 1990s. The vast majority of sunflower
the United States is 32.8% and 10.5%, respectively. Canada was production area in the United States is in the central Plains
the top linseed producer in the world in the 2000s, dropping states (from Dakotas to Texas, with North Dakota being the
from 34.8% in the 1990s due to the identification of low-level predominant production area state).
presence (LLP) of genetically engineered flax seeds. Mexico has been growing sunflower since early 1970s. Dur-
ing the period of 200312 the country on an average has
harvested sunflower from only 564 ha (0.002% of Mexican
Disposition and Consumption
arable land) annually (Table 3); this is <10% of the produc-
Canada is a net exporter of linseed. Of the average annual tion area when compared with the acreage of this crop in the
695 570 tonnes of linseed produced in Canada and the 1970s and 1980s. The area of sunflower harvested in Mexico
13 000 tonnes imported into Canada, 178 000 t was used has varied widely since 1970s; the crop almost vanished in
domestically and 530 570 tonnes was exported, primarily for mid-2000s (Table 3). The greater part of the sunflower pro-
industrial oil uses (Table 9). Of the linseed used domestically duction area is in central Mexico.
in Canada, only small amounts of linseed (<5%) were used For the entire North American continent, sunflower was
for food, while 74% was used for other uses combined (seed harvested from an average 0.9 Mha (0.4% of North American
crushed for industrial oil and meal production for feed), and arable land) annually during the period of 200312 (Table 4);
17% was used for planting seed (Table 9). An average of this constitutes about 3.6% of the total sunflower production
304 000 tonnes was retained on farm, for either seed or tem- area in the world. The area harvested in North America has
porary storage (Statistics Canada 2014). Canada is a major varied greatly in the last 50 years (Table 4). Of the total
exporter of linseed to the world (530 570 tonnes), with sig- sunflower harvested area in this continent during 200312,
nificant exports to Europe, the United States, Japan, South Canadas share is 7.2%, the United States 92.7%, and Mexico
Korea, and China. Canada currently supplies nearly 60% of only 0.07%.
the linseed used in the United States.
The United States is a net importer of linseed but exports
Production
80% linseed to either Belgium or Canada. Because linseed is a
relatively minor bioproduct crop, internal disposition of flax is Canada has produced on an average 93 610 tonnes of sun-
not tracked by USDA or FAO. flower annually during the period of 200312 (Table 5). The
408 GRAINS AROUND THE WORLD | Grain Production and Consumption: Oilseeds in North America
production of sunflower in Canada has varied greatly from a Exercises for Revision
low of <20 000 tonnes to a high of >150 000 tonnes (Table 1)
reflecting the variation in harvested area. The average yield of Describe the contrasts in the origins and cultivation of these
sunflower in Canada has been 1.5 tonnes ha1. major oilseed species in North America.
The United States has produced on an average 1.3 Mt of Explain the role of plant breeding in the improvement of
sunflower annually during the period of 200312 (Table 6) these crop species.
with an average yield of 1.6 tonnes ha1 (similar to the What features distinguish oilseed production, quality, and
Canadian average yield). Compared to the 1990s, the country consumption in your region from these factors in North
has produced about 20% less sunflower in the last 10 years. America (or choose one of the North American countries)?
Mexico has produced on an average 887 tonnes of sun- Compare and contrast the utilization of the products of
flower annually during the period of 200312 (Table 7). The these four major oilseeds.
production of sunflower in this country has varied greatly; this
is primarily due to the variation in production area (Table 1).
Exercises for Readers to Explore the Topic Further
The average yield of sunflower in Mexico has been
1.6 tonnes ha1, very similar to the yields for Canada and the
United States.
Flax is a minor but significant crop in North America, where
it is primarily grown for its seed. Find out about the possi-
For the entire North American continent, average annual
ble use of the traditional product of flax as a fiber.
sunflower production was 1.4 Mt during the period of
200312 (Table 8). This is about 23% reduction in production
Find out the differences in composition and properties
between the oils obtained from these four oilseed species.
when compared with the production in the 1990s. In terms of
North American production proportions, Canada has pro-
Find out the differences in composition and properties
between the residues (meal) left after oil removal obtained
duced 6.9%, the United States 93.0%, and Mexico <0.1% of
from these four oilseed species.
the total sunflower produced in North America during
200312. In terms of world sunflower production during
Obtain details of oilseed-production areas of North Amer-
ica for each of the species. How similar (or different) are the
200312, North America has produced 4.2% of all the sun-
respective growing sites in each of these countries?
flowers produced in the world (Canadas and Mexicos share is
<0.4% and the United Statess share is 3.9%).
See also: Agronomy of Grain Growing: Canola: Agronomy;
Soybean: Agronomy; Breeding of Grains: Canola/Rapeseed:
Disposition and Consumption Genetics and Breeding; Soybean: Germplasm, Breeding, and Genetics;
Fats: Healthy Fats and Oils; Lipid Chemistry; Grain Composition
Of the average annual 115 560 tonnes of sunflower produced
and Analysis: The Composition of Food Grains and Grain-Based
and the 21 823 tonnes of sunflower imported in Canada dur-
Products; Grain Harvest, Storage and Transport: Canola:
ing the period of 200009, 57 106 tonnes was used domesti-
Harvest, Transport, and Storage; Grains Around the World: Grain
cally and 82 127 tonnes was exported (Table 9). About 89% of
Production and Consumption: Cereal Grains in North America; Non-
the sunflowers used domestically in Canada were utilized in
food Products from Grains: Cereal Grains as Animal Feed; Oilseed
processing and food manufacture, 9% was used as animal feed,
and Legume Processing: Canola: Processing; Soybean:
and a small amount was used as planting seed (2%) (Table 9).
Processing; Soybean: Soy Concentrates and Isolates; The Oilseeds:
Of the average annual 1.3 Mt of sunflower produced and
Canola: Overview; Linseed: Overview; Oilseeds: Overview; Soybean:
the 71 636 tonnes of sunflower imported in the United States
Overview; Sunflower: Overview.
in the 2000s, 1.2 Mt was used domestically and 147 185 tonnes
was exported (Table 10). About 70% of the sunflowers used
domestically in the United States were utilized in processing Further Reading
and food manufacture, 28% were used as animal feed, and 1%
were used for planting seed (Table 10). Robbelen G, Downey RK, and Ashri A (eds.) (1989) Oil Crops of the World, p. 554.
Mexicos average annual domestic supply of sunflower in Toronto: McGraw-Hill.
Slinkard AE and Knott DR (eds.) (1995) Harvest of Gold: The History of Field Crop
the 2000s was about 26 300 tonnes of which only 1% was Breeding in Canada, p. 367. Saskatoon, Saskatchewan: University Extension Press/
produced within the country and the rest being imported University of Saskatchewan.
(25 472 t) (Table 11). All domestic supply of the sunflower in
Mexico was used in processing and food manufacture
(Table 11). Relevant Websites
Of the average annual 1.5 Mt of sunflower produced and http://www.canolacouncil.org/ Canola Council of Canada.
the 118 931 tonnes of sunflower imported in the North Amer- http://www.canadasunflower.com/ National Sunflower Association of Canada (NSAC).
ican countries during the period of 200009, about 84% of this http://faostat3.fao.org/faostat-gateway/go/to/download/Q/*/E Food and Agriculture
supply was used domestically (Table 12). Over 70% of the Organization of the United Nations.
http://www.sunflowernsa.com/ National Sunflower Association.
sunflower used domestically in North America was used for http://www.statcan.gc.ca/start-debut-eng.html Statistics Canada.
processing and food manufacture, 27% was used as animal http://www.uscanola.com/ U.S. Canola Association.
feed, and 1% was used for planting seed (Table 12). http://usda.gov United States Department of Agriculture (USDA).
Grain Production and Consumption in Oceania: Australia and Pacific Countries
CW Wrigley, University of Queensland, Brisbane, QLD, Australia
RJ French, University of Western Australia, Crawley, WA, Australia
2016 Elsevier Ltd. All rights reserved.
Table 1 Production, in kilotonnes, of grains according to categories improvement of grain quality for all grain species. A world-first
in crop year 2012 for countries of Oceania where there is significant innovation of that time (1890s) was the development of a very
production small-scale flour mill, suited to evaluating the milling quality
of the small samples of grain coming from Farrers wheat
Country Cereal grains Oilseeds Pulses
breeding (Figure 5). Subsequent years brought the excess pro-
Australia 43 372 5291 2956 duction of wheat that justified Farrers vision of export to
Fiji 9 0.3 2.4 England, largely as a result of the development of new wheat
Micronesia 0.3 varieties (Table 3). These contributions by Farrer were recog-
New Caledonia 3 nized by the issue of $2 bank note when Australia changed to
New Zealand 1168 5 23 decimal currency in 1966 (Figure 2).
Papua New Guinea 16 1.4 4 However, the greatest expansion in wheat production
Solomon Islands 4 6
occurred in the second half of the twentieth century, with
Vanuatu 0.8 3
production expanding from about 2 million tonnes (Mt) in
Source: http://faostat.fao.org. 1940 to a high of about 25 Mt in 1999/2000. Nevertheless,
production has fluctuated on occasions due to drought. The
high of 2000 contrasts with the drought year of 2002/2003,
when the national wheat crop was only a little over 10 Mt, less
Table 2 Production, in kilotonnes, of grain crops for the 2012 crop
than half of the crops either side (about 24 Mt in both 2001/
year in Australia, New Zealand, and Papua New Guinea (PNG)
2002 and 2003/2004). Similar reductions in production were
Crop Australia New Zealand PNG experienced for all grains in the 2002/2003 drought year. More
recently, fluctuations have not been so great (Table 4) and total
Cereals wheat production has continued to grow, with the production
Barley 8221 439 of nearly 30 Mt in 2012.
Maize 451 211 10
Millet 45
Oats 1262 18
Phalaris, canary seed 6 Diversification of Markets for Australian Wheat
Rice, paddy 919 1.2
Rye 40 Further improvements in the quality of wheat varieties and in
Sorghum 2239 4.5 management practices through the early twentieth century
Triticale 285 extended opportunities for grain exports. Initially, these
Wheat 29 905 489 exports were to England, where the grain was used for tradi-
Oilseeds tional products, mainly bread. This situation changed dramat-
Cottonseed 1694 ically in the second half of the twentieth century. In recent
Linseed 7 2.4
decades, Australia has produced about four times more than its
Peanuts 25 1.4
domestic needs, so that 7080% of the wheat harvest is
Rapeseed 3427 2.7
Safflower 5 exported. Exported wheat goes to a wide range of countries,
Soybeans 86 including Egypt, Indonesia, Iran, Iraq, Japan, and China.
Sunflower seed 47 In the early days of this expansion, there was minimal
Pulses 4 diversification of the crop on the basis of grain quality. Fair
Beans (various) 475 average quality was the basis of assessment of quality for most
Chickpeas 673 of the crop, although the region of origin of the FAQ wheat
Lentils 463 3 provided a good indication of quality (Figure 6). The 1960s
Lupins 982 brought the realization that export markets for Australian
Peas 343 20
wheat included a diversity of foods that could then be classed
Vetches 20
as exotic (to Australians), such as Arabic flat breads in the
Gaps indicate insignificant production. Middle East and a range of noodles and steamed breads in
Source: http://faostat.fao.org. Asian countries. Furthermore, in countries such as Indonesia,
even the manufacture of leavened bread involved non-
traditional methods that required wheat of quality different
Farrers use of crossbreeding was partly aimed at developing from the English market.
wheats with grain quality suited for export to England by Accordingly, experts were brought to cereal chemistry lab-
combining the dough quality of Canadian Fife wheats with oratories in Australia from the new export markets to demon-
the drought tolerance of wheats introduced from countries strate the production of these exotic products. Figure 7 shows
such as India, Palestine, and South Africa. This vision led him an expert from the Middle East with experimental Arabic
to enlist the help of the agricultural chemist, Frederick Guthrie, pocket breads, from both normal-scale baking and small-
to evaluate the milling and baking quality of the small samples scale baking. As a result of seeking this expertise, it was possible
of his many crossbred lines. for wheat breeders to select new varieties that would suit these
Thus, there was a start to the development of small-scale diverse quality requirements. The resulting research led to the
test methods and (probably) the worlds first breederchemist establishment of specifications for the wide range of wheat-
association, a combination that is now an integral part of the based products made around the world. With the publication
GRAINS AROUND THE WORLD | Grain Production and Consumption in Oceania: Australia and Pacific Countries 411
Wheat terminals
Australian premium white wheat area
Predominantly Australian premium
white and hard wheat area
Predominantly hard wheat area
Northern
Territory
Queensland
Western Australia
South Australia
New South
Wales
Victoria
Tasmania
Figure 1 Wheat-growing areas of Australia. Reproduced from Australian Crop Report, 2007, Australian Bureau of Agricultural and Resource
Economics, Canberra.
Figure 2 The $2 bank note, introduced with Australias decimal currency in 1966, featured William Farrer the wheat breeder responsible, in the period
18901910, for providing farmers with new varieties adapted to Australian conditions.
of these findings, advantages of the knowledge have flowed to crops (Table 2). These pulses have only emerged as important
the importing countries involved and to other exporters. crops since the 1980s. Expansion of production has depended
Another form of diversification came with the challenge significantly on the growth of export markets. Recent expan-
presented to wheat growers by the sandy soils of Western sions in wheat and pulse production have warranted the
Australia, due to the lack of soil nitrogen. An efficient solution enlargement of terminal storage facilities in Perth, Western
to this problem was the introduction of grain-legume crops in Australia (Figure 8).
agronomic rotation with wheat. The ability of the legumes to Another form of agricultural diversification accompanied
fix atmospheric nitrogen led to the provision of a natural the downturn in wheat prices during the 1980s. In addition to
source of nitrogen fertilizer, plus the welcome harvest of growing grain legumes, a valuable crop was supplied in the
other grains, particularly lupins and various pea and bean form of canola (rapeseed). Production of canola continued,
Table 3 Historical developments in the Australian wheat industry
Raven
Aroona
WW15
2* Condor Meering
WW80 2* Brochis S Batavia
Banks
PWTH Janz
Cook
Winglen 4* Cunningham
3Ag3
Timgalen WW15 3* Sunco
AKMS 3Ag14
Sunvale
Madden 5*
SUN9E-27 4*
VPM1
Gabo Sunstate
2* Machete
PavonS Hartog 4*
Kite Amber mutant 2* Sunbrook
2* Blade
TPP Sonora 64
Spica
Yaqi54 BT-Schomburgk Suneca
MEC3 Ciano
Barunga
Sonora 64 Molineux Wilgoyne
Sabre Frame Gallo
Dagger
Wren
Insignia Spear Stiletto
RAC177 3* Gaboto
3* Dollarbird
Condor Vereanopolis Kalyansona
Farro Lunga
WW33B WW33G Cocamba
4* Ouyen
TA3PNB3P TM56
AUS10894 Takari
Figure 4 Pedigrees of recent prominent varieties of wheat grown in Eastern and Western Australia. Pedigrees are read from left to right. For example,
Raven was crossed with the line WW15 to produce the variety Aroona. The line WW15 was crossed with WW80 and then crossed again (double
backcross, 2*) to produce the variety Condor. The variety Meering was selected from Condor. Condor was a parent entering into the pedigrees of Banks
and Cook. A sister line of Condor was a parent that also entered into the pedigrees of Banks and Cook.
GRAINS AROUND THE WORLD | Grain Production and Consumption in Oceania: Australia and Pacific Countries 415
Figure 6 Types of wheat quality according to the fair average quality system of the mid-twentieth century in Australia. Map provided by John Moss.
416 GRAINS AROUND THE WORLD | Grain Production and Consumption in Oceania: Australia and Pacific Countries
Table 5 Mean yields of crops in tonnes per hectare for the 2012 crop
year
Source: http://faostat.fao.org.
Oats of the pea varieties grown in the early twentieth century, Dun,
was still being grown in the 1980s. This began to change with
Most of the oat crop (Tables 2, 4, and 5) is used within
the work of John Gladstones on lupin at the University of
Australia. It may serve on-farm as a forage crop to be grazed
Western Australia and later in the WA Department of Agricul-
when plants are young, with the option of allowing the plants
ture beginning in the 1950s, in which he took what was
to mature and produce harvestable grain. Some varieties are
essentially a wild plant, Lupinus angustifolius, and by gradually
suited to milling and processing for food purposes, but much
incorporating genes for low seed alkaloids, soft seed, non-
of the oat crop is used as animal feed in its various forms
shattering pods, white flowers, and early flowering, created
forage, hay, and grain.
the modern crop-type narrow-leafed, or Australian Sweet,
lupin beginning with the release of Uniwhite in 1967. Field
Sorghum pea breeding resumed in Australia in the 1970s, and chickpea
and lentil breeding began in the 1980s. All of these pulses are
Sorghum is a feed grain. Much of the crop is exported for this
susceptible to devastating diseases, and improving disease
purpose. The main customers are Japan and Taiwan. Produc-
resistance has been an important objective of these breeding
tion is mainly confined to the warmer subtropical parts of
programs, as well as improving adaptation to Australian
Australia.
conditions.
Pulses in Australia are grown in rotation with cereals and
Cereal Rye improve production by providing fixed N and reducing cereal
Flour produced from rye grain is used in Australia for disease levels. Production expanded rapidly in the 1980s and
European-style breads, biscuits, crispbreads, multigrain break- 1990s as cereal production intensified, whereas legume-based
fast cereals, muesli bars, pet food, and special dietary foods that pastures, which had formerly played a similar rotational role,
provide higher lysine, pentosan, and b-glucan content than declined in response to the changing economic environment
other cereals. Production is minor in volume. of Australian farming.
There are geographic differences in pulse production
across the country. Narrow-leafed lupin is the predominant
Triticale pulse in Western Australia; white or albus lupin (L. albus) is
The wheatrye hybrid triticale has enjoyed reasonable popu- grown mainly in southern New South Wales, desi chickpea in
larity in Australia. Annual production varies, recently peaking northern New South Wales, and the other species in South
in 2010 at about half a million tonnes (Table 4), making Australia, Victoria, and southern New South Wales. Produc-
Australia the worlds fourth largest producer of triticale, tion of lupin, field pea, and lentil declined between 2000 and
depending on seasonal fluctuations. Triticales attraction has 2012 in response to changing grain prices relative to wheat
mainly been as an alternative to wheat for the stockfeed indus- and canola, which has displaced pulses to some extent.
try, which is the major domestic end user. However, efforts Increasing weed control costs as herbicide-resistant weeds
have been made to develop niche markets for human food. become more prevalent has contributed to declining lupin
production in Western Australia, and more frequent drought
during the past decade has also been important. On the other
Maize hand, production of chickpea has more than quadrupled in
Maize is a relatively minor crop in Australia, with annual the same period.
production ranging up to half a million tonnes (Table 4), A high proportion of Australian pulse production is
less than the production figures for the major cereal grains. exported. Field pea, lentil, and chickpea mostly go to the
However, grain yields are good. About 10% of the maize crop Indian subcontinent where they are used in human food,
is exported. About one-quarter of the crop is used domestically although a significant proportion of field pea is used domesti-
as human food, mainly for breakfast cereals, sweet corn/vege- cally in pig and poultry rations, especially in Eastern Australia.
table uses, corn-based foods, and industrial manufacture of Australia produces more than 70% of the worlds lupin and
corn starch. The remainder is used domestically as stockfeed. about 50% of this is exported. Most goes to Europe or northern
Industrial use of maize is minor compared to northern- Asia where it is used for animal feed. Australian lupin is also
hemisphere maize use, because Australian starch production used for aquaculture feed, and there has been considerable
is mainly based on wheat. Maize processing into starch relates effort over the past decade to develop human food uses for
mainly to special products such as high-amylose starch. lupin, including fiber and protein additives, which can be used
in a wide range of manufactured foods. However, these are
currently only minor uses.
Pulses
N
Crop yield still increasing by year 2070
Figure 11 Predicted changes in crop yields for the Australian state of Victoria, based on modeling of rising levels of CO2 together with other
expectations, such as heat episodes, rainfall, and known soil types. Illustration, courtesy of G. OLeary and coworkers.
canola is providing lines capable of producing omega-3 fatty Identification of Varieties of Food Grains; Wheat: Grading and
acids. The mining of novel genes is being facilitated by the Segregation; Wheat: Marketing; Non-food Products from Grains:
establishing the Australian Grains Genebank in Horsham, Vic- Fuel Alcohol Production; The Cereal Grains: An Overview of the
toria. It houses about 300 million seeds from around the Family of Cereal Grains Prominent in World Agriculture; Barley: An
world, for use in current and future plant breeding programs. Overview of a Versatile Cereal Grain with Many Food and Feed Uses;
Wheat: An Overview of the Grain That Provides Our Daily Bread; The
Oilseeds: Canola: Overview; Cottonseed: Overview; Wheat
Exercises for Revision
Processing: Gluten and Modified Gluten.
What features distinguish grain production, quality, and
consumption in your region from these factors in Oceania
(or choose one of the Oceanian countries).
What advantages are claimed for Australian export wheat?
Further Reading
What are the reasons for determining the variety identity in
the Australian wheat harvest? Anonymous (1994) Australian Grains. Melbourne, VIC: Morescope Publishing.
Name some of the improvements made to Australian OBrien L and Blakeney AB (eds.) (2000) An Introduction to the Australian Grains
wheats by breeding and selection. Industry. Melbourne, VIC: Royal Australian Chemical Institute.
OBrien L, Morell M, Wrigley C, and Appels R (2001) Genetic pool of Australian wheats.
What uses of barley and wheat are described?
In: Bonjean AP and Angus WJ (eds.) The World Wheat Book, pp. 603640. Paris:
Lavoisier Publishing.
OLeary G, Christy B, Weeks A, et al. (2011) Downscaling global climatic predictions
Exercises for Readers to Explore the Topic Further to the regional level: A case study of regional effects of climate change on
wheat crop production in Victoria, Australia. In: Shyam S, Yadav RJR, Hatfield JL,
What factors could account for the differences in grain Lotze-Campen H, and Hall AE (eds.) Crop Adaptation to Climate Change,
pp. 1226. Chichester: Wiley.
yields for the various grain species in Table 5, contrasting
Quail KJ (1996) Arabic Bread Production. St. Paul, MN: American Association of Cereal
Australia, New Zealand, and world averages? Chemists.
Find out about the development and expansion of wheat in Wrigley CW (1978) W.J. Farrer and F.B. Guthrie: The unique breeder-chemist
your region. How did their early times contrast with the combination that pioneered quality wheats for Australia. Records of the Australian
origins of the Australian wheat industry? Academy of Science 4: 725.
Wrigley CW (1981) Farrer, William James. In: Nairn NB and Serle AG (eds.) Australian
How would starch from wheat (as is usual in Australia) Dictionary of Biography: 18911939, pp. 471473. Melbourne: Melbourne
differ from the commercial production of starch from University Press.
maize in many northern-hemisphere countries in produc- Wrigley CW (1983) Guthrie, Frederick Bickell. In: Nairn NB and Serle AG (eds.)
tion methods and in quality? Australian Dictionary of Biography: 18911939, vol. 9, pp. 143144. Melbourne:
Find out about the relative advantages of the oilseeds men-
Melbourne University Press.
Wrigley CW (2005) Contributions by Australians to wheat quality research. In: OBrien L
tioned with respect to the amount and quality of the oils and Blakeney AB (eds.) An Introduction to the Australian Grains Industry,
produced. pp. 3341. Melbourne: Royal Australian Chemical Institute.
How does the value of the respective residues after oil Wrigley CW and Rathjen A (1981) Wheat breeding in Australia. In: Carr DJ and
removal affect the overall economics of oilseed utilization? Carr SGM (eds.) Plants and Man in Australia, pp. 96135. New York: Academic
Press.
Wrigley CW, Tomoskozi S, and Bekes F (2011) Hungarian-Australian
collaborations in flour milling and test milling over 120 years. Cereal
See also: Grain Harvest, Storage and Transport: Barley: Research Communications 39(2): 216225. http://dx.doi.org/10.1556/
Harvesting, Storage, and Transport; Grain Marketing and Grading: CRC.39.2011.2.5.
Grain Production and Consumption: South America
OR Larroque, CSIRO Agriculture Flagship, Canberra, Australia
JC Catullo, INTA, Buenos Aires, Argentina
2016 Elsevier Ltd. All rights reserved.
Argentina 1121 274 3384 227 236 971 18 749 1687 4189
Bolivia, Plurinational 53 52 331 6 64 178 105 1012 182 175
State of
Brazil 92 3275 13 375 175 2615 703 24 078 72 45 2071
Chile 17 11 127 97 24 3 23 260
Colombia 3 116 510 3 488 13 33 6
Ecuador 25 46 370 1 1 365 9 44 8
French Guiana 3
Guyana 3 132
Paraguay 61 812 70 24 2826 104 572
Peru 151 81 493 11 2 36 380 1 151
Suriname 55
Uruguay 64 5 105 23 180 32 952 5 373
Venezuela, Bolivarian 55 664 245 243 23 62 1
Republic of
World 48 401 30 040 170 865 9462 101 162 759 40 410 104 283 24 681 3842 218 197
Country/yield 2003 and 2012 Maize 03 Maize 12 Wheat 03 Wheat 12 Rice 03 Rice 12 Soybean 03 Soybean 12
Argentina 4180 340 24 059 492 1520 4429 51 019 3078 13 656
Bolivia, Plurinational 49 61 915 6 37 454 401 2206 238 235
State of
Brazil 280 3 60 774 387 12 035 1834 69 757 95 110 5414
Chile 98 20 1430 465 125 6 108 1438
Colombia 5 133 1643 4 2001 59 72 11
Ecuador 18 13 1004 1 1 1583 13 72 7
French Guiana 10
Guyana 5 582
Paraguay 53 3178 368 148 8040 165 1421
Peru 210 91 1578 12 42 2825 3 220
Suriname 227
Uruguay 210 3 455 47 1405 122 2119 6 1167
Venezuela, Bolivarian 47 2421 1334 481 38 54
Republic of
World (total) 129 693 23 173 870 060 21 114 80 713 984 58 578 260 075 36 418 13 652 453 350
100
90
80
70
60
% 50 Wheat
Rice (milled)
40 Maize
30
20
10
0
Argentina Bolivia Brazil Chile Colombia Ecuador Guyana Paraguay Peru Suriname Uruguay Venezuela South
America +
(Total)
Figure 1 Share of main cereals in the South American diet. Source: FAOSTAT (2009).
According to the same source, the country has 33 000 bakeries Wheat is the top commodity available for consumption
where artisan continental bread is baked daily. Argentina is (2009) with 406 kcal per capita per day. Maize is the second
currently the fifth largest world exporter of wheat flour. (302 kcal per capita per day) and rice third (280 kcal per capita
Argentinas undernourishment figures are not significant, per day). The prevalence of undernutrition is very high at 24%
comprising <5% of the population in 201012. (201012).
Dry bean production in 2012 was 60 000 tonnes.
Bolivia is also the largest producer and exporter of quinoa, a
Bolivia, Plurinational State of
high-protein grain native to South America that was the staple
food of the Incas. The UN declared 2013 the Year of Quinoa.
Landlocked Bolivia is the least developed country of the region,
Unfortunately, higher international prices for the crop are
with a large proportion of its population living in rural areas
leaving the poorest segments of the population unable to
(32.54% in 2012). Arable land has increased in recent years
purchase quinoa anymore.
from 2.59 million hectares in 1996 to 3.84 million hectares in
2011. This increase was mainly made at the expense of forest
cover. Agriculture employs 40%of the workforce (down from
45% in 1997). While the GDP has doubled between 2005 and
2010, the share of agriculture value added in total GDP has Brazil
decreased, continuing a trend that saw its figures going down
from nearly 20% in 1995 to just above 10% in 2010. The South American giant is the largest economy of the region
In recent decades, this Andean country has consolidated a and the major grain producer. Brazilian agriculture has chan-
strong growth in oilseed crop production, particularly soybean ged dramatically in the last 30 years. From being a major food
with planted areas exceeding 1 000 000 hectares and total pro- importer (Figure 3), the country also became a top grain
duction of 2 298 857 tonnes in 201112. Soybean is the largest exporter (Figure 1). Brazil is the first of the big exporters to
agricultural export and the third largest Bolivian export. be in a tropical area and not a temperate area (like it is the case
Santa Cruz de la Sierra is the main producing area with an with the United States, Canada, Australia, Argentina, and EU).
average production yield of 16002400 k ha1. There are two The arable land has increased from 57 860 000 ha in 1996
crops per year: the most important in summer (November/ to over 70 000 000 ha in 2011. It corresponds to 8.3% of the
December) and also in winter (June/July). total country area. Furthermore, it has over 400 000 000 ha of
The second most important grain crop is maize potentially arable land, the largest area for any country in the
(1 006 000 tonnes in 2012) followed by sorghum world according to FAO. Most of this land is in the Brazilian
(478 000 tonnes) and paddy rice (440 000 tonnes). savannah (the Cerrado).
GRAINS AROUND THE WORLD | Grain Production and Consumption: South America 423
35 000
32 986
30 000
26 832
25 000
20 000
15 806
Thousands of tonnes
15 000 14 355
10 820
10 000 9487
8411
5010
5000 4417
0
Soybean Soybean cake Maize Soybean cake Soybean Maize Wheat Soybean Soybean oil
Brazil Argentina Argentina Brazil Argentina Brazil Argentina Paraguay Argentina
Figure 2 Major grain commodities exports South America. Source: FAOSTAT (2011).
The most recent data show that primary agriculture (151 kcal per capita per day). Brazils traditional, basic meal
accounted for 5% of Brazils GDP and agricultural population consists of rice and beans (feijoada). Brazil is the second
nears 20 000 000 people (12% of total population). world producer of dry beans, behind India. Production figures
Brazil is second largest producer of soybean oilseed for 2012 were nearly 3 000 000 tonnes. Consumption of
(88 000 000 tonnes) and the third largest producer of maize 16.30 kg per capita per year was recorded for 2009.
(70 000 000 tonnes) in the world, behind the United States Although the importance of breakfast cereals in the region
and China. In 2011, maize exports accounted for is limited, Brazil and Venezuela show some increase in con-
9 486 914 tonnes (second in the region and third in the sumption. From 2011 to 2016, it is predicted that Brazils
world), and soybean exports were just below market will increase by 27%. Increase in consumption is asso-
33 000 000 tonnes (number one exporter in the world). ciated with upper and middle-upper income segments of the
The country is the sixth largest importer of wheat population. On the other end of the spectrum, breakfast cereals
(5 740 753 tonnes in 2011) although it also exports are still very expensive for lower-income earners when com-
(2 350 000 tonnes in 2011). Brazil is the third largest producer pared with traditional breakfast meals.
of pasta in the world. The prevalence of undernutrition has dropped from 12% in
Rice is a staple food in the country (352 kcal per capita per 19992001 to 7% in 201012. The Zero Hunger Program
day), so its consumption is correlated with the population launch in 2001 by the then-presidential candidate Lula da
growth. Most of Brazilian rice is cropped in uplands, particu- Silva has successfully helped in the improvement of these
larly in the northern and midwestern regions. Production of figures. Family farming, the main pillar of food security in
rice (milled) was 8 200 000 tonnes in 2011. Brazil is the ninth the country, is responsible for the domestic production of
world producer. 70% of beans, 46% of maize, 34% of rice, and 21% of wheat.
Other grain commodities available for consumption are Brazil legislation now permits the use of GMOs. Since the
wheat (384 kcal per capita per day), soybean oil (336 kcal per full lifting of the ban in 2005, Brazil has turned into the second
capita per day), maize (209 kcal per capita per day), and beans largest producer of GM crops in the world, behind the United
424 GRAINS AROUND THE WORLD | Grain Production and Consumption: South America
7000
6000 5740
5000
4000
Thousands of tonnes
3000 2820
1908
2000
1684
1535
1299 1215
1014 1004
1000
0
Wheat Maize Maize Wheat Wheat Maize Wheat Soybean cake Soybean cake
Brazil Colombia Peru Peru Colombia Venezuela Venezuela Colombia Peru
Figure 3 Major grain commodity imports into South American countries. Source: FAOSTAT (2011).
States. Over 80% of the soybean crop is GM, and they are Wheat is the top overall commodity available for consump-
planning to bring to the market a GM bean with resistance to tion (2009) with 850 kcal per capita per day. Maize is the
the golden mosaic virus. second grain and third overall (211 kcal per capita per day)
and rice the third and eighth, respectively (90 kcal per capita
per day). Cereals account for 42.3% of the total food supply
(2009). Chileans consume an average of 86 k of bread per
Chile
capita per year (200110), making them the largest consumers
of bread in the region and second in the world, behind
Located on the western coast of the continent, facing the
Germany. The most consumed types are the soft bread marra-
Pacific, the Republic of Chile occupies a narrow strip of land
quetas and the flat round hallullas. Chile fortifies basic food
between the sea and the Andes mountains.
(in this case, wheat flour) with micronutrients (iron).
The share of agriculture value added in total GDP is very
Chiles undernourishment figures are not significant, com-
little (<5%) and the rural population is just above the 10%
prising <5% of the population in 201012. In 199092, 8.1%
mark. The arable land has diminished in the last years, from
of the population was undernourished.
2 000 000 hectares in 1996 to 1 320 000 hectares in 2011.
The main grain crop is wheat (1 575 822 tonnes in 2011).
A quarter of this production comes from 40 000 small farmers
(subsistence group, no or little alternative crop to grow). Colombia
A little over 550 farmers produce 50% of the wheat and around
5000 farmers are considered medium size producers. Maize Situated in the northwestern part of South America, Colombia
(1 493 292 tonnes in 2012) is the second crop in importance. is the fourth largest country of the region, after Brazil,
Chile is the top producer of oats (450 798 tonnes in 2012) in Argentina, and Peru. The arable land fluctuates around 2% of
the region and within the top five in the world. the total area of the country. The eastern plains region called
To fulfill internal demand, wheat (655 527 tonnes), maize the Altillanura has potential for future agricultural expansion.
(668 151 tonnes), and sorghum (636 112) are three of the The share of agriculture value added in total GDP has
most imported commodities in the country (2011). abruptly dropped from the turn of the millennium. In 2011,
GRAINS AROUND THE WORLD | Grain Production and Consumption: South America 425
Colombias agriculture contributed nearly 7% of total GDP. Dry beans production averaged 13 000 tonnes for 201012.
Nevertheless, agriculture continues to be an important source The main grain commodities available for consumption in
of employment (24% of the economically active population). 2009 were rice (milled) at 431 kcal per capita per day and
The most important grain crop is rice (paddy wheat (295 kcal per capita per day). Soybean oil availability
2 000 000 tonnes; milled 1 320 000 tonnes in 2011), which for consumption was 120 kcal per capita per year. Cereals
ranks sixth among the top ten commodities produced in share was 34.4% in total food supply (2009).
Colombia. Most of the crop is grown under irrigation. The The prevalence of undernutrition was one of the highest in
harvested area averages just over 400 000 ha in recent years. the region with 18% for the period 201012, slightly down
Maize contributed with a production of 1 700 000 in 2011. from an average of 21% (19992009).
The harvested area remains steady at 500 000 ha in recent
years. Around 10% of the planted area uses biotech seed and
is expected to increase in the future. French Guyana
Colombia is the second largest importer of the region,
behind Brazil. Maize imports have been well above Agriculture is concentrated in coastal areas (10 000 hectares of
3 000 000 tonnes in recent years (3 900 000 tonnes in 2013) arable land). Rural population accounts for 23% of the total
positioning the country as the largest maize importer in population.
South America. Ninety-five percent of maize imports (mainly The main and almost only grain crop grown in the country
yellow) are used in the feed industry. is rice, with a production of 9587 tonnes in 2011 (130th in the
Soybean is being used in rotations with maize, but produc- world). Production has dramatically decreased in the last
tion is not significant (86 634 tonnes in 2012) decade (31 000 tonnes in 2001).
Wheat imports average 1 500 000 tonnes per year. Bread
production demands 75% of the wheat, while the pasta indus-
try requires 15% and the cookies and pastry industry the Guyana
remaining 10%.
Rice (358 kcal per capita per day), maize (333 kcal per Annual crops are grown on the narrow coastal strip. Arable
capita per day), and wheat (222 kcal per capita per day) are land represents 2% of the total country area. Rice is almost the
the main commodities available for consumption. Colombia is only grain produced, with nearly 600 000 tonnes harvested in
the third largest producer of beans in South America 2011. There are two crops per year.
(133 000 tonnes in 2012) and ranks fourth in consumption Agricultural value added in total GDP decreased from
with 3.5 kg per capita per annum. around 40% in 1995 to just over 20% in 2010.
Prevalence of undernutrition (201012) remains steady The main grain commodity available for consumption in
at 13%. 2009 was rice (796 kcal per capita per day) followed by wheat
(483 kcal per capita per day). Cereals account for 48.3% of the
total food supply (2009), the highest in the region.
Ecuador
this crop in detriment of maize (third place, 3 345 877 tonnes), The share of agriculture value added in total GDP was
wheat (fifth place, 1 461 020 tonnes), rice, and sorghum. Para- 10.8% in 2012.
guay is the fourth most important exporter of soybeans in the The main crop is rice, which accounts for almost all the
world. harvested grain (paddy rice 220 000 tonnes) according to FAO
Maize is the top commodity available for consumption statistics for 2012.
(2009) with 509 kcal per capita per day. Wheat is the second Rice availability for consumption (2009 data) was 629 kcal
(216 kcal per capita per day) and beans third (54 kcal per per capita per day of the milled equivalent. Wheat accounted
capita per day). Per capita consumption of beans ranks for 413 kcal per capita per day and soybean oil for 263 kcal per
second in the region (5.8 kg). capita per day. Cereals share in total food supply was 41.6%.
The prevalence of undernutrition is very high at 26%
(201012), a sharp increase from figures of 13% to 17%
recorded in the previous 10 years. Uruguay
14
12.3
12
10
8.4
kg per capita per year
8.2
7.9
8
6.2
6
4.8
3.9
4
3
0
Venezuela Chile Peru Argentina Brazil Bolivia Ecuador Colombia
Figure 4 Pasta consumption in South American countries (2011). Source: www.internationalpasta.org.
428 GRAINS AROUND THE WORLD | Grain Production and Consumption: South America
Climate changes in the region may negatively impact from Brazil says yes to genetically modified food, Mexico says no http://www.
an economic and social perspective if not properly addressed. businessweek.com/articles/2013-10-30/brazil-says-yes-to-genetically-modified-
foods-dot-mexico-says-no (accessed 12/12/13).
Nutrition Profile Bolivia http://www.childinfo.org/files/nutrition/DI%20Profile%20-%
20Bolivia.pdf (accessed 12/12/13).
Exercises for Revision Nutrition Profile Paraguay http://www.childinfo.org/files/nutrition/DI%20Profile%20-%
20Paraguay.pdf (accessed 15/12/13).
What pseudocereals are native to South America? Why is Argentina so efficient? http://cornandsoybeandigest.com/tillage/why-argentina-
so-efficient (accessed 15/11/13).
What is the social cost of the agricultural economic boom in Dros JA (2004) Managing the soy boom: Two scenarios of soy production expansion in
the region? South America. Amsterdam: AIDEnvironment.
What strategy is Brazil using to mitigate hunger in its The miracle of the cerrado http://www.economist.com/node/16886442
population? (accessed 17/12/13).
Colombia is one of the major importers of maize in the Ecuador 2013 maize production estimated at record levels http://www.thecropsite.com/
news/15111/ecuadors-2013-maize-production-estimated-at-record-levels
region. What is it mainly used for? (accessed 4/1/14).
How is Ecuador planning to increase its maize production? Current status of adoption of no-till farming in the world and some of its main benefits
Why are breakfast cereals not popular in the region? http://www.fao.org/ag/ca/CA-publications/China_IJABE.pdf (accessed 14/11/13).
Where is El Cerrado located and what is its importance? Brazil tries to fill the potholes in its path to growth http://www.ft.com/cms/s/0/
7dd98ed6-059f-11e3-ad01-00144feab7de.html#axzz2xV094L00
Name the countries where wheat is not the main staple (accessed 31/3/14).
cereal in the region. Pasta Consumption http://www.internationalpasta.org (accessed 1/11/13).
Find a recipe for Feijoada and Pabellon on the Net. What Family Agriculture in the Mercosur http://www.reafmercosul.org/reaf
grain components do they share? (accessed 13/12/13).
What were the main economic and political reasons behind Sunny side up for Latin America breakfast cereals http://www.just-food.com/analysis/
sunny-side-up-for-latin-america-breakfast-cereals_id97594.aspx (accessed 17/12/
the 2008 farmers strike in Argentina? 13).
GMOs a global debate: Brazil, second largest GMO producer in world http://www.
theepochtimes.com/n3/162906-gmos-a-global-debate-brazil-second-largest-gmo-
Exercises for Readers to Explore the Topic Further producer-in-world/?photo2 (accessed 18/12/13).
Value Added (%GDP) in Colombia http://www.tradingeconomics.com/colombia/
agriculture-value-added-percent-of-gdp-wb-data.html (accessed 12/12/13).
Sustainability of the soybean production models Vergara W, Rios A, Trapido P, and Malarin H (2014) Agriculture and future climate in
Brazil agriculture before and after lifting the ban on GMOs Latin America and the Caribbean: Systemic impacts and potential responses. Inter-
No-tillage and postharvest technology in the region American Development Bank Discussion Paper No. IDB-DP-329.
Contents
Research Organizations of the World: CGIAR
Research Organizations of the World: Europe and North America
Research Organizations of the World: Asia-Pacific, CentralSouth America, and AfricaMiddle East
Research Organizations of the World: Global Trends and the Commercial Sector
Scientific Societies Associated with Grain Science
This article includes some content from the previous edition article by F.J.B Reifschneider & S. Hussain, volume 3, pp 2636, 2004, Elsevier Ltd.
Topic Highlights used for irrigation, and agriculture frequently accounts for 70%
of freshwater withdrawals and agricultural activities are one of
The CGIAR international agricultural research centers pro- the leading causes of climate change, contributing 20% of all
vide a significant contribution to grains research, particu- greenhouse gases.
larly for the developing world. Globally, the agricultural sector is set to face pressing chal-
The CGIAR centers have traditional strengths in germplasm lenges in the future:
development, agronomy, quality assessment, biotechnol-
ogy, pathology, natural resource management, and World population is expected to reach nine billion by 2050.
Over the same time frame, demand for food is set to more
economics.
than double, with most of the increased demand coming
In 201112, much of the research undertaken within the
from developing countries.
CGIAR system was organized into major research programs.
Five CGIAR Research Programs have a primary emphasis on Erosion, salinization, compaction, and other forms of soil
degradation affect 30% of the worlds irrigated lands, 40%
improvement of yields and profitability of grain crops,
of rainfed agricultural lands, and 70% of rangelands.
dryland cereals, grain legumes, maize, rice, and wheat,
while other programs focus on farming systems, crop nutri- Most of the worlds freshwater is used for irrigation and
water use is expected to increase by 50% by the 2030s.
tional characteristics, production sustainability, soil and
water management, and climate change. Arable land per person in developing countries was 0.32 ha
in 196163, but is expected to drop to 0.16 ha by 2030.
Cereal productivity growth increases (i.e., rises in cereal
Learning Objective yields per ha) have slowed.
such as droughts and floods) and human-induced conflicts However, the benefits of the green revolution were spread
and civil wars can all trigger famines. unevenly and largely bypassed Africa. On average, crop yields
The 1960s marked a period when deep-rooted pessimism in Africa are one-third of those achieved by Asian farmers. Less
about food availability surfaced anew. However, scientists than 4% of Africas arable land is irrigated. There is enormous
working with farmers successfully averted the worst Malthu- potential for growth-led productivity in sub-Saharan Africa
sian specter by ensuring abundant agricultural yield. The story where agriculture provides 70% of employment, 40% of
of wheat improvement in England is compelling. It took nearly exports, and one-third of gross domestic product. However,
1000 years for wheat yields to increase from 0.5 to 2 tonnes productivity benefits for Africa are now starting to emerge,
per hectare (mt ha 1), but only 40 years to climb from 2 to particularly for maize, roots, and tubers.
6 tonnes ha 1, illustrating the ability of modern grain science
to increase agricultural productivity.
Grains Research Centers in the CGIAR
Mobilizing Grain Science for Development
CGIAR achieves its mission through the work of 15
international agricultural research centers (IARCs, Table 1)
The Consultative Group on International Agricultural Research
that work with national agricultural research systems, the pri-
(CGIAR) was established in 1971. A group led by The Rocke-
vate sector, and civil society. The IARCs are autonomous insti-
feller Foundation, Ford Foundation, Food and Agriculture
tutions, linked by a common commitment to promoting
Organization, The World Bank, and others created the
agricultural development in developing countries; 13 IARCs
CGIAR. The founding objective was based on a profound belief
are headquartered in developing countries.
in the ability of science, especially grain science, to increase
Improving the sustainable productivity of food grains
food supply in low-income countries facing severe food scar-
(including cereals, legumes, and pulses) is a major objective
city. At that time, the central idea of mobilizing grain science
of CGIAR research. Nine of the IARCs have specific crop
internationally to meet food and development needs, within a
improvement mandates. More specifically, the CGIAR centers
public goods framework, was novel.
have traditional strengths in germplasm development, provid-
The CGIAR system is supported by the World Bank and a
ing much of the germplasm for development of the modern
range of national governments, usually through their overseas
varieties that are now utilized throughout most of the devel-
development assistance programs. In 2013, the CGIARs
oping world (as well as in some developed countries with
research investment exceeded $US 1 billion for the first time
similar production environments such as Australia). To sup-
and represented the single largest effort in mobilizing science
port this work, centers have strengths in agronomy, quality
for agricultural development. The CGIAR is best known for the
assessment, biotechnology, pathology, natural resource man-
green revolution, a term used to refer to the set of agricultural
agement, and economics.
innovations (new high-yielding varieties of wheat, rice, and
For the purposes of this paper, only eight IARCs have crop
maize, increased investments in irrigation, easy availability of
improvement mandates that correspond to the definition of
farm inputs such as fertilizers and pesticides, improved grain
grains cited earlier, the exception being root and tuber crops,
distribution, and food policies in developing countries) that
including potato. Even though Bioversity (formerly known as
helped boost grain yields and incomes of farmers in poor,
International Plant Genetic Resources Institute) does not have
agrarian economies. The green revolution transformed agricul-
a plant breeding program, its mandate to serve as a world
ture in developing countries especially in India, Mexico,
center for conserving plant genetic resources is germane to
Pakistan, and the Philippines.
the focus of this article.
Cereal crop yields and food production more than doubled
between 1960 and 1985. The new varieties of rice, wheat, and
maize increased daily calorie supply in developing countries by
25%, from under 2000 calories per person in the early 1960s to The International Agricultural Research Centers
2500 calories per person by the 1990s. Without the productiv-
ity growth in basic food crops, the number of poor and hungry Several of the centers have a grain science focus, including the
would have been far greater. Centro Internacional de Mejoramiento de Maz y Trigo (The
Critics of the green revolution have argued that the new International Maize and Wheat Improvement Center
crop technologies favored farmers in well-endowed areas, (CIMMYT) www.cimmyt.org). CIMMYT is based in Mexico
worsened pollution through excessive use of fertilizers and but has programs in Latin America, Africa, Central and West
pesticides, threatened biodiversity, promoted monocultures, Asia, and North Africa. Three centers carry out studies on rice
and required higher input costs from impoverished farmers. improvement. The International Rice Research Institute (IRRI,
While a comprehensive discussion of these issues is beyond the www.irri.org) is the main one, and its work aims to develop
scope of this article, subsequent studies and analyses have new rice varieties and crop management techniques to help
increased understanding about the effects of the new technol- farmers improve the yield and quality of rice in a sustainable
ogies. A range of subsequent policy interventions (e.g., way. The semidwarf rice varieties and management packages
increased credit, improved marketing and seed distribution developed by IRRI were central to the successes of the green
facilities, land reforms, and strengthened emphasis on pesti- revolution in the 1960s and 1970s. Two other centers under-
cide safety, and others) have helped poor farmers to benefit take rice breeding and management in Latin America and
from green revolution technologies. Caribbean (International Center for Tropical Agriculture
SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: CGIAR 431
Year Host
Center founded country Mandate crops
International Center for Tropical Agriculture (CIAT) 1967 Colombia Cassava, beans, rice, tropical forages
International Maize and Wheat Improvement Center 1966 Mexico Maize, wheat (bread and durum wheat, triticale)
(CIMMYT)
International Center for Agricultural Research in the Dry 1975 Syriaa Barley, wheat (durum species), lentil, chickpea, faba bean,
Areas (ICARDA) forage legumes
International Crops Research Institute for the Semi-Arid 1972 India Sorghum, pearl millet, groundnut, chickpea, pigeon pea
Tropics (ICRISAT)
International Institute of Tropical Agriculture (IITA) 1967 Nigeria Cassava, yams, maize, soybean, cowpea
International Plant Genetic Resources Institute (now called 1974 Italy Genetic resources, cocoa, coconut/Musa species
Bioversity)
International Rice Research Institute (IRRI) 1960 Philippines Rice
International Potato Center (CIP) 1970 Peru Potato, sweet potato
West Africa Rice Development Association (now called 1975 Cote Rice
AfricaRice) dIvoireb
a
Currently Lebanon.
b
Currently Benin.
(CIAT) ciat.cgiar.org in Cali, Colombia) and West Africa Evolving Research Agenda
(AfricaRice, www.africarice.org, in Cotonou, Benin).
Other CGIAR centers with a strong grains focus include the The CGIARs role and mission has evolved over time and this is
International Crops Research Institute for the Semi-Arid Tro- reflected in its structure. When the CGIAR was founded, four
pics (ICRISAT, www.icrisat.org) in Patancheru, India (focussing IARCs (CIAT, CIMMYT, IITA, and IRRI) were brought under its
on sorghum, pearl millet, chickpea, pigeon pea, and peanut), umbrella. Thus, at that time, it was largely an alliance of grains
and the International Center for Agricultural Research in the research organizations, and in the 1960s, the research agenda
Dry Areas (ICARDA, www.icarda.cgiar.org) long established in was clearly around maize and rice productivity.
Syria, but now headquartered in Beirut, Lebanon (focussing on During the 1970s, new IARCs were established to pioneer
barley, wheat, lentil, faba bean, chickpea, and forage legumes). improvements in key food crops (legumes, roots, tubers, and
The CGIAR centers have traditional strengths in germ- other cereals), livestock health, and ecological regions (dry, semi-
plasm development, providing much of the germplasm for arid, and tropical regions). In the 1980s, the mandate (and struc-
development of the modern varieties that are now utilized ture) was expanded to encompass food policy and strengthening
throughout most of the developing world (as well as in some of national agricultural research systems and in the 1990s further
developed countries with similar production environments expanded to conduct research on pressing issues involving agro-
such as Australia). To support this work, centers have forestry, forestry, water management, and aquatic resources.
strengths in agronomy, quality assessment, biotechnology, During the 1990s and early-mid-2000s, the research pro-
pathology, natural resource management, and economics. At grams undertaken in the CGIAR were organized around five
CIMMYT, wheat breeding and maize breeding (agronomy focus areas:
and quality evaluation programs) are backed up by programs
in economics, natural resource management, and biotechnol- Increasing productivity (of crops, livestock, fisheries, for-
estry, and the natural resource base)
ogy. ICRISAT has a geographic emphasis on South Asia and
sub-Saharan Africa, and its research addresses water scarcity Strengthening national systems (joint research, policy sup-
port, training, and knowledge sharing)
through natural resource management (rainwater conserva-
tion and utilization) and plant breeding to improve water-use Protecting the environment (by developing new technologies
that make more prudent use of land, water, and nutrients and
efficiency and drought tolerance. ICARDA has a geographic
help reduce agricultures adverse impacts on ecosystems)
emphasis on central West Asia and North Africa, but its germ-
plasm is utilized worldwide. ICARDA focuses on agriculture Saving biodiversity (collecting, characterizing, and conserv-
ing genetic resources)
in tropical dry areas, while ICRISAT emphasizes temperate
and subtropical dry areas. It also has an emphasis on water- Improving policies (with a major impact on agriculture,
food, health, the spread of new technologies, and the man-
soil management in integrated farming systems for low-
agement and conservation of natural resources).
rainfall areas. The International Institute of Tropical Agricul-
ture (IITA) with headquarters in Nigeria focuses on germ- In summary, over 50 years, the CGIARs research portfolio
plasm (including cowpea, maize, and soybean), pest evolved from the original focus on increasing productivity in
management, and crop-based farming systems for small- individual critical food crops. The current approach recognizes
holder farmers of sub-Saharan Africa. Some work on proces- that saving biodiversity, protecting the environment, improving
sing quality is carried out by the centers, but little work on food policies, and strengthening scientific capacities of develop-
grain storage and other postharvest technologies. ing countries through training and knowledge sharing are all key
432 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: CGIAR
components in the drive to enhance sustainable agricultural formation of the CGIAR and its forerunners over 50 years
productivity. The CGIARs focus on the fundamentals remains ago. Another continuing challenge is the level of skills and
as strong as ever: Agricultural growth and increased farm pro- resourcing within the National Agricultural Research and
ductivity in developing countries creates wealth, reduces poverty Extension Services (NARES) as this affects the ability of
and hunger, and protects the environment. national organizations in poorest countries to be able to effec-
Recognizing the complexity of the challenges faced in devel- tively partner. Centers with an ecoregional focus, such as
oping world agriculture and the importance of establishing ICARDA, have traditionally had a more extensive program of
sufficient capacity, the CGIAR was reorganized effective 2010. collaboration with the NARES in their region than those with a
The CGIAR Consortium (www.cgiar.org/cgiar-consortium) global crop mandate. The NARES themselves have recognized
integrates the research of the 15 centers to increase the effec- the importance of leveraging off CGIAR research and trying to
tiveness and relevance of the work of the CGIAR; enhance the do so in a more structured manner than in the past. For
impact of the research by setting common objectives and plan- example, the Asia-Pacific Association of Agricultural Research
ning concerted action; and to take advantage of opportunities Institutions (APAARI) has recently developed a strategy for
for the CGIAR to become more relevant, more effective, and strengthening partnerships in South Asia between the NARES
more efficient. and the CGIAR (www.apaari.org/publications/nars-cgiar-pro-
In 201112, much of the research undertaken within the ceedings.html).
CGIAR system was organized into major research programs Various approaches have been used to develop or
(www.cgiar.org/our-research/cgiar-research-programs), strengthen relationships between centers and the NARES in
which typically involve several CGIAR centers as well as uni- developing countries. One of the most effective and enduring
versities, NGOs, and government agencies from both devel- has been development of mechanisms by each of the grains
oping and developed countries. The CGIAR Consortium centers to ensure that developing country breeders are trained
coordinates CGIAR Research Programs across research at their centers and that grains breeding programs have access
centers, which are members of the CGIAR Consortium and to materials. This has typically involved provision of germ-
partner. plasm and advanced lines (but not usually finished varieties),
Five CGIAR Research Programs have a primary emphasis on although the level of crossing from the materials made avail-
improvement of yields and profitability of grain crops: able by the centers depends on the resources available in the
particular NARES as well as the extent of further work required
Dryland cereals (such as barley, sorghum, finger, and pearl
to ensure that the material is sufficiently locally adapted.
millets) in South Asia and Africa
Translation (or extension) of agronomic research results
Grain legumes (addressing major biotic and abiotic stres-
from CGIAR centers to the NARES is necessarily more
ses, nitrogen fixation, and opportunities to use grain
complex by definition, it involves practice change rather
legumes to intensify cropping systems)
than having the technology embedded in the seed. Apart
Maize (focussing on the productivity of maize-based crop-
from the expected approaches of short- and long-term training
ping systems)
and development of publications and other written materials,
Rice (focussing on genetic diversity, development of
a challenge has been to develop longer-term partnerships that
new varieties, rice-based farming systems, quality, and
achieve impact while not distracting from the core function of
marketing)
knowledge generation at the centers.
Wheat (wheat-based cropping systems, pest, diseases, and
To facilitate such partnerships, a number of individual
nutrient management, yield, and seed systems)
national and multilateral donors have funded major partner-
In addition, two programs (Integrated Systems for the Humid ship projects, which through the provision of restricted
Tropics and Dryland Systems) aim to improve the productivity, funding (now called bilateral funding) can fund the exten-
profitability, sustainability, and resilience of entire farming sion of CGIAR crops research results through formal, funded
systems; one program (Agriculture for Nutrition and Health) partnership projects. For example, the African Development
includes a thrust on grains nutritional composition, while a Bank is currently supporting a major collaboration between
further program underpins the CGIAR crop (particularly grain) AfricaRice, ICARDA, and IITA to enhance the productivity and
germplasm collections. Further two programs aim to improve income of four priority value chains (cassava, maize, rice, and
production sustainability and environmental integrity and wheat) on a sustainable basis. The importance of partnerships
adapt to and mitigate climate change namely, the CGIAR with the NARES has also been recognized in the design of
Research Programs on Climate Change, Agriculture and Food most of the new CGIAR Research Programs and most projects
Security and Water, Land and Ecosystems. that involve not only several CGIAR centers but also the
NARES and universities in both developed and developing
countries.
Finally, capacity building of the NARES organizations has
Collaboration with National Agricultural Research itself been a focus of the CGIAR over many years. This led to
and Extension Systems the formation of a specialized CGIAR center, the International
Service for National Agricultural Research (ISNAR, in the
As comparatively small, research-oriented centers focussed on Netherlands), although it merged into the IFPRI in 2004.
international public goods, the tension between addressing Some controversy remains, however, as to whether the
this mission and achieving practical impacts at the national CGIAR is weakening the NARES capacity rather than building
level and in the shorter term has been present since the it as in some regions the most able, internationally trained
SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: CGIAR 433
scientists are hired to work for the CGIAR institutes. In addi- improved germplasm, and a unique set of wild crop relatives.
tion, where the NARES are weak, there has in the past been a These include wheat, barley, oats, and other cereals; food
tendency for the CGIAR to assume the NARES role of develop- legumes such as faba bean, chickpea, lentil, and field pea;
ing and testing technologies at plot and field scale. This returns and forage crops, rangeland plants, and wild relatives of each
us to the challenge of demonstrating impact versus seeing the of these species.
CGIAR as an intermediary.
Rice
Selected Impacts of CGIAR Grains Research IRRI began rice improvement in the 1960s. Expanding on the
successes in wheat improvement, IRRI scientists successfully
There have been several published analyses of the overall introduced semidwarfism into two indica rice varieties (IR5
impacts of CGIAR research, and most of them have shown an and IR8). The latter variety would lay the foundations of the
overwhelmingly positive return on investment. In the initial green revolution in Asia. IRRI has made substantial contribu-
studies, the point was made that in some countries, the bene- tions toward improvement of rice varieties. Of the estimated
fits from modern varieties developed by the CGIAR centers had 2040 rice varieties released over 40 years by national rice
favored poor urban food consumers and large farmers rather research systems of South and Southeast Asia, 219 were direct
than poor smallholder farmers, although it was pointed out IRRI lines. Of the 2021 varieties, 31% had originated from one
that small farmers do tend to adopt modern varieties but later or more parents developed at IRRI. This number rises to 46%
than large farmers. Evenson and Gollin published an extensive when the IRRI-developed varieties are included. In the 1970s,
study of CGIAR crop research impacts across a wide range of the release of IRRI-related varieties peaked at 60%, subse-
crops and continents. Renkow and Byerlee assessed the quently stabilizing at 40% of all varieties released. Since that
impacts of more recent GIAR research that published since time, the increase in rice production has been enormous. More
2000 and argued that impacts of CGIAR crops genetic recently, Scuba rice, which can survive under water for 2 weeks,
improvement research have continued to be strongly positive is protecting the harvests, incomes, and food security of poor
even some decades after the green revolution. Lastly, Raitzer farmers and consumers across monsoon Asia.
and Kelley attempted to undertake a benefitcost meta-analysis AfricaRice conducts rice improvement research in West and
of investment in CGIAR research and concluded benefitcost Central Africa, focussing on three distinct rice ecologies:
ratios of between 1.9 and 17.3, depending on the stringency of upland, rainfed lowland, and irrigated land. Using interspecific
demonstration of causality of benefits from particular research hybridization techniques, Africa Rice successfully combined
activities. However, they concluded that the true benefits the ruggedness of the local African species of rice (Oryza gla-
arising from the CGIAR is probably in excess of even the berrima) with the phenomenally high productivity traits of
upper bounds of these results as only a small subset of system Asian rice (O. sativa) (common rice) that was the mainstay of
impacts have been quantified. the green revolution. The new rice varieties, NERICA or New
Rice for Africa, resist droughts and pests and are able to thrive
in poor soils. CIAT has a mandate to promote sustainable
Wheat
increases in rice production and productivity in Latin America
CIMMYT has a global mandate for wheat improvement (bread and the Caribbean region.
and durum species). In the late 1960s, about one-third of all
wheat varieties in developing countries were CIMMYT crosses.
Maize
By the 1990s, these numbers rose to about half from CIMMYT
crosses and another quarter from varieties that had a CIMMYT CIMMYT has a global mandate for maize improvement, and it
parent. Worldwide, around 90% of all spring bread wheat targets lowland tropical, subtropical, midaltitude, and tropical
releases had at least one CIMMYT ancestor; the percentage for highland environments throughout the developing world.
durum wheat was even higher with nearly all spring durum Mexico, where CIMMYT headquarters are located, is a center
wheat releases having a CIMMYT ancestor. In 1970, semidwarf of origin for maize. The use of CIMMYT germplasm has
wheat varieties were important only in Asia. By the 1990s, increased over time, with about two-thirds of public sector
semidwarf wheat varieties covered 80% of world wheat area, varietal releases containing CIMMYT germplasm. Estimates of
with adoption rates of 90% and higher in Asia and Latin economic benefits from CIMMYTs maize research vary, but
America. More recently, across Afghanistan, Bangladesh, based on the earlier results (21.2 Mha planted to CIMMYT-
Ethiopia, Egypt, Nepal, and Pakistan, high-yielding wheat vari- related varieties) and a conservative estimate of farm-level
eties resistant to Ug99, a highly virulent disease, have protected maize prices (US$120 per tonne, on import parity basis),
the livelihoods and food security of 500 000 farming families. gross benefits range from US$1.3 billion to US$4 billion per
ICARDAs wheat improvement research focuses on spring year. These benefits take into account both germplasm
bread and durum wheats in the West AsiaNorth Africa region. improvements and better crop management practices.
ICARDA grain scientists have made major advances in identi- CIMMYT also developed quality protein maize (QPM), a
fying new sources of resistance to abiotic stresses (drought, product of 30 years of research. QPM looks and tastes like
heat, and cold) and biotic stresses (rusts, Septoria diseases, normal maize but has nearly twice the amount of lysine and
and Hessian fly), in broadening genetic diversity and in select- tryptophan, two amino acids essential for protein synthesis in
ing for grain quality. ICARDAs genebank holds over 135 000 humans. QPM is now being planted on 1 Mha in 20 countries,
accessions from over 110 countries: traditional landraces, boosting food, nutrition, and income security. More recently,
434 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: CGIAR
drought-tolerant maize has increased farmers yields by The CGIAR, with greater emphasis on the centers working
2030%, benefiting 20 million people in 13 African countries. together with each other and with partners in developed and
IITA has a regional maize improvement mandate mainly tar- developing countries to leverage complementary skills and
geting humid tropical and moist savannah zones of Western resources expect a number of significant impacts from the
and Central Africa. Maize accounts for more than 20% of new CGIAR Research Programs. Some of the planned impacts
domestic food production in Africa. from the new programs related to grains (www.cgiar.org/our-
research) include the following:
Other Crops By 2035, research on rice will increase farmers yields and
lower prices for poor consumers, lifting 150 million people
Two significant achievements involving ICRISAT should be
out of poverty and reducing the number of undernourished
mentioned. Breeding and disseminating micronutrient-rich
people in Asia by 62 million.
crop varieties through biofortification are potentially a cost-
effective strategy to increase micronutrient intake such as iron By 2020, 12 million households in Africa will have access to
sustainable irrigation, thanks to research on water, land,
and zinc. India has some of the worlds worst malnutrition rates,
and ecosystems.
resulting from a lack of essential vitamins and minerals in the
diet. This affects the survival and development of women and By 2022, research will help increase harvests of grain
legumes a key source of protein for the poor in low-
children in particular. About three quarters of children under
income countries in five regions, improving their nutrition
three and half of women suffer from anemia, mostly caused by
from 2.1 million tonnes of extra protein.
iron deficiency. Pearl millet is a nutritious but underutilized
staple crop, eaten by 50 million people in dry rural areas of By 2018, 50 million people will have access to staple food
crops specifically bred to be rich in key vitamins and
India. As well as being rich in B vitamins, calcium, iron, potas-
minerals namely, iron, zinc, and vitamin A in an effort
sium, magnesium, and zinc, it is well adapted to drought, poor
to combat malnutrition.
soils, and high temperatures. It is much higher in iron than
wheat, rice, or maize. ICRISAT is developing drought-tolerant
pearl millet varieties with even higher levels of iron. This is part Exercises for Revision
of a larger biofortification initiative coordinated through Har-
vestPlus and IFPRI and reviewed in Saltzman et al. Which CGIAR centers have a mandate for rice research?
Another significant impact involves partnership with agri- What has been the main shift in research organization in
business on the microdosing of fertilizer to resource-poor the CGIAR centers over the last 35 years?
farmers in sub-Saharan Africa. This involves using only What are some of the approaches that have been used to
810 kg of nitrogen per hectare but applying it precisely to the facilitate the extension of research results from CGIAR cen-
roots three to 4 weeks after planting. In Zimbabwe, ICRISAT has ters to the NARES?
recorded yield increases of 3050% among farmers adopting the What has been implemented to provide a back up in the
technique. With evidence that precision agriculture techniques event of a CGIAR centers genebank being destroyed
can work, the challenge is creating appropriate enabling envi- through war or civil unrest?
ronments to encourage take-up. In addition, ICRISAT has been
active in developing agribusiness incubators, but the involve- Exercises for Readers to Explore the Topic Further
ment of other centers in agribusiness has been variable.
What are the main research activities of IRRI? How are they
funded and what is their relationship with major research
Conserving Biodiversity organizations in the developed world?
How do developed countries benefit from the results of
For more than 30 years, CGIAR scientists have been collecting, CIMMYT research?
characterizing, and conserving biodiversity. Currently, the Which centers provide the enabling science in natural
CGIAR holds over half a million samples of crop, forage, and resources management to underpin the work of the crops-
agroforestry genetic resources in 11 gene banks around the specific centers? How do they collaborate to support each
world. Since the 1980s, CGIAR gene banks have distributed other?
well over one million germplasm samples worldwide, and the
vast majority of requests came from developing countries. In
1994, CGIAR signed an agreement with FAO, placing the col- Acknowledgments
lections in public trust, available to researchers worldwide.
Recognizing the challenge of creating a sustainable funding Some of the text, particularly in the Current and Future
base for conservation activities, the Global Crop Diversity Trust Challenges and Mobilizing Grain Science for Development
(GCDT) was established in association with the CGIAR to sections, has been adapted from an article for the first edition
support global conservation of biodiversity. More recently, of the encyclopedia, by F.J.B. Reifschneider and S. Hussain,
the Svalbard Global Seed Vault has been established as a secure formerly of the CGIAR office in Washington DC.
seedbank in Norway, only 1300 km from the North Pole. It
maintains duplicate samples of seeds held in other genebanks
worldwide, as an attempt to insure against the loss of seeds in See also: Agronomy of Grain Growing: Implication of Climate
other genebanks during large-scale crises. Changes; Wheat: Agronomy; Breeding of Grains: Barley: Genetics
SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: CGIAR 435
and Breeding; Maize: Breeding; Rice: Breeding; Soybean: Germplasm, Conway G (1999) The Doubly Green Revolution Food for All in the 21st Century.
Breeding, and Genetics; Wheat Breeding: Exploiting and Fixing Genetic London: Penguin Books.
Evenson RE (2003) In: Evenson RE and Gollin D (eds.) Crop Variety Improvement and
Variation by Selection and Evaluation; Food Grains and the
its Effect on productivity The Impact of International Agricultural Research,
Consumer: Cultural Differences in Processing and Consumption; pp. 71108. Wallingford, UK: CABI Publishing.
Grains Around the World: Grain Production and Consumption: Hazell P (2003) Green revolution. In: Mokyr J (ed.) The Oxford Encyclopedia of
China and South-East Asia; Grain Production and Consumption in Economic History UK: Oxford University Press.
Oceania: Australia and Pacific Countries; Grain Production and Hazell P and Ramasamy C (1991) The Green Revolution Reconsidered: The Impact of
High-yielding Rice Varieties in South India. Baltimore, MD: Johns Hopkins
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Grain Crops: An Overview; The Cereal Grains: An Overview of the Lipton M and Longhurst R (1985) Modern Varieties, International Agricultural
Family of Cereal Grains Prominent in World Agriculture; Wheat- Research and the Poor. CGIAR Study Paper Number 2, Washington DC, USA: The
Based Foods: Cereals: Overview of Uses: Accent on Wheat Grain. World Bank.
Lipton M and Longhurst R (1989) New Seeds and Poor People. Baltimore, MD: Johns
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Newman LF (1990) Hunger in History: Food Shortage, Poverty, and Deprivation.
Further Reading Cambridge, UK: Basil Blackwell.
Pingali PL (2012) Green revolution: Impacts, limits, and the path ahead. Proceedings of
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Agricultural R&D Policy, p. 203. Washington, DC: American Enterprise Institute. 109: 1230212308.
Baum WC (1986) Partners Against Hunger: Consultative Group on International Raitzer DA and Kelley TG (2008) Benefit-cost meta-analysis of investment in the International
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Bouman BAM, Hengsdijk H, Hardy B, Bindraban PS, Tuong TP, and Ladha JK (2002) Renkow M and Byerlee D (2010) The impacts of CGIAR research: A review of current
Water-wise rice production. In: Proceedings of the International Workshop on evidence. Food Policy 35: 391402.
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Research Organizations of the World: Europe and North America
JH Skerritt, University of Queensland, St Lucia, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights the United States, France, and Germany, with Canada and
particularly the United Kingdom having smaller public sector
While maintaining a systems and sustainability focus, there roles. There is also a greater proportion of agricultural (includ-
has been renewed emphasis on research directed at improv- ing grains) research carried out in universities in Europe and
ing grain yields in many countries. North America compared with developing countries, where it
The formation of networks between grain research investors, is largely within government-funded research institutes. Grain
research institutes, and industry increasing in Europe and R&D carried out by commercial and industry organizations is
North America. reviewed in a separate article, so the emphasis of this article is
A greater proportion of agricultural (including grains) on major public sector (government and university) grain
research is carried out in universities in Europe and North science organizations.
America compared with developing countries, where it is
largely within government-funded research institutes.
Significant increase in commercial involvement in grain
The United States
breeding and biotechnology, particularly where strong intel-
lectual property protection and end point royalty systems
are in place. Farmer organization-led grain research is also Although the United States has very large private sector (seed
becoming more important. companies; milling, baking, brewing, and oilseed processing
companies; and biotechnology companies) involvement in
Research to underpin grain utilization for animal feed and
grain R&D, government investments are also significant. A
for biofuels is important in several countries.
wide range of temperate grain crops are targeted especially
not only wheat, barley, maize, and oilseeds but also pulses and
subtropical rice, with most resources dedicated to maize and
Learning Objectives wheat research. State and federal governments invest roughly
equally in agricultural research.
To understand the structure and major roles of public sector The Agricultural Research Service (ARS, www.ars.usda.gov)
grain research organizations within Europe and North is the principal research agency of the (national) US Depart-
America. ment of Agriculture (USDA), and together with the land
grant universities, they form the main research centers that
conduct grain research. The development of a single strategic
Introduction plan and national program structure by USDAARS in recent
years has increased the amount of coordination between the
Compared to the long history of cultivation of grains (many 800 or so individual USDA projects underway in different
thousands of years), the use of research principles to address centers. Crop Production and Protection is the third national
constraints is a comparatively recent approach. Agricultural program described in USDAARSs 201217 strategic plan
societies were established in the United Kingdom, France, and delivers science-based information, genetic resources,
and Germany in the mid-1800s (e.g., The Royal Agricultural and technologies for increased crop productivity, economically
Society of England (1839), La Societe des Agriculteurs de and environmentally sustainable methods of crop production,
France (1850), and Deutsche Landwirtschafts-Gesellschaft and protection from plant diseases and pests. There are six
(1885)) and by the mid-to late nineteenth century, agriculture subprograms, each relevant to grain research: Plant Genetic
experiment stations were appearing in Europe and North Resources, Genomics and Genetic Improvement; Plant Biolog-
America. The tradition of government-funded grain research ical and Molecular Processes; Plant Diseases; Crop Protection &
is the longest established in Europe and North America, Quarantine; Crop Production; and Methyl Bromide
although similar initiatives were started in the late nineteenth Alternatives.
century in a number of European colonies. The two major goals under area 3 are firstly to Protect,
Government-funding bodies have shown a greater interest expand, and enhance the United States crop genetic resource
in assessing the economic and social impacts of research that base, increase scientific knowledge of crop genes, genomes,
they fund research funding is increasingly viewed as an biological processes and systems, and deliver economically
investment rather than as grants. The most significant trend and environmentally sound technologies that improve the
in terms of research intensity is the very significant growth over production efficiency, quality, health and value of the nations
the last couple of decades in the level of grain (especially crops and secondly to Develop knowledge, strategies, sys-
cereals) R&D carried out by the private sector or alternatively tems and technologies that maximize the production efficiency
funded by the private sector but carried out in universities or of our annual, perennial, greenhouse and nursery cropping
government organizations. Most publicly funded agricultural systems. Develop new technologies and tools contributing to
research in North America and Europe is concentrated in improving these systems to meet current and future food crop
production needs of diversified consumers, while ensuring Pullman, WA (Pacific West Area) Wheat Genetics, Quality
economic and environmental sustainability and production Physiology and Disease Research, Land Management and
efficiency, health, and value of our nations crops. The Natural Water Conservation Research, and Root Disease and
Resources and Sustainable Agricultural Systems program clus- Biological Control Research
ter is also relevant. Raleigh, NC Soybean and Nitrogen Fixation Research and
Some USDA centers are free-standing multidisciplinary Plant Science Research Laboratories
research centers, while smaller ones are typically colocated on Shafter, CA Western Integrated Cropping Systems
university campuses. Many centers have the mandate to carry Research
out research directed at the particular agroecological zone in Sidney, MT Agricultural Systems Research Unit
which the institute is based. For example, wheat is grown in Stillwater, OK Wheat, Peanut, and Other Field Crops
many parts of the United States, but there are different quality Research, Plant Science and Water Conservation Research
types suited for different locations soft red winter wheat in Stoneville, MS Crop Genetics & Production Research Unit
the Midwest, Virginia, and North Carolina; hard white and St Paul, MN Midwest Area Cereal Disease Laboratory and
some hard red winter wheat in Pacific Northwest; hard red Plant Science Research
winter varieties in the Great Plains, from Texas to Montana Stuttgart, AR Dale Bumpers National Rice Research Center
on the Canadian border; and spring varieties in northern Tifton, GA Crop Genetics and Breeding Research and
central states such as Minnesota, North and South Dakota, Crop Protection and Management Research
and Montana and appropriately located USDA centers Urbana, IL Soybean/maize Germplasm, Pathology, and
develop germ plasm and agronomy packages for the different Genetics Research
types of varieties. West Lafayette, IN Crop Production And Pest Control
Some of the USDA centers carrying out grain research are Research
based in the following: Wooster, OH Corn, Soybean and Wheat Quality Research
Wyndmoor, PA Crop Conversion Science & Engineering
Aberdeen, ID Small Grains and Potato Germplasm Research Unit
Research
Albany, CA (Western Regional Research Center) Genomics USDA also has Economic Research Service (www.ers.usda.
and Gene Discovery and Crop Improvement/utilization gov), which carries out research and communication activities
Research in many areas relevant to grains, including farm-level risk
Ames, IA Corn Insects and Crop Genetics Research management, commodity outlook research, and economic
Beltsville, MD (Beltsville Agricultural Research Center) and statistical research on marketing and production sustain-
Sustainable Agricultural Systems Laboratory, Instrumentation ability. The USDA National Institute of Food and Agriculture
and Sensing Laboratory, National Germplasm Resources (NIFA, www.nifa.usda.gov) replaced the Cooperative State
Laboratory, Soybean Genomics and Improvement Labora- Research, Education, and Extension Service in 2009 and is
tory, and Food Quality Laboratory intended to consolidate all federally funded agricultural
College Station, TX (Southern Plains Agricultural Research research (including federal support to land grant universities)
Center) Crop Germplasm Research and Insect Control and have overarching federal coordination of extension work
and Cotton Disease Research done through USDA.
Colombia, MO Plant Genetics Research The land grant college system was established in 1862
Corvallis, OR Forage Seed and Cereal Research the name is derived from the fact that the colleges were origi-
Fargo, ND Cereal Crops Research and Hard Spring and nally endowed by grants of public lands in the expanding
Durum Wheat Quality Laboratories western United States. The 1887 Hatch Act established the
Fort Collins, CO National Center for Genetic Resources agricultural experiment station system to enable these colleges
Preservation to undertake research, and the 1914 SmithLever Act estab-
Ithaca, NY Plant Genetic Resources, Plant Protection lished the cooperative extension system. This unique system
Research, Plant, Soil and Nutrition Research has the advantage of linking state government R&D and exten-
Lane, OK (South Central Agricultural Research Laboratory) sion activities to university research. The greatest emphasis for
Genetics and Production Research state government efforts is on breeding new cultivars and
Lincoln, NE Grain, Forage, and Bioenergy Research adapting crop production approaches to local conditions and
Lubbock, TX Cropping Systems Research Laboratory and on education. The universities usually have experiment sta-
Plant Stress and Germplasm Development Research tions at different locations in their state, again often corre-
Madison, WI Cereal Crops Research sponding to different agroecological zones but also requiring
Manhattan, KS (Center for Grain and Animal Health researchers to carry out a local outreach function. There are
Research) Grain Quality and Structure Research, Stored usually a balance of applied grain research on constraints to
Product Insect Research, and Hard Winter Wheat Genetics productivity in a particular state and a proportion directed
Research toward more fundamental research. Cooperative extension
Peoria, IL (National Center for Agricultural Utilization systems from land grant universities have a state development
Research) Cereal Products and Food Science Research, mandate and often involve location of farm advisors in
Plant Polymer Research, Mycotoxin Research, New Crops counties throughout the state. The continued involvement of
and Processing Research, Crop Bioprotection Research, and the government and public funding in agricultural extension in
Bacterial Foodborne Pathogens & Mycology Research the United States is in contrast with the winding back or exiting
438 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: Europe and North America
of state or national government staff from extension roles in universities are usually predominantly state-funded, but with
other countries such as the United Kingdom, Australia, and significant federal funding as well.
New Zealand. A summary of some of the main US universities In 2011, the US government commenced the presidents
that carry out grain research is provided in Table 1. Land grant global hunger and food security initiative (www.feedthefuture.
Colorado State University, Fort Collins, CO Genomics, biotechnology, dryland farming systems, wheat breeding, precision agriculture
(www.agsci.colostate.edu)
Cornell University, Ithaca, NY Genetics, biotechnology, basic plant science, maize breeding, seed science, cereal pathology, cereal
(www.cals.cornell.edu) product rheology. Wheat variety development, genetics research, disease resistance, and crop
management. Particular emphasis on breeding methodologies, disease resistance, comparative
genomics, association mapping, allele characterization, and genomic selection methods
Iowa State University, Ames, IA (www.ag. Corn and soybean breeding, nonfood crop products, corn quality, basic plant sciences, crop genomics,
iastate.edu) seed technology
Kansas State University, Manhattan, KS Wheat, soybean, and sorghum breeding; cereal (wheat, corn, sorghum, and millet) agronomy and
(www.k-state.edu) physiology, entomology, cereal chemistry, milling and baking, livestock feeds. Wheat genetics,
development of genomics resources and high throughput genotyping technologies, wheat breeding,
germ plasm development
Michigan State University, East Lansing, MI Grain crop breeding, cereal chemistry, genomics
(www.maes.msu.edu)
Montana State University, Bozeman, MT Winter wheat, barley, durum, pulses, canola, and soybean breeding, precision agriculture, integrated
(www.montana.edu/agriculture) pest management
North Carolina State University, Raleigh, NC Bioinformatics, integrated pest management
(www.cals.ncsu.edu)
North Dakota State University, Fargo, ND Basic research on wheat quality and processing; milling; pasta production; brewing; farming systems,
(www.ag.ndsu.nodak.edu) pathology, and entomology
Oklahoma State University, Stillwater, OK Baking, grain protein chemistry, oilseed chemistry, plant diseases
(www1.dasnr.okstate.edu)
Ohio State University, Columbus, OH Maize production systems, cereal and oilseed quality, soft red winter wheat and soybean agronomy and
(www.oardc.ohio-state.edu) breeding, soy protein functionality
Pennsylvania State University, University Starch chemistry, entomology
Park, PA (www.research.cas.pse.edu)
Purdue University, West Lafayette, IN Plant stress physiology, biotechnology, integrated pest management, precision agriculture, new grain
(www.agriculture.purdue.edu/arp) products, carbohydrate research
Rutgers University, New Brunswick, NJ Cereal processing, plant biotechnology
(www.foodsci.rutgers.edu/caft)
NJ Agricultural Experiment Station
(www.cook.rutgers.edu)
Texas A&M University, College Station, TX Precision agriculture, farming systems, plant genomics, grain food processing
(www.agresearch.tamu.edu; www.
agrprogram.tamu.edu)
University of Arkansas, Fayetteville, AR Rice, wheat, oat, and soybean breeding; agronomy and physiology, rice processing
(www.uark.edu/admin/aes)
University of California, Davis (www.ucanr. Genetic resources conservation, genetics of cereals and legumes, agronomy, wheat breeding,
org; www.caes.ucdavis.edu) entomology, plant pathology, rice breeding
Breeding, Marker-Assisted selection, Positional cloning of agronomic genes, Dissection of flowering
and disease resistance regulatory pathways
University of Illinois, UrbanaChampaign, IL Maize and soybean breeding and agronomy, cereal diseases, starch chemistry
(www.web.aces.uiuc.edu)
University of Idaho, Aberdeen, ID (www. Grain breeding and agronomy (wheat, oilseeds)
ag.uidaho.edu)
University of Minnesota, St. Paul, MN Cereal structure, wheat and barley breeding and pathology, soybean diseases, precision agriculture,
(www.maes.umn.edu) cereal food processing
University of Missouri, Colombia, MO Tillage, soybean, sorghum, and corn cropping systems and soybean breeding, basic grain science,
(www.cafnr.missouri.edu) maize genome mapping
University of Nebraska, Lincoln, NE Processing of grain starches, cereal breeding, genomics. Wheat breeding and genetics of winter hardy,
(www.unl.edu) disease and insect resistant, high-yielding cultivars. Fundamental research in wheat virology, fungal
disease control, biochemistry, statistical approaches, and breeding methodology including
transformation.
Washington State University, Pullman, WA Grain crops breeding and genetics, pathology/entomology, soil science, processing
(www.cahe.wsu.edu)
SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: Europe and North America 439
gov/approach), which includes the Feed the Future research Centre, Quebec (for cool, wet production systems), Atlantic
strategy. This is a major international development assistance Cool Climate Crop Research Centre (St Johns, Newfound-
initiative managed through USAID. Investments under this land), Crops and Livestock Research Centre (Charlottetown,
strategy range from longer-term research to address major Prince Edward Island), Brandon Research Centre, and
global challenges to applied and adaptive research guided by Semiarid Prairie Agricultural Research Centre (Swift Current,
host-country priorities for nearer-term impact. A number of US Saskatchewan).
research laboratories within USDA and the university sector are Several universities carry out grain research in Canada,
involved given the central role of grains in food security, it is often working closely with Agriculture and Agri-Food Canada
not surprising that several initiatives support grain research. scientists. These include the following:
Within three broad research themes (advancing productivity,
transforming production systems, and enhancing food safety
University of Saskatchewan (www.usask.ca) with research
emphasis on wheat, barley, oat, canola, and legume (lentil,
and nutrition), these include work on wheat stem rust, abiotic
pea, dry bean, and chickpea) breeding and agronomy, plant
stress tolerance, genetic resources, marker-assisted technolo-
genetic engineering, molecular marker development, and
gies, legume productivity, and management of mycotoxin con-
malting and brewing science.
tamination in stored grains.
University of Alberta, Edmonton (www.afns.ualberta.ca),
studies grain and oilseed crops for Western Canada
breeding, disease resistance, agronomy, and livestock feed
utilization. In wheat, it focuses on conventional breeding
Canada and genetic research with an emphasis on adaptation of
wheat to the short growing season on the Canadian prairies
The majority of wheat breeding and research in Canada is
and agronomy and breeding of grains for organic agricul-
conducted by the public sector, although private sector breed-
tural environments.
ing is gaining momentum and Syngenta has released wheat
varieties in Canada since 1993. Several publicprivate partner-
University of Manitoba, Winnipeg (www.umanitoba.ca/
afs/plant_science), carries out research on canola and
ships exist between industry and most publically funded
wheat breeding and postharvest technology, grain storage
research institutions. Agriculture and Agri-Food Canada
technology, cereal chemistry, baking technology, pulse
(www.agr.gc.ca/science and innovation/research centers)
food science, functional foods, and cereal agronomy
carries out grain research at several of its 19 research centers
including precision agriculture, genetics and wheat breed-
across the country. Cereal research is carried out mainly at
ing, agronomic production systems, pathology, breeding
the Winnipeg (Manitoba) Cereal Research Centre, which is co-
methodologies, and end-use quality.
located with the University of Manitoba and emphasizes six
areas: cereal diseases, cereal germ plasm and genomics, pulse
University of Guelph, Ridgetown, Ontario (www.ridge-
townc.com/index.cfm), emphasizes grain crops breeding
germ plasm, nutrition and functional foods, bioprospecting
(soybean, barley, wheat, and canola), agronomy, pathol-
from grains, and grain storage research. The Eastern Cereal
ogy, and genome mapping.
and Oilseed Research Centre (Ottawa, ON) develops new vari-
eties of spring and winter wheat, barley, corn, oats, and soy-
McGill University, Quebec, Ontario (www.mcgill.ca), is
involved with basic plant science, biotechnology, and
beans and has the central biodiversity mandate for Canada.
some grain food processing research.
There is a strong emphasis on gene isolation, gene transfer,
and the study of gene expression in grain crops, bioprospecting The National Research Council (NRC) Plant Biotechnology
for new bioactives, and study of Fusarium resistance. Institute in Saskatoon (www.pbi.nrc.ca) carries out basic
Wheat breeding is also carried out at the Semiarid Prairie research in genomics and proteomics and has a special inter-
Agricultural Research Centre (Swift Current, Saskatchewan, est in the modification of oil composition and of agronomic
along with breeding of other cereals) and the Lethbridge traits of oilseeds and in cereal and legume molecular marker
Research Centre (Alberta). The Lethbridge Research Centre and transformation technology. Some individual provinces
also carries out bean and forage crop breeding and research also have agencies that carry out applied agriculture and
on water, soil, crop, and manure management for prairie con- food research. For example, the Field Crop Development
ditions. There is an emphasis on disease and disease resistance Centre is funded in part by the province of Alberta and
and agronomy. The Brandon Research Centre (Manitoba) is conducts breeding and development programs focussed on
the main site of barley breeding along with a strong soil science feed and malting barley, spring and winter triticale, and
focus, while the Lacombe Research Centre (Alberta) under- winter wheat. The Canadian Grains Commission in Winni-
takes oat-varietal improvement research, along with barley peg, Manitoba, regulates grain handling and establishes and
breeding (disease resistance focus) and farming systems maintains grain quality standards for Canada. Its Grain
research. The Saskatoon Research Centre (Saskatchewan) is Research Laboratory (www.grainscanada.gc.ca/cgc-ccg/grl-lrg/
the main center of breeding programs for canola and mustard grlm-mlrg-eng.htm) has emphasis on research to understand-
for prairie conditions. Some canola breeding and breeding for ing the genetic, environmental, structural, and biochemical
other short-season grain crops also take place at the Lacombe basis on end-use quality of major grains, how grading factors
Research Centre, while the Greenhouse and Processing Crops affect end-use qualities, and investigating new uses for Cana-
Research Centre in Harrow, Ontario, also carries out soybean dian grain. The Canadian International Grains Institute
breeding. Agronomy research for grain production is under- (www.cigi.ca), also in Winnipeg, has a technical training
taken at the Soils and Crops Research and Development and information dissemination role.
440 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: Europe and North America
related to cereals. At INRA Montpellier, there is an ongoing institute that is now part of Wageningen University and Research
research on processing technology, grain crop production Center, carry out grain research, including the Agrotechnology
systems, plant protection, and genetic resource conservation and Food Sciences Group and Plant Research International
as well as more basic studies on grain crop development. (www.pri.wur.nl/uk) who focus on crop protection, crop ecol-
Several French universities have Ecole Nationale Superieure ogy, agricultural systems, genetics, and reproduction. The Neth-
Agronomique, for example, in Paris Grignon, (the Institut erlands Organisation for Applied Scientific Research (TNO)
National Agronomique (INA-PG)), Montpellier, Rennes, conducts research on cereal processing. Wageningen University
Nancy, and Toulouse. ARVALIS Institut du vegetal (www.arva- (www.wageningenur.nl/en/wageningen-university.htm) is the
lisinstitutduvegetal.fr/en) is responsible for applied research main university research center for grain research. In Belgium,
on cereals, maize, pulses, potato, and forage crops. The activ- there are relevant government agricultural research centers
ities of this technical institute, which is run and financed by in Gent and Gembloux, while at least four universities carry
farmers, include sustainable cropping systems, precision farm- out some aspects of grain production and processing research
ing, remote sensing, satellite images, biotechnology, farm eco- Ghent University, Catholic University of Leuven, Gembloux
nomics, quality assurance, safety quality, traceability, human Agricultural University, and Catholic University of Louvain.
food, animal feed, and technology transfer. CETIOM (www. The Italian government Ministry of Agricultural, Food and
cetiom.fr) is responsible for similar topics on oil crops. Cer- Forestry Policies has several research institutes covering grain
eales Vallee (www.cereales-vallee.org) is a government research, including various Research Institutes for Cereal
competitiveness cluster established in 2005. It supports the Research (ISC, www.sito.entecra.it/portale/cra_dati_istituto.
development of applied cereal research projects through col- php?id205&linguaEN) and the Industrial Crops, Plant
laboration between a number of public and private partners in Nutrition, Plant Pathology and Food and Nutrition
France. There are four programs that in turn address sustain- (IMRAN). The NRC (CNR, www.cnr.it) has a separate system
able cereal production, cereals for human nutrition and health, of research institutes, and some of these also carry out grain
cereals for animal feed, and cereals for agromaterials. research. Many major states in Italy host universities that have
Breeding of new cereal cultivars in Germany is aimed at agriculture faculties, and most of these carry out at least some
increased yields, improved resistances to biotic and abiotic grain research the Universities of Turin, Milan, Padua, Udine,
stress, and high quality and like many other European coun- Bologna, Piacenza, Florence, Pisa, Ascoli, Perugia, Viterbo,
tries is now conducted by private breeding companies, with Naples, Bari, Potenza, Reggio Calabria, Catania, Palermo, and
government researchers carrying out prebreeding activities. Sassari.
Cereal research is conducted at the following: The Cereal Institute, National Agricultural Research Foun-
dation (NAGREF, www.nagref.gr), Thessaloniki, Greece, is the
Universities, for example, the Technical University of
Greek government research institute responsible for cereal
Munich, Technical University of Berlin, Martin Luther Uni-
breeding and cereal food product development. However, the
versity Halle-Wittenberg, Justus Liebig University Giessen,
Ministry of Agriculture has a separate research service with a
Humboldt State University, and the Universities of Bonn,
network of institutes in different parts of the country. At
Gottingen, Hohenheim, Kassel, Kiel, Rostock, and
least three universities carry out some grain research work
Paderborn.
Agricultural University of Athens, Aristotle University of
Federal research institutes, for example, the Leibniz Insti-
Thessaloniki, and the Mediterranean Agronomic Institute at
tute of Plant Genetics and Crop Plant Research (IPK, www.
Chania. In Hungary, the Ministry of Agriculture and Rural
ipk-gatersleben.de/en); the Julius Kuhn Institute (JKI, www.
Development manages the Central Food Research Institute
jki.bund.de/en); Federal Centre for Cereal, Potato, and
(KEKI) and the Agricultural Biotechnology Center (MBK), as
Lipid Research (BAGKF, Detmold, www.mri.bund.de/en/
well as the Cereal Research Institute in Szeged. The Hungarian
de/max-rubner-institut/geschichte/bundesanstalt-fuer-
Academy of Sciences has several institutes relevant to grain
getreide-kartoffel-und-fettforschung.html); the Federal
science the Agricultural Research Institute, the Research Insti-
Research Centre for Plant Breeding (BAZ, www.genres.de);
tute for Soil Science and Agricultural Chemistry, and the Plant
and the Institute of Agricultural Engineering (ATB,
Protection Institute. Universities with research areas most rel-
Potsdam-Bornim, www.atb-potsdam.de/en.html).
evant to grain science are the Technical University of Budapest
Max Planck Institute for Plant Breeding Research (MPIPZ,
(grain processing) and the Universities of Agricultural Sciences
www.mpipz.mpg.de/2169/en).
at Godollo, Debrecen, and Pannon.
German state research centers in agriculture several of
In Austria, the Federal Ministry of Agriculture, Forestry,
these carry out grain research, sometimes in conjunction
Environment and Water Management has research institutes
with universities (e.g., Bavarian State Research Centre for
for general agriculture and an institute of cereal processing,
Agriculture and State Plant Breeding Institute of the Univer-
applied soil science, and agricultural engineering. There is also
sity of Hohenheim (Baden-Wurttemberg)).
the University of Natural Resources and Life Sciences in Vienna.
The proWeizen initiative was recently founded and is coordi- The International Association for Cereal Science and Technol-
nated by the German Federation of Private Plant Breeders (GFP). ogy (www.icc.or.at) is headquartered in Vienna and has
Its aim is to coordinate and foster German wheat research. It evolved from being a professional organization of cereal scien-
consists of leading research institutes and private companies and tists to also hosting a number of EU (Seventh Framework)-
organizations involved in wheat breeding and growing. funded grain research projects. These include projects on
In the Netherlands, two institutes within the Dienst Land- food-related disease prevention, bioactives and food supple-
bouwkundig Onderzoek (DLO), a private nonprofit research ments from cereal grains, food allergens and allergy risk
442 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: Europe and North America
management, and mycotoxin analysis. In the Czech Republic, After separation from Russia in 1991, national agricultural
relevant Ministry of Agriculture research institutes include the research systems developed in most of the former Soviet
Food Research Institute, Research Institute of Crop Production, republics using the institutional assets left by the former
Research Institute for Soil and Water Conservation, and USSR; several of these have a crop emphasis although resources
Research Institute of Agricultural Engineering. There are semi- for research are very limited and it has been difficult (and often
commercial institutes responsible for cereals (Agricultural undesirable) to maintain the staff numbers and infrastructure
Research Institute Kromeriz Ltd.), oilseed crops (Oseva Ltd.), left behind. They have had to develop their priority setting to
and brewing and malting. Two universities Mendel University match specific requirements of their various agroecological
of Agriculture and Forestry and Czech University of Agriculture zones and the changed policy/economic environment includ-
are relevant to grain research. In Spain, the Ministry of Agri- ing a greater emphasis on markets rather than a central procure-
culture, Fisheries and Food runs the National Institute for ment and distribution system.
Agriculture and Food Research and Technology (INIA, www.
inia.es), while the Polytechnic University of Madrid has a large
faculty involved with research in agriculture. There is also an
INIA in Portugal, while agricultural research is carried out at the Exercises for Revision
Technical University of Lisbon, the Universidade de Tras-os-
Montes e Alto Douro, and Universidade dos Acores. What are some of the major institutional trends in grain
In Sweden, the government Swedish Institute for Food research in North America and Europe over the last
Research and the Swedish Institute for Food and Biotechnol- decade?
ogy (SIK, a joint industry/government body) carry out grain How do the major grain research priorities differ between
processing research. Production research and basic scientific now and 30 years ago?
investigations within universities include the Swedish Univer- What are the major priorities globally for biofuel research?
sity of Agricultural Sciences at Svalov and Lund University. Name three of the countries within Europe and North
Three main institutes in Norway carry out grain research. America that have the highest proportion of publicly
Within the government Ministry of Agriculture, there is the funded agricultural R&D.
State Agricultural Research Station (SFL) and the Agricultural Why are agricultural universities in the United States called
University of Norway at Aas, while the Norwegian Food land grant colleges?
Research Institute (Matforsk) is a nonprofit organization. In
Finland, the Ministry of Agriculture and Forestry has the Agri-
cultural Research Centre of Finland and Plant Breeding Insti-
tute, while VTT Biotechnology in Espoo carries out cereal Exercises for Readers to Explore the Topic Further
technology research in close collaboration with industry. The
University of Helsinki is the main higher education center Explore the major grain research projects funded under the
dealing with grain research. In Denmark, the Ministry of Seventh Framework Programme of the European Union.
Food, Agriculture and Fisheries has Institutes of Agricultural What are the common themes?
Sciences and Agricultural Engineering. The Royal Veterinary What have been the major achievements of the N.I. Vavi-
and Agricultural University also carries out grain research. lov Institute of Plant Industry in Russia over the last
Activities related to the Carlsberg laboratory are described in century?
the article Research Organizations of the World: Global What areas of grain research from the Netherlands major
Trends and the Commercial Sector. agricultural university in Wageningen have had global
In Russia, the Russian Academy of Agricultural Sciences impact?
(www.euracadagri.com/eng/members/russia.php) comprises
many dozen institutes. These include All-Russia Research Insti-
tutes (e.g., All-Russia Research Institute of Plant Quarantine See also: Agronomy of Grain Growing: Plants: Diseases and
(Bykovo), Plant Protection (St. Petersburg), and Biological Pests; Soybean: Agronomy; Wheat: Agronomy; Breeding of Grains:
Protection of Plants (Ekaterinburg), which carryout entomol- Barley: Genetics and Breeding; Canola/Rapeseed: Genetics and
ogy research related to grains) and regional research institutes Breeding; Maize: Breeding; Soybean: Germplasm, Breeding, and
in former republics. The latter often cover a range of agricul- Genetics; Wheat Breeding: Exploiting and Fixing Genetic Variation by
tural disciplines and some have an almost 100-year history; Selection and Evaluation; Genetics of Grains: Development of
many have significant grain breeding and seed distribution Genetically Modified Grains; Genome Mapping; Genomics of Food
programs. Three of the All-Russia Research Institutes most Grains; Grain Marketing and Grading: Barley: Grading and
relevant to grains are the Institute of Legumes and Groat Marketing; Wheat: Grading and Segregation; Wheat: Marketing;
Crops (Streletskoye), Institute of Sorghum and Cereals (Zerno- Grains Around the World: Grain Production and Consumption:
grad), and N.I. Vavilov Institute of Plant Industry (www.vir. Cereal Grains in North America; Grain Production and Consumption:
nw.ru). The latter institute, based in St. Petersburg, but with a Europe; Grain Production and Consumption: Oilseeds in North
number of experimental stations throughout Russia, is the America; Grain Production and Consumption: Overview; Non-food
main internationally renowned Russian institute emphasizing Products from Grains: Biodiesel; Cereal Grains as Animal Feed; Fuel
the conservation and study of plant genetic resources, includ- Alcohol Production; Scientific Organizations Related to Grains:
ing grain crops. Most states have state agrarian universities, and Scientific Societies Associated with Grain Science; The Basics: The
there are a small number of specialist agrarian academies Grain Crops: An Overview; The Cereal Grains: An Overview of the
including Moscow Agricultural Academy. Family of Cereal Grains Prominent in World Agriculture.
SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: Europe and North America 443
Further Reading Pardey RG and Beintema NM (2001) Slow Magic: Agricultural R&D a Century after
Mendel. Washington DC: International Food Policy Research Institute.
Fox JA and Ward LL (2016) Grain production and consumption: Cereal McVetty PBE, Lukow OM, Hall LM, Rajcan I, and Rahman H (2016) Grain production
grains in North America. In: Wrigley C, Seetharaman K, Corke H, and and consumption: Oilseeds in North America. In: Wrigley C, Seetharaman K,
Faubion J (eds.) Encyclopedia of Food Grains, 2nd edn., vol. 1, pp. 391400. Corke H, and Faubion J (eds.) Encyclopedia of Food Grains, 2nd edn., vol. 1,
Elsevier Ltd. pp. 401408. Elsevier Ltd.
Grassini P, Estraidge KM, and Cassman KG (2013) Distinguishing between yield Ray DK, Mueller ND, West PC, and Foley JA (2013) Yield trends are insufficient to
advances and yield plateaus in historical crop production trends. Nat. Commun. double global crop production by 2050. PLoS One 8(6). http://dx.doi.org/10.1371/
4: 2918. http://dx.doi.org/10.1038/ncomms3918. journal.pone.0066428.
Lindhaeur MG (2016) Grain production and consumption: Europe. In: Wrigley C, Skerritt JH (2016) Research Organizations of the World: CGIAR. In: Corke H, Wrigley C,
Seetharaman K, Corke H, and Faubion J (eds.) Encyclopedia of Food Grains, Faubion J, and Seetharaman K (eds.) Encyclopedia of Food Grains, 2nd edn., vol. 1,
2nd edn., vol. 1, pp. 383390. Elsevier Ltd. pp. 429435. Elsevier Ltd.
Research Organizations of the World: Asia-Pacific, CentralSouth America, and
AfricaMiddle East
JH Skerritt, University of Queensland, St Lucia, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
In some cases, the alliances cover most areas of grains research; in University of Western Australia grain crop science and
others, specialized areas such as biotechnology. Examples rele- molecular genetics
vant to grains research include the Queensland Alliance for Agri- Charles Sturt University agronomy, irrigation, and plant
culture and Food Innovation (QAAFI, involving the University of pathology
Queensland, www.qaafi.uq.edu.au); the EH Graham Centre for Curtin University agronomy, precision agriculture, and
Agricultural Innovation (involving Charles Sturt University, grains processing
www.csu.edu.au/research/grahamcentre); the Primary Industries Murdoch University molecular markers for barley and
Innovation Centre in NSW (involving the University of New wheat, grain legume genome mapping, transgenics,
England, www.dpi.nsw.gov.au/research/alliances/primary-indus- diseases, and viruses
tries-innovation-centre); AgriBio in Victoria (involving La Trobe University of New South Wales grain storage engineering
University, www.latrobe.edu.au/agribio); the South Australia University of South Australia agricultural machinery
Research and Development Institute, colocated with the Univer- research and design
sity of Adelaide (www.sardi.sa.gov.au); the Western Australian University of Southern Queensland plantpathogen
State Agricultural Biotechnology Centre, located at Murdoch interactions, cereal genetic engineering, marker-assisted
University (www.sabc.murdoch.edu.au); and the Tasmanian breeding, and canola breeding
Institute of Agriculture (www.tia.tas.edu.au). While Queensland University of Tasmania cereals, grain legumes, and oil-
and Victoria retain reasonable capacity, an increasing proportion seeds as break and rotation crops
of extension services, formerly provided by the state governments,
Over the last 20 years, there has been a significantly increased
are now provided to farmers on a commercial basis by the private
emphasis on establishment of more formal links between
sector and/or incorporated farmers (growers) groups.
research organizations. This has been driven to a significant
Within the national research organization CSIRO (www.
extent by the advent of the Cooperative Research Centres
csiro.au), much research (including grains research) is now
(CRCs, www.crc.gov.au) program and encouragement by fun-
being conducted in cross divisional flagship programs. The
ders. At present (January 2014), there are only two CRCs that
Food Futures Flagship is using advanced genetics to create dif-
have relevance to grains research (Future Farm Industries CRC
ferentiated grain, food, and feed products, focussing on wheat
and the Plant Biosecurity CRC) many fewer than earlier in
barley with improved processing quality and oilseeds with
the program. The Grains Research & Development Corpora-
enhanced oil content and improved nutritional value. Other
tion (www.grdc.com.au) facilitates and manages funding for
flagships carrying out grains research include biosecurity,
research and assists in communicating results. Government
sustainable agriculture (natural resource management and farm-
and industry R&D levies co-fund this research.
ing systems focus), and climate adaptation flagships. A small
The main New Zealand organization undertaking grains
amount of grains-related research is conducted outside flag-
research is Plant & Food Research (www.plantandfood.co.nz),
ships, by the Divisions of Plant Industry; Animal, Food and
one of several government-owned Crown Research Institutes.
Health Sciences; Ecosystem Sciences; and Land and Water.
Its program includes wheat and barley genomics, breeding and
Some independent nongovernment grains research organi-
production systems, and food processing.
zations exist including GrainGrowers (www.graingrowers.
Universities with research interests in grains include Massey
com.au), their expertise including milling, near-infrared reflec-
University (Palmerston North, www.massey.ac.nz) with a
tance technology, baking science and technology, and Asian
research emphasis on food science and engineering. Lincoln
wheat foods. The Australian Grains Centre (www.cbh.com.au/
University (Christchurch, www.lincoln.ac.nz) carried out
our-business/operations/grain-quality/australian-grain-centre.
research on grain foods composition, crop agronomy, seed
aspx) and Agrifood Technology (www.agrifood.com.au)
science and technology, and some biotechnology. Although
emphasize testing services.
wheat is one of the major cereals, the growing environment is
Relevant grains research is carried out at several universities.
very different from that in Australia, requiring rather different
This includes the following:
varietal characteristics. Further information on grain produc-
University of Adelaide bread and durum wheat and barley tion and consumption in Oceania is discussed elsewhere in
prebreeding, molecular markers, cereal root diseases, cereal this Encyclopedia.
pathology, biotechnology, weed management, malting,
and brewing biochemistry
University of Sydney prebreeding for rust resistance in
North Asia
barley and wheat, cereal diseases, cereal genomics, preci-
sion agriculture, soil science, and crop modeling In Japan, much of the grains research is carried out in institutes
University of Melbourne research to underpin the sustain- managed under the Agriculture, Forestry, and Fisheries
ability of temperate grain crop production, with emphasis Research Council of the Ministry. National Agriculture and
on agronomy and farming systems, and protection of nat- Food Research Organizations (www.naro.affrc.go.jp/english/
ural resources index.html) Institute of Crop Science in Tsukuba conducts
University of New England agronomy and soil science, research on the breeding of rice, wheat, barley, soybean,
plant nutrition, crop protection, weed science, and inte- sweet potato, and industrial crops such as sesame and on the
grated pest management development of new technologies for breeding. Research activ-
University of Queensland production systems and post- ities also cover the understanding of basic mechanism of agro-
harvest handling of cereals, grain legumes and oilseeds, weed nomic and physiological traits regarding crop quality,
management, plant protection, and molecular biology photosynthesis, and productivity. The NARO Agricultural
446 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organisations of the World
Research Center, also headquartered in Tsukuba, focuses on The Crop Science crop germplasm resources cluster addresses
rice and soybean systems, agronomy, soil fertility, crop nutri- genetics and breeding, molecular biology, cultivation and
tion and rotations, pest management, and biofuels. There are physiology, molecular breeding, functional genomics, proteo-
four main regional research centers. The National Institute of mics, metabonomics, and bioinformatics participating grains
Agrobiological Sciences (www.nias.affrc.go.jp/index_e.html, institutes include the Institute of Crop Sciences, Biotechnology
also headquartered in Tsukuba) is the largest agricultural Research Institute, China National Rice Research Institute, and
research institute in Japan for basic life sciences, and a number the Oil Crops Research Institute. The Agricultural Resources
of its programs focus on grain crop/genomics and genetic and Environment cluster addresses plant diseases, insect pests,
research. Major universities with agriculture faculties include weeds and rodents, dryland and water-efficient agriculture, soil
Tokyo University of Agriculture and Technology, Tohoku Uni- sciences, and plant nutrition, and participating grains institutes
versity, Kyoto University, Kyushu University, Hokkaido Uni- include the Institute of Plant Protection, Institute of Environ-
versity, Kagoshima University, Yamaguchi University, Tottori ment and Sustainable Development in Agriculture, Biotech-
University, and Nagoya University. Several universities carry nology Research Institute, the Farmland Irrigation Research
out research in grains processing. Institute, China National Rice Research Institute, and the
The Korean Rural Development Administration of the Min- Agro-Environmental Protection Institute. Grains research is
istry of Agriculture, Food and Rural Affairs has a series of also undertaken within the Agricultural Mechanization and
national research institutes, including the National Institute Engineering; Agro-Product Quality, Safety, and Processing;
of Crops Science (www.nics.go.kr/english) with a focus on and Agricultural Information and Economics clusters. The
rice, wheat, maize, soybean, and maize breeding, agronomy Agricultural Science and Technology Innovation Program
and applied biotechnology, and postharvest technology; (ASTIP), launched in early 2013, funds four crosscutting objec-
National Academy of Agricultural Science in Suwon (www. tives supporting long-term and interdisciplinary research,
naas.go.kr/English/naas_main.asp) with relevant work on capacity building, expanding research support facilities and
crop protection and agro-ecology; National Institute of Agri- infrastructure, and fostering international cooperation over
cultural Biotechnology (www.niab.go.kr); and several provin- the next 13 years.
cial agricultural and extension institutes. Grains research is The Chinese Academy of Sciences (www.english.cas.cn),
strongly rice-focussed, but with an increasing environmental under the Ministry of Science and Technology, also carries out
protection and biotechnology emphasis. Of the ten Flagship some research relating to grains, at the Institute of Botany and
Korean National Universities, eight have agriculture colleagues Institute of Genetics and Developmental Biology (Beijing; basic
that undertake grains research to varying extents, including plant biology and biotechnology), Institute of Zoology (Beijing;
Chonbuk, Chonnam, Chungbuk, Chungnam, Gyeongsang, pest management), Institute for Soil Research (Nanjing),
Kangwon, Kyungpook, and Seoul National Universities. National Centre for Genetic Research (Shanghai; rice genomics),
China has the largest national agricultural research systems Chengdu Institute of Biology (cereal breeding), Institute of Soil
(NARS) in the world and also the most geographically and and Water Conservation (Yangling; agronomy), NW Institute of
administratively decentralized. The government has placed a Plateau Biology (Qinghai; crop assessment), and Shijiazhuang
high priority on developing research capacity and infrastruc- Institute of Agricultural Modernization (Hebei; farming systems
ture over the last decade, and private sector expenditure in for the North China plain) among others.
research has also increased. Because of the large range of pro- Several provincial academies of agricultural science have
duction environments, research on both temperate and tropi- strengths in grains research including Jiangsu, Sichuan, Guang-
cal grains is important, and some institutes undertake dong, Fujian, Henan (for wheat), Shandong, Liaoning, and
pioneering grains research, for example, in the development Gansu. The State Administration of Grain (www.chinagrain.
of high-yielding rice varieties, rice functional genomics, and gov.cn/english/Scientific%20Research.html) also carries out
transgenic crops. In recent years, the Chinese government has grain storage research, including through the Grain Storage
been making major investments in R&D physical infrastructure Research Institute in Chengdu. In recent years, much of the
and equipment, and there has been a move to retain or attract increase in research capacity has come from the universities.
back eminent scientists from the Chinese diaspora. There is an Each province has at least one agricultural university, but the
increasing shift in emphasis in China from research aimed at three strongest universities in agricultural science are Nanjing
increasing grain production to increasing quality and ensuring Agricultural University, Zhejiang University, and Huazhong
that production is environmentally sustainable, especially in Agricultural University. Other agricultural universities carrying
view of the increasing pressure on water resources for agricul- out grains research are South China Agricultural University
ture in China. (Guangzhou), Shenyang Agricultural University, Southwest
Grains research is carried out in five main types of organi- Agricultural University (Chongqing), and Northwest A&F Uni-
zations. The Chinese Academy of Agricultural Sciences (CAAS, versity (Yangling). Further information on grain production
www.caas.net.cn/en) under the Ministry of Agriculture has the and consumption in China is discussed elsewhere in this
largest number of grains research projects. There are 32 Encyclopedia.
research institutes (employing over 5000 research staff directly
affiliated with CAAS; another nine are hosted jointly with
universities and provincial governments), with a growing
emphasis on environmental safety and sustainability, rather
South Asia
than just higher yields. CAAS has recently consolidated its India also has one of the largest NARS in the world and a
vast array of research projects into eight disciplinary clusters. traditional strength in crops research, particularly grain
SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organisations of the World 447
sciences, given the ongoing national focus on food security. Pakistan Agricultural University at Faisalabad. Sindh
Most grains research is done within the Indian Council of Agriculture University, the University of Arid Agriculture
Agricultural Research (ICAR, www.icar.org.in) and state (Rawalpindi), and the Northwest Frontier Agricultural Univer-
agricultural universities. Of the 45 ICAR Central Research Insti- sity (Peshawar) are all smaller, but with a similar range of
tutes, most of the grains research institutes are within the Crop activities. The Ministry of Science and Technology manages
Science Division. Major thrusts include development of grain the Pakistan Council of Scientific and Industrial Research,
varieties resistant/tolerant to a range of abiotic and biotic which carries out grains postharvest research. There is also a
stresses, hybrid technology, application of molecular tech- separate National Institute for Biotechnology and Genetic Engi-
niques, plant genetic resources conservation, and farming sys- neering (Faisalabad) as well as the NARC Institute of Agricul-
tems research. The main ICAR institutes carrying out grains tural Biotechnology and Genetic Resources. State departments
research include of agriculture in Balochistan, Punjab in particular, carry out
some grains research.
the Indian Agricultural Research Institute (emphasis on
Elsewhere in South Asia, the Bangladesh Agricultural
more strategic research);
Research Council has a central Bangladesh Agricultural
the Indian Institute of Pulses Research;
Research Institute (which carries out significant amounts of
the National Bureau of Plant Genetic Resources;
grains research) together with satellite institutes covering soils
the National Research Centres for Groundnut, Rapeseed
and rice. The largest university conducting grains research is
Mustard, Sorghum and Soybean;
Bangladesh Agricultural University.
the National Centre for Integrated Pest Management;
the National Research Centre on Plant Biotechnology;
the Central Institute of Post-Harvest Engineering and Southeast Asia
Technology;
The major Indonesian government agricultural research and
the Central Soil Salinity Research Institute;
development agency is the Indonesian Agency for Agricultural
the Central Arid Zone Research Institute;
Research and Development (IAARD, www.en.litbang.deptan.
the Central Research Institute for Dryland Agriculture;
go.id). Research institutes working on grains (unlike planta-
the Indian Institute of Soil Science;
tion crops) are publicly funded and those focussing on grains
Directorates of Wheat Research, Sorghum Research, Oil-
include the Centers for Food Crops R&D (Bogor, W Java), Rice
seeds Research, Soybean Research, Rice Research, and
Research (Subang, W Java), Agricultural Engineering R&D
Weed Science Research.
(Serpong, W Java), and Postharvest R&D (Bogor, W Java).
Several smaller project directorates covering biological control, Typically smaller research institutes work on legume and
oilseeds, rice, wheat, and maize. All India Coordinated tuber crops (Malang, E Java), cereals (particularly maize)
Research Projects attempt to link institutes as a mechanism to (Makassar, S Sulawesi), and soils (Bogor, W Java). Adaptive
build nationwide coordinated interdisciplinary research net- research is carried out by Assessment Institutes for Agricultural
works increasing the level of interinstitutional collaboration Technology (under IAARD, but located in each province).
is currently a high priority of the Indian government. Major agricultural universities include Bogor Agricultural Uni-
The second significant resource is the 55 state agricultural versity (IPB), Gadjah Mada University, Hasanuddin University,
universities (SAUs). Some of the SAUs with strong grains University of Mataram, Sam Ratulangi University, Sebelas
research programs include Acharya NG Ranga AU (Andhra Maret University, Andalas University, Haluoleo University,
Pradesh), CCS Haryana AU, GB Pant AU (Uttar Pradesh), Brawijaya University, and Padjadjaran University.
Punjab AU, and Tamil Nadu AU. The Council for Scientific Under the Philippine Department of Agriculture (DA),
and Industrial Research (CSIR) has the Central Food Techno- there is a national rice research center (PhilRice in Munoz,
logical Research Institute (CFTRI, Mysore, www.cftri.com) Luzon, www.philrice.gov.ph), addressing both research and
has a department that researches cereal (especially wheat) technology dissemination, and the Bureaus of Plant Industry,
milling, baking, and biochemistry. A number of private sector Soil and Water Management (www.bswm.gov.ph), and Post-
companies undertake research into crop breeding and plant harvest Research and Extension (which includes an emphasis
protection. Further information on grain production and on grain storage research). The Food and Nutrition Research
consumption in India is discussed elsewhere in this Institute (www.fnri.dost.ph) and the research planning and
Encyclopedia. funding body, the Philippine Council for Agriculture, Aquatic
Pakistan also has one of the largest numbers of grains and Natural Resources Research and Development (www.
researchers in the developing world. The main institute of the pcaarrd.dost.gov.ph), sit under the Department of Science
Pakistan Agricultural Research Council (PARC, www.parc.gov. and Technology. There are also Regional Integrated Agricul-
pk), the National Agricultural Research Centre in Islamabad, tural Research Centers there is one organization in each of
covers soil fertility; wheat, rice, sorghum, and millet; farm the 15 regions, but some have 23 research stations and several
machinery; water resources; agricultural biotechnology; verte- focus on grain crops. The universities have a far greater share of
brate pests; crop diseases; crop production; land resources; research capacity than DA. The strongest university in agricul-
plant genetic resources; technology transfer; and seed sciences. ture including grains research is the University of Philippines at
There are a separate tropical agricultural research center at Los Banos, which has specialized institutes of plant breeding
Karachi and four arid zone research centers in different regions and of farming systems research. These include Benguet
of Pakistan. There are several agricultural universities in State University, Central Luzon State University, Visayas State
Pakistan that carry out grains research, the largest being the University College of Agriculture, Don Mariano Marcos State
448 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organisations of the World
University, Central Mindanao University, and the University of Maize Research Institute, and Mekong Rice Research Institute.
Southern Mindanao. There are also several others with some The Department of Plant Protection of MARD is also becoming
strengths in agriculture (Mindanao State University, Bicol Uni- an important applied research organization. The main univer-
versity, and Central Visayas State College). Some of the univer- sities with research interests in grain science include Hanoi
sities have regional research and development centers and also Agricultural University, Hanoi University of Science (also
have quite good extension/outreach systems. While significant known as Hanoi College of Sciences), Nong Lam University,
private sector research enterprises are present in horticulture, Hue University of Agriculture and Forestry, Thai Nguyen Uni-
grains research remains in the public domain. versity, and Can Tho University. Further information on grain
The Thailand Department of Agriculture (DOA, www.doa. production and consumption in Southeast Asia is discussed
go.th/en) is responsible for crops and farm mechanization elsewhere in this Encyclopedia.
research, while the Department of Agricultural Extension
(DOAE) is responsible for transfer of technology to farmers.
While there are experiment stations in every region of the
Central and South America
country, most research is carried out in Bangkok. DOA has
programs covering agricultural chemistry, agricultural engi- The major grains R&D capacity in Central America is within
neering, plant pathology and microbiology, entomology and Mexico. Initially, it was focussed in several of the Instituto
zoology, soil science, botany, and weed science, crop protec- Nacional de Investigaciones Forestales, Agricolas y Pecuarias
tion, and specialist rice and field crops institutes (there is (INIFAP, www.inifap.gob.mx) crops institutes based on differ-
reasonable research capacity in maize and legumes research). ent agroecological zones (www.inifap.gob.mx), but it is now
The Land Development Department conducts research in rela- spread across a range of universities and other government
tion to land improvement and soil and water conservation. agencies. Grains food research and biotechnology is carried
Grains postharvest technology is carried out within the out in the Consejo Nacional de Ciencia y Tecnologia
Thailand Institute of Scientific and Technological Research (CONACYT, www.ciad.mx). Grains research is carried out in
(TISTR, www.tistr.or.th/tistr_en) in Bangkok. One of the the Universidad Autonoma Chapingo, the Instituto Polit-
three research centers with National Science and Technology ecnico Nacional, and over a dozen regional universities.
Development Agency is Genetic Engineering and Biotechnol- In South America, Brazil has a well-developed NARS, with
ogy (BIOTEC, www.biotec.or.th/en), which carries out some more agricultural researchers than any other country in Latin
rice biotechnology work. Kasetsart University in Bangkok, America. Many centers fall under EMBRAPA (Empresa Brasile-
Khon Kaen University, Chiang Mai University, Prince of ira de Pesquisa Agropecuaria, www.embrapa.br), a specialized
Songkla University (Hat Yai), and King Mongkuts Institute of science agency of the Agriculture Ministry. Brazil has the largest
Technology in Bangkok are all involved in grains research, the NARS of any Central or South American country, followed by
latter having a special focus on grains postharvest technology. Mexico; Brazil also has an advanced university system. In addi-
In Malaysia, agricultural R&D spending as a percentage of tion, 16 of Brazils 26 states operate agricultural research insti-
production value (research intensity) remains comparatively tutes, with Sao Paulo the largest. There are separate EMBRAPA
high. The Rice and Industrial Crops Research Centre of the institutes relating to genetic resources and biotechnology
Malaysian Agricultural Research and Development Institute (CENARGEN), upland rice and beans (CNPAF), corn and sor-
(MARDI, www.mardi.gov.my) researches rice, maize, and ghum biotechnology (CNPMS), soils technology transfer
groundnuts as well as several nongrain crops. The most rele- (CNPS), soybean and sunflower (CNPSO), wheat, barley,
vant activity is development of high-quality rice varieties with oats, soybeans, and biotechnology (CNPT), temperate climate
disease resistance and accompanying agronomy. The food agricultural research (CPACT, irrigated rice), and food technol-
technology research center does work on the development of ogy (CTAA), as well as at least 12 regional centers. There are
rice-based snacks and other food products. The major univer- also nonprofit centers Foundation Center for Wheat Experi-
sities carrying out grains science research is the Universiti Putra mentation and Research (FUNDACEP, www.fundacep.com.
Malaysia and Universiti Kebangsaan Malaysia. br), the Central Agricultural Cooperative of Technology Devel-
Most grains research in Vietnam is conducted under opment and Economics (COODETEC; soy, maize, and wheat),
the Ministry of Agriculture and Rural Development (MARD, and the Rice Institute of the Rio Grande do Sul (IRGA). Several
www.agroviet.gov.vn). Other relevant ministries are education universities carry out grains research the federal universities
and training (university sector), science and technology (www. of Goias (www.agro.ufg.br), Lavras, Paraiba, Pelotas (www.
most.gov.vn), which has some excellent crop biotechnology faem.upfel.edu.br), Rio Grande do Sul, Rural do Rio de Janeiro
and plant protection research facilities, and the Ministry of (www.la.ufrrj.br), Santa Catarina (www.cav.udesc.br), Sao
Industry (which has an Oil Plant Research Institute with Carlos (www2.ufscar.br), Santa Maria, Vicosa, and Brasilia as
the mandate for peanut, soybean, sunflower, and sesame). well as several state universities (Santa Catarina State, Paulista,
While most institutes are located in either Hanoi or Ho Chi Campinas, Sul de Santa, and Sao Paulo). Some Brazilian com-
Minh City, many maintain research centers in more remote panies, such as Agroceres, DuraFlora, Mogiana, and Braskalb,
locations. The key MARD institutes in grain science are the carry out breeding and agronomy research to support seed
Food Crops Research Institute, National Institute for Plant distribution businesses.
Protection, National Institute for Soils and Fertilizers, Institute The Argentine Government Instituto Nacional de Tec-
of Agricultural Engineering and Post Harvest Technology, Insti- nologia Agropecuaria (INTA, www.inta.gov.ar) is under the
tute of Agricultural Science in the South, Vietnam Agricultural Agriculture Ministry, and capacity has increased significantly
Science Institute, Agricultural Genetics Institute, National over the last decade due to significant increase in government
SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organisations of the World 449
investment. There are also a large number of programs and (www.sasa.ukzn.ac.za), the Free State (www.ufs.ac.za), and
centers relevant to grains research under CONICET (Consejo Stellenbosch (www.acadmic.sun.ac.za/agric/). In the 1990s,
Nacional de Investigaciones Cientificas y Tecnicas, www.con- there has been a significant shift in research emphasis in
icet.com.ar), which is under the Culture and Education Minis- South Africa to accommodate the needs of the wider popula-
try. This includes a number of crop breeding, agronomy, and tion, rather than just white commercial agriculture, to include
natural resource (CADIC and CEFYBO), plant protection emerging farmers and subsistence farmers in former home-
(CEAVE and IFEVA), postharvest technology (CEFOBI and lands. Private companies such as Monsanto and Pannar carry
CRILAR), and plant physiology and molecular biology out seed and fertilizer research.
(INFIVE and INGEBI) centers. Several national universities In Zimbabwe, most crops research is carried out by the
have agronomy science faculties Buenos Aires, Buenos Aires Department of Research and Specialist Services in the Minis-
Province, Catamarca, Comahue, Cordoba, Cuyo, Entre Rios, try of Lands (www.drss.gov.zw) and Agriculture, Mechani-
Jujuy, La Pampa, La Plata, Litoral, Lomas de Zamora, Mar del zation and Irrigation Development, with the University of
Plata, Noredeste, Rio Cuarto, Rosario, Santiago del Estero, de Zimbabwe (www.uz,az.zw) maintaining reasonable strength.
Sur, de Salta, Tucuman, and de Formosa. The Kenya Agricultural Research Institute (KARI, www.kari.
The Chilean Ministry of Agriculture has an Instituto de org) is comparatively strong in crops and livestock and the
Investigaciones Agropecuarias (INIA, www.inia.cl), which Kenyan government agricultural research system one of the
undertakes some grains research, although overall agricultural strongest in Africa. In the grains area, mandate crops include
research is much more limited than in Brazil or Argentina. There cereals (wheat, maize, sorghum, millets, and rice) and grain
are also agronomy departments at several universities Austral legumes with efforts ranging from plant breeding for adapta-
University (www.agrarias.uach.cl), the University of Conception tion to different agroecological zones and stress resistances,
(www.chillan.udec.cl), the University of La Frontera, the agronomy, postharvest technology, and cooking quality.
University of La Serena (www.userena.cl), the University of Almost all Kenyan grain production is rain-fed. The breeding
Magallanes, the University of Talca, the University of Tarapaca and postharvest research is backed up by a soil and water
(www.uta.cl), Universidad de Chile, and Universidad Ibero- management research program. The Kenya Industrial R&D
americana de Ciencias as well as the Catholic universities Institute (www.kirdi.go.ke) also carries out some postharvest
(Temuco (www.uctemuco.cl), Valparaiso (www.ucv.cl), and research. Some grains research is carried out at various uni-
Maule). In Colombia, the main government crops research versities (Jomo Kenyatta University of Agriculture and Tech-
agency is CORPOICA (www.corpoica.gov.co); but much nology (www.jkuat.ac.ke), Egerton University (www.egerton.
more research is now carried out through the producer associa- ac.ke), Moi University (www.mu.ac.ke), the University of
tion, Asociacion Nacional de Acuicultores de Colombia Eastern Africa (www.uaeb.ac.ke), and the University of Nai-
(ACUANAL); there are research groups working on rice. Apart robi (www.ounbi.ac.ke). The Ethiopian Institute of
from the campuses of the national university in Bogota, other Agricultural Research (www.eiar.gov.et) has national and
universities with grains agronomy faculties include Cordoba regional centers emphasizing maize, plant protection, soils,
(www.unicordoba.edu.co) and Tolima (www.ut.edu.co). The and agricultural mechanization. Crops research is carried
main research agency in the Ecuador agriculture ministry is out at Hawassa University (www.hu.edu.et), Haramaya
INIAP (Instituto Nacional Autonomo de Investigaciones Agro- University (www.haramana.edu.et), Jimma University (www.
pecuarias); universities in most regional centers have small ju.edu.et), and Mekelle University (www.mu.edu.et). In
agronomy departments. In Uruguay, the government Instituto Uganda, most grains research capacity is within the main
Nacional de Investigacion Agropecuaria (INIA, www.inia.org. government National Agricultural Research Organization
uy) is the largest crops research institute. Further information (NARO, www.naro.go.ug) and at Makerere University (www.
on grain production and consumption in South America is agric.mak.ac.ug).
discussed elsewhere in this Encyclopedia. In Nigeria, the Ministry of Agriculture through the Agricul-
tural Research Council of Nigeria (www.arcnigeria.org) has a
series of research institutes National Cereals Research Insti-
tute, Lake Chad Research Institute (www.icresmaid.org), and
Africa and the Middle East
the Institute for Agricultural Research (www.abu.edu.ng). Post-
South Africa has the best-developed NARS of the African harvest technology research is carried out under the Ministry of
countries, although its capacity has declined over the last Industry at the National Stored Products Research Institute
decade. South Africa maintains reasonable research capacity (www.nspiilofin.com), while the Ministry of Science and Tech-
in maize (the main grain crop), sorghum, and some experi- nology has established a National Centre for Genetic Resources
ence in wheat and a little in barley. The Agricultural Research and Biotechnology (www.nacgrab.gov.ng). Universities with
Council (www.arc.agric.za) is the main agricultural research relevant research programs include the University of Ibadan
structure and has a series of research institutes, several carry- (www.agric.ui.edu.ng), University of Nigeria (www.unn.edu.
ing out grains research particularly the Grain Crops Institute ng), Bayero University, Ahmadu Bello University (www.abu.
and the Small Grains Institute, with some other institutes edu.ng), and Obafemi Awolowo University (www.oauife.edu.
carrying out supporting natural resource management ng). In Zambia, the Zambia Agricultural Research Institute
research. The nine provincial departments also carry out (www.zari.gov.zm), Postharvest Food Processing and Nutri-
selection and agronomy research to varying degrees. There tion Institute (www.nsirir.org.zm), and the University of Zam-
are some well-resourced grains research programs at the Uni- bia (www.unza.zm) are the main grains research providers. In
versities of Pretoria (www.web.ip.ac.za), KwaZulu-Natal Tanzania, it is the Directorate of Research and Development of
450 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organisations of the World
the Ministry of Agriculture and Food Security (www.agricul- How do the priorities of grains research for the Indian state
ture.go.tz) along with the Sokoine University of Agriculture agricultural universities differ when they are based in dif-
(www.suanet.az.tz); in Botswana, the Department of Agricul- ferent geographic zones of India?
tural Research (www.dar.gov.bw), while in Ghana, the Crops Being a small country, why doesnt New Zealand simply
Research Institute of the Council for Scientific and Industrial adopt the grain varieties and agronomy research findings of
Research (www.cropsresearch.org), along with the University its larger neighbor, Australia?
of Ghana (www.ug.edu.gh) and the Kwame Nkrumah Univer-
sity of Science and Technology (www.knust.edu.gh), are the
main crops research providers. Further information on grain
production and consumption in Africa is discussed elsewhere See also: Agronomy of Grain Growing: Implication of Climate
in this Encyclopedia. Changes; Wheat: Agronomy; Breeding of Grains: Barley: Genetics
The Israeli NARS is reasonably well resourced. Within the and Breeding; Canola/Rapeseed: Genetics and Breeding; Maize:
Agricultural Research Organization (www.agri.gov.il), the Breeding; Rice: Breeding; Soybean: Germplasm, Breeding, and
Institute for Plant Sciences has programs on cereal crop agron- Genetics; Wheat Breeding: Exploiting and Fixing Genetic Variation by
omy to overcome drought, disease, and heat stress. There are Selection and Evaluation; Food Grains and the Consumer: Cultural
separate institutes for plant sciences, soil water and Differences in Processing and Consumption; Grains Around the
environmental sciences, postharvest and food sciences, and World: Grain Production and Consumption: China and South-East
agricultural engineering, which also carry out grains-related Asia; Grain Production and Consumption in Oceania: Australia and
research. Universities carrying out grains research include Pacific Countries; Grain Production and Consumption: Overview; Grain
Technicon (Israel Institute of Technology (Haifa)) and the Production and Consumption: South America; Grain Production and
Ben-Gurion University of Negev. Consumption: Africa; Production and Consumption of Grains: India;
Egypt also has a large government agricultural research Scientific Organizations Related to Grains: Scientific Societies
system, within the Agricultural Research Center (ARC, www. Associated with Grain Science; The Basics: The Grain Crops: An
arc.sci.eg), within the Ministry of Agriculture and Land Recla- Overview; The Cereal Grains: An Overview of the Family of Cereal
mation. In ARC, there are separate programs for plant protec- Grains Prominent in World Agriculture; Wheat-Based Foods:
tion, plant pathology, food technology (bread and pasta Cereals: Overview of Uses: Accent on Wheat Grain.
research), agricultural genetic engineering, and farming sys-
tems research relating to wheat, barley, maize, rice, sorghum,
legumes, and oil crop production. Universities with agriculture
faculties include Alexandria, Cairo, Ain Shams, Al-Azhar, Suez Further Reading
Canal, Zagazig, Assiut, Mansoura, Minia, Menoufia, Tanta, and
South Valley. Further information on grains research in other Beintema NM and Stads G-J (2010) Public agricultural R&D investments and capacities
West Asian and North African countries is discussed elsewhere in developing countries: Recent evidence for 2000 and beyond. Background note for
the Global Conference on Agricultural Research for Development, Montpellier,
in this Encyclopedia.
France, March 2831.
Beintema N and Stads G-J (2011) African Agricultural R&D in the New Millennium:
Progress for Some, Challenges for Many. Washington, DC: International Food
Exercises for Revision Policy Research Institute.
Cassas J, Sohl M, and Hafaz H (1999) The National Agricultural Research Systems in
the West Asia and North Africa Region. Aleppo, Syria: ICARDA, p. 278.
What is the largest national agricultural research system in Corke H and Cai YZ (2016) Grain production and consumption: China and South East
the world? Asia. In: Corke H, Wrigley C, Faubion J, and Seetharaman K (eds.) Encyclopedia of
In what regions of the world is grains research capacity Food Grains, 2nd edn., vol. 1, pp. 374382. Elsevier Ltd.
thought to have decreased over the last decade? Flaherty K, Stads G-J, and Srinivasacharyulu A (2013) Benchmarking Agricultural
Name three countries where the involvement of the private Research Indicators Across Asia-Pacific. Washington, DC: International Food Policy
Research Institute.
sector in grains breeding has increased significantly over the Khatkar BS, Chaudhary N, and Dangi P (2016) Production and Consumption of Grains:
last decade. India. In: Corke H, Wrigley C, Faubion J, and Seetharaman K (eds.) Encyclopedia of
What role does the national Australian government have in Food Grains, 2nd edn., vol. 1, pp. 367373. Elsevier Ltd.
grains R&D delivery? Pardey PG and Alston JM (2006) Agricultural R&D in the Developing World: Too Little,
Too Late? Washington, DC: International Food Policy Research Institute.
What are the two major organizations or systems undertak- Roseboom J, Cremers M, and Lauckner B (2001) Agricultural R&D in the Caribbean: An
ing grains research in India? Institutional and Statistical Profile. The Hague: International Service for National
Agricultural Research, ISNAR Research Report No. 19.
Taylor JRN (2016) Grain production and consumption, Africa. In: Corke H, Wrigley C,
Exercises for Readers to Explore the Topic Further Faubion J, and Seetharaman K (eds.) Encyclopedia of Food Grains, 2nd edn., vol. 1,
pp. 359366. Elsevier Ltd.
Wrigley CW and French RJ (2016) Grain production and consumption in Oceania:
How do agricultural research in Brazil contributed to the Australia and Pacific Countries. In: Corke H, Wrigley C, Faubion J, and Seetharaman
export success of its grains industries? K (eds.) Encyclopedia of Food Grains, 2nd edn., vol. 1, pp. 409419. Elsevier Ltd.
Research Organizations of the World: Global Trends and the Commercial Sector
JH Skerritt, University of Queensland, St. Lucia, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights development) aimed to support their flour milling and bakery
product businesses (including breakfast cereals), but some
Greater emphasis on demand-driven rather than science- carry out more basic research and their staff contribute actively
driven research to research conferences and the scientific literature. Major
Increasing emphasis on topics such as production sustain- examples in the United States include ConAgra Foods in
ability, croplivestock systems, systems modeling, and the Omaha, NE (www.conagrafoods.com), General Mills in Min-
use of techniques from molecular and information sciences neapolis, MN (www.generalmills.com), Kellogg Company in
but also a return to treatment of research on grain produc- Battle Creek, MI (www.kelloggcompany.com), and Nabisco,
tivity and yield. New Jersey (www.snackworks.com). Nabisco Research carries
Specialization into particular areas common as well as out fundamental studies on dough rheology and physical
clearer identification by funders and institutes about how properties. Riceland Foods (www.riceland.com, Stuttgart, AR,
near to market the United States) and Riviana Foods (www.riviana.com,
Greater collaboration between disciplines and between Houston, TX, the United States) are two large manufacturers
research institutes as well as increased private sector has of rice- and soy-based food ingredients and undertake applied
greater involvement in agricultural R&D research to support their markets. The National Starch and
Major commercial groups involved in grains and grain Chemical Company, Bridgewater, NJ (www.nationalstarch.
product research com), focuses on cereal starch-based polymers and supports a
significant research program including work on new adhesives,
specialty food starches, and biodegradable packaging mate-
Learning Objectives rials. Cargill has wheat flour milling and maize wet milling
divisions and provides grains-based feeds for livestock.
To obtain a better understanding of global trends in grains Through Cargill Health and Food Technology (www.cargill-
research and grains research undertaken by the commercial foods.com), it focuses on nutraceuticals research, especially
(biotechnology, seed, crop protection, milling and baking, on developing products derived from soy.
brewing, and cereal foods) sector. In Japan, two large milling companies, Nisshin Flour Mill-
ing (www.nisshin.com/english/group) and Nippon Flour
(www.nippn.com), have played an important role over the
last few decades in increasing the consumption of wheat in
Grains Research in the Commercial Sector
Japan. They also carry out a significant amount of research. In
the United Kingdom, Rank Hovis (www.rankhovis.co.uk),
The private sector has greater involvement in agricultural R&D,
ADM Milling (www.adm.com/en-US/MillingUK/Pages/
especially in linkages between seed business and biotechnol-
default.aspx), Associated British Foods, Allied Bakeries, and
ogy/life science concerns. Pardey and Beintema had calculated
British Bakeries are among the largest milling and baking
that since the mid-1990s, in developed countries, expenditures
companies, and they conduct some in-house research and
from the private sector in agricultural research have exceeded
technology development. Barilla (www.barillagroup.com/cor-
public expenditure.
porate/en/home/lavora-con-noi/organization-and-job-fami-
While bakery and grains processing companies have always
lies/research-development-quality.html) has a large cereal
engaged in research to support new product development, a
products research center in Italy, while in Australia, Goodman
particular focus of their research over the last decade has been
Fielder (www.goodmanfielder.com.au) and GWF (www.geor-
in the development of grain food products that promote health
gewestonfoods.com.au) have active technology development
and well-being. Despite the health benefits, there are chal-
programs. Other companies within the oilseed crushing and
lenges to increasing consumption of whole grain foods includ-
animal feeds industry also have applied research programs.
ing taste preferences and convenience of these foods. In
addition, there is a growing tendency for increasing numbers
of people to avoid cereals in their diets because of perceived
health issues, even though the documented prevalence of Baking Industry Associations
celiac disease and cereal allergy is comparatively low. Develop- Industry associations are also an important provider of
ment and marketing of gluten-free foods has thus become a research and technical services in the grains industry. In the
significant business opportunity. United Kingdom, Campden BRI (www.campdenbri.co.uk),
Chipping Campden (Gloucestershire) and Nuffield (Surrey),
is a membership-based organization that provides research
Grains Processing Companies
and consultancy services for cereal-processing companies.
Most of the research carried out by these companies is con- Leatherhead Food Research (www.leatherheadfood.com)
cerned with practical applications (e.g., new product started as a UK food-industry association but has evolved
Table 1 Major international life sciences and seed companies carrying out grains research
Monsanto Company Maize, soybean breeding, hybrids, herbicide, insect and wilt resistance, agrochemicals, seeds, www.monsanto.com
St. Louis, MO, USA and genomics
Agracetus (part of Crop plant transformation, therapeutic protein expression in plants www.agracetus.com
Monsanto)
Middleton, WI, USA
ADM (Archer Daniels Food ingredients; oilseeds; soy-based foods; feed ingredients; bakery science; nutraceuticals; www.adm.com
Midland) biofuels; extrusion research
Decatur, IL
Pioneer Hi-Bred Seeds, grain additives www.pioneer.com
(a Du Pont Company) Crop management, precision farming
Des Moines, IA, USA Grain seeds, especially hybrid maize (also millet, rice, sorghum, wheat)
BASF Crop protection products www.basf.de/en;
Limburgerhof, Germany Transgenic crops with stress tolerance, high oil content, and pathogen tolerance www.basf.com
Raleigh, NC, USA
Bayer Crop Science Crop protection www.
Akeno and Yuki, Japan bayercropscience.
Frankfurt, Germany com
Kansas City, MO, USA
Lyon, France
Syngenta Crop protection, seeds, and crop genetics www.syngenta.com
Jealotts Hill, UK Genomics, biotic stress tolerance, fungal diseases, insect and nematode control, marker-
Research Triangle Park, assisted breeding, maize, rice and soybean breeding, and transgenics
USA
Saint-Sauveur, France
Uberlandia, Brazil
Beijing, China
Dow Agrosciences Plant genetics and biotechnology; agrochemicals; seeds (hybrid and GM), traits (sunflower, www.dowagro.com
Indianapolis, IN, USA corn, and soybean) and oils; crop protection
Mycogen Breeding of sorghum, hybrid corn, sunflower, soybean, and canola www.mycogen.com
Indianapolis, IN, USA
Cargill Cereals for animal feed, biofuels; novel traits; oilseed breeding; biotechnology www.cargill.com
Ft Collins, CO, USA
Elk River MN, USA
Minneapolis, MN, USA
Vilvoorde, Belgium
KWS UK Wheat, barley, maize, and canola varieties www.kws-uk.com
Thriplow, UK
Limagrain Cereal breeding; seeds; plant genome analysis; transgenesis; biotechnology; maize, wheat, www.limagrain.com
Auvergne, France rapeseed, sunflower, soybean, barley, peas; baking technology
Biogemma Maize, wheat, rapeseed, sunflower breeding plant genome analysis; drought tolerance; www.biogemma.fr
Clermont Ferrand and disease resistance
Mondonville, France
planning exercises have been in favor in a number of commer- Changes in Research Subject Matter
cial and government organizations across the globe. It is unclear
Grains agronomy programs have had to develop a stronger
to what extent individual creativity has been affected by these
research emphasis on sustainability of production in recent
trends certainly, there may be less scope for curiosity-driven
years, responding to a shift in community values as well as serious
grains research. The greater focus on markets by farmers and
scientific concerns about the long-term impact of some grain
producing for a particular market is reflected in research.
farming practices. Concerns about effects of irrigation, salinity,
An overarching trend has been the greater level of cooper-
acidification (especially with certain legume crops), loss of soil,
ation between research organizations; funding networks such
and of soil structure have led to an increase in research on reduced
as those established by the European Commission and Austra-
tillage, water conservation, and weed control and maintenance of
lian Government CRC system have facilitated this trend, as
soil fertility. Molecular breeding approaches have been imple-
well as organizational changes in major government depart-
mented through the development and release of new varieties a
ments such as US Department of Agriculture. However, it is
move from the laboratory to the field over the last decade.
probably true that US grains research is less well coordinated at
Impacts on climate variability and change and adaptation
a national level although locally, the close interactions
to variable climates have proved a significant focus for grains
between a number of USDA-ARS facilities and land grant uni-
research over the last decade. However, the shift in research
versities are very productive. The increased importance of net-
resources from productivity to other areas has been controver-
works in many areas of science is a response to the increased
sial Alston et al. demonstrated that a shift in research invest-
cost of carrying out research plus an increasing realization that
ment in productivity or yield-related outcomes has coincided
problems are multidimensional and are best addressed
with a plateauing in global yields for major cereals. Indeed, the
through multidisciplinary approaches. There has also been
major factor underpinning sustainable agriculture has been the
pressure from governments and funding bodies for research
ability to feed the worlds increasing population through
providers to network more, in part to avoid duplication of
enhanced yields, and with the exception of a few communities,
research and also to ensure that particular subprojects are
grains, especially cereals, are the central element of almost
carried out by the most qualified group. In the developing
diets. Since the 1940s, there has not been a significant increase
world, development banks such as the World Bank and Asian
in the area of land available for agriculture. However, it is true
Development Bank have funded reform programs that encour-
that with the expansion of irrigation systems and with new
age rationalization in the number of research providers as well
varieties and increased fertilizer use in the middle of the twen-
as greater coordination. In addition, since the spike in global
tieth century, there was an expansion of grain-growing areas.
grains prices in 200708, development agencies and banks
Since the 1980s, the expansion of irrigated areas has largely
have had a resurgence of interest in funding agriculture includ-
ceased and so production increases must come from increases
ing grains-related research.
in yield. In suitable ecologies, there has been a shift toward
Scientist-to-scientist collaboration across institutional and
more frequent harvests, especially in irrigated areas that may
national boundaries has been greatly facilitated by the commu-
accommodate three irrigated rice crops per year, as well as also
nications revolution the ability to e-mail (or more recently, use
improved rainfed rotations, for example, between a winter
cloud computing for) large amounts of data and draft manu-
cereal and a legume, rather than leaving the soil fallow over
scripts for immediate receipt and prompt publication online
summer. On the other with the recognition that breeding
has totally changed the possibilities for collaboration compared
targeted toward ideal high yield environments does not pro-
with 20 years ago. Globalization in industry through both
vide varieties that perform well under normal farm conditions,
development of alliances between commercial research pro-
a greater focus on breeding for representative environments
viders in different countries and increased emphasis on grains
and on abiotic stress tolerance has emerged.
breeding for export markets has also become a more impor-
Therefore, the key contributor to sustainability in grain
tant force. Markets are becoming more differentiated so there
production will continue to be productivity; increases are
is an increased emphasis on grain quality with targets for
important for both developing and developed country farmers,
breeding efforts being additional yield and disease resistance
as the terms of trade for farmers continue to become poorer. An
and on research aimed at improving the understanding of the
even greater challenge to developing countries is how North
genetic, chemical, and structural factors underpinning grain
American and European governments shield their grain
quality.
farmers from market realities through subsidies.
An increased focus on impact assessment, better under-
Livestock are becoming a very significant consumer of
standing of the concept of spillover but recognition of the lag
grains worldwide; the trend is especially noticeable in devel-
between successful R&D and adoption claimed to be 1525
oping countries, where meat consumption has increased sig-
years for much public agricultural R&D. While such lag periods
nificantly through the 1980s and 1990s, from a low base.
have been documented for early and mid-twentieth-century
Many of these animals are now grain-fed, so research has had
innovations such as hybrid maize and semidwarf cereals,
to underpin the increased importance of grains for feed rather
respectively, the lag period may persist it is two decades
than food applications. This research is also driven by
since the first genetically engineered grain crops were grown
increased intensification of monogastric animal production,
commercially, and yet for food grains, these still represent only
including in developing countries. In many countries, farmers
a minority of grains produced globally. So grains research
are not solely grain growers or cattle or sheep farmers, but
requires a longer-term view to be taken, both by aid donors
manage a mixed enterprise. There has been a more active
who fund grains R&D in developing countries through
development of systems-based research to address their needs.
commercial investors in advanced economies.
SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Global Trends and the Commercial Sector 455
Another grains research area focused on Africa and South been farmers themselves. Farmer-driven groups now often
and South East Asia is biofortification. There are several inter- have a key role in identifying research priorities as well as
national projects focused on biofortification of staple cereals being a vehicle for dissemination of research results in a
with micronutrients such provitamin A, iron, and zinc. Some number of countries, farmer groups directly manage on-farm
approaches involve the use of genetic engineering technology adaptive research trials on technologies such as rotations, weed
to transfer biosynthetic genes (e.g., for vitamins) into the grain control, and reduced tillage, but some are also adopting a
species, while other programs focus on exploration of natural wider perspective. Realizing that both small and large farms
variation for the desired micronutrient in a grain species or in are often diversified enterprises, grains research has a much
primitive relatives of a target grain species followed by intro- greater emphasis on farming systems than in previous years.
gression of the desired characteristics into high-yielding, high- The influence of farmers is important in fitting grains research
quality locally adapted varieties. into a wider farming systems context. Organizations that use
Consumer trends provide critical direction for many of the farmers levies to fund research such as the Home Grown
commercial milling and baking organizations that conduct Cereals Authority in the United Kingdom or the Australian
research, including on product development. The trend toward Grains Research and Development Corporation have also
cooking less at home and convenience foods is continuing; been instrumental in encouraging greater farmer involvement.
there is increasing popularity of gluten-free diets (whether or Grains science is carried out within formal institutional
not they are indicated on medical grounds) and greater con- structures, most of which have evolved from a range of histor-
sumption of weight loss and diet snacks that have improved ical and political situations, rather than having been developed
taste. In addition, a focus on high-protein/low-fat and low- with a consideration of whether the institutional structures fit
carbohydrate grain products and emergence or niches for some the needs of the twenty-first century. In many state- or
ancient grains such as spelt and nongrains/pseudocereals such provincial-based departments, there remains a close collabora-
as quinoa and buckwheat used in traditionally grain foods tive linkage between cereal breeding and cereal chemistry, and
have also driven industry cereal foods product research and the scientists involved carry out a service function as well as
development. In parallel, whole grain and whole fiber, spe- independent research often creating a tension between the
cialty baked goods, have emerged as an increasingly important time spent on the two, but with the advantage that the research
sector for bakery research. Biofuel research continues to be objectives are informed by genuine industry problems.
important in certain countries as demands grow for example, There are few grains science institutions globally that cover
in 2010/11, 40% of the US corn crop and 14% of soybean oil all of cereals, legumes, and oilseeds. Instead, there may be
were used for biofuels in the United States. research clusters within university agronomy or agriculture
faculties or grains researchers within plant science or crops
research organizations. It is more usual for postharvest tech-
Institutional Trends
nology institutions or grain storage research laboratories to
Another major change in the nature of agricultural research consider various issues relating to a wider range of cereals
since the 1980s, particularly in the United States and Europe, and other grains together, on the basis that the storage and
has been the shift in balance of research to a greatly increased handling of grains as durable commodities have common
role for the private sector. And within government systems, the research challenges. Over the last decade, there has been a
greatly increased resourcing and impact of Chinese, Indian, decrease in postharvest grains research globally, given that
and Brazilian grains researchers and their work are being felt there is a prevailing view that the main priority is now adop-
globally. Different countries also have different types of tion of established technologies.
involvement of universities in grains breeding, although in Some institutes do cover several cereals or grains more
the process of training the next generation of grains breeders, broadly, although usually with a focus on those crops relevant
some universities have developed significant breeding pro- for their country or targeted region. For example, wheat and
grams and have become a mainstay of breeding efforts for a barley are often studied together. Some of the equipment and
country. Several interdependent factors have given rise to techniques are common, while others specialized for exam-
greater commercial involvement in grains breeding, including ple, while Falling number is used as a measure of postharvest
the increased importance of plant breeders rights, develop- sprouting for both wheat and barley, dough rheology testing is
ment and application of biotechnology in grains science, specific to wheat. Specialized grains science departments are
shrinkage of government support for agricultural research, found in a small number of US and European universities, but
and the increased importance of intellectual property protec- their numbers are decreasing. Arguments for this model are the
tion. In some cases, privatization of former public sector plant critical mass of researchers and ability of students to train in
breeding institutes has taken place. aspects from grains breeding to postharvest technology. On the
In some parts of the world, such as Europe, there has been other hand, such departments can lack linkages to emerging or
a long history of significant plant breeding efforts being discovery science and to the broader range of disciplines in
carried out by industry; in others, such as Australia (with the agriculture and science.
exception of hybrid cereals), almost all plant breeding was One of the lessons of modern molecular biology is that
formerly carried out by government and universities. Nowa- techniques such as those in functional genomics are often
days, Australia and several other countries separate the respon- readily transferable between grains, other crops, microorgan-
sibilities for prebreeding and breeding between public and isms, and animals. This aspect has very much been exploited in
private sector organizations, respectively. The other industry recent years as whole genome sequencing becomes faster and
group that has become more involved in grains research has much less expensive, and its use to understand disease
456 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Global Trends and the Commercial Sector
resistance and quality associations becomes more widespread. and the Consumer: Consumer Trends in Grain Consumption;
The high degree of synteny tells us that gene structure and Genetically Modified Grains and the Consumer; Grains and Health;
function are conserved across cereals and the great similarity Genetics of Grains: Development of Genetically Modified Grains;
of their physiological and biochemical processes often makes it Genome Mapping; Genomics of Food Grains; Processing of Grains:
simple to utilize research ideas and techniques between differ- Cereals: Breakfast Cereals; The Basics: The Grain Crops: An
ent grain crops. Overview; The Cereal Grains: An Overview of the Family of Cereal
There are no definitive answers to the issue of an optimal Grains Prominent in World Agriculture; Wheat-Based Foods:
institutional structure for grains research other than to assume Breads; Cookies, Biscuits and Crackers: Formulation, Processing and
that earlier structures need not necessarily be more important Characteristics; Pasta.
than ever for breeders, agronomists, crop protection scientists,
and researchers of processing quality to interact. Finally, insti-
tutional structures can coevolve with the evolution of technol-
ogy. With the growth of networks of researchers collaborating Further Reading
nationally and globally and able now to exchange large data
sets through cloud computing, the establishment of formal Alston JM, Pardey PG, and Smith V (1999) Paying for International Productivity.
institutional structures and a prerequisite for geographic co- Baltimore/London: Johns Hopkins University Press.
Alston JM, Beddow JM, and Pardey PG (2009) Agricultural Research, productivity and
location to foster collaboration has become less important. But
food prices in the long run. Science 325: 12091210.
humans will remain social animals, and it is hard to envisage a Cuesta-Marcos A, Kling JG, Belcher AR, Filichkin T, Fisk SP, Graebner R, et al. (2016)
world where face-to-face collaboration will become irrelevant. Barley: genetics and breeding. In: Corke H, Wrigley C, Faubion J, and
Seetharaman K (eds.) Encyclopedia of Food Grains, 2nd edn., vol. 4, pp. 287295.
Elsevier.
Exercises for Revision DePauw R and OBrien L (2016) Wheat Breeding: Exploiting and Fixing Genetic
Variation by Selection and Evaluation. In: Wrigley C, Seetharaman K, Corke H, and
Faubion J (eds.) Encyclopedia of Food Grains, 2nd edn., vol. 4, pp. 279286.
What are some of the factors leading grains research to be Elsevier Ltd.
more demand-driven? Evenson RE and Gollin D (2003) Assessing the impact of the Green Revolution,
How has greater collaboration between research institutes 19602000. Science 300: 758762.
been encouraged by governments and funding agencies? Henry RJ (2016) Genomics. In: Corke H, Wrigley C, Faubion J, and Seetharaman K
(eds.) Encyclopedia of Food Grains, 2nd edn., vol. 4, pp. 360364. Elsevier Ltd.
What type of research do grains processing companies in Jetter KM and Cassady DL (2006) The availability and cost of healthier food alternatives.
Europe and North America predominantly carry out? American Journal of Preventative Medicine 30: 3844.
What opportunities stimulated agrochemical companies to Jones JM and Sheats DB (2016a) Grains and Health: Misinformation and
get involved in grains breeding? Misconceptions. In: Corke H, Wrigley C, Faubion J, and Seetharaman K (eds.)
What are the main areas of subject matter emphasis for
Encyclopedia of Food Grains, 2nd edn., vol. 2, pp. 2328. Elsevier Ltd.
Jones JM and Sheats DB (2016b) Consumer Trends in Grain Consumption.
commercial grains research and development in India and In: Corke H, Wrigley C, Faubion J, and Seetharaman K (eds.) Encyclopedia of Food
Africa? Grains, 2nd edn., vol. 2, pp. 2934. Elsevier Ltd.
What factors have led to greater commercial involvement in Pardey RG and Beintema NM (2001) Slow Magic: Agricultural R&D a Century after
grains breeding? Mendel. Washington, DC: International Food Policy Research Institute.
Pray C, Gisselquist D, and Nagarajan L (2011) Private investment in agricultural
research and technology transfer in Africa. Agricultural R&D: Investing in Africas
future, ASTI-FRA Conference, Accra, Ghana, 57 December, Working paper 13.
Exercises for Readers to Explore the Topic Further Pray CE and Nagarajan L (2012) Innovation and Research by Private Agribusiness in
India. Washington, DC: International Food Policy Research Institute IFPRI
Explore the similarities and differences between focus areas Discussion Paper 01181.
for wheat and barley research. Why are some areas similar Ray DK and Foley JA (2013) Increasing global crop harvest frequency: Recent trends
and future directions. Environmental Research Letters 8: 044041. http://dx.doi.org/
yet others quite different? 10.1088/1748-9326/8/4/044041 10 pp.
How do priorities for research relating to irrigated wheat Saltzman A, Birol E, Bouis HE, et al. (2013) Biofortification: Progress toward a more
and rice differ from those for rainfed (or dryland) wheat nourishing future. Global Food Security 2: 917.
and rice? Skerritt JH (2016a) Research organizations of the world: Europe and North America.
Compare and contrast the role, organization, and business In: Corke H, Wrigley C, Faubion J, and Seetharaman K (eds.) Encyclopedia of Food
Grains, 2nd edn., vol. 1, pp. 436443. Elsevier Ltd.
models of farmers organizations undertaking grains Skerritt JH (2016b) Research organizations of the world: Asia-Pacific, Central-South
research in a developed country and a developing country. America, and Africa-Middle East. In: Corke H, Wrigley C, Faubion J, and
Seetharaman K (eds.) Encyclopedia of Food Grains, 2nd edn., vol. 1. pp. 444450.
Elsevier Ltd.
See also: Barley, Rice and Maize Processing: Barley: Malting; Tiwari VK, Faris JD, Friebe B, and Gill BS (2016) Genome mapping. In: Corke H,
Breeding of Grains: Barley: Genetics and Breeding; Canola/ Wrigley C, Faubion J, and Seetharaman K (eds.) Encyclopedia of Food Grains,
2nd edn., vol. 4, pp. 365375. Elsevier Ltd.
Rapeseed: Genetics and Breeding; Maize: Breeding; Soybean: US Energy Information Administration (2012) Biofuels Issues and Trends, October
Germplasm, Breeding, and Genetics; Wheat Breeding: Exploiting and 2012, US Department of Energy, Washington (www.eia.gov/biofuels/issuestrends/
Fixing Genetic Variation by Selection and Evaluation; Food Grains pdf/bit.pdf).
Scientific Societies Associated with Grain Science
SC Nelson, AACC International (AACCI), St. Paul, MN, USA
CW Wrigley, The University of Queensland, Brisbane, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights succeeding text, but the range covered is by no means compre-
hensive. In addition to scientific societies, there are also trade
Large numbers of scientists and technologists are involved associations, formed for the purposes of interaction at the
with food grains worldwide. technical and commercial levels and for lobbying purposes. A
These people have formed scientific associations, based on list of many scientific and trade associations of the food indus-
their specific interests, to facilitate interactions and infor- try has been assembled by www.stagnitomedia.com and www.
mation sharing. hoovers.com.
These interactions involve working collaborations and the The activities of the scientific associations include the orga-
exchange of ideas via conferences, publications, standard- nization of conferences at which lectures are presented, as
izing analytic methods, and explaining grain science to the shown in Figure 1. In addition to lectures, there are poster
general public. presentations, involving the author standing beside the
Two such associations, originally based in the United States description of a research project on a large poster board for
and in Europe (the AACCI (American Association of Cereal discussion with conference delegates (Figure 2).
Chemists International) and the ICC (International Associ- Conferences also serve the purpose of providing opportu-
ation for Cereal Science and Technology)), have become nities for personal interactions to occur; these face-to-face sit-
international. uations complement the less personal interactions by phone
Other associations have regional memberships. and Internet at other times. Trade exhibits (Figure 3) at con-
Some of these associations specialize in specific aspects ferences provide information about the latest equipment
such as seeds, breeding, agronomy, soil science, and grain- and resources. These learned societies use conference time for
related health benefits. making awards to their members who have excelled in aspects
of the science and technology relevant to the societys activities.
Figure 4 shows the recent presentation of several awards
Learning Objectives at a meeting of AACCI. The Thomas Burr Osborne Medal
(Figure 5) is one of these awards.
Other activities include the dissemination of information
Achieve understanding of the need for and the activities
of scientific associations that have been established to via journals, books, and other publications; the establishment
facilitate collaboration and exchange of ideas between of professional standards of conduct and training; and the
scientists and technologists involved in the breeding, development of standard methods of analysis.
growing, harvesting, marketing, and processing of food A related activity is the provision of check samples
grains and nutritionists and consumer groups. characterized subsamples, such as grain or flour, that are pro-
vided to many cooperating laboratories for them to establish
that standard methods of analysis are providing similar results
across various regions.
Introduction The provision of reliable scientific information is becoming
an important function, thereby advising decision-making bod-
The industries that grow, harvest, and process grain are big ies and the general public; these activities may even border on
businesses worldwide, involving large numbers of people, lobbying when it becomes apparent that misinformation is
including many scientists and technologists. It is important prevailing.
that all these professionals should have opportunities to inter-
act with one another, to exchange ideas, to learn from one
another, and to develop grain-testing methods that are
accepted internationally. The Range of Societies
In addition, they need to interact more broadly with con-
sumers, the media, and government agencies. Opportunities Prominent among these learned societies are those relevant to
for these various activities are provided by specialist societies the quality and processing of grain. In particular, AACCI has a
and associations (learned societies), with grain scientists as worldwide membership of scientists involved in grain produc-
members, formed for the purposes of interacting with one tion and processing. The ICC, based in Vienna, Austria, is an
another in their areas of expertise. international association committed to international coopera-
Many such scientific societies have been formed for grain tion through disseminating information and developing stan-
scientists, covering the wide range of aspects of grain science, dard methods relevant to grain production and processing.
from breeding, to agronomy and farming, to the chemistry of There are also national bodies with similar objectives, such as
the harvested grain, to marketing, processing, and retailing. the Chinese Cereals and Oils Association (CCOA), based in
Some of these societies are selected for description in the Beijing, China, and the Australasian Grain Science Association
Figure 1 A major activity of scientific associations is the organization of Figure 4 Scientific societies honor members who excel in their field of
conferences at which lectures are presented to inform members about study by the presentation of awards.
recent developments in their field of grain science.
AACC International (AACCI) segments of the food industry. They are responsible for pro-
ducing some of the most popular and profitable products, such
AACCI is one of the largest international nonprofit organiza- as new types of cracker, cookies, pretzels, or other snack foods.
tions dedicated to creating opportunities for providing and Bringing such products into the market involves design, devel-
exchanging knowledge and advancing innovations across the opment, and marketing skills. Breads, tortillas, frozen doughs,
broad disciplines of cereal grain science by facilitating research, and dry mixes all rely on cereal chemists for their success in the
education, collaboration, technical service, and advocacy efforts. marketplace.
AACCI was founded in 1915 for the purpose of standardizing Other AACCI members are involved in the upstream of the
methods of analysis among cereal laboratories. Today, the food industry. They may be associated with plant-breeding
associations mission is to be the global authority in cereal grain programs, with responsibility for ensuring that the grain from
science and technology. The association now provides a compre- new varieties is suited for specific marketing and processing
hensive and respected collection of grain science methods. All requirements. Research activities are prominent for other cereal
methods go through a rigorous approval process by an AACCI chemists, with the objectives of elucidating the chemical and
technical committee. AACCIs technical committees are com- molecular bases of grain quality, thereby laying a foundation
posed of scientists chosen for their specific areas of expertise. for the development of improved testing methods for grain
Today, as industry issues arise, AACCI strives to provide quality. Other research activities are likely to lead to the devel-
science-based approaches and responses to address current opment of new grain types with better processing and nutri-
needs and trends. The diverse membership provides the creative tional attributes.
and scientific strength to help provide creative solutions and
quality recommendations in reaction to a dynamic industry
and world environment. Through its activities in methods val- AACCI Resources
idation, AACCI is acknowledged as a scientific authority when
proposing definitions and providing comments to regulatory AACCI has provided resources in grain science for a century,
agencies worldwide. Key definitions developed include dietary supporting members professional needs in private industry,
fiber, whole grain, oat bran, glycemic carbohydrates, and aleu- academia, and government. The association provides an ave-
rone. Additionally, AACCI has provided comments on whole nue of bringing together and disseminating scientific informa-
grain label statements; proposed guidance on food allergens, tion and technical research on cereal grains and related
food labeling, and nutrition with respect to gluten-free foods; materials and their processing and utilization. This informa-
and health claims relating to the consumption of certain oat tion is made available through annual meetings, through con-
products to a reduced risk of coronary heart disease. In 2013, tinuing education courses, and in publications. The range of
AACCIs Whole Grains Working Group approved its character- published materials includes scientific journals, books, elec-
ization of a whole grain. The characterization states that a tronic newsletters, and data provided on the website.
whole grain food product must contain 8 g or more of whole AACCI publishes Cereal Chemistry, a journal with peer-
grain per 30 g of product. Volunteers have also organized ad reviewed, original research on raw materials, processes, and
hoc internal committees to examine key issues affecting the products that relate to the utilization of cereal grains, oilseeds,
science and developed action plans to address these issues. and pulses and analytic procedures and methods in the grains
AACCIs strategic initiatives include (1) generating and dis- area. AACCI also publishes Cereal Foods World, which includes
seminating the highest quality of science and information on feature articles and original research that focus on advances in
cereal grains and their products; (2) driving scientific commu- grain-based food science and the application of these advances
nication, collaboration, and innovation; (3) providing profes- on current practices in baking, snack foods, breakfast foods,
sional development for its members; (4) facilitating the and other grain-based products. In addition to these two jour-
interface of the many disciplines involved in cereal and grain nals, AACCI has published more than 80 titles on various food-
science; and (5) ensuring a vital organization in future years. In science topics. Some titles are technically focused while others
2015, AACCI commemorated its centennial year by examining are more general. The AACCI handbook series offers a single
the major advances made in cereal grain science through the source of practical information for the major ingredients used
associations 100 years. AACCI produced a series of feature in food processing.
articles in its publication, Cereal Foods World, examining the AACCIs website offers members the opportunity to obtain
advances by category and explaining the importance of these information and resources in one common location. The web-
advances in science today. Centennial activities also included site features an online catalog of books, special reports, mem-
the creation of a timeline of historic advances and key moments bership directory, a calendar of events, and online symposia.
in AACCI history, which is currently available on AACCIs AACCI journals and electronic copies of publications can be
website. AACCIs 2015 Centennial Meeting included additional found in the AACCI online Grain Science Library. This content
scientific content and a speaker who looked to how todays platform delivers a large and growing collection of resources,
understanding of cereal grain science will inform the future of including Cereal Chemistry, Cereal Foods World, and AACCI
the science and its impact on the world. PRESS online books. AACCIs continuing education programs
offer professional development services for food industry pro-
fessionals at any level in a variety of food-related industries.
Core services include short courses taught by experts that offer
The Activities of AACCI Members
comprehensive training and tools for real-world applications.
Many AACCI members are involved in food manufacturing AACCIs continuing education programs also focus on emerg-
using grains, one of the fastest-growing, most dynamic ing issues and strategies for the future success in the food
460 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Scientific Societies Associated with Grain Science
industry. Members receive special rates on technically focused, an apolitical forum for all cereal scientists and
hands-on training. Courses are offered globally. technologists;
a publisher of international standard methods and a scien-
tific journal;
AACCI Meetings an organizer of major national and international events in
the field;
AACCIs annual meetings are attended by an international a promoter of international cooperation on a global,
audience of more than 1000 grain-based professionals, provid- regional, and national level;
ing a rich scientific program and networking opportunities for a significant player in the coordination and participation in
the global grain industry. Presenters have the opportunity to international research projects.
share the latest research on topics such as biotechnology, func-
tional foods and ingredients, health and nutrition, structure
and function of food components, and quality enhancements. Objectives
The annual meeting also provides students with numerous The ICC is a nonpolitical, nonprofit-making organization.
opportunities to display their research, including product Membership is open to all interested countries, corporations,
development and research paper competitions. The annual and institutions that are prepared to offer their cooperation
meeting features a trade show, typically composed of approx- and to support the aims and ideals of the association.
imately 250 exhibits.
AACCI recognizes excellence through its awards program
(Figure 4). AACCI awards include the AlsbergFrenchSchoch A Short History of the ICC
Memorial Lectureship, the C. W. Brabender Award, the Edith A.
The ICC was founded in 1955 on the occasion of the third
Christensen Award, the Excellence in Teaching Award, AACCI
International Bread Congress in Hamburg, Germany, as the
Fellowships, the Phil Williams Applied Research Award, the
International Association for Cereal Chemistry (ICC).
Texture Technologies Quality Research Awards, the Thomas
The original objective of the ICC was the development of
Burr Osborne Medal (Figure 5), the William F. Geddes Memo-
internationally approved and accepted standard testing pro-
rial Award, the Young Scientist Research Award, and Honorary
cedures for cereals and flour. Today, the ICC is one of the
Memberships. In addition, the AACCI Foundation provides
foremost international organizations in the field of grain sci-
scholarships and fellowships annually for students majoring
ence dedicated to international cooperation, the dissemination
in disciplines related to cereal science. Student and young pro-
of knowledge, and the improvement in safety and quality of
fessionals receive travel assistance grants to attend the annual
cereal-based foods. The association has its headquarters and its
meeting through the Student Travel Award program and the
General Secretariat in Vienna, Austria. At present, some 30
Cecil F. Pinney Travel Award.
countries from five continents are represented in the ICC.
Members have the opportunity to attend local section meet-
ings, connecting them to other professionals within their own
region. AACCI also has several subject-matter divisions focused ICC Mission Statement
on basic research and development areas. The Student Associ-
ation, in conjunction with the Professional Development ICC is the preeminent international association in the field of
Panel, provides students with professional development cereal science and technology, committed to international
opportunities such as showcasing their research (with lectures cooperation through the dissemination of knowledge, con-
and poster presentations; Figures 1 and 2), interacting with ducting research, and developing standard methods that con-
well-respected members of the scientific community through tribute to advance innovation and improve food quality, food
mentorships, and building professional leadership skills safety, and food security for the health and well-being of all
through the competitions mentioned previously. Students people.
also have opportunities to attend hands-on workshops includ-
ing the AACCI Breakfast Cereals and Europe Sections Euro-
pean Young Cereal Scientists and Technologists Workshop and National Cereal Chemistry Societies
the Young Cereal Chemists Meeting.
Many countries have formal and informal associations of pro-
fessionals in grain science. In addition, international associa-
tions, such as the AACCI, have regional sections and branches
The International Association for Cereal Science that provide opportunities for members to hold local
and Technology meetings.
wider range of grain scientists. The CCOA has over 6000 indi- commemorates the pioneering wheat research of Australian
vidual members and has over 570 organization members. The Frederick Bickell Guthrie, who formed the worlds first
CCOA is a member of the ICC. The CCOA is divided into four chemistbreeder collaboration with the wheat breeder William
subassociations, namely, grain storage, grain food, edible-oil Farrer commencing in the 1890s.
processing, and animal-feed production. The CCOA publishes Throughout the life of AGSA and the Cereal Chemistry
a journal, the Journal of the Chinese Cereals and Oils Association. Division, presentations at annual meetings have been recorded
The Chinese version is published every 2 months, while the in published form, in recent years under the titles Cereals 2016,
English version appears annually. Cereals 2015, Cereals 2014, etc. Details of these publications
and other activities are provided on the AGSA website (http://
ausgrainscience.org.au).
Australian and New Zealand Grain Scientists
The Australasian Grain Science Society (AGSA) is currently the Societies Involved with Grain Growth
main association for cereal chemists and other grain scientists
The International Seed Testing Association
in Australia, New Zealand, and nearby regions. AGSA, now a
stand-alone society, has grown out of the Cereal Chemistry The ISTA is a worldwide, nonprofit association, whose main
Division, which was a part of the RACI, based in Melbourne, activity is to provide methods and services for the testing of
Australia. In addition, many Australian grain scientists are seed traded internationally. The primary purpose of the
members of the AACCIs Downunder Section. association is thus to develop, adopt, and publicize standard
AGSA provides opportunities for scientists and technolo- methods for sampling and testing seeds of all types, not only
gists to exchange information and to interact with colleagues the edible grains. ISTAs secondary purpose is to promote all
internationally, especially at annual meetings. These annual areas of seed science and technology. These purposes are
meetings continue the series of annual Australian Cereal largely served by the publication of relevant handbooks and
Chemistry Conferences, which commenced in 1950, so that bulletins, scientific journals (Seed Science and Technology
the 2016 annual meeting is the 66th conference in this series. and Seed Symposium Abstracts), proceedings of symposia and
At regular intervals, these meetings are combined with other workshops, and training booklets.
international societies, especially the AACCI and the ICC. The primary ISTA instrument for promoting uniformity of
Other activities include workshops and training seminars seed-testing procedures is the International Rules for Seed Test-
on specific aspects of grain science, the provision of standard ing, a publication that is updated annually with amendments
methods of analysis, and facilities for checking samples with and additions resulting from ISTA meetings. The ISTA Secre-
known analytic results by which laboratories are able to com- tariat is based in Bassersdorf, Switzerland.
pare the results of their analytic methods with others.
AGSA has several awards to acknowledge special contribu-
The American Societies of Agronomy, Crop, and Soil Science
tions to grain science. The Guthrie Medal (Figure 6)
Founded in 1907, the American Society of Agronomy is
dedicated to the development of agriculture enabled by sci-
ence, in harmony with environmental and human values. The
society supports scientific, educational, and professional activ-
ities to enhance communication and technology transfer
among agronomists and those in related disciplines on topics
of local, regional, national, and international significance.
The Crop Science Society of America is an educational and
scientific organization composed of more than 5000 members
dedicated to the advancement of crop science. The motto is
Plant science for a better world. Founded in 1955, the society
is international in scope with members in more than 100
countries who are advancing the discipline of crop science by
acquiring and disseminating information about crops in rela-
tion to genetics and plant breeding, crop physiology and pro-
duction, germplasm resources, and environmental quality. An
important function of the society is the production of its main
journal Crop Science.
The Soil Science Society of America, established in 1936, is
the professional home for over 6000 professionals throughout
the world involved in soil science. Their claim is Soils sustain
Figure 6 The Guthrie Medal, originally awarded by the Cereal Chemistry
life. The primary purpose of the society is to advance the
Division of the Royal Australian Chemical Institute, is now awarded of the discipline and practice of soil science by study and education
Australasian Grain Science Association. It is awarded for outstanding concerning soils in relation to crop production, environmental
contributions in the field of grain science. The medal commemorates an quality, ecosystem sustainability, bioremediation, waste man-
Australian cereal chemist of the 1890s and 1900s. agement and recycling, and wise land use.
462 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Scientific Societies Associated with Grain Science
Societies Involved with Grain Analysis communities. Membership helps shape this foundations
objectives and foster integral conversations that create practical
AOAC International change. The Grains for Health Foundation actively seeks both
AOAC International is a globally recognized, independent, sponsor members and partner members.
third party, not-for-profit association with 130 years of expe- Grains for Health Foundation goals include redesigning the
rience in validating and approving analytic methods for foods food supply by facilitating the development, delivery, and
and agriculture. It has over 3000 members worldwide. consumption of grain-based foods in order to promote a
The organization was founded in 1884 as the Association of healthy body weight, reduce chronic disease, and curb health-
Official Agricultural Chemists, under the auspices of the US care costs. These bridges ignite the necessary conversations
Department of Agriculture. Its initial role was to adopt uniform around making healthy, grain-based foods accessible.
methods of analysis for fertilizers. In the following year, a Grains for Health Foundation measures its success to the
convention established the AOAC as an independent organi- extent that contributions help build and maintain the crucial
zation, but membership was restricted to analytic chemists in relationships needed between the whole grain food supply and
state and federal positions of the US government, and this the researchers and policy makers that regulate the distribution
membership requirement remained for a century. In 1965, to and education surrounding grain-based foods. Measures of
recognize the expansion of AOACs scope of interest beyond success include the following:
agricultural topics, the associations name was changed to the
Association of Official Analytical Chemists.
Foods with healthy grains are available in your grocery,
school, or business.
During the 1970s, membership was extended to scientists
outside the United States and to nongovernmental scientists,
Health-care professionals recommend whole grains as a
means of reducing chronic illness.
with many members working in industrial laboratories. Dur-
ing the 1980s and 1990s, the scope of analyses expanded and
Policy makers design legislation leading to healthier grain
products.
there was increasing demand for quality control of laboratories
and international laboratory accreditation. Consequently, the
Scientists create grains able to feed our communities.
known for 130 years, while eliminating reference to a specific Grains for Health Foundation undertake strategic and transfor-
scientific discipline or profession. mative work on all levels of the grain chain from connecting
The scope of the AOAC includes methods not only for the growers to scientists to politicians to distributors to educators
analysis of grains but also for a much wider range of materials. to health-care professionals. The cross-disciplined work builds
AOAC provides a number of key publications (especially its a vast community needed to create the positive change in
Official Methods of Analysis), hosts technical meetings and con- public health so you can be assured access to healthy, grain-
ferences, and offers training courses in the areas of laboratory based foods.
management, quality assurance, accreditation, statistics, and
measurement uncertainty.
The HEALTH GRAIN Forum
This association was founded in May 2010 to continue promo-
The American Oil Chemists Society
tion of science and communication for production and con-
The AOCS (Your Global Fats and Oils Connection) is another sumption of healthy cereal foods, as a follow-up of the
American scientific association that has become international, HEALTHGRAIN Forum of the European Union. There are cur-
growing and expanding in scope since its origins in 1915. This rently members from 19 countries. The organizational struc-
scope now includes oil- and fat-related commodities, oilseeds, ture includes a board and four task forces to plan activities,
oilseed meals, and edible fats. Grains are thus prominent in interacting via telemeetings on a regular basis. Two annual
this range of materials for analysis, but the scope goes beyond workshops are organized to provide opportunities for
grains, including other sources of edible fats, such as fish and interaction.
animals. The AOCS technical services are based in Champaign, The mission of the HEALTHGRAIN Forum is to promote
IL, the USA. science-based concepts fully, unlocking the health-promoting
potential in the entire grain food production chain to obtain
healthy, convenient, and appealing foods. The vision is to
assist consumers in health maintenance worldwide, help
Societies Relating to Food Grains and Health
reduce health-care costs, and provide added value for compa-
Grains for Health Foundation nies in the production chain. The motto is The best of the grain
for consumer well-being.
The people working with Grains for Health Foundation
include scientists, business leaders, educators, and health pro-
fessionals working together at the grassroots level encouraging
Celiac Societies
all sectors, disciplines, and communities to use their expertise
to improve public health. Celiac societies have been formed in many countries and
The Grains for Health Foundation provides publications, regions by people with celiac disease and related forms of
resources, and events within the grains and health dietary intolerance to gluten. These are not associations of
SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Scientific Societies Associated with Grain Science 463
professionals only, but rather of people with celiac disease, other examples can you find? Think, for example, of asso-
together with their relatives and friends. They may, in turn, ciations involving breeders, geneticists, grain growers,
rely on scientists and medical workers to provide advice in the health professionals, nutritionists, and consumers.
forms of publications, consultation, and conference presenta- Find out the costs of membership, of conference atten-
tions for the benefit of the members. A few web addresses are dance, and of other forms of involvement for some of
provided in the list in the succeeding text, but there are many these associations. Try to assess the value for money in
more such societies worldwide. terms of professional benefits to be gained by members.
To what extent do you reckon that benefits depend on
involvement via publications (only), conference atten-
Future Prospects dance (only), and/or volunteering for direct participation
in association affairs?
Into the future, grain science professionals will continue to Look for indications that these associations have recently
need opportunities to interact and collaborate. Scientific soci- been involved in advising decision-making bodies with
eties can continue to provide scientists not only with validated respect to their areas of expertise.
and archived information but also with the essential feature of
contact with other scientists. While todays technology pro-
vides new options for frequent and instant communications, See also: Food Grains: Intolerance, Allergy and Diseases:
it continues to be important for scientists to meet face to face Celiac Disease; Grain Composition and Analysis: Standardized
for effective dialogue. Some means of interaction will progres- Test Methods for Grains and Grain-Based Products; Scientific
sively change, largely due to changes in the range of novel Organizations Related to Grains: Research Organizations of the
electronic media. Already, traditional means of print publica- World: Asia-Pacific, CentralSouth America, and AfricaMiddle East;
tion are being duplicated or replaced by various electronic Research Organizations of the World: CGIAR; Research Organizations
means. The value of face-to-face meetings and conferences of the World: Europe and North America; Research Organizations of the
cannot be readily replaced. World: Global Trends and the Commercial Sector.
HAROLD CORKE
University of Hong Kong, Hong Kong, Peoples Republic of China
KOUSHIK SEETHARAMAN{
JON FAUBION
Kansas State University, Manhattan KS, USA
{
deceased
The following articles are US government works in the public domain and are not subject to copyright:
No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including
photocopying, recording, or any information storage and retrieval system, without permission in writing from the publisher. Details on
how to seek permission, further information about the Publishers permissions policies and our arrangements with organizations such as
the Copyright Clearance Center and the Copyright Licensing Agency, can be found at our website: www.elsevier.com/permissions.
This book and the individual contributions contained in it are protected under copyright by the Publisher (other than as may be noted
herein).
Notice
Knowledge and best practice in this field are constantly changing. As new research and experience broaden our understanding, changes in
research methods, professional practices, or medical treatment may become necessary.
Practitioners and researchers may always rely on their own experience and knowledge in evaluating and using any information, methods,
compounds, or experiments described herein. In using such information or methods they should be mindful of their own safety and the
safety of others, including parties for whom they have a professional responsibility.
To the fullest extent of the law, neither the Publisher nor the authors, contributors, or editors, assume any liability for any injury and/or
damage to persons or property as a matter of products liability, negligence or otherwise, or from any use or operation of any methods,
products, instructions, or ideas contained in the material herein.
Preface xvii
Tributes to Former Co-Editors xix
Editors Biography xxiii
Editorial Advisory Board xxv
Contributors xxvii
VOLUME 1
The Basics 1
The Grains that Feed the World 1
CW Wrigley, H Corke, and J Faubion
Grain and Plant Morphology of Cereals and how characters can be used to identify varieties 51
J Wyatt
Maize: Overview 99
MP Scott and M Emery
Wheat: An Overview of the Grain That Provides Our Daily Bread 105
CW Wrigley
v
vi Contents
African Rice (Oryza glaberrima): A Brief History and Its Growing Importance in Current
Rice Breeding Efforts 140
JT Manful and S Graham-Acquaah
Barley: An Overview of a Versatile Cereal Grain with Many Food and Feed Uses 147
AM Stanca, A Gianinetti, F Rizza, and V Terzi
Grain Production and Consumption in Oceania: Australia and Pacific Countries 409
CW Wrigley and RJ French
Research Organizations of the World: Asia-Pacific, CentralSouth America, and AfricaMiddle East 444
JH Skerritt
viii Contents
Research Organizations of the World: Global Trends and the Commercial Sector 451
JH Skerritt
VOLUME 2
Celiac Disease 83
P Koehler, H Wieser, and KA Scherf
Proteins 98
The Protein Chemistry of Cereal Grains 98
F Bekes and CW Wrigley
The Enzymes Associated with Food Grains and Their Functions in Processing 138
DJ Mares, K Mrva, and GB Fincher
Carbohydrates 161
Carbohydrate Metabolism 161
RN Chibbar, S Jaiswal, M Gangola, and M Baga
Fats 248
Lipid Chemistry 248
L Day
Mycotoxins 290
MT Fletcher and BJ Blaney
Units of Grain Science and Trade: Equivalence between the US, Chinese, and Metric Units 308
W Huang and CW Wrigley
VOLUME 3
Wheat-Based Foods 1
Cereals: Overview of Uses: Accent on Wheat Grain 1
CF Morris
Breads 8
TR Moore
Noodles: Starch 64
LS Collado and H Corke
Pasta 79
M Sissons
Tortillas 90
LW Rooney and SO Serna-Saldivar
Non-wheat Foods 97
Maize: Foods from Maize 97
SO Serna-Saldivar
Contents xi
Biodiesel 245
Y Yan
The Gluten Proteins of the Wheat Grain in Relation to Flour Quality 375
F Bekes, MC Gianibelli, and CW Wrigley
VOLUME 4
Wheat: Marketing 9
RL Cracknell and RM Williams
The Nature, Causes, and Control of Grain Diseases in the Major Cereal Species 74
RA McIntosh, PM Williamson, and CW Wrigley
Contaminants of Grain 83
J McLean and CW Wrigley
xiv Contents
Appendix 1. Nutrient-Composition Tables for Grains and for Grain-Based Products 433
SF Schakel, N Van Heel, and J Harnack
Appendix 2. Lists of Standardized Testing Methods for the Analysis of Grain and Grain-Based Foods 450
AR Bridges and CW Wrigley
Appendix 3. Grains, Foods, and Ingredients Suiting Gluten-Free Diets for Celiac Disease 460
JM Jones, V Zevallos, and CW Wrigley
Glossary 467
Index 491
PREFACE
The grain-producing plants are essential to life on earth. They are the primary means by which sunlight, the
primary source of the earths energy, is captured via photosynthesis to turn carbon dioxide and water into
carbohydrates and thus into the wider range of the biochemicals needed by all life forms.
The summary is simple:
The suns energy CO2 H2O
#
Grain-bearing plants
which provide:
Food, Fats, Fuel and Fiber for ourselves
Feed, Forage, Fodder for our animals
Food for Fish Farms
As its title indicates, the Encyclopedia of Food Grains concentrates on the food uses of grains, but details are also
provided about the wider roles of grains. Greater detail about the use of grains for animal feeding is already
available via the on-line Feedipedia encyclopedia (http://www.feedipedia.org/).
Worldwide, however, it is our food that is our obsession first and foremost. We see our health (or
otherwise) to be intimately related to our diet. Therefore nutrition is a major accent of this second edition,
occupying the whole of the second volume, following the first volumes description of the basic aspects of the
world of grains. The third and fourth volumes work their ways back up the value-added grain chain to describe
grain processing and production, thence to breeding and genetics.
The sequence of articles in the first edition (the Encyclopedia of Grain Science, 2004) followed an alphabetical
order. In contrast, the distinct ordering of articles in the second edition assembles fundamental concepts in
Volume 1 The World of Food Grains. Moving into the later volumes, greater complexity and erudition are
evident. Thus, unusual terms such as Transcriptomics, Aeciospore and Allelopathy are more likely to be
found in later volumes than in the first. In any case, reference to the glossary and the index should help in
elucidating such terms.
This second edition is a considerable expansion on the content of the first edition, with its three print volumes
and 168 articles. Now, some twelve years later, the four print volumes of the Encyclopedia of Food Grains provide
xvii
xviii Preface
216 articles, many newly written for the new edition. Those articles that have been reprinted from the first
edition have been updated. In all cases, authors and reviewers are world renowned experts in their respective
fields.
This second edition has been more than three years in its development. In the early stages, the editors were
Harold Corke, Chuck Walker and Colin Wrigley (the three editors of the first edition). Sadly in 2012, Chuck
Walker passed away, leaving a gap that was difficult to fill.
Chuck was replaced by Koushik Seetharaman, who worked avidly in developing the concepts for this second
edition, including his contributions to a meeting of the editors and Elsevier staff in Sydney, January, 2013.
Sadly, Koushik suffered a heart attack in June, 2014. However, his contributions to the second edition during
this critical development stage were such that Elsevier decided that his name should remain as a co-editor.
Tributes to Professors Walker and Seetharaman are provided below.
In July, 2014, Professor Jon Faubion was appointed as a co-editor. Dr Faubion is the Charles Singleton
Professor of Baking and Cereal Science in the Department of Grain Science, Kansas State University. He is a
more than worthy replacement under such tragic circumstances.
The production of an encyclopedia involves a wide range of experts with expertise in a diversity of areas of
knowledge. We acknowledge with grateful thanks the groups of contributors:
Members of the Editorial Advisory Board;
Authors who have shared their special knowledge and experience;
Elsevier staff who have provided extensive advice and administrative services, especially Donna de
Weerd-Wilson, Simon Holt and Rachel Gerlis (successive Supervisory Staff) and Gemma Tomalin,
Gemma Taft, Joanne Williams and Mark Harper (successive Content Project Managers).
The Editors
Harold Corke, Jon Faubion, Koushik Seetharaman, and Colin Wrigley
TRIBUTES TO FORMER CO-EDITORS
Chuck Walker was one of the three editors of the first edition of Elseviers Grains
Encyclopedia (the Encyclopedia of Grain Science, 2004). He was again appointed
as an editor of this second edition. He participated in the planning stages for the
second edition until his untimely death on 26 April 2012. At the time of his
death, Chuck was a Professor Emeritus at the Department of Grain Science and
Industry at Kansas State University, Manhattan, KS, USA.
Charles Eugene Walker was born on 17 December 1936, in Winterset, Mad-
ison county, IA, USA, during the great depression. He was the eldest child of
Albert Cecil and Bessie Irene Headley Walker, poor farmers in Winterset, Mad-
ison county, IA, USA. He walked more than a mile to attend rural one-room
grammar schools in Warren county and graduated from high school in New
Virginia, IA, USA.
Better known as Chuck in later life, Dr Walker received a BS degree in
Chemical Engineering from Iowa State University, Ames, IA, USA, in May 1959. At a mixer party in the fall
of 1957, he met Shannon Rose Philp, a pretty freshman student from Loup city, NE, USA. This was just after a
summer spent working alone on a fire-lookout tower for the US Forest Service, in the wilds of Clearwater
National Forest, in Northern Idaho. They were married on 1 June 1958 and she became his wife of 54 years. She
later rewarded him with two sons, Alan Eugene and Tomas Charles, and they adopted a daughter, Bekianne.
Upon graduation, Chuck worked in flour milling research at the General Mills Central Research Laboratories
in the Minneapolis area from 1959 to 1962. He then accepted a fellowship to graduate school at North Dakota
State University in Fargo from 1962 to 1965, majoring in Cereal Chemistry. Chuck received his PhD in
Chemistry in the summer of 1966. For his PhD dissertation, he developed a micro brewery and studied barley
proteins and their influence on beer quality a rather ironic choice of topics, considering he remained a strict
teetotaler all his life.
While finishing his thesis, he joined the faculty of the North Dakota State College, Valley city, ND, USA
(19651974), teaching various physical science and chemistry courses to undergraduate students. Feeling the
pull of industry, he moved south to work for the Fairmont Foods Central Research Laboratory in Omaha, NE,
USA (19741980), where he did research, product development, and technical assistance on baking, pizza,
snack, and dairy products, eventually reaching the position of Associate Director of Research.
Dismemberment and sale of this Fortune-500 company provided him an opportunity to return to academia
at the University of Nebraska, Lincoln, NE, USA (19801987), where he developed a Cereal Technology
Program within the Department of Food Technology. He developed and taught various food science and
technology courses to both undergraduate and graduate students, and developed a vigorous research program.
He became Interim Department Head and Food Processing Center Director.
In December 1987, Chuck joined the Department of Grain Science and Industry at Kansas State University as
the Bakers National Educational Foundations Baking Science Professor. There he maintained a busy program
xix
xx Tributes to Former Co-Editors
in research, teaching, graduate student advising, publishing, traveling and consulting, specializing in baking
technology.
During his career, Chuck published more than 100 technical papers, and advised about 50 MS and PhD
graduate students. He was heavily involved in applied rheology (mixing and flow characteristics) of flour
products. In recognition of his commitment to international graduate students, Mr Roland Temme, TMCO,
Lincoln, NE, USA, established an international graduate student fellowship in his honor.
Chuck entered 50% phased-in retirement at age 65 in 2002 and retired in 2005, only to be hired back 1/
10-time afterward to assist with seminars and the departments centennial recognition in 2010. He continued
to support the departments activities on a volunteer basis.
Chucks research and consulting travels took him to more than a dozen countries, including many extended
trips to Australia, eventually earning permanent resident status in Australia a country which had fascinated
him since childhood. Most of his time in Australia was spent with the Bread Research Institute in Sydney. He
also made several extended trips to China, serving as guest lecturer at ShanDong Agricultural University, TaiAn,
and at Henan University of Technology, Zhengzhou.
Chuck always commented that the best legacy a person could leave is the knowledge he has shared. A
significant way in which his knowledge has been shared is represented by his contributions to the first and
second editions of Elseviers Encyclopedia of Food Grains.
In August 2012, Koushik Seetharaman was appointed as the third editor of this
second edition of Elseviers Grains Encyclopedia, replacing Professor Walker.
Tragically, Koushiks many contributions to the Encyclopedia of Food Grains were
cut short by his untimely death at 48 years in June 2014.
Koushik received his BS at the Gujarat Agricultural University, Anand, Guja-
rat, India. His MS training was at Cornell University, Ithaca, NY, USA, and his
PhD research was carried out at Texas A&M University, College Station, TX, USA.
At the time of Koushiks appointment as encyclopedia editor (August 2012),
he was Associate Professor and Cereals Chair in the Department of Food Science
at the University of Guelph, Canada. Research in his laboratory spanned the
cereal-grain value chain, focusing on grain quality, processing and ingredient
interactions, and on consumer acceptability and health.
His research on grain quality at Guelph involved collaboration with cereal-breeding companies, grain
farmers of Ontario and local processors, by analyzing Ontario wheats for functionality and nutritional
attributes. Special accents were on antioxidant activity, and on fiber and phenolics contents. Interactions
with breeders and processor related to the chemistry, sensory properties and consumer acceptance of whole
grain products, with specific focus on red versus white wheats in different product matrices.
Koushik had a special interest in the behavior of starch in water-poor systems, such as dough and baked
products. His research group had demonstrated that the constituent polymers of wheat starch are a series of
biopolymers ranging from linear to branched, depending on several factors including genetics and growth
environment. Moreover, the proportion of these polymers further defines their functional properties.
Research funding had been provided by the Canadian Agricultural Adaptation Council with contributions
from the Ontario Cereal Industry Research Council, Kellogg, Kraft, Kraft Canada Mill, Dow AgroSciences, C&M
Seeds, Grain Farmers of Ontario and Brabender GmbH.
Several months before his death, Koushik had been appointed as an Associate Professor to the Department
of Food Science and Nutrition at the University of Minnesota, St Paul, MN, USA, there taking the position of
General Mills Cereal Chair. In that capacity, Koushik had again initiated collaborations with breeders and
geneticists, millers and processors, with a focus on what product quality means for the consumer. His research
focused on interactions of grain biopolymers starch and gluten in particular and the impact on processing,
product attributes and consumer health.
Tributes to Former Co-Editors xxi
His group included three postdoctoral research associates, five doctoral students and one MS student. They
were collaborating extensively with research groups around the world including University of Milan, Italy;
University of Copenhagen, Denmark; Lille University, France; ONIRIS, France; Tamilnadu Agricultural Uni-
versity, India; Iowa State University, USA; African Rice Research Institute, Benin; University of Guelph, Canada;
and Memorial University, Newfoundland, Canada.
Koushiks research background and his collaborations placed him well to contribute to the preparation of
the grains encyclopedia, both by his our writing and via his contacts to engage world-authority contributors.
The grain-science community world-wide was shocked by Koushiks sudden passing. Koushik left a wife
Debra Freedman and an eight-year-old son, Samuel.
Koushiks contribution to the encyclopedia was such that Elsevier decided that his name should remain as a
co-editor even though he did not live to see the end result of his contributions.
Colin W. Wrigleys 55 years in grain-science research have earned him international recognition in
the form of several international and Australian research awards. His work is described in about
600 research publications, including several patents, a series of eight books on Australian cereal
varieties, and many edited books. He was Editor-in-Chief of the first edition of the Encyclopedia of
Grain Science (2004).
His research interests have centered on the characterization of cereal-grain proteins in relation to
processing quality. This has involved developing new methods of protein fractionation, including
gel isoelectric focusing and its 2D combination with gel electrophoresis, leading into proteomic
mapping. Other diagnostic methods developed relate to the evaluation of grain quality in wheat
and barley, such as better methods for variety identification and for characterizing quality in starch
and sprouted grain (as co-patentee of the Rapid ViscoAnalyser). Research involvement has also
included elucidation of grain-quality variation due to environmental factors (heat stress, fertilizer
use, CO2 levels and storage conditions).
In 2009, Wrigley was appointed a Member of the Order of Australia (AM) for service to primary
industry, particularly to grain science as a researcher, and to the development of methods for
improving wheat quality.
He is currently an Honorary Professor at the University of Queensland, Brisbane, Australia.
Harold Corke is a professor in the Food and Nutritional Sciences program at The University of
Hong Kong, and a Chutian Scholar Distinguished Foreign Professor in the Glyn O. Phillips
Hydrocolloid Research Center at HUT, Hubei University of Technology, Wuhan, China. In more
than 22 years in Hong Kong, he has had responsibility for teaching a diverse array of food science
courses, including the major courses Grain Production and Utilization, Food Safety and Quality
Management, Food Product Development, and general education courses Feeding the World, and
Food: Technology, Trade and Culture. He is author or co-author of 180 refereed journal articles,
and his 22 PhD graduates have gone on to successful careers in academia and industry around the
world. He is on the editorial boards of Journal of Food Composition and Analysis, LWT Food Science and
Technology and Journal of Cereal Science, and with Colin Wrigley and Chuck Walker was one of the
editors of the first edition of Encyclopedia of Grain Science, published in 2004. Corke is active in
consulting in grain processing and food safety in Asia and Eastern Europe.
Jon Faubion is the Charles Singleton Professor of Baking and Cereal Science in the Department of
Grain Science, Kansas State University. Over his 35 year career, he has had the sole or shared
responsibility for teaching nine different cereal or food science courses at Texas A&M University,
The University of Minnesota and Kansas State. In 2011, he received the Excellence in Teaching
Award from the American Association of Cereal Chemists International. He is a senior editor of
Cereal Chemistry, and a member of the Board of Directors of the American Association of Cereal
Chemists International and an executive editor of Journal of the Science of Food and Agriculture. Prior
to rejoining the department faculty in 2005, he directed the Applied Technology and Sensory
Science Groups for the research and development arm of The Schwan Food Company.
xxiii
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EDITORIAL ADVISORY BOARD
xxv
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CONTRIBUTORS
E Akhunov BJ Blaney
Kansas State University, Manhattan, KS, USA The University of Queensland, St Lucia, Australia
PC Annis JE Bock
CSIRO Entomology, Canberra, ACT, Australia University of Guelph, Guelph, ON, Canada
GA Annor F Bonomi
University of Ghana, Accra, Ghana University of Milan, Milan, Italy
CA Atkins AR Bridges
The University of Western Australia, Crawley, WA, AACC International, St Paul, MN, USA
Australia
H Brinch-Pedersen
W Atwell Aarhus University, Slagelse, Denmark
Bill Atwell Consulting, Champlin, MN, USA
G Bultosa
M Baga Haramaya University, Dire Dawa, Ethiopia; Botswana
University of Saskatchewan, Saskatoon, SK, Canada College of Agriculture, Gaborone, Botswana
JS Bao W Bushuk
Zhejiang University, Hangzhou, PR China University of Manitoba, Winnipeg, MB, Canada
VJ Barthet YZ Cai
Canadian Grain Commission, Winnipeg, MB, Canada China National Seed Group, Beijing, PR China;
SA Barton Huazhong Agricultural University, Wuhan, PR China;
Oxford Agricultural Trials Ltd, Oxford, UK The University of Hong Kong, Hong Kong, PR China;
Hubei University of Technology, Wuhan, PR China
IL Batey
Sunset Cereal Services, Meadowbank, NSW, Australia EF Caldwell
AACC International, Roseville, MN, USA
F Bekes
FBFD PTY LTD, Beecroft, NSW, Australia KJ Carpenter
University of California, Berkeley, CA, USA
AR Belcher
Oregon State University, Corvallis, OR, USA JC Catullo
INTA, Buenos Aires, Argentina
JN BeMiller
Purdue University, West Lafayette, IN, USA SP Cauvain
BakeTran, Witney, OX, UK
T Beta
University of Manitoba, Winnipeg, Manitoba, Canada S Ceccarelli
Via delle Begonie 2, Ascoli Piceno, Italy
JL Black
John L Black Consulting, Warrimoo, NSW, Australia GK Chandi
University of Guelph, Guelph, ON, Canada
MH Blanco
(Retired) Plant Breeder and Geneticist, Encinitas, N Chaudhary
CA, USA G. J. University of Science and Technology, Hisar, India
xxvii
xxviii Contributors
RN Chibbar JE Dexter
University of Saskatchewan, Saskatoon, SK, Canada Canadian Grain Commission, Winnipeg, MB, Canada
M Chisi B Dobraszczyk
Golden Valley Research Station, Chisamba, Zambia Reading Science Centre, Reading, UK
S Cloutier D Domingues
Agriculture and Agri-Food Canada, Ottawa, ON, General Mills Inc., Minneapolis, MN, USA
Canada
C Dowd
LS Collado General Mills Inc., Minneapolis, MN, USA
The University of the Philippines Los Banos, Laguna,
LM Duizer
Philippines
University of Guelph, Guelph, ON, Canada
NC Collins
RW Duncan
School of Agriculture Food and Wine, The University of
University of Manitoba, Winnipeg, MB, Canada
Adelaide, Glen Osmond, SA, Australia
KG Duodu
H Corke
University of Pretoria, Pretoria, South Africa
The University of Hong Kong, Hong Kong, Pokfulam,
PR China; Hubei University of Technology, Wuhan, SR Eckhoff
PR China (Retired) University of Illinois, Urbana, IL, USA
RL Cracknell M Edney
Crackers Consulting, Mount Eliza, VIC, Australia Canadian Grain Commission, Winnipeg, MB, Canada
GB Crosbie M Emery
Crosbie Grain Quality Consulting, East Fremantle, WA, Iowa State University, Ames, IA, USA
Australia
W Erskine
JS Croser University of Western Australia, Crawley, WA, Australia
University of Western Australia, Crawley, WA, Australia
WD Eustace
A Cruickshank (Retired) Kansas State University, Manhattan, KS, USA
DAFF Queensland, Warwick, QLD, Australia
A Evans
A Cuesta-Marcos Tate and Lyle, Hoffman Estates, IL, USA
Oregon State University, Corvallis, OR, USA
E Evans
MG DEgidio The Tassie Beer Dr, Lindisfarne, TAS, Australia
Unita di ricerca per la Valorizzazione Qualitativa dei
JD Faris
Cereali, Rome, Italy
USDA-ARS Cereal Crops Research Unit, Fargo, ND,
D Dan Ramdath USA
Agriculture and Agri-Food Canada, Guelph, ON, Canada
J Faubion
P Dangi Kansas State University, Manhattan, KS, USA
G. J. University of Science and Technology, Hisar, India
AM Fehily
JK Daun Tinuviel Software, Warrington, UK
(Deceased)
T Filichkin
C David Oregon State University, Corvallis, OR, USA
ISARA Lyon, Lyon, France
GB Fincher
L Day Australian Research Council Centre of Excellence in Plant
AgResearch Ltd., Palmerston North, New Zealand Cell Walls, University of Adelaide, Glen Osmond, SA,
Australia
D Deneke
South Dakota State University, Brookings, SD, USA SP Fisk
Oregon State University, Corvallis, OR, USA
R DePauw
Agriculture and AgriFood Canada, Swift Current, SK, MT Fletcher
Canada The University of Queensland, St Lucia, Australia
Contributors xxix
RJ Fletcher A Gunaratne
University of Queensland, Gatton, QLD, Australia The University of Hong Kong, Hong Kong, PR China
F Fleurat-Lessard YN Guragain
INRA, Mycology and Food Safety Research Unit, Kansas State University, Manhattan, KS, USA
Villenave dOrnon, France
RCE Guy
JA Fox Campden and Chorleywood Food Research Association,
Kansas State University, Manhattan, KS, USA Chipping Campden, UK
RJ French C Hall III
The University of Western Australia, Crawley, WA, North Dakota State University, Fargo, ND, USA
Australia
LM Hall
B Friebe University of Alberta, Edmonton, AB, Canada
Kansas State University, Manhattan, KS, USA
EG Hammond
BJ Furman Iowa State University, Ames, IA, USA
San Jose State University, San Jose, CA, USA
R Haraszi
M Gangola Campden BRI, Gloucestershire, UK
University of Saskatchewan, Saskatoon, SK, Canada
J Harnack
P Gelinas University of Minnesota, Minneapolis, MN, USA
Food Research and Development Centre, Agriculture and
D Hauge
Agri-Food Canada, Saint-Hyacinthe, QC, Canada
Grains for Health Foundation, St. Louis Park, MN, USA
SM Ghazani
AMR Hayes
University of Guelph, Guelph, ON, Canada
St. Catherine University, Lafayette, MN, USA
MC Gianibelli
PM Hayes
(Retired)
Oregon State University, Corvallis, OR, USA
A Gianinetti
L Helgerson
Consiglio per la ricerca in agricoltura e lanalisi
Oregon State University, Corvallis, OR, USA
delleconomia agraria, CRA-GPG, Fiorenzuola dArda,
Italy RJ Henry
University of Queensland, Brisbane, QLD, Australia
BS Gill
Kansas State University, Manhattan, KS, USA D Herb
Oregon State University, Corvallis, OR, USA
JS Godber
LSU Agricultural Center, Baton Rouge, LA, USA I Herencia
Universidad Politecnica de Madrid, Madrid, Spain
A Goldstein
University of Minnesota, St. Paul, MN, USA E Hernandez
Texas A&M University, College Station, TX, USA
R Graebner
Oregon State University, Corvallis, OR, USA KB Hobson
New South Wales Department of Primary Industries,
S Graham-Acquaah
Tamworth, NSW, Australia
Africa Rice Center, Cotonou, Republic of Benin
M Hochhalter
RA Graybosch
North Dakota State University, Fargo, ND, USA
USDA-ARS, University of Nebraska Lincoln, Lincoln,
NE, USA RD Horsley
North Dakota State University, Fargo, ND, USA
S Grundas
Bohdan Dobrzanski Institute of Agrophysics of the Polish W Huang
Academy of Sciences, Lublin, Poland Jiangsu Academy of Agricultural Sciences, Nanjing,
PR China; Jiangnan University, Wuxi, Jiangsu, PR China
TJ Gulya
USDA-ARS, Northern Crop Science Laboratory, Fargo, Y Huang
ND, USA China Agricultural University, Beijing, PR China
xxx Contributors
V Humphrey-Taylor P Koehler
(Retired) New Zealand Institute for Crop Food Research Deutsche Forschungsanstalt fur Lebensmittelchemie,
Limited, Christchurch, New Zealand Leibniz Institut, Freising, Germany
T Hymowitz R Korczak
University of Illinois at Champaign-Urbana, Urbana, University of Minnesota, St. Paul, MN, USA
IL, USA
SG Krattinger
S Iametti University of Zurich, Zurich, Switzerland
University of Milan, Milan, Italy
L Krishnamurthy
C Isaak International Crops Research Institute for the Semi-Arid
University of Manitoba, Winnipeg, MB, Canada Tropics, Patancheru, Andhra Pradesh, India
MS Izydorczyk S Kumar
Canadian Grain Commission, Winnipeg, MB, Canada International Center for Agricultural Research in the Dry
Areas (ICARDA), Rabat, Morocco
P Jacques
Tufts University, Boston, MA, USA OR Larroque
CSIRO Agriculture Flagship, Canberra, Australia
S Jaiswal
University of Saskatchewan, Saskatoon, SK, Canada JW Lawton
J Jane (Retired) ARS-USDA
Iowa State University, Ames, IA, USA EA Lee
LA Johnson University of Guelph, Guelph, ON, Canada
Iowa State University, Ames, IA, USA EC Leonard
JM Jones AgriKnowHow, Maitland, SA, Australia
St. Catherine University, Arden Hills, MN, USA JS Li
A Juhasz China Agricultural University, Beijing, PR China
Centre for Agricultural Research of the Hungarian WD Li
Academy of Sciences, Martonvasar, Hungary Shanxi University, Taiyuan, PR China
BO Juliano Z Li
Philippine Rice Research Institute Los Banos, Laguna, CSIRO Food Futures National Research Flagship,
Philippines Canberra, ACT, Australia; CSIRO Plant Industry,
RS Kadan Canberra, ACT, Australia
(Deceased)
R Lindberg
GP Kadkol Minneapolis Heart Institute Foundation, Minneapolis,
NSW Department of Primary Industries, Calala, NSW, MN, USA
Australia
MG Lindhauer
LW Kannenberg Max Rubner-Institut (MRI), Detmold, Germany
University of Guelph, Guelph, ON, Canada
K Liu
TN Khan Agricultural Research Service, US Department of
University of Western Australia, Crawley, WA, Australia Agriculture, Aberdeen, ID, USA
BS Khatkar Q Liu
G. J. University of Science and Technology, Hisar, India Agriculture and Agri-Food Canada, Guelph, ON, Canada
JG Kling S Lu
Oregon State University, Corvallis, OR, USA National Chung Hsing University, Taichung, Taiwan
EJ Knights OM Lukow
New South Wales Department of Primary Industries, Agriculture and Agri-Food Canada, Winnipeg, MB,
Tamworth, NSW, Australia Canada
Contributors xxxi
BL Ma NJ Mendham
Agriculture and Agri-Food Canada, Ottawa, ON, University of Tasmania, Hobart, TAS, Australia
Canada
R Menon
C-Y Ma The General Mills Bell Institute of Health and Nutrition,
The University of Hong Kong, Hong Kong, PR China Minneapolis, MN, USA
A MacLeod TE Michaels
Canadian Grain Commission, Winnipeg, MB, Canada University of Minnesota, St. Paul, MN, USA
L MacLeod DM Miskelly
Dairy Innovation Australia, Werribee, VIC, Australia Westcott Consultants P/L, Goulburn, NSW, Australia
CK Madsen
CS Moffat
Aarhus University, Slagelse, Denmark
Curtin University, Bentley, WA, Australia
RJ Mailer
ES Monyo
Australian Oils Research, Lambton, NSW, Australia
ICRISAT, Nairobi, Kenya
JT Manful
Africa Rice Center, Cotonou, Republic of Benin TR Moore
AIB International, Manhattan, KS, USA
AG Marangoni
University of Guelph, Guelph, ON, Canada MK Morell
International Rice Research Institute, Los Banos,
DJ Mares Philippines
University of Adelaide, Glen Osmond, SA, Australia
CF Morris
L Marquart USDA-ARS Western Wheat Quality Laboratory,
University of Minnesota, St. Paul, MN, USA; Grains for Pullman, WA, USA
Health Foundation, St. Louis Park, MN, USA
LA Morrison
A Marti
Oregon State University, Corvallis, OR, USA
Universita degli Studi di Milano, Milan, Italy
B Maschoff WK Mousa
Grains for Health Foundation, St. Louis Park, MN, USA University of Guelph, Guelph, ON, Canada; Mansoura
University, Mansoura, Egypt
A Mathiowetz
University of Minnesota, St. Paul, MN, USA; Rochester, K Mrva
MN, USA University of Adelaide, Glen Osmond, SA, Australia
JJ Mayko L Munck
Canola Council of Canada, Mundare, AB, Canada University of Copenhagen, Frederiksberg C, Denmark
CM McDonough R Munns
Texas A&M University, College Station, TX, USA CSIRO Agriculture, Canberra, ACT, Australia; The
University of Western Australia, Crawley, WA, Australia
RA McIntosh
The University of Sydney, Cobbitty, NSW, Australia PA Murphy
Iowa State University, Ames, IA, USA
JD McKeehen
Cereal Partners Worldwide, Minneapolis, MN, USA ED Nafziger
J McLean University of Illinois, Urbana, IL, USA
The University of Newcastle, Callaghan, NSW, Australia SC Nelson
PBE McVetty AACC International (AACCI), St. Paul, MN, USA
University of Manitoba, Winnipeg, MB, Canada T Nleya
B Meints South Dakota State University, Brookings, SD, USA
Oregon State University, Corvallis, OR, USA
L OBrien
A Meldrum The University of Sydney, Narrabri, NSW, Australia;
Pulse Australia, Perth, WA, Australia Solheimar Pty Ltd, Narrabri, NSW, Australia
xxxii Contributors
DK OToole I Rajcan
City University of Hong Kong, Hong Kong, PR China University of Guelph, Guelph, ON, Canada
EA Oelke L Ramsden
University of Minnesota, St. Paul, MN, USA University of Hong Kong, Hong Kong, PR China
RP Oliver KD Rausch
Curtin University, Bentley, WA, Australia University of Illinois at UrbanaChampaign, Urbana, IL,
USA
DE Ortiz
W. K. Kellogg Institute, Battle Creek, MI, USA A Regina
MA Pagani CSIRO Food Futures National Research Flagship,
Universita degli Studi di Milano, Milan, Italy Canberra, ACT, Australia; CSIRO Plant Industry,
Canberra, ACT, Australia
TS Palav
Rich Products Corporation, Buffalo, NY, USA MN Riaz
Texas A&M University, College Station, TX, USA
L Pallas
Yanco Agricultural Institute, Yanco, NSW, Australia D Richard-Molard
(Retired) INRA, Nantes, France
GH Palmer
Heriot-Watt University, Edinburgh, UK F Rizza
Consiglio per la ricerca in agricoltura e lanalisi
RG Palmer delleconomia agraria, CRA-GPG, Fiorenzuola dArda,
Iowa State University, Ames, IA, USA Italy
RF Park MJ Robertson
The University of Sydney, Narellan, NSW, Australia CSIRO Sustainable Ecosystems, St. Lucia, QLD,
GM Paulsen Australia
Kansas State University, Manhattan, KS, USA LW Rooney
LC Paulson Texas A&M University, College Station, TX, USA
The Bama Companies, Tulsa, OK, USA WL Rooney
DS Petterson Texas A&M University, College Station, TX, USA
Department of Agriculture, Nedlands, WA, Australia CM Rosell
RA Porter Institute of Agrochemistry and Food Technology (IATA-
University of Minnesota-NCROC, Grand Rapids, CSIC), Paterna, Valencia, Spain
MN, USA AS Ross
L Privalle Oregon State University, Corvallis, OR, USA
Bayer CropScience, Research Triangle Park, Durham, SJ Roy
NC, USA School of Agriculture Food and Wine, The University of
KV Probst Adelaide, Glen Osmond, SA, Australia
Kansas State University, Manhattan, KS, USA; Grain HD Sapirstein
Processing Corporation, Muscatine, IA, USA University of Manitoba, Winnipeg, MB, Canada
KJ Quail AK Sarkar
Grain Growers Limited, North Ryde, NSW, Australia Canadian International Grains Institute, Winnipeg, MB,
RCN Rachaputi Canada
University of Queensland, Kingaroy, QLD, Australia
A Sarker
H Rahman International Center for Agricultural Research in the Dry
University of Alberta, Edmonton, AB, Canada Areas (ICARDA), New Delhi, India
S Rahman SK Sathe
Monash University Malaysia, Bandar Sunway, Malaysia Florida State University, Tallahassee, FL, USA
MN Raizada SF Schakel
University of Guelph, Guelph, ON, Canada University of Minnesota, Minneapolis, MN, USA
Contributors xxxiii
KA Scherf VA Solah
Deutsche Forschungsanstalt fur Lebensmittelchemie, Curtin University, Perth, WA, Australia
Leibniz Institut, Freising, Germany
AM Stanca
MP Scott University of Modena and Reggio Emilia, Reggio Emilia,
Iowa State University, Ames, IA, USA Italy
K Seetharaman FL Stoddard
(Deceased) University of Helsinki, Helsinki, Finland
GJ Seiler K-C Tan
USDA-ARS, Northern Crop Science Laboratory, Fargo, Curtin University, Bentley, WA, Australia
ND, USA
AS Tatham
SO Serna-Saldivar Cardiff Metropolitan University, Cardiff, UK
Centro de Biotecnologa FEMSA, Escuela de Ingeniera y
JRN Taylor
Ciencias, Monterrey, Mexico
University of Pretoria, Hatfield, South Africa
S Shea Miller
V Terzi
Eastern Cereals and Oilseeds Research Centre,
Consiglio per la ricerca in agricoltura e lanalisi
Agriculture and Agri-Food Canada, Ottawa, ON,
delleconomia agraria, CRA-GPG, Fiorenzuola dArda,
Canada
Italy
DB Sheats
K Tiefenbacher
St. Catherine University, St. Paul, MN, USA
(Retired)
PR Shewry
VK Tiwari
Rothamsted Research, Harpenden, UK; University of
Kansas State University, Manhattan, KS, USA
Reading, Reading, UK
SM Tosh
RD Shillito
Agriculture and Agri-Food Canada, Guelph, ON, Canada
Bayer CropScience, Durham, NC, USA
P Tosi
JP Shroyer
University of Reading, Reading, UK
Kansas State University, Manhattan, KS, USA
LC Trugo
KJ Shroyer
Federal University of Rio de Janeiro, Rio de Janeiro, Brazil
Kansas State University, Manhattan, KS, USA
SE Ulrich
Qingyao Shu
Washington State University, Pullman, WA, USA
Zhejiang University, Hangzhou, PR China
EH Unger
KHM Siddique
Eagle Rock Solutions, Inc., Idaho Falls, ID, USA
University of Western Australia, Crawley, WA, Australia
PV Vadlani
J Singla
Kansas State University, Manhattan, KS, USA
University of Zurich, Zurich, Switzerland
SA Valencia-Chamorro
M Sissons
Escuela Politecnica Nacional, Quito, Ecuador
NSW Department of Primary Industries, Calala, NSW,
Australia; Tamworth Agricultural Institute, Tamworth, V Vamadevan
NSW, Australia University of Minnesota, St. Paul, MN, USA; Agriculture
and Agri-Food Canada, Guelph, ON, Canada
JH Skerritt
University of Queensland, St Lucia, QLD, Australia N Van Heel
University of Minnesota, Minneapolis, MN, USA
DJ Skylas
Grain Growers Limited, North Ryde, NSW, A Vandenberg
Australia University of Saskatchewan, Saskatoon, SK, Canada
DL Smith SK Vasal
McGill University, Ste-Anne-de-Bellevue, QC, Canada BISA, Punjab, India
xxxiv Contributors
Contents
Functional Foods: Overview
Functional Foods: Dietary Fibers, Prebiotics, Probiotics, and Synbiotics
Nutrition: Soy-Based Foods
The World Health Organization (WHO) recommends to the food to which a component has been added to provide benefits;
public to move toward healthful diets. The functional-food a food from which a component has been removed by techno-
expansion across the globe is enormous with the current market logical or biotechnological means so that the food provides
size estimated from 7 to 190 billion USD. The ever-increasing benefits not otherwise available; a food in which a component
health-care costs and public awareness on roles of diet on health has been replaced by an alternative component with favorable
are contributing factors for such expansion. Functional-food properties; a food in which a component has been modified by
development involves evaluation of the roles, safety, and con- enzymatic, chemical, or technological means to provide a bene-
sumer acceptance of bioactive compounds. For foods that have fit; a food in which the bioavailability of a component has been
no history of consumption as regular diets (novel foods), the modified and combination of any of the above.
evaluation and regulatory requirements are rigorous and are In Japan, FOSHU refers to foods consumed as part of an
required to be substantiated by scientific evidence for their ordinary diet, containing functional ingredients and exerting
efficacy and safety. As a result, even though a number of novel health or physiological effect. FOSHU requires that safety
foods were developed in the past, their wider expansion was assessment, health functions, and claims must be approved
limited. by the Ministry of Health, Labour, and Welfare. Functional
Various grain-based functional foods are today on the mar- foods are also required to have three other fundamental fea-
ket in the form of whole grain snacks, baked foods, extruded tures: (1) nutritional functions, (2) sensory functions, and (3)
products, ready-to-eat breakfast cereals, fermented cereals, bev- physiological functions such as regulation of biorhythms, the
erages, and gruels. Other products are: dietary fibers, inulin- nervous system, the immune system, and body defense beyond
type fructans, b-glucans, carotenoids, o-3 fatty acids, plant nutrient functions. As of April 2011, in Japan, 955 products
sterols, and/or stanols-fortified products. There are also indig- were approved as FOSHU.
enous foods that are a heritage of a given community and their Based on extensive literature review and expert consultation,
record as safe diet is lost in antiquity and yet most of them A functional food is, or appears similar to be, a conventional
fulfill what is expected of functional foods. food. It is part of a standard diet and is consumed on a regular
In this article, functional food definitions, distinction from basis, in normal quantities. It has proven health benefits that
related concepts, recognized bioactive ingredients, dietary reduce the risk of specific chronic diseases or beneficially affect
sources, health benefits, and processing principles are target functions beyond its basic nutritional functions.
described giving emphasis on grains. Read also functional Even though there are slight variations, the following are
foods: dietary fibers, prebiotics, probiotics, and synbiotics as reflected on definitions given by different sources: (1) func-
these are not covered here. tional foods should be consumed as part of regular safe diet
not as drugs; (2) they are required to bear functionally bioac-
tive adequate ingredient(s) beyond basic nutrients; (3) diet
Functional Food: Concepts and Definitions with its functionally bioactive ingredient(s) should impact
positive health in an individual by either (i) decreasing occur-
So far, there is no global consensus on the definition of func- rences of disease(s), (ii) decreasing disease-causing factor(s), or
tional foods. The Codex Alimentarius Commission only issued (iii) promoting positive human physiological functions for
guidelines (CAC/GL 231997) on nutrition and health claims optimal health such as by maintaining body homeostasis
to be used through consultation with country policy. However, and/or through bolstering body immune systems; (4) they
food regulatory agencies and various professional societies in should improve quality of life; and (5) if there is no history
different countries have described closely related concepts/ of consumption as a regular diet, they must pass a strict regu-
working definitions. Definitions offered from various sources latory evaluation process for their safety and for beneficial
for functional foods, features, and requirements on what con- claim(s) declared substantiated with sound scientific evidence.
stitute to be nutrition, health, and structure/function claims
are listed under the Further Reading section.
According to the ADA, all foods are functional at some Other Concepts Related to Functional Foods
physiological level because they provide nutrients or other
Nutraceuticals
substances that furnish energy, sustain growth, or maintain/
repair vital processes. However, functional foods are recog- These are distinct from functional foods and defined as a
nized to provide additional health benefits that may reduce product isolated or purified from foods generally sold in
disease risks and/or promote optimal health. Functional foods medicinal forms (i.e., as pills, powder, syrups, or other medic-
include: (1) conventional foods or whole foods, (2) modified inal forms) that have specific health benefits.
foods, (3) medical foods, and (4) foods for special dietary use.
According to Functional Food Science in Europe, Functional
Dietary Supplements
foods are those satisfactorily demonstrated to affect beneficially
one or more target functions in the body, beyond adequate These are products other than tobacco intended to supplement
nutritional effects, in a way that is relevant to either an improved diet, which contain one or more of the following dietary ingre-
state of health and well-being and/or reduction of risk of disease dients: vitamin, mineral, herb or other botanical, amino acid,
when consumed as part of normal food pattern. Thus, func- concentrate, metabolite, constituent, extract, or combinations
tional foods include: a natural, unmodified food; a food in of these ingredients; they are ingested in pill, capsule, tablet, or
which one of the components has been enhanced through spe- liquid form; they are not represented for use as conventional
cial growing conditions, breeding, or biotechnological means; a food or as a sole item of diet.
FOOD GRAINS AND WELL-BEING | Functional Foods: Overview 3
Novel Foods phytosterols and triterpene alcohols) in rice (Figure 2). Alkyl-
resorcinol metabolites are currently studied for use as
These are defined as raw foods or food materials with no
biomarkers of whole-grain intake. Bran also contains vitamins,
history of safe use, for example, genetically modified foods;
phytosterols, minerals, proteins, lipids, carotenoids, tocols,
foods produced by algae, fungi, or microorganisms; foods
and phytic acids. Consumption of grain foods along with
isolated from plants and animals without a history of safe
their bran is beneficial due to the presence of various bioactive
use; and foods that have been processed where the process
compounds. Functional roles documented for phenolics
has dramatically changed the food.
include: bolstering of cellular antioxidant defenses, protection
from cardiovascular disease (CVD) risks, brain wellness, anti-
Bioavailability neurodegenerative, antiulcer, anticarcinogenic, antimutagenic,
anti-inflammatory, antimicrobial properties; and improve-
This refers to a fraction of a given compound and/or its metab- ment of eye health and vision, muscle performance, and
olite that reaches systemic circulation and involves gastrointes- immune responses.
tinal digestion, absorption, metabolism, and tissue Phenolics are generally partially bioavailable and their
distribution. Bioavailability encompasses bioaccessibility and properties are influenced by their structures and solubility.
bioactivity. Most soluble phenolics are found unbound, whereas insolu-
bles are found bound to cell wall polysaccharides or proteins
Bioaccessibility where they form stable complexes. Insoluble phenolics are not
absorbed in the small intestine, but they are fermented by
This refers to the quantity or fraction of a bioactive that is colon microbiota yielding largely ferulic acid followed by
released from a food matrix in the gastrointestinal tract and p-coumaric acid. Within cells, phenolics suppress reactive-
becomes available for absorption. It includes digestive trans- oxygen (ROS) and nitrogen species (RNS) by donating hydro-
formations, absorption/assimilation into intestinal epithelium gen atoms or electron, quenching free radicals, and activating
cells, and also the presystemic metabolism (both intestinal and endogenous antioxidant enzymes. They also chelate metal pro-
hepatic). oxidants. Oxidative stress can happen when generation of
oxidants exceeds the natural antioxidant systems. Oxidative
stress causes damages to DNA, RNA, lipids, proteins, and car-
Bioactivity
bohydrates and leads to different types of chronic diseases. In
This refers to a specific response, biomarker, or effect observed cereal grains, anti-oxidant potentials follow pigmented grains
upon exposure to the bioactive substance. It includes tissue (tannin sorghums, black rice, brown and black sorghums, teff,
uptake and the subsequent physiological response. millets, blue maize) > non-pigmented grains (white-sorghum,
wheat, rice). Even though, phenolics are shown to suppress
cancers in in vitro in cells and in vivo in animal models, studies
Recognized Bioactive Compounds in Functional Food in human intervention are less clear. The synergetic effects are
Phenolics (Polyphenolics) believed to have a role. Phenolics are also used as antimicrobial
food preservatives, food colorants, and flavor imparters. Tan-
Phenolics are ubiquitous in plants and more than 8000 com- nins, despite having high antioxidant activity, are antinutri-
pounds are known. Dietary phenolics can be broadly divided tional and inhibit digestive enzymes and mineral absorptions.
into flavonoids (anthocyanins, flavonols, flavones, flavanones,
isoflavones, flavan-3-ols, condensed tannins or proanthocya-
nidins, lignans, chalcones, coumarins, coumestans, and aur- Phytoestrogens
one) and nonflavonoids (simple phenols, phenolic acids
(hydroxybenzoates and hydroxycinnamates), and stilbenes) Phytoestrogens are nonsteroidal plant compounds with struc-
(Figure 1). Flavonoids are the largest group and more than tures and biological activities similar to steroid hormone estro-
6000 compounds have been identified. Hydrolyzable tannins gen, 17b-estradiol (Figure 3). Functional roles documented
are derivatives of gallic acids. They exert deleterious effects and include: influence on growth and functioning of female and
may be found at trace levels in some edible foods. Flavonoids male reproductive tissues; maintenance of skeletal and central
often exist as glycosides with sugar molecules attached often at nervous systems; cardio-protection; suppression of colon,
position 3, sometimes at 5, 7, 40 , 30 , and 50 and hydroxyls at 40 , breast, and prostate cancers and skin aging; and relief from
5, and 7 on the flavonoid basic structure. Flavonoids without menopausal symptoms. At low concentration, they act like
sugars are called aglycones. estrogens (agonists) and at high doses as blockers (antago-
Phenolic acids and various flavonoids in whole grains and nists). Phytoestrogens are potential endocrine disrupters and
bran fractions (pericarp, aleurone layer, testa, and germ) are become cytotoxic at high dose. Groups of compounds recog-
extensively documented. Bran is removed during cereal grain nized as phytoestrogens are lignans, isoflavones, coumestans,
milling for production of refined flours. Among phenolic and stilbenes.
acids, ferulic acids are dominant (about 80% to 90%) in cereal
grains. Sorghum is known to have unique 3- Lignans
deoxyanthocyanins. Other unique phenolics documented in These are 18-carbon skeleton compounds composed of two
grains are alkylresorcinols in rye, wheat, triticale, and barley, phenylpropanoid units with various functional groups con-
avenanthramides in oats, and g-oryzanols (ferulic cid esters of nected to benzene rings (Figure 4). Secoisolariciresinol,
4 FOOD GRAINS AND WELL-BEING | Functional Foods: Overview
OH O OH
Derivatives of simple phenol C Derivatives of hydroxybenzoic acid
R1 Phenol: R1=R2=R3=H Gallic acid: R1=H, R2=R3=R4=OH
R1 Gentisic acid: R1=R4=OH, R2=R3=H
Catechol: R1=OH, R2=R3=H
Salicylic acid: R1=OH, R2=R3=R4=H
R2 R3 Resorcinol: R1=R3=H, R2=OH R4 p-(OH) Benzoic acid: R =R =R =H, R =OH
R2 1 2 4 3
Phloroglucinol: R1=H, R2=R3=OH
Protocatechuic acid: R1=R4=H, R2=R3=OH
OH R3
Syringic acid: R1=H, R2=R4=OCH3, R3=OH
C Derivatives of hydroxycinnamic acid Vanillic acid: R1=R4=H, R3=OH, R2=OCH3
R1 O Cinnamic acid: R1=R2=R3=R4=H
o- Coumaric acid: R1=OH, R2=R3=R4=H
m- Coumaric acid: R1=R3=R4=H, R2=OH
R2 R4
p- Coumaric acid: R1=R2=R4=H, R3=OH
R3 Caffeic acid: R1=R2=H, R3=R4=OH
Ferulic acid: R1=R2=H, R3= OH, R4 =OCH3
Sinapic acid: R1=H, R2=OCH3, R3=OH, R4=OCH3
Phenolic acids
Stilbene
3`
2`
4`
8
1 O
7 O 2 5` O O
6` Flavan-4-o1
6 3 OH OH
5 4 OH
Flavonoid basic structure Flavan-3-o1 Flavonol
O O
OH
O
OH
+ Flavone Flavanone
O O
HO O
O O
OH
Anthocyanidin Isoflavone O
OH O Aurone
O O
O O O Lignan
Chalcone Coumarin
Coumestan
O
Basic structures of main class flavonoids and related structures (aurone, lignan)
OH
OH
HO O
R OH n:0 to many
OH OH
OH
OH
HO O
HO +O
OH
OH
OH
OH
OH HO O
Sorghum 3-deoxyanthocyanidins
(Apigenindin, R=H and Luteolinidin, R=OH) OH
OH Proanthocyanidins
R1 O
OH R2
N
H
O C OH R3
HO R Avenanthramides (more than 35 forms are identified)
Alkylresorcinols (R = alkyl chain, Avenanthramide 2p, R1=R3=OH, R2=H
odd carbon C15 to C27) Avenanthramide 2f, R1=R3=OH, R2=OCH3
Avenanthramide 2c, R1=R2=R3=OH
O
CH3O C OR
O
CH3O C O
HO
Gamma-oryzanol (ferulic acid esters,
HO
R = phytosterols or triterpene alcohols) Gamma-oryzanol (campesteryl ferulate)
OH OH O
OH
H H
H H H H H H
HO HO HO
17 beta-Estradiol Estriol Estrone
HO OH
O OH
H
H CH2O
CH3O CH2OH CH3O HO OH
H OH
H
HO CH2O O
HO CH2OH
OH
OH
OCH3 OCH3
Secoisolariciresinol diglycoside
Secoisolariciresinol OH
OH
O
H O
O H
CH3O O
O HO
O O
HO Sesamolin
O O
Matairesinol O OH O
O
O H
OCH3 H O
O O
OH Sesamol O
Sesamin O
H H
HO OH HO
O
OH
Enterolactone O
Enterodiol
OH OH
matairesinol, sesamin, and sesamolin are typical examples. function and maintenance of menopausal health in women, and
When two glucose molecules are attached to the hydroxyl also suppress CVD by lowering total cholesterol and LDL
groups of the propanol chain of secoisolariciresinol, the com- cholesterol and raising HDL in the blood vessels. However, the
pound is called secoisolariciresinol diglucoside (SDG). Lig- isoflavone content of soybean products varies with processing.
nans have higher antioxidant potential than vitamin E. Water- and alcohol-extracted soybean protein concentrate and
Lignans are implicated to have potential use in hormone soybean oils have insignificant levels of isoflavones.
replacement therapy, to maintain good cognitive function in
postmenopausal women, and to have anticancer activities Coumestans
(breast, prostate, colon, and skin cancers). They may contrib- Coumestans are less abundant than isoflavones in human
ute to the reduction of hypercholesterolemia, atherosclerosis, diets. They are known to have stronger estrogenic activity
and diabetes. SDG is converted by bacteria in the human than isoflavones. Typical examples are coumestrol and 40 -
colons to mammalian lignans enterodiol and enterolactone, methoxycoumestrol (Figure 6). Coumestrol and genistein have
which are also antioxidants. Enterodiol and enterolactone are higher binding affinities to ERb than other phytoestrogens.
purported to reduce growth of cancer tumors, especially Coumestrol has been reported to inhibit bone resorption and
hormone-sensitive ones. Grain sources of lignans include oil- stimulate bone mineralization by in vitro studies. Among grains,
seeds (flax, rapeseed, and sesame), whole-grain cereals (wheat, chief sources are soy sprouts, mung bean, and sunflower seeds.
oats, rye, barley, and millets), and legumes (soybean). Flaxseed
is the richest source of SDG and also contains other lignans Stilbenes
(matairesinol, pinoresinol, lariciresinol, and isolariciresinol). Stilbenes are C6C2C6 compounds with H,OH, and OCH3
Baked products and salad dressings fortified with flax and as substituents on their two benzene (C6) rings (Figure 7).
sesame seeds can be considered as functional foods rich in Stilbenes are known to be anticarcinogenic, antioxidative,
lignans antimutagenic, antitumor, and antiosteoporotic. Diets rich in
resveratrol, which exists in both trans and cis isomers of the
Isoflavones stilbene, are purported to prevent CVD, neurodegenerative dis-
Isoflavones are often found as glycosides. Common isofla- orders such as Parkinsons and Alzheimers disease, and various
vones include daidzin, genistin, biochanin A, and formonone- cancers. Resveratrol has stronger antioxidant activity than propyl
tin (Figure 5). The glycosides are converted to aglycones by gut gallate, vanillin, phenol, 2,6-di-tert-butyl-p-hydroxytoluene
microflora. Among isoflavones, studies with genistein on (BHT), and a-tocopherol. Among grains, stilbenes are found in
breast cancer cell lines show cell proliferation at low doses Fabaceae and Poaceae. Other good sources are peanuts, grape
and inhibition at high concentrations. Isoflavones are also skins, and notably some sorghum varieties.
recognized as mild antioxidants.
Isoflavones are exclusively documented in leguminosae of
which soybean is the richest source. Other sources are kidney, Phytosterols
navy, pinto, red, small white, and mung beans and chickpeas,
split peas, peanuts, sunflower seed, and walnut. Raw soybeans Phytosterols are natural components of human diets, are cho-
can contain 1.24.2 mg g1 dry weight isoflavones, while high- lesterol-like and lipophilic, and comprise sterols and stanols.
protein soy flour may contain up to 1.11.4 mg g1 dry weight. Stanols are hydrogenated products of sterols. Of more than
Diets rich in soybeans are implicated to improve cognitive abil- 250 sterols and stanols known to exist in nature, only six
ities, prevent osteoporosis and various cancers, improve immune (campesterol, campestanol, stigmasterol, sitosterol, sitostanol,
and brassicasterol) are dominant in seed oils (rapeseed,
soybean, corn, and sunflower oils), other grains (corn, rye,
R1 O R2 Isoflavones
R1 R2 R HO
Geinstein OH OH R OH
HO O Daidzin H OH
Biochanin A OH OCH3 R HO
Formononetin H OCH3 Stilbenes,R = H, OH or OCH3 Resveratrol
OH O OCH3
O
HO O HO O O
O
Coumestrol 4`-M ethoxycoumestrol
R 26
18 R 20
12 20 23
11 21 20 22 25 21 24
17 16 21 20 22 23 24 25 26 23 26 25
19 13 24 22 27
1 9
H 14 15
2 27 Brassicasterol
27
10 8
Cholesterol/stanol Sitosterol/stanol
5 7
HO 3
4 6 18 R20
12
Sterols 11 21 20 22 23 25 26
17 16 21 20 22 23 25 26
19 24
1 9
H 13
14 15
24
2 27
27
10 8 Stigmasterol/stanol
5
Campesterol/stanol
7
HO 3
4 6
(a) Stanols
C O
Sitosteryl stearate
O
H
H3CO
HO CH CHCOO
Sitostanyl ferulate
H
CH2OH
O
OH O
HO Steryl glycoside (SG)
OH
O
C O CH H
2
O
OH O
HO Acylated steryl glycoside (ASG)
(b) OH
Figure 8 (a) Structure of plant stanols and plant sterols. (b) Conjugated structure of plant stanols and plant sterols.
wheat, barley, millets, rice, oats, and peanuts), and tree oils humans remain largely unknown. Plant sterols/stanols are
(Figure 8(a)). In plant tissues, sterols and stanols can exist as also implicated to have mild anti-inflammatory effects and
conjugates often esterified with fatty acids, oleic, or linoleic in are not toxic at physiological concentrations.
five forms (Figure 8(b)).
Plant sterols and stanols are known to attenuate absorption
of low-density lipoprotein cholesterol (LDLC), a risk factor for Carotenoids
premature atherosclerosis and CVD. Consumption of 2 g d1
of plant sterols or stanols as part of diet is known to lower Carotenoids are lipid-soluble tetraterpene hydrophobic
LDLC plasma levels by about 10%. Consumption of low cho- compounds, which impart yellow, orange, or red colors, and
lesterol and saturated fat along with high stanols or sterols can found associated with lipid portions in human cells. Carotenoids
reduce LDL by 20%. Statin medication along with sterols or are divided into two groups (Figure 9): carotenes (hydrocarbons)
stanols is more effective in lowering LDL than doubling statin and xanthophylls (oxygenated carotenes). Carotenoids bear con-
dose. Ranking toward blocking of cholesterol absorption is as jugated double bonds in their structures, which are responsible
follows: free-form stanols > stanol esters > free-form sterols > for color formation and reactive oxidant stabilization. Caroten-
sterol esters due to structural and metabolic reaction differ- oids are antioxidants, and functional foods processed from carot-
ences toward gut microorganisms. The precise mechanisms of enoids are helpful for prevention of CVD and cancers. b-carotene,
cholesterol-lowering effect by individual compounds in a-carotene, and b-cryptoxanthin are provitamin A compounds.
8 FOOD GRAINS AND WELL-BEING | Functional Foods: Overview
H3C CH3
CH3 CH3
CH3
CH3 CH3 H3C
H3C CH3 Beta-cryptoxanthin
H3 C CH3
HO CH3 CH3
CH3
CH3 CH3 H3C OH
CH3 Lutein*
H3C
H3C CH3
HO CH3 CH3
CH3
CH3 H3C
CH3 CH3 O
H3C CH3
OH Retinoic acid
CH3
CH3 CH3 O CH3 CH3
H3C CH3 H3C CH3
H OH
Retinal Retinol
CH3 CH3
Of more than 700 carotenoids identified in nature, 24 occur protective against age-related macular degeneration and cata-
commonly in human foods and 40 have been detected in racts, a leading cause of eye blindness. Cereal-based lutein and
human milk, serum, and tissues. zeaxanthin-containing functional foods processed from yellow
Among grains, chief sources are yellow maize, durum maize, durum wheat, golden rice, einkorn, and maize bran
wheat, einkorn, and golden rice. Carotenoid contents can include high-lutein bread, high-lutein cookies, high-lutein
vary depending on varieties, cultivation conditions, geographic muffins, corn tortilla, corn chips, and extruded products.
locations, part of plant utilized, maturity stage, food proces-
sing, and storage conditions used. Heating, cooking, and
mechanical disruption and the presence of fats/oils can
enhance bioavailability of carotenoids. Carotenoid activities Omega (v)-3 Fatty Acids
can be destroyed by isomerization and oxidative reactions.
Lutein and zeaxanthin are not synthesized by humans. They Omega-3 fatty acids are long-chain polyunsaturated fatty acids
can be supplied from grains (yellow maize, durum wheat, (PUFA) (C18-C22) commonly designated as o-3 fatty acids
golden rice, and einkorn), vegetables (basil, parsley, spinach, (Figure 10). The o-3 and o-6 fatty acids are essential nutrients.
kale, leek, and red pepper), yellow egg yolks, and human milk. Alpha-linolenic acids (ALA, o-3) derived from plant sources
Along with their metabolite meso-zeaxanthin, both are are precursors for physiologically important longer-chain
FOOD GRAINS AND WELL-BEING | Functional Foods: Overview 9
O
C OH
ALA (C18:3 9,12,15)
O
C OH
EPA (C20:5 5,8,11,14,17)
O
C OH
DHA (C22:6 4,7,10,13,16,19)
PUFAs: eicosapentaenoic (EPA, o-3) and docosahexaenoic acid such best candidate because it is always consumed as whole
(DHA, o-3). grain often fermented as injera with insignificant antinutrients
Omega-3 fatty acids are components of cell membranes. in it and in many respects it supplies better nutrients for celiac
Their documented health benefits include suppressions of patients. Other approaches to gluten-free bread quality
CVD, cancer, type 2 diabetes, depression and stress, inflamma- improvements are the use of gluten-free flours and starches,
tory diseases, and (6) Alzheimers disease. They are also bene- incorporation of nutritional ingredients, and use of structure
ficial in infant nutrition for normal brain, eye, and mammalian modifier additives and processing technologies that will
nervous system development and function. Omega-3 fatty change the allergenic glutens.
acids are available as supplements from pharmaceuticals. Other bioactive compounds to be noted include tocols
However, consumers prefer o-3 fatty acids from diets. In this (tocopherols and tocotrienols) from cereal germ and aleurone
respect, there is high opportunity for o-3 fatty acid-containing layers; betaine and choline from wheat, barley and rye germ,
functional foods. The main source of ALA is flaxseed. Other and brans; and benzoxazinoids from whole-grain wheat and
sources are canola, soybean oils, and walnuts. In flaxseed, rye. Some bioactive compounds known for their antinutri-
bioavailability of ALA is as follows: oil > milled > whole tional effects tend to portray beneficial effects under specific
flaxseed. Despite fish oil being a major source of EPA and conditions. They include glucosinolates from rapeseed, phytic
DHA, it is sensitive to autoxidation, and there is also a possi- acids from germ and aleurone layers of grains, and lectins
bility for methyl mercury toxicity since fish has potential for (hemagglutinins) from legumes and whole cereal grains.
contamination with this toxin. Where consumption of fish oil
is limited and methyl mercury toxicity is a possibility, EPA and
DHA are required to be supplied from ALA even though con-
Conclusions
version of ALA to EPA and DHA is known to be inefficient.
Baked foods like bread and injera and various cereal grain
Opportunities for grain-based functional foods are high
snacks processed by incorporating flaxseed oils and/or milled
because grains are sources of many bioactive compounds that
flaxseed flours are an alternative way of supplying o-3
are located in specific anatomical structures. However, bio-
fatty acids.
availability can be influenced by many factors including
forms of existence, processing technology, and food matrices
that prevail during consumption. Functional foods are recog-
Gluten-Free Foods nized to be consumed as varied regular diets. However, the
absence of a definition that has received global consensus
In some genetically predisposed individuals, gluten proteins potentially poses various risks such as inaccurate health claims,
from wheat, rye, barley, and triticale when ingested will lead to adverse reactions, poor quality control during manufacturing,
celiac disease (CD). Oats are also not considered free, because allergic reactions, and carcinogenicity. Novel foods in particu-
of potential contamination and the possibility that some peo- lar may pose unprecedented risks because they do not have a
ple who are intolerant to gluten are not immune to it. CD long history as diets to populations of different genetic back-
involves interactions between genetic susceptibility (HLA- ground. A distinction between novel and functional foods may
DQ2 and HLA-DQ8 alleles), gluten proteins, and an help to reduce such effects. Misuse of functional foods can
immunologic response that causes small intestinal mucosal lead to adverse effects such as anti-nutritional, thyroid toxicity,
damage characterized by complete loss of absorptive villi to drug interactions, allergic reactions and geno-toxicity/
maldigestion and malabsorption of nutrients. Such effects lead carcinogenicity.
to chronic diseases like skin lesions, anemia, diarrhea, weight A given diet is believed to bear more than 25 000 bioactive
loss, osteoporosis, fatigue, infertility, and cancer. About 1% of constituents. Minimum dietary intake that would lead to desir-
the world population is estimated to be gluten-intolerant. able health effect and maximum intake to avoid adverse effects
There is no drug for CD, and the remedy is a complete adher- are yet not available for most bioactive compounds. A distinc-
ence to a gluten-free diet (GFD). The Codex Alimentarius Com- tion is yet to be clearly established for most bioactive com-
mission and FDA have set standards for GFD to be pounds whether a desirable outcome is caused by a single
<20 mg kg1 gluten. Gluten proteins are vital for production compound or by synergic effects. Since they are consumed as
of aerated baked products to which consumers are attracted. regular varied diets for health promotion, designing functional
The use of baked products from gluten-free cereal grains (teff, foods for synergic effects of bioactive compounds seems ben-
maize, rice, sorghum, and the millets) particularly those pro- eficial. While designing for an optimum supply for those
cessed from whole grains is a remedy for CD. Teff grain is one health effects is established, caution is also required not to
10 FOOD GRAINS AND WELL-BEING | Functional Foods: Overview
affect other components used to be served in a given regular Diseases: Celiac Disease; Cereal Allergens; The Gluten-Free Diet;
diet. With an accurate definition in place, functional foods can The Basics: Grain: Morphology of Internal Structure.
be better regulated to improve health and well-being.
Learning Objectives
Dietary Fibers and Whole Grains
To achieve understanding of the definitions of DFs, whole
grain foods, prebiotics, probiotics, and synbiotics. Dietary Fibers
To achieve understanding of DF and prebiotic grain sources. DFs are a structurally diverse group of compounds. Various
To impart principles of DFs, prebiotics, probiotics, and regulatory bodies and professional societies American Asso-
synbiotics with respect to the processing of functional foods. ciation of Cereal Chemists International (AACCI), Institute of
To explain linkages of the intake of DFs, prebiotics, probio- Medicine (IOM), Food Standards Australia and New Zealand
tics, and synbiotics to health. (FSANZ), FAO/WHO Codex Alimentarius Commission, Euro-
pean Food Safety Authority, and Health Canada have offered
definitions for DF primarily based on the physiological roles
Introduction and analytical methods used to isolate and quantify them.
According to Codex, DFs are carbohydrate polymers with 10
Functional foods processed from DFs are linked to various or more monomeric units (decision on whether to include
health benefits. Soluble fibers and b-glucans in particular are carbohydrates of 39 monomeric units should be left up to
known to modulate hyperlipidemia, hyperinsulinemia, national authorities), which are not hydrolyzed by the endog-
hyperglycemia, and obesity, which are risk factors for chronic enous enzymes in the small intestine of humans and comprise
diseases including cardiovascular diseases (CVDs), diabetes the following:
mellitus, and cancers. Soluble fibers are rapidly fermented by
(1) Edible carbohydrate polymers naturally occurring in the
gut microbiota. Adequate intake (AI) of insoluble fibers is also
food as consumed. When derived from a plant origin, DF
linked to the reduction of colon cancers and improved gastro-
may include fractions of lignin and/or other compounds
intestinal health (GIH), reduced CVD, and obesity. Cereal
protein fractions, phenolic compounds, waxes, saponins,
grain fibers and legume cotyledons are included in bread,
phytates, cutin, vitamins, phytosterols, minerals, etc.
extruded products, cakes, cookies, and fermented and other
when such compounds are associated with the polysac-
ready-to-eat breakfast cereal (RTE) products. Consumption of
charides in the plant cell walls and extracted together.
whole grain foods is encouraged for AI of DF. Health-
(2) Carbohydrate polymers, which have been obtained from
promoting effects of DF are imparted by nonstarch carbohy-
food raw materials by physical, enzymatic, or chemical
drates, resistant starches, and closely associated/attached
means and which have been shown to have a beneficial
compounds. Those DFs that selectively promote growth and/
physiological effect to health as demonstrated by generally
or activities of probiotics and promote host health are pre-
accepted scientific evidence to competent authorities.
biotics. Arabinoxylans, fructans, inulin, resistant starches, b-
(3) Synthetic carbohydrate polymers, which have been shown
glucans, and a-galactooligosaccharides are fermented by pro-
to have a beneficial physiological effect to health as dem-
biotics to short-chain fatty acids (SCFAs): butyrate, propionate,
onstrated by generally accepted scientific evidence to com-
and acetate that are absorbed in the colon to furnish additional
petent authorities.
energy for the host. Prebiotics are also known to improve the
absorption of calcium and magnesium and to suppress colon The Codex definition is expected to be used in all jurisdictions.
cancers and absorption of triglycerides and cholesterol. Cellu- Currently, there is wider consensus that the AACCI definition is
lose and water-insoluble arabinoxylans are major contributors reconciling with Codex definition. AACCI defines DF as edible
to laxative effects and fecal bulking. Barley, wheat, rye, oats, parts of plants or analogous carbohydrates that are resistant to
triticale, maize, einkorn, chickpea, and lentils are notable digestion and absorption in the human small intestine with
sources of prebiotics. Intake of prebiotics is also achieved complete or partial fermentation in the large intestine. DFs
through consumption of whole grains like teff, millets, include polysaccharides, oligosaccharides, lignin, and associated
sorghums, and brown rice. Dietary intake of probiotic bacteria plant substances. DFs promote beneficial physiological effects
mainly from the genera of Lactobacillus and Bifidobacterium is including laxation and/or blood cholesterol and glucose attenu-
linked to improved immune capacity and GI health and to ations. DFs thus comprise the following:
i. Nonstarch polysaccharides and resistant oligosaccha- secretion of saliva and gastric juice resulting in an expansion
rides: cellulose, b-glucans, hemicellulose (arabinoxy- of the stomach and increased satiety; and (3) reducing absorp-
lans and arabinogalactans), polyfructoses (inulin and tion efficiency of the small intestine. Diets with high DF con-
oligofructans), galactooligosaccharides, gums, tents are known to have slow nutrient digestion and gastric
mucilages, and pectins emptying rates. This action leads toward attenuation of blood
ii. Analogous carbohydrates (indigestible dextrins/resis- glucose and helps in the control of type 2 diabetes mellitus.
tant maltodextrins from corn and other sources and Some DFs are fermented (prebiotic) completely or partially by
resistant potato dextrins), synthesized carbohydrate the symbiotic microorganisms of the large intestine to SCFA-
compounds (polydextrose, methylcellulose, and like butyrate, which is the preferred energy source for mucosa
hydroxypropyl methylcellulose), and resistant starches cells. Inulin and oligofructans are not used by potential path-
iii. Lignins ogenic bacteria but are the preferred substrate for beneficial
iv. Substances associated with the nonstarch polysaccha- Bifidobacteria and thus help to maintain and restore the balance
rides and lignin complex in plants (waxes, phytates, of healthful gut microbiota. Beta-glucans and the presence of
cutins, saponins, suberin, polyphenols, tannins, magnesium in whole grains and/or bran-rich DF are known to
vitamins, minerals, alkylresorcinols, carotenoids, beta- lead to improved insulin sensitivity to suppress type 2 diabetes
ine, etc.) and other metabolic syndrome diseases. Psyllium (or
The classification of DFs as soluble (gums, oligosaccharides, ispaghula; Plantago ispaghula or Plantago ovata) fibers and oat
pectins, water-soluble arabinoxylans, b-glucans, galactoman- and barley b-glucans are used in functional foods targeted at
nans, psyllium fibers, and alginates) and insoluble (cellulose, lower cholesterol and blood sugars for improved cardiovascu-
lignins, and water-insoluble arabinoxylans) also continues to be lar health, control of type 2 diabetes, and weight reduction and
in use. DFs for human nutrition are largely derived from cereal maintenance. Cereal grain DFs are generally recognized to be
grains 44%, vegetables 21%, fruits 13%, and legumes 10%. The more potent than fruits and vegetable fibers in the many
outer pericarp of cereal grains, endosperm cell walls, seed coat beneficial physiological roles of the DF functional foods.
and cotyledons of legumes, psyllium husks, and fruit and vege- Excessive DF intake through functional foods or other regular
table skins are rich sources of DF. Diets rich in bran (pericarp, diets may lead to deleterious effects because it contributes to
aleurone layer, and testa) and germ are known to have high reduced absorption of some vitamins, minerals, proteins, and
antioxidant activities than diets based on refined white flours energy. It may also lead to diarrhea, flatulence, bloating, and
largely due to associated DFphenolics, tocols, and carotenoid abdominal discomfort. Children less than 1 year old and older
antioxidants. Functional foods containing DFs are currently people, in particular, are advised not to consume higher than
widely available in the form of breads; extruded products like recommended DF levels. The recommended AI for total DF is
pasta, cakes, and cookies; and fermented and other RTE prod- 1925 g day 1 for children aged 18 years, 3138 g day 1 for
ucts. Although incorporation of DF to some levels has produced boys aged 918 years, 3830 g day 1 for males greater than 19
products comparable to those processed from refined flours, years old, 26 g day 1 for girls aged 913 years, 2521 g day 1
increased DF contents have been associated with reduced con- for females greater than 19 years old, 28 g day 1 at pregnancy,
sumer acceptance (color, texture, and taste). Currently, improve- and 29 g day 1 during lactation.
ments being researched include the use of enzymes and other
physicochemical means to modify fiber structures and incorpo-
Whole Grains
ration of fibers from fruits and vegetable skins.
The beneficial physiological effects conferred by AI of DF Whole grains are defined by AACCI in 1999 as consisting of
include attenuation of blood cholesterol and blood sugars, the intact, ground, cracked or flaked caryopsis, whose principal
suppression of CVD diseases (coronary heart disease, stroke, anatomical components the starchy endosperm, germ, and
and hypertension), weight control, improved insulin sensitiv- bran are present in the same relative proportions as they exist
ity and suppression of type 2 diabetes mellitus, improvement in the intact caryopsis. Thus, whole grains are good sources of
of GIH, improved laxation, suppression of cancers such as various nutrients, starch, proteins, lipids, DF, vitamins (thia-
colon and breast cancers, improved immunity, and contribu- mine, riboflavin, niacin, pantothenic acid, vitamin B6, folate,
tion to the overall longevity. The physiological roles of DF can and vitamin E), minerals, and phytochemicals such as various
vary depending on nature of the compound. Properties of DF phenolic compounds, phytosterols, tocols, lignans, and carot-
such as viscosity, molecular weight, molecular size, solubility, enoids. AI of whole grain foods has been associated with GIH
water-holding capacity, and fermentation characteristics by gut and reduced risk of various metabolic syndrome-related dis-
microbiota are known to play a role in the physiological effects eases (obesity, diabetes, hypertension, and dyslipidemia) and
of DF-based functional foods. Increased intake of soluble fibers various cancers. For example, consumption of two to three
is known to suppress hyperglycemia and to improve insulin servings per day (about 48 g) of whole grains is preventive of
sensitivity in nondiabetic and diabetic individuals. Cardiovas- the risks of CVD, type 2 diabetes mellitus, overweight, and
cular protective roles of DF are imparted by attenuation of obesity. Whole grain intake results in greater intake of diverse
blood cholesterol levels specifically by interfering and limiting nutrients. Although whole grain has been defined, it remains
cholesterol fractions transported by LDL. The viscous charac- challenging for researchers, industry, regulatory authorities,
teristics of fiber are implicated to have a dominant role in this and consumers to clearly identify what constitutes whole
regard. Energy load reduction and weight control are imparted grain foods. Following a review of evidence from various epi-
by (1) displacing available energy and nutrients from the diet; demiological, interventions, animals, and human clinical stud-
(2) increasing chewing, limiting intake, and promoting ies on relationship of whole grain intake and the many benefits
FOOD GRAINS AND WELL-BEING | Functional Foods: Dietary Fibers, Prebiotics, Probiotics, and Synbiotics 13
on health and wellness, an expert panel of whole grain working given by FAO/WHO (2001) as Live microorganisms which
committee in 2014 has recommended that 8 g of whole grain when administered in adequate amounts confer a health ben-
per 30 g serving (27 g per 100 g), without a fiber requirement, efits on the host was recommended in August 2014 by Inter-
be considered a minimum content of whole grains that is national Scientific Association for Probiotics and Prebiotics
nutritionally meaningful and that a food providing at least (ISAPP) to be used with minor rewording as Live microorgan-
8 g of whole grains per 30 g serving be defined as a whole- isms that, when administered in adequate amounts, confer a
grain food. This definition also reconciles with the one offered health benefit on the host. Major bacterial strains commonly
by AACCI that whole grain food product must contain 8 g or used in probiotics are lactic acid bacteria (LAB) from the genera
more of whole grain per 30 g of product. Sources of whole of Lactobacillus and Bifidobacterium (Table 1). Other LAB
grain functional foods include whole wheat and rye flours, reported are from Pediococcus, Lactococcus, Leuconostoc,
barley and whole barley flour, whole grain maize meal, Streptococcus, and Weissella. Certain yeast and fungal strains
grits and popcorn, teff, sorghum, oat groats, whole grain are also recognized for probiotic effects. The ISAPP restricted
millets, brown rice, quinoa, amaranth, buckwheat, and wild the term probiotic to be used only on products that deliver live,
rice. Thus, whole grain foods (whole grain snacks and safe microorganisms with suitable viable counts ( 106 colony-
extruded, fermented, and baked foods) can be regarded as forming units per gram of a product) of well-defined strains
functional foods rich in DF and phytochemicals. Many African with a reasonable expectation of delivering benefits for well-
traditional diets and other indigenous foods around the globe being of the host. Ability to survive, proliferate, and colonize
are based on whole grains being consumed as boiled, cooked, the gut is also a criterion although some cultures lack such
fermented, and baked as in injera from grain teff in Ethiopia. ability like traditional yogurt starter cultures (S. salivarius
Such diets can be described as functional foods and they have subsp. thermophilus and L. delbrueckii subsp. bulgaricus), which
been serving communities since folklore. are recognized as probiotics because of their production of
b-galactosidase that helps lactose digestion in maldigesters.
ISAPP has divided live cultures into two: (1) probiotic (non-
Probiotics, Prebiotics, and Synbiotics
oral probiotic foods, probiotic drugs, probiotic medical foods,
nonoral probiotics, defined microbial consortia, probiotic die-
Human organs particularly the gastrointestinal tract (GIT) are
tary supplement, probiotic infant formula, and probiotic ani-
known to be host for vast numbers of microbes ( 1015 bacte-
mal feed) and (2) not probiotic (fermented foods with
rial cells), which are designated as microbiota, microflora, or
undefined microbial content and undefined consortia includ-
normal flora. These microbiota are today extensively exploited
ing fecal microbiota transplant).
for GIT health and beyond under probiotic and synbiotic
The various beneficial functions of probiotics are (1)
functional foods. Consumption of such diets is actually not
structural and histological such as the development of healthy
new because a range of naturally occurring prebiotics
structure and morphology of the gut; (2) metabolic such as in
and probiotics primarily from the genera Lactobacillus,
the synthesis of B vitamins, vitamin K, amino acids, SCFA,
Bifidobacterium, and Saccharomyces are consumed throughout
antioxidants, and bacteriocins and in the biotransformation of
the world in the indigenous fermented foods since folklore.
bile, which help to modulate glucose and cholesterol
Probiotics and prebiotics are also available commercially in the
homeostasis; and (3) protective from colonization by pathogens
form of dietary supplements and clinical therapeutics to be
such as through production of antimicrobial compounds, by
delivered orally or nonorally.
competing for nutrients and attachment sites on gut lining. The
various health benefits documented for probiotic diets include
Probiotics
suppression of (1) obesity and type 2 diabetes through modu-
The word probiotics is derived from the Greek term pro bios lating glucose metabolism; (2) allergic inflammation and atopic
meaning for life. The widely accepted definition of probiotics dermatitis; (3) liver disease; (4) hypercholesterolemia and CVD
Species Strain
Lactobacilli L. acidophilus, L. bulgaricus, L. casei, L. delbrueckii ssp. bulgaricus, L. fermentum, L. johnsonii, L. crispatus, L. salivarius,
L. bifidus, L. reuteri, L. plantarum, L. helveticus, L. casei subsp. rhamnosus, L. gallinarum, L. brevis, L. gasseri,
L. cellobiosus, L. vitulinus, L. collinoides, L. cremoris, L. ruminis, L. dextranicum, L. lactis, L. rhamnosus, L. curvatus, L. faecium,
L. paracasei
Bifidobacteria B. bifidum, B. adolescentis, B. brevis, B. longum, B. animalis, B. infantis, B. thermophilum, B. breve, B. essencis, and B. lactis
Bacillus B. coagulans (GanedenBC30), B. lactis DR10, B. licheniformis, B. subtilis R0179, B. subtilis (natto) OUV23481
Pediococcus P. acidilactici, P. pentosaceus, P. halophilus
Lactococcus L. lactis subspp. lactis and cremoris
Leuconostoc L. mesenteroides subsp. dextranium, paramesenteroides, or lactis
Streptococcus S. diacetilactis, S. cremoris, S. lactis, salivarius subsp. thermophilus, S. faecium, S. equinus
Weissella W. cibaria and confusa
Yeast Saccharomyces cerevisiae, S. cerevisiae var. boulardii
Fungi Aspergillus oryzae, Scytalidium acidophilum
14 FOOD GRAINS AND WELL-BEING | Functional Foods: Dietary Fibers, Prebiotics, Probiotics, and Synbiotics
through modulating cholesterol, LDL-cholesterol, and Synbiotics are a combination of probiotics and prebiotics
triglyceride metabolisms; (5) various cancers such as colon, when administered together. There is synergy between probio-
bladder, and cervical cancers; (6) inflammatory bowel disease; tics and prebiotics resulting in more effectiveness than using
(7) diarrhea; (8) lactose intolerance; (9) renal diseases; (10) either one because it favors competitive advantage for probiotic
hormonal immune response failure; and (11) ingestion of rapid adaptation in the GIT. Synbiotics are currently available
toxic compounds. Although probiotics containing dairy prod- as combinations of bifidobacteria and fructo-oligosaccharides
ucts are dominant, probiotic-fermented cereals and soy bever- (FOS), lactobacillus GG and inulins; bifidobacteria, lactobacilli
ages, flakes, baked goods, soups, and traditional fermented and FOS or inulins.
cereals are also available. Microencapsulation of probiotics is
currently being researched to enable incorporation into various
Functional Foods for Lactose Intolerance
grain products. Fermented grain products containing probiotics
have the following advantages: the possibility of grain compo- Lactose intolerance is the inability to utilize lactose from milk
nents to be prebiotic sources, multimedia characteristics of the because of limited lactase enzyme production in the small
grain products, and their lactose- and cholesterol-free natures. intestine in some individuals. Infants can produce adequate
lactase enabling them to utilize lactose of milk, which is the
sole carbon source for the newborn. The ability to generate
Prebiotics
lactase enzymes diminishes with age. For individuals with
The current consensus definition given is A dietary prebiotic is a limited lactase production, lactose will be transferred to the
selectively fermented ingredient that results in specific changes, large intestine intact where it is hydrolyzed and fermented. The
in the composition and/or activity of the gastrointestinal micro- hydrolyzed products cause secretion of water into the intestinal
biota, thus conferring benefit(s) upon host health. Compounds lumen to balance the osmolarity pressures. This accumulation
recognized as prebiotics include inulin-type fructans (inulin of water produces dehydration and electrolyte imbalance on
fructooligosaccharides, and oligofructose), lactulose, galactooli- the systemic side and watery stools on the intraintestinal sides.
gosaccharides, isomaltooligosaccharides, lactosucrose, xylooli- The fermentation of the sugar adds gaseous components (car-
gosaccharides, glucooligosaccharides, sugar alcohols, resistant bon dioxide, hydrogen, and methane) leading to bloating,
starches, and modified starches. Resistant starches, fructans, ara- cramping, and flatulence. The limited calcium supply through
binoxylans, arabinoxylan oligosaccharides, and b-glucans are inadequate milk intake may lead to osteoporosis. Probiotics
reported in various grains such as in wheat, rye, maize, barley, containing Lactobacillus strains are known to recover the lactose
oat, and maize. tolerance. The various nondairy probiotic functional foods and
Prebiotic food products are available in the market as bev- drinks developed include the following:
erages (soy milk, smoothies, and juice), bakery products
(1) Soy-based (nonfermented soy-based frozen desserts, fer-
(snack bars, biscuits/cookies, waffles, and pancakes), breakfast
mented soymilk drink, and soy-based stirred yogurt-like
cereal (extruded cereals and instant oatmeal), confectionery
drinks).
(chocolate and gummy candy), dairy products (yogurt, ice
(2) Cereal-based puddings; rice-based yogurt; oat-based drink;
cream, and frozen yogurt), desserts (pudding and jelly/jello),
oat-based products; yosa, an oat-bran pudding; mahewu, a
fruit products (fruit preparations), and infant and toddler
fermented maize beverage; maize-based beverage;
foods and in specialty liquid nutrient supplements. Inulin-
probiotic beverages fermented from wheat, rye, millet,
type fructans are popular prebiotics. Even though fructans are
maize, teff, sorghum, and other cereals; malt-based drinks;
present in grains, good sources are from Jerusalem artichoke,
boza fermented from cereals, maize, and sorghum; millet
chicory, garlic, and onion.
malt-fermented probiotic beverages; and millet or sor-
Prebiotics are fermented to SCFA acetate, propionate, and
ghum flour-fermented probiotic beverage.
butyrate. Other bacterial end products include lactate, ethanol,
(3) Fruit- and vegetable-based drinks; fermented banana pulp
succinate, formate, valerate, caproate, isobutyrate, 2-methyl
and fermented banana juice; beet-based drink; tomato-
butyrate, and isovalerate. Bacterial fermentation takes place
based drink; many dried fruits; green coconut water; pea-
in the cecum and colon, where SCFAs are absorbed, stimulat-
nut milk; cranberry, pineapple, and orange juices; ginger
ing absorption of vitamins, water, and minerals (calcium and
juice; grape and passion fruit juices; cabbage juice; carrot
magnesium). The SCFAs have been known to have a protective
juice; noni juice; onion puree; probiotic banana puree;
effect on the intestinal epithelium. The colonic bacteria prefer
nonfermented fruit juice beverages; and blackcurrant juice.
butyrate as their sole source of energy and are also metabolized
(4) Others (starch-saccharified probiotic drink, probiotic
by the colonic epithelium where it regulates cell growth and
cassava-flour product, meat products, and dosa (rice and
differentiation. Butyrate is known to induce secretion of
Bengal gram)).
mucin, antimicrobial peptides, and other factors, which rein-
forces the defense barrier in the colon. Butyrate limitation
leads to inflammation characteristics of the ulcerative colitis. Conclusions
Acetate and propionate are found in portal blood and are
eventually metabolized by the liver (propionate) or peripheral Various DFs are available from whole grains, which are also
tissues, particularly muscle (acetate). Acetate and propionate sources of other nutrients and phytochemicals. The health-
might also play a role as modulators of glucose metabolism. promoting effects of DF (GIH, cancer prevention, and suppres-
Absorption of such SCFAs is implicated to result in low glyce- sion of metabolic syndrome-related diseases) are widely
mic responses and to ameliorate insulin sensitivity. acknowledged. Functional foods containing DF are available
FOOD GRAINS AND WELL-BEING | Functional Foods: Dietary Fibers, Prebiotics, Probiotics, and Synbiotics 15
as baked, extruded, fermented foods and beverages. Many How would you provide further characterization of specific
components of DF have prebiotic effects. The health- DF components from grains for their health benefits and
promoting effects of prebiotics are known by their selective prebiotic effects?
growth promotion of probiotics, fermentation to SCFA, pre- Suggest further improvements that may be made to tradi-
vention of various cancers, and enhancement of mineral tional fermented grain foods to promote their probiotic
absorption. They also suppress CVD, obesity, and diabetes. value.
Prebiotic functional foods are available as beverages, whole
Explore possibilities for developing grains using conventional
grains, fermented cereal foods, bakery products, and RTE,
breeding or through genetic manipulation to provide desirable
infant, and toddler foods and in specialty liquid nutrient sup-
DF profiles for prebiotic potential, like inulin-type fructans and
plements. Probiotics are known to have various healthful
b-glucans.
effects (enhancement of gut immunity, anti-inflammation,
prevention of pathogenic infection, and suppression of irrita-
ble bowel syndrome (IBS), cancers, CVD, diabetes, rheumatoid
arthritis, liver diseases, lactose intolerance, and diarrhea). See also: Beverages from Grains: Fermentation: Foods and
Currently, nondairy probiotic functional foods and drinks Nonalcoholic Beverages; Bioactives and Toxins: Bioactives:
such as soy-based foods, fermented cereals, and extruded and Antioxidants; Bioactive Compounds in Wheat Bran; Carbohydrates:
baked foods are on the rise. Microencapsulation is one option Cereals: Chemistry and Physicochemistry of Non-starchy
to incorporate probiotics in grain products. A combination of Polysaccharides; Glycemic Index, Glycemic Load, and Their Health
prebiotics and probiotics as synbiotics has synergetic health- Benefits; Health Effects of b-Glucans Found in Cereals; Resistant
promoting effects. Many traditional fermented grain foods Starch and Health; Food Grains and the Consumer: Cultural
with live or active cultures if fully characterized have huge Differences in Processing and Consumption; Fortification of Grain-
potential to be used as probiotic functional diets. Develop- Based Foods; Genetically Modified Grains and the Consumer; Grains
ment of grains with desirable DF profiles of prebiotic potential and Health: Misinformation and Misconceptions; Grains and Health;
and improved phytonutrients can advance prebiotic, probiotic, Labelling of Grain-Based Foods; Food Grains and Well-being:
and DF functional foods. The bioactives used in DF, whole Functional Foods: Overview; Nutrition: Soy-Based Foods; Food
grains, and prebiotic functional foods are concentrated in the Grains: Intolerance, Allergy and Diseases: Celiac Disease;
outer portions of the grains, which may be contaminated by Non-wheat Foods: Soybean: Soy-Based Fermented Foods; The
toxicants that can pose food safety challenges. Good agricul- Basics: Grain: Morphology of Internal Structure.
tural practices, good manufacturing practices, hazard analysis
critical control points, and adequate grain drying, cleaning,
scouring, and/or removal of very small portion of outermost
layers of grains during milling can suppress such food safety
challenges and also preserve functional ingredients. Further Reading
AACCI Dietary Fiber Definition Committee (2001) The definition of dietary fiber. Cereal
Foods World 46(3): 112126.
Anderson JW, Baird P, Davis Jr. RH Jr., Ferreri S, Knudtson M, Koraym A, Waters V,
Exercises for Revision and Williams CL (2009) Lead article: Health benefits of dietary fiber. Nutr. Rev.
67(4): 188205.
Binns N (2013) Probiotics, Prebiotics and the Gut Microbiota. Belgium: ILSI Europe.
Provide definitions of DFs, whole grain foods, prebiotics,
Birkett AM and Cho SS (2013) Cereal fiber and health: Current knowledge. Cereal
probiotics, and synbiotics. Foods World 58(6): 309313.
List up to seven different compounds regarded to be DFs. Cho SS and Finocchiaro ET (2010) Handbook of Prebiotics and Probiotics Ingredients
List up to four different compounds regarded to be Health Benefits and Food Applications. Boca Raton, FL: CRC Press, Taylor &
prebiotics. Francis Group.
Corona-Hernandez R, Alvarez-Parrilla E, Lizardi-Mendoza J, Islas-Rubio AR, de la
Provide genera of microbial species widely used as
Rosa LA, and Wall-Medrano A (2013) Structural stability and viability of
probiotics. microencapsulated probiotic bacteria: A review. Comprehen. Rev. Food Sci. Food
Provide essential characteristics of probiotics. Saf. 12: 614628.
Provide at least two health beneficial effects of DFs, whole Dwivedi S, Sahrawat K, Puppala N, and Ortiz R (2014) Plant prebiotics and human
grain foods, prebiotics, probiotics, and synbiotics. health: Biotechnology to breed prebiotic-rich nutritious food crops. Electron. J.
Biotechnol. 17: 238245.
List up to four indigenous fermented foods consumed in Ferruzzi MG, Jonnalagadda SS, Liu S, Marquart L, McKeown N, Reicks M,
your communities believed to have the effects of prebiotics, Riccardi G, Seal C, Slavin J, Thielecke F, van der Kamp JW, and Webb D
probiotics, and synbiotics. (2014) Developing a standard definition of whole-grain foods for dietary
List potential safe diets for lactose-intolerant individuals. recommendations: Summary report of a multidisciplinary expert roundtable
discussion. Adv. Nutr. 5: 164176.
Foschia M, Peressini D, Sensidoni A, and Brennan CS (2013) Review the effects of
dietary fibre addition on the quality of common cereal products. J. Cereal Sci.
58: 216227.
Exercises for Readers to Explore the Topic Further Granato D, Branco GF, Nazzaro F, Cruz AG, and Faria JAF (2010) Functional foods and
nondairy probiotic food development: Trends, concepts, and products. Comprehen.
Rev. Food Sci. Food Saf. 9: 292302.
What innovations in grain-processing technology can you Hill C, Guarner F, Reid G, Gibson GR, Merenstein DJ, Pot B, Morelli L, Canani RB,
envisage to preserve grain bioactive ingredients used to Flint HJ, Salminen S, Calder PC, and Sanders ME (2014) Consensus statements:
deliver optimum DFs, prebiotics, and probiotics? The International Scientific Association for Probiotics and Prebiotics consensus
16 FOOD GRAINS AND WELL-BEING | Functional Foods: Dietary Fibers, Prebiotics, Probiotics, and Synbiotics
statement on the scope and appropriate use of the term probiotic. New York: Slavin J, Tucker M, Harriman C, and Jonnalagadda SS (2013) Whole grains:
Macmillan, Expert Consensus Document, 11, 506-514. Definition, dietary recommendations, and health benefits. Cereal Foods World
Jones JM (2014) CODEX-aligned dietary fiber definitions help to bridge the fiber gap. 58(4): 191198.
Nutr. J. 13(34): 210.
Marsha AJ, Hill C, Rossa RP, and Cotter PD (2014) Fermented beverages with health-
promoting potential: Past and future perspectives. Trends Food Sci. Technol.
39: 135145. Relevant Websites
Poutanen K, Sozer N, and Valle GD (2014) Review: how can technology help to deliver
more of grain in cereal foods for a healthy diet? J. Cereal Sci. 59: 327336. http://wholegrainscouncil.org/whole-grains-101/existing-standards-for-whole-grains
Saad N, Delattre C, Urdaci M, Schmitter JM, and Bressollier B (2013) An overview of the Whole Grains Council.
last advances in probiotic and prebiotic field. LWT- Food Sci. Technol. 50: 116. http://www.isapp.net/Home ISAPP.
Nutrition: Soy-Based Foods
AM Fehily, Tinuviel Software, Warrington, UK
2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition, volume 2, pp. 348354, 2004, Elsevier Ltd.
Table 1 Vitamin and mineral composition of soybeans (lectins, goitrogens, and digestive enzyme inhibitors). These
substances are, however, inactivated by appropriate cooking of
Amount per 100 g as consumed the beans or, in the case of goitrogens, counteracted by ade-
Sodium 1 mg quate iodine intake.
Potassium 510 mg
Calcium 83 mg Lectins
Magnesium 63 mg Lectins, also known as haemagglutinins, are present in raw
Phosphorus 250 mg soybeans and in other legumes. They are, however, heat labile
Iron 3.0 mg and are inactivated when the beans are properly cooked. In
Copper 0.32 mg experimental animals these polymeric proteins have been
Zinc 0.9 mg shown to cause damage to red blood cells and intestinal
Manganese 0.7 mg
mucosa, and thereby impaired nutrient utilization and loss of
Selenium 5 mg
body weight. In humans, these compounds can result in nau-
Iodine 2 mg
Vitamin A (retinol equivalents) 1 mg sea, vomiting, diarrhoea, and abdominal pain.
Vitamin D 0 mg
Vitamin E 1.13 mg Goitrogens
Thiamin 0.12 mg Enlargement of the thyroid gland has been shown to occur in
Riboflavin 0.09 mg rats fed soybean meal. The goitrogenic agent in soybeans is
Niacin equivalents 2.7 mg unknown. However, the effect is counteracted by adequate
Vitamin B6 0.23 mg iodine intake.
Vitamin B12 0 mg
Folate 54 mg
Digestive enzyme inhibitors
Pantothenate 0.18 mg
Raw soybeans contain certain proteins (protease inhibitors)
Biotin 25.0 mg
Vitamin C Trace which react with digestive enzymes (trypsin, chymotrypsin or
salivary, and pancreatic a-amylase), thereby interfering with
Data from Royal Society of Chemistry and Ministry of Agriculture Fisheries and Food the digestion of protein and starch. In humans, raw soy or
(1991) McCance and Widdowsons The Composition of Foods, 5th edn. Cambridge, isolated protease inhibitors increase levels of cholecystokinin
UK: Royal Society of Chemistry. Crown copyright is reproduced with the permission of (CCK) and pancreatic secretion. It is thought that chronic
the Controller of Her Majestys Stationery Office. pancreatic stimulation may lead to pancreatic hypertrophy,
hyperplasia, and possibly to cancer. In rats, raw soy has been
Folate shown to increase risk of pancreatic cancer. However, protease
inhibitors present in raw soy are inactivated by heat and are
Vitamin B6
therefore not a problem in cooked beans. For example, in
Niacin
countries such as Japan where soy foods are widely consumed,
Riboflavin
the incidence of pancreatic cancer is similar to or less than that
Thiamin for Western countries where soy forms a relatively small com-
Zinc ponent of the total diet.
Phosphorus
Potassium Phytic acid
Magnesium Phytic acid is present in soybeans and also in other fiber-
Iron containing foods. It has been suggested to reduce the absorp-
Calcium tion of calcium, iron, zinc, and vitamin D from the diet.
However, these effects have been observed in in vitro experi-
0 10 20 30 40 50 ments and not in in vivo studies, even at fiber intakes at the
Figure 1 Vitamin and mineral composition of soybeans in relation to upper limit of the normal human consumption range. In
reference nutrient intakes (% of reference nutrient intake (RNI) provided addition, levels of micronutrients tend to be higher in fiber-
by 100 g). (Reproduced from (2003) Encyclopedia of Food Sciences rich foods than in fiber-poor foods. Thus, adverse effects of
and Nutrition, 2nd edn, p. 5393. Elsevier Ltd.) phytic acid on nutrient absorption are likely to be unimportant
at the levels of fiber normally consumed in the human diet.
Fiber
Soybeans are a valuable source of fiber in the diet: 6 g of non- Isoflavones
starch polysaccharides (NSP) per 100 g. Of this, 44% is soluble
fiber and 56% insoluble fiber. One hundred grams of soybeans Soybeans contain isoflavones, a group of compounds that are
provides 33% of the daily dietary reference value (UK) for fiber. structurally similar to oestradiol (phyto-oestrogens). Soybeans
contain higher amounts of the isoflavones genistein and
daidzein (see Figure 2) and smaller amounts of glycitein,
primarily in the form of glycosides: genistin, daidzin, and
Antinutrients
glycitin (total isoflavone content of 180 mg per 100 g). The
Raw soybeans contain a number of substances which may have glycosides are hydrolyzed in the gut by a bacterial enzyme,
detrimental effects on digestion and other metabolic processes glucose being removed to produce the aglycone forms
FOOD GRAINS AND WELL-BEING | Nutrition: Soy-Based Foods 19
lipids tends to be greater among those with the highest plasma development of atherogenesis may be delayed: inhibition of
cholesterol levels at baseline. the migration and proliferation of smooth muscle cells; inhi-
A great deal of research has been done to identify the bition of platelet activation and aggregation; and reduction in
component(s) of soy protein that is responsible for its effect platelet serotonin uptake.
on blood lipids. Studies using a mixture of amino acids that In 1999, in response to the available evidence on the ben-
duplicate the amino acid profile of soy protein have found efits of soy protein, the USA Food and Drug Administration
that this does not have the same effect on blood lipids as (FDA) announced that it would allow food manufacturers to
the intact protein. However, studies of soy protein with iso- label products containing 6.25 g of soy protein per serving as
flavones intact have shown that this has a lipid lowering helping to reduce risk of heart disease, as part of a balanced diet
effect, whereas soy protein with isoflavones removed has low in fat and saturates.
no significant effect. It has therefore been suggested that
the isoflavones present in soy protein are largely responsible
for the lipid lowering effect of soy protein. The benefits of Soy and Cancer
isoflavones present in soy protein on cholesterol lowering
have been suggested to be mediated through upregulation Cancer is a major cause of morbidity and mortality. For exam-
of LDL-receptor activity. Nevertheless, studies of isolated ple, in the UK about one in three people will develop cancer at
and purified isoflavones have failed to show that these have some time during their life and cancer accounts for about one
a lipid lowering effect. in four of all deaths. There are marked differences in death rates
Soy may reduce risk of CHD through several mechanisms in from several types of cancer between Asian countries and many
addition to its ability to lower blood lipids (see Figure 3). For Western countries. The differences are more striking for
example, soy isoflavones are known to act as antioxidants and hormone-dependent cancers of the prostate, breast, and
have been suggested to reduce oxidative damage to LDL-C. colon/rectum. For example, the risk of dying from prostate
A decrease in oxidized LDL particles, which are considered cancer among Japanese men is only one-fifth that of men in
atherogenic, may reduce the risk of atherosclerosis. the USA and Japanese women have a breast cancer mortality
A study of young cynomolgus monkeys showed that the rate that is only a quarter of that of women in the USA.
size of atherosclerotic lesions was 70% less in those fed on a Countries where mortality from prostate and breast cancers
diet containing soy protein with isoflavones present compared is low have considerably higher intakes of soy than those
with those fed on a diet containing soy protein with isofla- countries where mortality from these cancers is high. In animal
vones removed. This suggests that isoflavones may inhibit models, studies investigating the effects of soybeans on pros-
atherogenesis. In another study of surgically postmenopausal tate cancer or breast cancer have shown reduced tumorigenesis.
cynomolgus monkeys, soy protein with isoflavones intact was In humans however, evidence is inconclusive. Epidemiological
found to reduce the progression of atherosclerosis. The magni- studies of prostate cancer conducted in Japan and in Japanese
tude of the effect was reported to be comparable to that of migrants to Hawaii have shown no significant effect of soy
postmenopausal oestrogen therapy. consumption. However, a recent study in the USA reported
Soybean isoflavones, in particular genistein, impact benefi- that men who drank one cup of soy milk per day had a risk
cially on vascular reactivity. Genistein has also been shown, of developing prostate cancer that was 70% lower than that of
in vitro, to have a number of other effects whereby the controls. For breast cancer, although some studies have shown
Figure 3 Mechanisms through which soy could reduce risk of coronary heart disease (CHD).
FOOD GRAINS AND WELL-BEING | Nutrition: Soy-Based Foods 21
that soy can help prevent breast cancer, others have found no problem in many Western countries, although not in Asian
significant effect. An increase in menstrual cycle length has countries. A number of factors are known to increase risk of
been observed in some studies, in response to soybean con- osteoporosis, including insufficient dietary calcium, low phys-
sumption, although no change was observed in others. Such an ical activity, and lack of oestrogen.
effect is of potential relevance to the hypothesis that soy may In women, bone mass reduces at a rapid rate in the first few
be protective against breast cancer, because some data indicate years after the menopause, due to the reduction in oestrogen.
that longer cycles, which are typical of Asian women, are Hormone replacement therapy (HRT) is well known to reduce
associated with a reduced risk of breast cancer. bone loss in postmenopausal women. However, HRT is not
For colo-rectal cancer data are also inconclusive: some stud- appropriate for all women. Since soy may have oestrogenic
ies showed no significant effect, some reported a protective effect effects, it has been hypothesized that it may provide an alter-
of soy, and others observed an increased risk with soy consump- native to HRT.
tion. For cancers at other sites, data tend to show that increased Soy protein containing isoflavones has been reported to
soybean consumption may be protective against lung and stom- reduce bone loss due to oestrogen deficiency in ovariectomized
ach cancer. Much more research is needed to confirm whether rats. Soybean isoflavones have also shown a protective effect
soy has a protective effect against cancer in humans. on bone loss in ovariectomized rats, suggesting that the bene-
Isoflavones appear to be the components of soy that would ficial effect of soy protein is due to isoflavones. The effect of
be most likely to account for any protective action of soy isoflavones on bone loss in rats has been reported to be similar
against cancer, especially in inhibiting the initiation stage of to that for oestrogen. The bone sparing effect of soy protein
carcinogenesis (see Figure 4). Genistein has been shown to isoflavones has been suggested to be due to a reduction in bone
suppress the growth of a wide range of cancer cells. It inhibits resorption and/or to an increased osteoblast activity (increas-
certain enzymes that could affect the onset of cancer or the ing bone formation).
growth of tumors: tyrosine protein kinase, mitogen-activated In humans, most but not all studies indicate that soy pro-
protein (MAP) kinase, and ribosomal S6 kinase. Genistein also tein containing isoflavones favorably affects bone turnover and
inhibits the DNA repair enzyme topoisomerase II and acts as bone mineral density in the lumbar spine of perimenopausal
an antioxidant, thus potentially preventing oxidative DNA and postmenopausal women. Treatment with ipriflavone, a
damage. It increases in vitro concentrations of transforming synthetic isoflavone, has also been reported to have bone
growth factor b, which is thought to inhibit the growth of conserving effects in postmenopausal women with low bone
cancer cells. In addition, genistein has also been shown to mass, confirming that the beneficial effect of soy protein is
inhibit angiogenesis, the formation of new blood vessels, an likely to be due to isoflavones.
abnormal event that occurs as part of the growth and expan- In addition to the effect of soy protein on bone mineral
sion of malignant tumors. However, many of these effects have density, it has also been suggested that soy protein, when
been shown with very high concentrations of genistein and not substituted for animal protein, may indirectly enhance bone
in cells treated with levels likely to be achieved in plasma of strength. Another effect of soy protein is that it helps to con-
human subjects eating soy foods. serve calcium by reducing urinary calcium excretion. This is
due to the lower sulfur amino acid content of soy protein.
Soy protein therefore appears to have modest beneficial
effects on bone density. However, studies to date have been
Soy and Bone Health short term and have involved only small numbers of subjects.
In addition, no study has investigated whether soy protein
Osteoporosis is a clinical condition in which there is a reduced containing isoflavones has an effect on fracture risk.
amount of bone per unit volume and an increased susceptibil-
ity to fractures, particularly fractures of the vertebrae, distal
forearm (Colles fracture) and hip. Of these, hip fracture is the
most severe, since patients require a prolonged hospital stay, Soy and Menopausal Symptoms
there is a high mortality rate (about 20% within six months of
the fracture) and of those who survive may suffer permanent Hot flushes (also known as hot flashes) are a common symp-
disability and dependency. Osteoporosis is a major health tom among menopausal women in Western societies, but are
Soy isoflavones
reported to be much less common in Japan. Whether this How would you design a soybean for optimum nutritional
difference is due to the higher consumption of soy in Japan is and nutraceutical composition.
not clear. Hormone replacement therapy (HRT) generally alle-
viates hot flushes and other menopausal symptoms such as
vaginitis. There is much research interest in the possibility that
See also: Agronomy of Grain Growing: Soybean: Agronomy;
soy may provide an alternative to HRT in this regard.
Food Grains and the Consumer: Genetically Modified Grains and
Evidence of benefit of soy in menopausal women is conflict-
the Consumer; Non-wheat Foods: Soybean: Soy-Based Fermented
ing. Some researchers have reported an improvement in the
Foods; Soybean: Soymilk, Tofu, and Okara; Oilseed and Legume
frequency of hot flushes in women taking soy protein daily.
Processing: Soybean: Processing; Soybean: Soy Concentrates and
Others have reported no effect of soy protein on the number of
Isolates; The Legumes and Pseudocereals: Beans: Overview;
hot flushes experienced but found a reduction in the severity of
Grain Legumes and Their Dietary Impact: Overview.
symptoms. Others have reported a reduction in both the inci-
dence and severity of hot flushes with soy protein or an isoflavone
extract compared with a control group. Others have reported no
difference in either the frequency or severity of hot flushes in the
intervention group compared with the control group.
Further Reading
Two studies have investigated the effects of phyto-oestrogen
supplements on vaginal cytology and found an increase in cell Anderson JJB, Anthony M, Messina M, and Garner SC (1999) Effects of
proliferation (an indication of oestrogenic activity) and reversal of phyto-oestrogens on tissues. Nutrition Research Reviews 12: 75116.
menopausal atrophy. Others have reported no significant effect. Anderson JW, Johnstone BM, and Cook-Newell ME (1995) Meta-analysis of the effects
of soy protein intake on serum lipids. New England Journal of Medicine
There are difficulties in interpreting the results of these
333: 276282.
studies. This is because there are differences in the amounts Bingham SA, Atkinson C, Liggins J, Bluck L, and Coward A (1998) Phyto-oestrogens:
of soy protein and isoflavones used and differences in the where are we now? British Journal of Nutrition 79: 393406.
duration of the studies. Interpretation is also complicated by Lissin LW and Cooke JP (2000) Phytoestrogens and cardiovascular health. Journal of
the fact that reported frequency and severity of symptoms American College of Cardiology 35: 14031410.
Messina MJ (1999) Legumes and soybeans: overview of their nutritional profiles and
tends to reduce in the control group as well as the intervention health effects. American Journal of Clinical Nutrition 70(suppl): 439S450S.
group. Thus, much more research needs to be done before soy Messina M and Messina V (2000) Soyfoods, soybean isoflavones and bone health: a
protein can be proposed as a potential alternative to HRT for brief overview. Journal of Renal Nutrition 10: 6368.
the control of menopausal symptoms. Messina M, Persky V, Setchell KDR, and Barnes S (1994) Soy intake and cancer risk: a
review of the in vitro and in vivo data. Nutrition and Cancer 21: 113131.
Potter SM (2000) Soy new health benefits associated with an ancient food. Nutrition
Exercises for Revision Today 35: 5360.
Royal Society of Chemistry and Ministry of Agriculture, Fisheries and Food. (1991)
McCance and Widdowsons. The Composition of Foods, 5th edn. Cambridge, UK:
Complile a summary of the effects of phytoestrogens on Royal Society of Chemistry.
young adult men and women.
What are the effects of high temperature processing on soy
nutritional characteristics?
Relevant Websites
Exercises for Readers to Explore the Topic Further
nutrition.org.uk http://www.nutrition.org.uk British Nutrition Foundation.
Investigate further the current state of knowledge on the food.gov.uk http://www.food.gov.uk Food Standards Agriculture Organisation.
effect of soy isoflavones on a medical condition of interest apps.fao.org http://apps.fao.org Food and Agriculture Organisation (FAO).
ajcn.org http://www.ajcn.org American Journal of Clinical Nutrition.
to you.
Food Grains and the Consumer
Contents
Grains and Health: Misinformation and Misconceptions
Consumer Trends in Grain Consumption
Cultural Differences in Processing and Consumption
Fortification of Grain-Based Foods
Genetically Modified Grains and the Consumer
Labelling of Grain-Based Foods
Grains and Health
Topic Highlights
Food and nutrition professionals need to be able to counter
Claims suggest that wheat is the product of genetic research unfounded theories about wheat and other grains with
and that we are eating genetically altered wheat. In reality, sound science and unbiased, critical thinking.
genetically modified wheat is not sold legally anywhere in
the world.
Claims suggest that wheat and its gluten have changed and
that is why celiac disease, an autoimmune disorder, is on Learning Objectives
the rise. While celiac disease rates are increasing, and theo-
ries abound, as to why. To identify current controversies surrounding wheat, grains,
Claims suggest that wheat is addictive. There are no data to and health.
support claims that removing wheat from the diet results in To describe scientific evidence regarding wheat, grains, and
withdrawal symptoms. health.
Claims suggest that wheat causes uncontrollable overeat- To understand the important role that food and nutrition
ing. Existing data show that ingestion of wheat protein professionals play in dispelling common misinformation
causes the release of satiety hormones. and misconceptions about wheat and grains.
Claims suggest that wheat ingestion leads to a host of
mental health issues. There are scanty scientific data to
support these claims.
Claims suggest that wheat causes obesity and diets that Introduction
eliminate wheat and other carbohydrates are the most effec-
tive way to lose weight. In reality, any significant weight loss Recently, grain consumption, especially wheat, has been
that occurs with these diets is due to the reduction in total blamed for obesity, diabetes mellitus, other chronic diseases,
calories from any source, not just wheat or carbohydrates. food addiction, and impaired brain function. Popular press
Claims suggest that the elimination of wheat from the diet books such as Wheat Belly by William Davis, the Paleo Diet by
results in the cure or improvement of many health condi- Loren Cordain, and Grain Brain by David Perlmutter and other
tions. In reality, many of these conditions are made better authors and food bloggers claim that eliminating grains is the
by weight loss and not specifically the elimination of wheat. key solution to improving health. A review of nutrition and
Claims suggest that the nutrient contribution of wheat and medical literature suggests a different scenario. This article pro-
grains can easily be obtained from other food groups. While it vides a review of the scientific evidence regarding grains and
is true that these nutrients are found in other foods, studies health to dispel misinformation and misconceptions. This
on nutritional quality indicate that diets without whole and piece will focus on wheat because it has been subject to the
enriched grains are often lacking in these nutrients. most misinformation.
Wheat Breeding and Genetics Ancient wheats such as einkorn are held as models of what
we should eat because they have not been hybridized and
Wheat originated in the Middle East thousands of years ago. because of their protein makeup. In fact, one author says that
According to the Food and Agriculture Organization (FAO), einkorn contains 28% protein compared with average protein
wheat is grown on more land area than any other commercial contents of 1215% in modern wheats and that ancient wheats
crop and continues to be the most important food grain source did not cause the symptoms that new varieties do. However,
for humans. Together, rice, wheat, and maize provide over half studies carried out at the University of Saskatchewan where
of the worlds calories. ancient and other varieties of wheat were grown on adjacent
Popular press authors and food bloggers suggest that wheat plots actually show that the crude protein content of einkorn is
is the product of genetic research, and today, we are eating 18% and that of other ancient and modern varieties had pro-
genetically altered wheat. They contend that wheat and its tein values ranging from 16.3% to 17.5%. This fits well with
gluten have changed and our bodies can no longer digest the the USDA World Wheat Collection, which shows approxi-
modified grains of today, and that is why people are having so mately a threefold variation in protein content of wheat from
much difficulty digesting wheat and why celiac disease, an 7% to 22%, with one-third of this under genetic control and
autoimmune disorder, is on the rise. two-thirds of this controlled by environmental conditions.
It is alleged that modern wheat has changed to be less
healthy because of breeding practices. In fact, wheat breeding
is an age-old practice. Significant advances in plant breeding Celiac and Related Disorders
occurred in the 1940s with the work of Norman Borlaug and
others, using traditional techniques (non-genetically modified Celiac disease is on the rise, as are other autoimmune diseases,
organisms). Lines selected provide increased yield through in Western countries. This fact is documented in the scientific
disease, drought, and insect resistance, especially in developing literature. Studies by Mayo Clinic gastroenterologist Joe
countries. Despite assertions to the contrary, these lines were Murray and colleagues compared celiac titers in blood samples
produced by crossing wheat varieties with other wheat varieties from the recent US Air Force recruits with stored blood
from all over the globe. In 1970, Dr Borlaug with his so-called samples taken from recruits more than 50 years ago. The
Green Revolution won the Nobel Peace Prize for his wheat and analysis showed 0.2% of recruits had the gene for celiac disease
grain breeding programs, which produced grains with high in 1950 compared with 0.9% of recent recruits, which is a
yields that grow under a wide variety of conditions and help quadrupling of the incidence rate. Data from Finland also
address world food supply challenges. show an increase from 1% to 2% or a doubling of the rate of
It is also alleged that genetic modification of wheat has celiac disease incidence in that population. While celiac disease
made it less healthy and has changed the gluten proteins. rates are increasing, it is uncertain whether this is due in part to
This is untrue for two reasons. First, no ill effects can come better identification and awareness of the disease, as well as a
from genetically modified wheat because there is none sold myriad of other dietary, immunologic, and environmental
legally anywhere in the world. Second, recent studies from changes, especially those that impact the microorganisms and
USDA labs and labs at the University of Saskatchewan and integrity of the gut. Some theories include the clean theory,
studies in Europe using grains from seed banks going back as poor diets overall, overuse of antibiotics and other drugs espe-
far as Canadian Red Fife in 1857 show wheat gluten has not cially acid-lowering drugs, lack of breast-feeding, and frequent
changed. use of Cesarean sections.
Concern has been raised about the use of hybrid dwarf Davis and other popular press authors posit that the reason
wheats. They now comprise 99% of the wheat grown world- for the increase in celiac disease is attributed to the fact that
wide. Short-straw naked (dwarf) wheats are one of the celiac disease-triggering proteins are expressed at higher levels
advances that were part of the green revolution. They increased in current wheat varieties than found 50 years ago. Such
yield for two reasons: (1) wheat varieties with long, spindly authors suggest that high-molecular-weight (HMW) glutenins
straws lodge, meaning that the kernel head is heavy and falls to and higher levels of gliadin of modern wheat are the problem.
the ground, making harvest difficult, and (2) shorter straw While studies show that the immune system reacts to break-
means that inputs, for example, water and nutrients, go into down products of HMW glutenins, data are lacking that show
making wheat kernels rather than straw. that these proteins trigger more reactions than those of ancient
Popular press authors imply that new wheat varieties and wheats or even wheat varieties from 50 years ago. Interestingly,
their proteins are unique or harmful in some way. Further, they gliadins are found in higher amounts in ancient versus modern
posit that 5% of the proteins in wheats are unique, meaning wheats.
they are found in neither parent, and that this unexpected Gluten-containing grains are also implicated in non-celiac
genetic rearrangement results in altered proteins with poten- gluten sensitivity (NCGS). This newly recognized disorder has
tially toxic effects. It is important to consider that plants can no medically agreed-upon test to confirm a diagnosis, and
only express proteins that they have the genetic code to pro- therefore, its existence and incidence are subject to much
duce. Creating a unique protein requires a mutation of the debate. The best estimates are that 46% of the population
DNA or RNA. While environmental conditions can promote may have NCGS. NCGS is thought to be present in about 5%
or inhibit the expression of certain proteins, it cannot code for of irritable bowel syndrome cases and may cause the rash,
proteins that are not in the genome. Therefore, hybridization dermatitis herpetiformis, which can be seen in some celiac
of wheat does not create unique proteins. patients.
FOOD GRAINS AND THE CONSUMER | Grains and Health: Misinformation and Misconceptions 25
known that schizophrenia incidence increases in the presence Grain Foods and the Glycemic Index
of any autoimmune disease or a history of severe infections. It
is thought that the antibodies produced can impact the brain. The glycemic index (GI) is a measure that compares the blood
In some studies, a subset of schizophrenia patients show ele- glucose response produced by 50 g of available carbohydrate
vated gliadin antibodies. However, classic celiac disease diag- from a food with the blood glucose response produced by 50 g
nostics, either antibodies or gene markers, were not found in of glucose. Popular press authors and food bloggers correctly
higher rates in schizophrenia patients than in the public at noted that wheat bread has a GI that is higher than table sugar.
large. However, GI is most often misunderstood because it is used to
A comprehensive review looking at the connection between compare very different amounts of food. For example, 50 g of
gluten and schizophrenia showed that gluten withdrawal table sugar (approximately three tablespoons) would yield
resulted in a drastic reduction or full remission of symptoms 50 g of available carbohydrate. In contrast, 50 g of available
but only among a small subset of schizophrenia sufferers. carbohydrate from whole wheat bread is much more than 50 g
Thus, in a small group of schizophrenia patients, removal of of bread since bread is not all carbohydrate and not all the
wheat may be helpful but not the miracle cure indicated by carbohydrate is available. Thus, it would take 144 g of whole
Davis and others. wheat bread (5.1 slices at 28 g per slice) or 111 g of white bread
(3.9 slices) to yield 50 g of available carbohydrate.
in many fruits or vegetables, and when found, its structure is Assess the evidence associated with grains, carbohydrates,
such that it does not have cholesterol-lowering properties. and brain function. How does this evidence compare with
Further epidemiological studies show that cereal fiber other nutritional impacts on the brain?
reduces the risk of certain gut cancers while fruit and vegetable Use the GI tables and look up the values for wheat and
fibers do not. These data suggest that balance as prescribed in grain staples such as sourdough and whole grain bread
USDA MyPlate provides the best diet. basmati rice and pasta. Explain why such products often
with the same ingredients have varying GIs. Then look at
the amount eaten for 50 g of carbohydrate and compare
Conclusion
that and the nutritional quality of indulgent and other
foods with lower GIs such as candy or chocolate cake.
Numerous health problems including obesity, diabetes melli-
Using this research, explain when the GI might be useful
tus, other chronic diseases, food addiction, and impaired brain
for food selection and when it might not yield optimal
function have been attributed to grains such as wheat. A review
nutrition choices.
of nutrition and medical literature reveals that there are little
evidence to support claims that wheat and grains cause adverse
health effects and chronic disease and much evidence to show See also: Appendix 3: Grains, Foods, and Ingredients Suiting
that a balanced diet that includes wheat and grains, especially Gluten-Free Diets for Celiac Disease; Carbohydrates: Glycemic
whole grains, reduces chronic disease risk and lowers overall Index, Glycemic Load, and Their Health Benefits; Food Grains and
mortality. the Consumer: Grains and Health; Food Grains: Intolerance,
Popular press authors and food bloggers claim that elimi- Allergy and Diseases: Celiac Disease; Cereal Allergens; The Gluten-
nating wheat grains is essential for optimal health for all. While Free Diet; Grain Composition and Analysis: The Composition of
it is true that wheat and other gluten-containing grains must be Food Grains and Grain-Based Products; The Cereal Grains: Wheat:
eliminated for those with reactions to gluten and wheat An Overview of the Grain That Provides Our Daily Bread.
because of celiac disease, allergies, or other conditions, it is
not true that wheat and grain elimination is best for all.
Further, it is also true that many diets contain too many calo-
ries and grain-based desserts and snacks contribute to excess
calories, and it is not true that the grains and wheats are the
Further Reading
culprit. Rather, it is the poor food choices that should be
addressed. Data for elimination of grains for all are, at best, Abdel-Aal E-SM, Hucl P, and Sosulski FW (1995) Compositional and nutritional
scanty and do not show scientific rigor. Food and nutrition characteristics of spring einkorn and spelt wheats. Cereal Chemistry 72: 621624.
professionals need to be able to counter unfounded theories Aune D, Chan DS, Lau R, et al. (2011) Dietary fibre, whole grains, and risk of colorectal
cancer: Systematic review and doseresponse meta-analysis of prospective studies.
about wheat and other grains with sound science and unbi-
BMJ 343: d6617.
ased, critical thinking. This article provides a review of the Benros ME, Nielsen PR, Nordentoft M, Eaton WW, Dalton SO, and Mortensen PB (2011)
scientific evidence regarding grains and health to dispel com- Autoimmune diseases and severe infections as risk factors for schizophrenia: A 30-
mon misinformation and misconceptions. year population-based register study. American Journal of Psychiatry
168: 13031310.
Bolotin D and Petronic-Rosic V (2011) Dermatitis herpetiformis. Part I. Epidemiology,
Exercises for Revision pathogenesis, and clinical presentation. Journal of the American Academy of
Dermatology 64: 10171024.
Burke SJ, Gibney MJ, ODwyer NA, and McCarthy SN (2005) The influence of cereal and
Take a specific wheat controversy and describe the scientific dairy consumption on the Irish diet: Implications for developing food-based dietary
evidence to support or refute this claim. guidelines. Public Health Nutrition 8: 227237.
Model a diet with and without grains. Calculate the nutri- Cascella NG, Kryszak D, Bhatti B, et al. (2011) The gluten connection: The association
tional value of the model diet using a computer analysis to between schizophrenia and celiac disease. Schizophrenia Bulletin 37: 94100.
Centers for Disease Control and Prevention (2014) Adult Obesity Facts. Atlanta, GA:
show what nutrients might be at risk. Show what foods CDC. Published online at: http://www.cdc.gov/obesity/data/trends.html
would need to be consumed in larger quantities. If this (accessed 1/14).
were done, what would happen to calories? Choi S, Disilvio B, Fernstrom MH, and Fernstrom JD (2009) Meal ingestion, amino
acids and brain neurotransmitters: Effects of dietary protein source on serotonin and
Calculate the calories for 3 days of diets with wheat or
catecholamine synthesis rates. Physiology and Behavior 98: 156162.
grains included and the calories of that same diet without
Corsica JA and Pelchat ML (2010) Food addiction: True or false? Current Opinion in
the wheat or grains. Show how much weight would likely Gastroenterology 26: 165169.
be lost with the elimination of these products. Corwin RL and Grigson PS (2009) Symposium overview Food addiction: Fact or
Go to a store and compare the cost, nutritional value, and fiction? Journal of Nutrition 139: 617619.
eating quality of products that are labeled as gluten- or wheat- Craig SA (2004) Betaine in human nutrition. American Journal of Clinical Nutrition
80: 539549.
free with their gluten- or wheat-containing counterparts. Crider K, Bailey LB, and Berry RJ (2011) Folic acid food fortification History, effect,
concerns, and future directions. Nutrients 3: 370384.
Exercises for Readers to Explore the Topic Further Dohan FC (1966) Wheat "consumption" and hospital admissions for schizophrenia
during World War II. A preliminary report. American Journal of Clinical Nutrition
18: 710.
Why is misinformation around diet so readily accepted?
Fasano A (2014a) Gluten-Related Disorders. Philadelphia, PA: Lippincott, Williams and
Show the evidence supporting the use of wheat and grains Wilkins, p. 10.
in preventing any one of the chronic diseases. Assess this Fasano A (2014b) Gluten-Related Disorders. Philadelphia, PA: Lippincott, Williams and
evidence in and evaluate evidence for their nonuse. Wilkins, p. 44.
28 FOOD GRAINS AND THE CONSUMER | Grains and Health: Misinformation and Misconceptions
Fewtrell M, Wilson DC, Booth I, and Lucas A (2011) Six months of exclusive Othman RA, Moghadasian MH, and Jones PJ (2011) Cholesterol-lowering effects of oat
breastfeeding: How good is the evidence? BMJ 342. http://dx.doi.org/10.1136/bmj. b-glucan. Nutrition Reviews 69: 299309.
c5955. Palosuo K, Varjonen E, Kekki OM, et al. (2001) Wheat omega-5 gliadin is a major
Foster-Powell K, Holt SH, and Brand-Miller JC (2002) International table of glycemic allergen in children with immediate allergy to ingested wheat. Journal of Allergy and
index and glycemic load values 2002. American Journal of Clinical Nutrition Clinical Immunology 108: 634638.
76: 556. Putnam J, Allshouse J, and Kantor LS (2002) U.S. per capita food supply trends: More
Genuis SJ and Bouchard TP (2010) Celiac disease presenting as autism. Journal of calories, refined carbohydrates, and fats. Food Review 25(3). http://ers.usda.gov/
Child Neurology 25: 114119. publications/foodreview/dec2002/frvol25i3a.pdf.
Geraedts MC, Troost FJ, Tinnemans R, Soderholm JD, Brummer RJ, and Saris WH Ravindra CN, Hucl P, Briggs C, Irvine C, Baga M, and Graf RJ (2014) Canadian
(2010) Release of satiety hormones in response to specific dietary proteins is ancestral and present wheat varieties have similar protein concentration. Abstr./
different between human and murine small intestinal mucosa. Annals of Nutrition poster. Canadian Nutrition Society. June 2014. St. Johns Newfoundland, Canada.
and Metabolism 56: 308313. Reicks M, Jonnalagadda S, Albertson AM, and Joshi N (2014) Total dietary fiber intakes
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Consumer Trends in Grain Consumption
JM Jones and DB Sheats, St. Catherine University, St Paul, MN, USA
2016 Elsevier Ltd. All rights reserved.
Topic Highlights balanced with the looming specter of inadequate food supplies
to feed the anticipated population of 2050.
Trend drivers in developed countries are different from Consumer attitudes and ideas change continually, but these
those in emerging markets and developing countries, but changes seem to occur at a faster rate in the age of social media,
each can influence the other. blogs, battling experts, 24 h news cycles, and talk shows,
Grain consumption worldwide is increasing, but direct all which report the latest research findings sometimes with
consumption is decreasing as wealth increases and animal varying degrees of accuracy. If these voices persist, the
foods are selected. information regardless of its veracity causes changes in
Gluten-free is needed for wheat allergy and celiac disease food and grain consumption.
and related disorders but is unnecessary and expensive. It Perception is reality and consumer beliefs and trends drive
may be lower in nutrients especially dietary fiber and folate the marketplace. Thus, it is important that those producing and
for those who do not need to adopt this diet. manufacturing grain in planning for the long term be able to
Low-carbohydrate, antigrain, and other fad diets impact (1) differentiate between fads and trends, (2) to tailor food
grain consumption. product development and market launches so that product
Underconsumption of whole grains and overconsumption introduction and supply meet consumer expectations, and
of grain-based desserts and snacks are trends that need to be (3) to predict changes in the future that allow innovation and
addressed in many regions. are ready with supply chain and production to meet these
The selection of alternate grains and sprouted grains is changed demands.
another emerging trend.
availability of a wide array of food products. Many jobs also Bureau and other government agencies. Publicly available
require travel that introduces the population to more tastes, databases play a key role in assessment of nutrient intakes
different grains, and different ways of using grains. Many and evaluation of the risks related to possible hazards in food
multiethnic cities have markets and restaurants that offer an (Figure 1).
array of grain products from amaranth cereal, to buckwheat
groats, to corn tortillas, to porridge (oat meal), to quinoa pilaf,
to rye crisp, to sorghum beer, to rice noodles, to breads from all
over the world. Trends in Worldwide Wheat Consumption
In the 1990s and the first decade of the 2000s, world wheat
Tracking Consumer Trends consumption continued to expand in response to rising popu-
lation and incomes. Rising wealth fosters increased consump-
Governments have traditionally tracked the supply and avail- tion of meat, meaning that more wheat and grains are being
ability of grain-based foods. This is done in a variety of ways. used as animal feed, thus boosting total usage. However, in
Government and international food and agriculture bodies some countries, direct human consumption of wheat is
such as WHO/FAO collect statistics that give information on decreasing. For example, wheat consumption in the United
types and varieties of grain and other products grown; cost and States has decreased since 1997. The pattern of wheat con-
availability of seeds and inputs; stores of grain; weather, sumption is shown in Figure 2. Per capita wheat use declined
environmental, and other factors that affect crop planting, from a high in 1879 when one in every four was on the farm
yield, and market availability; food disappearance and and many engaged in hard physical labor, making the per
consumption; health issues; and economic factors. Data are capita consumption 102.27 kg. For nearly 100 years, until the
usually country- or region-specific, but trends in one region 1970s, wheat use dropped to a low of 50 kg per capita as energy
may influence consumption in another region. required for both work and play decreased, wealth increased,
Food consumption data collected by governments, and diets became more diversified. By 1997, wheat use had
academia, and industry monitor what has been eaten. If col- rebounded to 66.72 kg per capita. The overall growth in per
lected in a consistent manner, they document changes in pat- capita use that occurred between 1973 and 1997 reflected
terns of food intake. These may also be useful in predicting changes that included (1) the boom in away-from-home eating
what will be needed in greater or lesser quantities in terms of and fast foods with their buns or other grain-based food items,
nutrition and the marketplace and may help distinguish (2) an increase in ready-to-eat options such as breakfast
between what might be a fad and a trend. cereals, (3) widespread dietary guidance that promoted diets
Trend watchers and marketers conduct and scan material low in fat and high in carbohydrates and grains for controlling
from books, media, blogs, and other social media, especially in coronary disease, and (4) a wider recognition of health benefits
areas that intersect with food such as cooking, health and stemming from eating bran, oat bran, and other high-fiber,
medicine, energy and the environment, employment and grain-based foods.
income factors, and agricultural economics. Pronouncements The 30-year pattern of increased wheat consumption ended
about nutrition by various authoritative bodies impact food in 1997. There are many reasons for this trend: (1) the decrease
choices and messaging about food. Government regulation in energy expenditure, (2) concern about increases in rates of
also impacts what may be sold and what may be said about overweight, obesity, and rising chronic disease, and (3) a num-
it. For example, in Mexico, breakfast cereals are being rated ber of missteps with the introduction of low-fat products, often
with a tax as a measure to address obesity. This causes cereal
manufacturers to reformulate to meet government guidelines
and to launch more high-fiber or reduced-sugar versions.
Understanding how these factors interact can help predict Production + beginning stocks + imports
consumer food trends.
minus
Trends in Grain Usage and Its Impact on Consumers
Pounds/person
150
140
130
120
110
100
1968 1972 1976 1980 1984 1988 1992 1996 2000 2004 2008 2012
Source: USDA, Economic Research Service calculations using data through the
second quarter of 2011 from U.S. Department of Commerce, Bureau of the
Census Flour Milling Products (MQ311A) and U.S. Department of Commerce,
Bureau of Economic Analysis Foreign Trade Statistic. Subsequent flour milling ERS
calculations are based on data from the North American Millers Association.
formulated with grains, that failed to either address overweight the HEALTHGRAIN project in the EU. Nordic efforts to
or add nutrient quality. increase rye and oats and other whole grains are part of a
Thus, around the turn of the millennia, consumers healthy eating advice. In Italy and France, there also has been
switched from low-fat diets to low-carbohydrate diets of vari- a modest increase in whole grain products with successful
ous types such as the Dr. Atkins Diet Revolution and the South launches of whole grain pastas and other products. The
Beach Diet. Some of these ketogenic diets allowed only 200 cal Whole Grains Council, which has international reach, has
from all carbohydrates, so that there was a decrease in con- documented a marked increase in the number of whole
sumption of all carbohydrates and grains. This switch reduced grain foods and a shift to whole grain offerings in all
per capita wheat and grain consumption. Consumer interest in categories.
low-carbohydrate diets spiked after 2000, although this trend Data show that consumers in Western countries eat too
continues because of medical literature showing that it is a many refined grain servings and too few whole grain servings
successful diet pattern for some dieters. such that their calories exceed their energy needs. Grain-based
ERS estimated per capita wheat flour use at 60.22 kg in desserts and snacks provide nearly as many calories as sugar-
2011. New allegations about wheat and wheat products from sweetened beverages in the US diet of children and teens. Since
blogs and books bashing gluten, wheat and wheat breeding, many of these grain foods offer little in the way of nutrition, it
and grains in general continue to keep wheat consumption at is critical that health professionals and the food industry work
lower levels than in 1997. together to both improve the nutritional contribution of such
products and promote their consumption in the correct serving
sizes and as special treats.
Whole Grains, Carbohydrate Quality and Glycemic Concern about the glycemic impact of foods including
Response as Trend Drivers wheat and grain products has caused some to use glycemic
index (GI) in as part of dietary advice, especially for diabetics.
The type of grain and wheat products is also changing. Since Either some governments have allowed (Australia) GI labeling
2000, emerging science around the health benefits of whole or some merchants (the United Kingdom) have voluntarily
grains was translated into dietary guidance in many countries labeled foods including grain-based foods with GI. Some gov-
with recommendations to substitute a portion of refined and ernment bodies such as Health Canada have decided that the
enriched grain staples with whole grain breads and cereals, for GI is not appropriate for food labeling because of problems
example, to make half your grains whole. Across many with measurement and lack of consumer understanding. When
continents, this effort is reflected in groups such as Australia used, dietary guidance in these countries encourages con-
and New Zealands Grains and Legume Nutrition Council or sumers to select products in the same category with GIs
32 FOOD GRAINS AND THE CONSUMER | Consumer Trends in Grain Consumption
deemed low to moderate. In some cases, foods carried a GI More than 90% of the worlds rice is consumed in Asia, where
label, and then, for a variety of reasons, the voluntary labeling it is a staple for a majority of the population, including some of
was withdrawn. Such labeling can impact the type of grain the worlds poorest populations. Developing countries have
foods selected and has become a driver in the selection of long depended and continue to rely on rices versatility, protein
many carbohydrate foods. availability, and high caloric value.
From the 1960s to the 1990s, Asias per capita rice con-
sumption went from 85 kg per year to nearly 103 kg. In that
Gluten-Free and Wheat and Grain Avoidance Trends same period, per capita consumption rose from 50 to 65 kg per
year. The rising per capita consumption plus the growing pop-
Gluten-free and the avoidance of wheat, gluten, and grain have ulation more than doubled global rice consumption during
become both a fad and a trend in some developed countries in this period from 150 to 350 Mt.
the last few years. The trend is also being exported and adopted Changes in the standard of living have altered rice
in some emerging economies. consumption patterns. Increased wealth in countries such as
Data published in 2008 changed the thinking in health India and China slowed the rate of growth in rice consumption
professions regarding the incidence of celiac disease. These due to a switch to foods with higher value such as meat and
data showed that the incidence, instead of being 1 in 2500, is produce. In some countries such as Japan and Taiwan, there is
around 1 in 133, and it is underdiagnosed. And data from the decreasing rice consumption due to Westernization of the
United States, Finland, and even Asia Pacific countries where diets, a decline of family size, increasing participation rate of
the overall incidence is low show that celiac disease incidence women in labor market, and aging populations. However, rice
is increasing. consumption continues to rise in Southeast Asia (the
Nonceliac gluten sensitivity has been proposed as a gluten- Philippines and Indonesia), South Asia (India and Bangla-
related disorder. The condition has no medically vetted diag- desh), and many parts of Africa. After Asia, the second largest
nostic test and no agreement about its incidence, but is charac- rise in rice consumption occurs in sub-Saharan Africa. Overall,
terized by a cluster of symptoms including gas and distention. the rising populations of emerging and developing economies
Beyond these conditions, the trend toward reduced wheat cause a rise in overall rice consumption. For those needing to
and grain consumption is driven by books and blogs suggest- avoid gluten, rice has become an important substitute.
ing that modern wheat and grains are addictive leading to Recently, concern about arsenic in rice has been raised as an
overconsumption of grains and are causing obesity and other issue and may impact consumption.
chronic diseases and health conditions. These same sources In summary, increasing wealth in some countries may shift
demonize not only high sugar intake but also wheat and consumers away from rice and grains to other foods afforded
grains, especially refined grains. Some make consumers believe by higher-income consumers, but the increases in population
that foods labeled as gluten- or wheat-free are more nutritious in many countries offset any decline in worldwide consump-
than their gluten-containing counterparts. tion. Worldwide rice production was 600 Mt in 2000. It is
Summing of all those who need to avoid gluten those predicted to increase 1.5 times by 2030, to 904 Mt with gains
with wheat allergy, celiac disease, and nonceliac gluten in production in nearly all areas.
sensitivity means that around 68% of the population
needs to avoid gluten. In 2014, as many as 30% of the North
American population said that they sometimes avoid wheat Maize, Oats, Sorghum, and Barley
and gluten. This is also a trend in Australia and New Zealand
with 10% of the population indicating that they are avoiding Maize (corn), sorghum, barley, and oats are used more as feed
gluten. In terms of markets, the United Kingdom has the grain than as human food. Maize contributes one-third of the
biggest gluten-free market in all Europe, but Germany has the worlds cereal output. Demand for maize as a feed grain will
largest market on the continent. Italy and Scandinavia also increase. Globally, maize production has increased by nearly
have a sizeable market. Yet, all of these countries have about 50% due to expansion in Asia and high-yielding varieties
1% of consumers with celiac disease. Compared to other including genetically modified corn (bT-corn). Differences in
European countries, sales of gluten-free foods are on the low consumer attitudes to the GM corn around the world have
side in both France and Spain. dramatically affected its acceptance. The challenge this creates
In terms of trends, the gluten avoidance fad will have an for international trade of maize and future production must be
impact for a while and will likely diminish. However, there will addressed and can sharply impact trends in its use as a feed or
be a baseline trend for those with celiac disease, allergies, and food grain.
other gluten issues that will remain and may increase. Concern Only 15% of maize is processed for human food to become
over carbohydrate quality will remain high, and changes in a variety of milled grain products and starch, sweeteners, corn
types of carbohydrates and grain-based foods will serve as oil, and alcoholic beverages. Direct use of maize as human
another trend driver. food is unlikely to change significantly, even in countries
where maize is an integral part of local diets.
Oats are an important foodstuff in northern climates
Rice because they tolerate cooler summers and more rain than
many grains. Oat production has remained steady for the
Rice supplies around 20% of the calories consumed worldwide past 10 years. The EU region is the largest producer of oats,
and is a staple for nearly half of the worlds 7 billion people. but Canada is the largest exporter. The United States is the
FOOD GRAINS AND THE CONSUMER | Consumer Trends in Grain Consumption 33
worlds largest importer of oats, even though it is the worlds accounting for at least half of this amount. Using ERSs loss-
fourth largest oat producer. Mexico is the worlds second larg- adjusted food availability data from 2005, it is estimated that
est oat importer, followed by Japan. more grain servings than recommended (8.1 oz-eq of grains
Per capita consumption of oats is high in former colonies of per person per day versus the 6 recommended) were con-
the United Kingdom with Canada and Australia topping the sumed. Instead of half of these as whole grains, nearly 90%
consumption charts. Russia, the United States, Poland, the was as refined grains. Consumption data for children and teens
United Kingdom, and Scandinavia are countries with a higher show that nearly as many calories emanate from consumption
consumption of oats than many countries. In the United of grain-based desserts and snacks as do from sugar-sweetened
Kingdom, oats comprise 3% of the crops cultivated, but half beverages. Thus, Americans, on average, overconsume refined
the oat crop is used as human food where about 50% of the grains with added sugar and fat and underconsume whole
population eat porridge (oatmeal). grain staples. Only a small percentage of the population was
Among the grains, oats is viewed as a healthy grain. This eating the recommended amount of whole grain. Efforts to
trend began with the oat bran craze of the late 1980s when bring total amount of grain eaten in line with recommenda-
research documented the health benefits of oats. Its status as a tions while substituting whole grains for refined grains must
whole grain and high nutritional marks, with its micronutri- continue in order to reverse current trends.
ents, phytochemicals, and protein content ranking high among It is difficult to predict what economic and agronomic
the grains, continue to lure consumers. conditions and policies, geopolitical climates, and technical
The real driver for oats consumption is its viscous fiber advances will affect food pricing and availability and costs
b-glucan. The data on its cholesterol-lowering ability are so especially with population, fuel, and environmental pressures.
well documented that it has a health claim in a number of Trend drivers such as new health research, social media with
countries. The first of many claims was in the United States in food fads and scares, changing social norms, adoption of foods
1997 with other countries following. Interest in the health from other countries, and other issues related to consumers
value of oats is expected to continue as consumers continue will continue to affect consumption but are hard to predict.
to want good-tasting, high-fiber whole grains. Emerging Trend watchers and available data sources can track trends and
research on benefits of oats for blood glucose control and may be used to help answer critical questions about food
satiety keeps consumers interested, and the production of choices.
gluten-free oats for celiacs is driving demand for oats.
The only concern is that oats is the only major world crop
where production has shown a steady decrease since 1970. The Summary
decline amounts to a total of about 60%.
Consumption data for grains and grain-based foods relate to a
number of trends. Major factors that affect trends around the
Barley
world include demographics (age, household size, household
Only 2% of the barley crop is used directly for human food. income, women in the labor force, amount of education,
Malt production for beer takes over 40% of the crop. Barley, geographic location, and ethnic background), diverse lifestyles,
like oats, contains the soluble fiber b-glucan, which has been and consumer attitudes (toward food habits and food prefer-
associated with maintaining healthy cholesterol levels and ences). Consumer trends that affect movement of grain-based
healthy heart. In 2005, FDA amended its health claim for foods through the food system include food marketplace
soluble oat fiber and coronary heart disease to include barley. trends, food processing/manufacturing trends, and agricultural
Studies indicate that barley may also have potential in modi- trends. Major trends in the food system include niche market-
fying glycemic response in humans. Also, the vitamin E of the ing, development of grocery stores to improve food shopping
outer layers of whole barley may be important. There is a trend convenience, increased demand for and development of con-
by consumers to romanticize ancient grains, and barley is venience foods, and the growing demand for food away from
considered part of this ill-defined category. Use of barley as home.
sprouted grains is also showing some consumer interest.
Exercises for Readers to Explore the Topic Further Kubo M and Purevdorj M (2004) Future of rice production. Journal of Food Distribution
Research 35(1): 128142, http://ageconsearch.umn.edu/bitstream/27145/1/
35010128.pdf (accessed January 2015).
Discuss impacts of inputs (water, fuel, fertilizer, etc.) on Lin BH and Yen ST (2007) The U.S. Grain Consumption Landscape: Who Eats Grain, in
trends in grain consumption in a region. What Form, Where, and How Much? ERR-50. U.S. Department of Agriculture,
Study the interaction of food trends such as the locavore Economic Research Service.
movement, the Paleo diet, and other meat-based antigrain Mohanty S (2013) Trends in rice consumption. http://irri.org/rice-today/trends-in-
global-rice-consumption (accessed January 2015).
diets and the organic movement on grain and food produc- Rice Outlook/RCS-14l/December 12, 2014 Economic Research Service, USDA.
tion and the ability to feed a growing world population. Wells HF and Buzby J (2008) Dietary Assessment of Major Trends in U.S. Food
Consumption, 19702005. Economic Information Bulletin No. 33. Economic
Research Service, U.S. Department of Agriculture.
See also: Carbohydrates: Glycemic Index, Glycemic Load, and Their World Agricultural Supply and Demand Estimates WASDE-537 January 12, 2015.
Health Benefits; Health Effects of b-Glucans Found in Cereals; Food
Grains and the Consumer: Genetically Modified Grains and the
Consumer; Grains and Health: Misinformation and Misconceptions; Relevant Websites
Grains and Health; Food Grains: Intolerance, Allergy and http://www.agr.gc.ca/eng/industry-markets-and-trade/statistics-and-market-
Diseases: Celiac Disease; Grains Around the World: Grain information/by-product-sector/crops/crops-market-information-canadian-industry/
Production and Consumption: Overview; The Cereal Grains: Barley: market-outlook-report/oats-situation-and-outlook-august-2010/?
An Overview of a Versatile Cereal Grain with Many Food and Feed Uses; id1378843401619.
Maize: Overview; Oats: Overview; Rice: Overview; Wheat: An Overview www.barleyfoods.org (accessed January 2015).
http://www.efsa.europa.eu/en/datex/datexfoodcdb.htm.
of the Grain That Provides Our Daily Bread. http://ers.usda.gov/data-products/food-availability-%28per-capita%29-data-system/.
aspx (accessed December 2014).
http://ers.usda.gov/data-products/food-availability-%28per-capita%29-data-system/
food-availability-documentation.aspx.
http://ers.usda.gov/topics/food-choices-health/food-consumption-demand.aspx.
Further Reading http://www.ers.usda.gov/topics/crops/wheat.aspx.
http://www.ers.usda.gov/topics/crops/wheat/usda-wheat-baseline,-2014-23.aspx.
Abbassian A (2007) Maize international market profile. http://siteresources.worldbank. http://www.ers.usda.gov/topics/international-markets-trade/global-food-markets/
org/INTAFRICA/Resources/257994-1215457178567/Maize_Profile.pdf (accessed international-consumer-retail-trends.aspx.
January 2015). http://irri.org/rice-today/trends-in-global-rice-consumption.
Economic Research Service (ERS), U.S. Department of Agriculture (USDA). Food http://www.just-food.com/management-briefing/germany-canada-australia-among-
Availability (Per Capita) Data System. http://www.ers.usda.gov/Data/ other-growth-markets_id123020.aspx.
FoodConsumption; http://ers.usda.gov/topics/food-choices-health/food- http://www.nationmaster.com/country-info/stats/Agriculture/Grains/Oats/
consumption-demand/food-consumption.aspx. Consumption-per-million (accessed January 2015).
Cultural Differences in Processing and Consumption
AMR Hayes, St. Catherine University, Lafayette, MN, USA
JM Jones, St. Catherine University, Arden Hills, MN, USA
2016 Elsevier Ltd. All rights reserved.
Consumption of cereal grain varies by cultural practice and Cereal grains are staple foods among many cultures. They can be
country, and each variety contributes unique nutrients that consumed separately or in combination with other foods. Each
can be affected by different processing techniques and prep- variety of cereal grain contributes unique nutrients, mainly pro-
aration methods. viding carbohydrates, protein, dietary fiber (DF), and vitamins
The processing techniques used in different countries and minerals such as B vitamins, vitamin E, magnesium, and
impact starch digestion and energy yield of cereal grains; iron. Additionally, different processing techniques can affect the
most minimally processed grains are digested slowly, and bioavailability of certain nutrients in cereal grains. The aims of
highly refined grains provide slightly more calories than this article are to discuss the place of grains in the diet, the effects
less refined types. of processing on grain nutriture, some specific types of grains,
Cereal grains are incomplete sources of protein but can be and the overall impact of cultural practices of grain consumption
paired with other foods containing the amino acids they on diet.
lack to provide a complete source of protein, as is evident in
the traditional dishes of many countries.
Processing can either inhibit or promote the bioavailability Cultural Differences in Consumption
of minerals and toxic compounds, affecting nutrient needs
in certain regions. Energy requirements
Wheat is the most common cereal grain in the majority of Cereal grains can provide over 50% of both energy and protein
Western countries; refined or enriched flours are popular in in parts of the developing world because they comprise the
the United States, Europe, Latin America, South Asia, and fundamental components of most diets. Since nutrient require-
much of Australia. ments, food availability, and cultural practices vary by country,
Rice contributes approximately half of the calories for half the role of cereal varies. For example, abundant supplies of
of the worlds population and is common in regions of food in developed countries result in cereals comprising a
South and Southeastern Asia, Japan, Latin America, the smaller percentage of the diet and energy intake. The protein
southern part of the United States, and the Philippines. contribution of cereal grains is less important for individuals
Discussions of cereal grains (teff (tef), oats, rice, corn/ in the developed world than those in the developing world.
maize, barley, rye, millets, sorghum, and triticale) and Exceptions may be for those who are vegetarian or flexitarian.
pseudocereal grains (amaranth, buckwheat, and quinoa) Cereal grains that are highly refined may contribute to more
provide insight into their histories and current uses. available carbohydrate and slightly more calories than unre-
Increasing globalization and promotion of specific foods, fined varieties. This is partly due to structures in the outer layers
such as those that are gluten-free or whole grain (WG), are of grain trapping some of the starch and making its form
altering the types and roles of cereal grains in the diet of resistant starch (RS) type 1. Germ and bran phytochemicals
many regions. may inhibit the activity of amylases and proteases, thus reduc-
ing the total number of calories available. High levels of tannin
in some grains such as tannin-containing sorghum varieties
can have a significant impact on calorie availability in coun-
Learning Objectives tries where this grain is a staple. Refined grains may be bene-
ficial in developing countries, where calories are in high
To identify general variations in energy, protein, and nutri- demand. Alternatively, in developed countries where excessive
ent requirements among cultures. caloric consumption is widespread, the inclusion of WGs
To recognize and understand some of the changes in cereal with their slightly reduced carbohydrate contribution may
grains that occur with certain processing techniques or be advantageous.
cultural preparation methods.
To gain insight into the diverse histories of distinct cereal
grains, their nutritional qualities, and some of their current Protein
uses. Each cereal grain is characterized by distinct nutritional quali-
To attain an understanding of cereal grains that promotes ties. When the cereal grain is a major food source in a culture or
the exploration for future applications addressing changes region, its nutritional characteristics are of notable importance.
in dietary habits. In developed countries, protein quality and quantity of grains
are less important because other protein-rich foods are more Lowering the pH as occurs through a sourdough process
available and constitute a higher percentage of the diet. changes the activity of the amylase and lowers the GR. Cultural
Processing methods impact protein nutritional quality of practices of adding acid, as with oil and balsamic vinegar on
cereal grains. Milling, fermentation, and germination increase bread or vinegar in Japanese sushi, also lower the glycemic
the digestibility and bioavailability of protein and certain impact. Using fats such as oils or butters on bread or using
essential amino acids. Germination also increases the synthesis legume and nut accompaniments also lowers the GR.
of the amino acids lysine and tryptophan. When cooking and
dehulling (debranning or decortication) accompany fermen-
Resistant starch
tation of grains such as sorghum and millets, protein digest-
Processing techniques can impact starch digestion. Processing
ibility is increased because antinutrients such as tannins are
can affect the degree to which a starch is digested and absorbed
reduced. In the case of tannins, this means that tannins bind-
and can increase or decrease the content of RS. The following
ing of protein and inhibition of digestive enzymes (proteases)
processes increase RS: extrusion at high temperatures, as in the
are reduced.
making of cornflakes, followed by cooling to induce crystalli-
Cereal grains are generally incomplete sources of protein, as
zation of the starch molecules. Thus, amylase enzymes do not
they are limited in the amino acid lysine. Corn (maize) is also
readily penetrate the starch, which means some starch is undi-
limited in tryptophan, while wheat, barley, and rye are low in
gested. Treatments of repeated heat and moisture, such as is
threonine. Through pairing cereal grains with other foods con-
practiced in Africa where maize porridges are repeatedly heated
taining the limiting amino acids, a complete protein can be
and cooled, also create RS (type 3). Cooked kernels and bul-
achieved. This protein complementation is exemplified
gurs, as is popular along the Mediterranean, trap starch in the
throughout the world by national dishes that pair cereal grains
grain matrix and make it unavailable to digestive enzymes
with legumes. Cereal grains contribute methionine, which is
resulting in RS (type 1). Microwave cooking shortens starch
generally limited in legumes, while legumes contribute lysine
chains in many foods and speeds digestion by having more
and other amino acids. Some regions use a grain-based dish
rapidly available glucose (RAG) and less RS. Germination and
with a very small amount of animal protein such as cheese or
fermentation not only may decrease RS but also may change
meat. The following are a few examples of popular dishes:
the levels of RAG and SAG
Ethiopia: Kik alicha wot (spiced cooked split peas) with The following processes increase RS:
injera (teff flatbread)
India/Sri Lanka: Chapati (cereal unleavened flatbread) with
Steam cooking
dhal (legumes, often lentils)
Autoclaving
Parboiling
Middle East: Pita wheat bread with hummus (garbanzo
Baking
beantahini spread)
China: Congee (rice porridge) with toppings of tofu, fish,
Extrusion cooking (some studies show it decreases it
slightly also)
and sometimes meat
Mexico: Corn tortillas, beans, and rice, sometimes with a
Coarse milling
little meat or cheese Cultural preferences for grain varieties or grain processing can
Southern Africa: Samp (porridge from coarse milled maize) dramatically impact carbohydrate availability and the GR. For
with whole common beans or cowpeas example, the amount of RS and SAG in rice varies by processes
The United States: Peanut butter on whole wheat bread such as parboiling and variety. High-amylose varieties, such as
The United States: Macaroni and cheese basmati preferred in India, have lower GR. Conversely, varie-
ties high in amylopectin, such as glutinous rice preferred in
parts of China and Japan, have high GR. Most forms of proces-
Carbohydrates, including DF
sing, notably parboiling, often reduce GR.
Starch
Energy from cereal grains is primarily from starch. Processing
impacts starch digestion and energy yield. When grain is milled Dietary fiber
into flour, energy yield is increased. This is important in areas A growing body of evidence suggests that inadequate DF intake
of nutrient and energy deficiency. Germination and fermenta- is associated with decreased gut motility and increased risks of
tion result in the partial degradation of starch, which promotes heart disease, diabetes, and certain types of cancer. Diets in
starch digestibility. Grains that are cooked whole or are mini- many countries are characterized by inadequate intake of DF
mally processed may be digested slowly, therefore slowing and WG. Extraction rate, or the degree of milling, impacts the
glucose release into the bloodstream, referred to as slowly nutritional quality of cereal grains. Higher rates retain more
available glucose (SAG). This may be advantageous for satiety bran and germ and therefore have a higher level of DF and
and control of blood glucose and lipid levels. grain phytochemicals. Countries that regularly use higher
Processing can alter the ability of digestive enzymes to extraction rates, such as Norway, have increased DF and grain
access the starch. For example, the compact structure of an phytochemical intakes.
al dente pasta impedes access of the digestive enzymes to the DF intake is higher in countries that regularly include WG
starch, lowering the glycemic response (GR), but not as much in the diet. For example, regular consumption of WG products
as is seen, for example, in a spatzle noodle from Germany. made from whole wheat, rye, oats, and barley as occurs in
With the porous structure of breads, even WG ones, amylases Scandinavia and parts of northern Europe contributes to
have easy access to the starch and thus elicit a rapid GR. higher DF intakes than in many European nations. In parts of
FOOD GRAINS AND THE CONSUMER | Cultural Differences in Processing and Consumption 37
Asia where white rice is the staple, the DF contribution of cereal to note that enrichment and fortification efforts can be negated
grains is frequently low. In a traditional rural Mexican diet with by cultural practices. For example, washing rice prior to cook-
its staple of WG corn tortillas and beans, DF intake is high, ing it or cooking it in large amounts of water, which is then
whereas in the urban areas, DF may be lower due to a West- discarded, washes away much of the vitamin-rich talc added to
ernized diet pattern. enrich the rice.
The specific types of cereal grain typically consumed by Fermentation and malting processes can alter nutrient qual-
various cultures also affect nutriture, as each grain offers vari- ity. Fermentation typically results in increased levels of
ous fiber types and phytochemicals. Rye, a common choice vitamin B, but the extent of increase depends on the type
among Germans and Scandinavians, is relatively high in arabi- of grain and the particular process. Malting or germination of
noxylans. Wheat, which is common in North America and grains and pulses has been found to increase vitamin C and
southern Europe, contains fewer arabinoxylans than rye, but phosphorus availability. Such practices are especially impor-
its cellulosic bran is associated with gut health and improved tant in parts of the world such as India and Africa.
laxation. b-Glucans, which are found in barley and oats, are Mineral status is largely impacted by the grain choice and
often consumed in northern Europe. In parts of the world processing of a certain culture. Ligands in grain, such as tan-
where high-tannin cereals such as sorghum are the staple, nins and phytates, bind iron, zinc, and other metals, rendering
nutrient availability, especially that of minerals, is compro- them less bioavailable and leading to mineral deficiencies. How-
mised. Additionally, some hull materials may be bowel irri- ever, absorption of some minerals can be improved through
tants or can damage the gut tissue (e.g., oat hulls with their processing techniques. Notably, milling with debranning
razor-sharp pieces). removes most of the tannin and some of the phytate, increasing
mineral bioavailability. The crushing and pulverization of the
Micronutrients grain may also increase bioavailability.
The bran and germ portions of cereal grain contain the major- Traditional lactic acid (sourdough-type) fermentation can
ity of the vitamins and minerals. Overmilling and high degrees facilitate the phytase reaction, releasing the tightly bound zinc
of extraction even as practiced in producing white bread flour in many grains. The longer the fermentation process, the lon-
can remove the aleurone layers of grains, along with germ-rich ger the phytase enzyme is allowed to degrade phytate, thus
vitamins, minerals, and other grain phytochemicals. Thus, improving mineral status. In the 1970s, hypogonadal dwarf-
limited extraction and the inclusion of WG foods as is part ism was shown to be due to zinc deficiency, due to diets where
of traditional cultural food patterns may help maximize 85% of the energy needs came from whole wheat pocket bread.
micronutrient amount contributed by grains. However, con- With efforts to increase the bioavailability of zinc in food and
sumption does not always equal absorption, as certain micro- improve overall diet, the incidence of this health condition has
components may be bound or unavailable. While milling been greatly reduced.
reduces the overall number of micronutrients, it also makes
them more absorbable.
Toxic compounds
Heat treatment of some grains not only increases shelf life
Aside from nutrient-binding phytates and tannins, cereals and
but also is critical to preventing rancidity of all fats including
other plant foods can contain toxic or antinutritional sub-
essential fatty acids, fat-soluble vitamins, and antioxidants
stances. However, processing techniques such as milling can
within the bran. Rice bran and oats are just examples of grain
remove a majority of these:
or grain components that are typically subjected to heat treat-
ment in order to inhibit lipase activity of the grain and germ. Protease and amylase inhibitors found in cereal bran are
However, severe heat treatments may destroy vitamins, bind lost or reduced during milling, germination, or fermenta-
nutrients, and potentially promote rancidity. tion processes. Grain soaking or heating as practiced in
B vitamins in grains are particularly affected by processing. some cultures may also reduce the activity of these
Since thiamin is among the most heat-labile of the vitamins, it inhibitors.
is easily reduced through heating in alkaline pH. Thus, the Saponins that occur, for example, in some pseudocereals
thiamin content of chemically leavened (baking soda) baked not only can cause growth inhibition and are known for
products, such as Irish soda bread, can be decreased by over their hemolytic activity but also may have anticancer
70%. Conversely, processing via germination and fermenta- impacts on the colon. Again, the cultural practices of soak-
tion can increase B vitamins. In the case of nixtamalization ing, fermentation, and germination may reduce these.
(the process of liming corn), the tightly bound niacin is Mold toxins can affect any grain, especially those grown in
released from niacytin to make it available to the body. Thus, humid climates or harvested with high moisture levels.
in countries where maize was traditionally limed, pellagra Grain grading and inspection processes used in various
(niacin deficiency) was not endemic. regions are in place to prevent various mycotoxins from
Enrichment and fortification are used in many countries to entering the food supply. Milling may remove mycotoxins
replace some of the vitamins and minerals lost during milling. in the outer grain layers but not those contaminating the
Such techniques are used to address nutrient deficiencies and endosperm. Industrial milling practices have steps that
their corresponding health consequences. For example, pella- remove unsound grains or incorrectly colored grains in
gra was eliminated in the southeastern United States by niacin early points of the milling process as an attempt to remove
enrichment of corn. In countries with mandatory folate forti- any infected grains. Fermentation and nixtamalization may
fication of grain products, birth defects such as spina bifida are further reduce levels of aflatoxins, due to either an opening
reduced, in some cases, by over 50%. However, it is important of the lactone ring or a loss of the toxins into soaking liquid.
38 FOOD GRAINS AND THE CONSUMER | Cultural Differences in Processing and Consumption
Cultural Uses of Specific Grains and Their Impact magnesium than wheat or farro. Like many ancient wheat
on Nutritive Qualities relatives, the lower yields and greater costs of einkorn and
farro make them less widely available. They can be used in
Wheat and Its Parent/Derivative Grains and Products any way that wheat is used, and they are now being sought as a
Wheat specialty grain for variety.
Wheat and its related grains such as durum spelt and other
wheat parents are used as flour, wheat berries, bulgur, cracked Kamut
wheat, and coarsely milled fractions such as farina or semolina. Kamut, a relative to durum (also known as khorasan), was
All types of wheat can be used for pastas, but durum makes the originally cultivated in the Fertile Crescent, and is still used
optimal product. along the Mediterranean. Given its naturally higher sugar con-
In most Western countries, more foods are made with tent, kamut has a sweet nutty flavor. It provides B vitamins,
wheat than with any other cereal grains. While WGs are used, selenium, and magnesium; is higher in zinc, magnesium, and
refined or enriched varieties are popular in the United States, selenium than modern wheat; and provides a higher amount
Europe, Latin American, South Asia, and much of Australia and of protein but a similar amino acid profile compared to hard
Africa. Breads, pastries and small cakes, breakfast cereals, and red wheat. Kamut can be used as whole kernels for pilafs or
pastas are just some of the ways wheat is used. Noodles are ground into refined WG flour to be used in breads, pasta, and
found around the world in various forms. In many countries, other grain-based foods.
they are made from durum wheat. Couscous, while often
mistaken for a grain, is a steamed semolina, which is a staple Spelt
of cooking in North African countries such as Morocco, Spelt is known as dinkel to Germans and farro (but different
Algeria, Tunisia, Libya, and Egypt. To produce couscous, from emmer, also called farro) to Italians. It dates back to 7000
yellow granules of semolina, which are made from durum BC. Like modern wheat, this ancient parent has winter and
wheat, are parcooked and then dried. Its culinary uses as a spring varieties as well as hard and soft types. It is mainly used
salad or pilaf base, an addition to soups, an accompaniment in breads and pastas. Its protein content is only slightly higher
for meats and vegetables, or a dessert make it like other small than modern wheat, but it is higher in selenium, phosphorus,
grains and some pastinas. Both plain and flavored varieties of and magnesium than most cereal grains. Spelt WG flours and
couscous exist. doughs contain significantly higher amounts of RS and give
less available starch. However, baking or other processes affect
this, with breads showing the highest starch availability and
Bulgur
cookies the lowest. All of the grains related to wheat contain
Whole wheat kernels that are broken to become cracked wheat.
gluten and must not be used in the preparation of products for
If they are soaked, precooked, and then dried, they become
the gluten-free market.
bulgur. Evidence indicates bulgur has been eaten by Arab,
Israeli, Egyptian, and Roman civilizations as early as
1000 BC, while the Chinese consumed bulgur as early as Teff (Tef)
2800 BC. Currently, bulgur can be sold as a pilaf or tabouli,
Teff, a tiny, slightly sweet-flavored grain, is so small its name
taboule, tabbouleh, or tabouleh mix.
means lost grain. The bran and germ compose a larger per-
centage of the grain than other grains, and it has high levels of
Grano RS. Teff provides fiber and B vitamins, has a higher overall
Grano is a specialty durum that has been cultivated in Italy mineral content than most other cereal grains, and is known
since the fifth century BC. Its origins in Sicily and Puglia make for its levels of iron and calcium. However, it may not be as
it a popular food choice on certain holidays, but it is a newly high in iron as was once thought. Currently, it is also being
emerging product in the United States. Firm and chewy in grown more widely in India, Australia, and Canada.
consistency, grano is made from specially selected durum Traditionally, teff is ground into flour and fermented to
wheat. Although it most resembles barley, its bran has been make spongy sourdough bread called injera (an Ethiopian
removed, making the starch more accessible. Grano can be flatbread resembling a spongy large pancake). Teffs use in
cooked like arborio rice in dishes similar to risotto and either some other countries has expanded into foods featuring
used as a hot accompaniment with pasta or served cold in ancient grains or multigrain blends, such as tortillas, pizza,
salads. crackers, pasta, pancakes, snacks, breads, cereals, and many
other products. Its application into gluten-free products is
Einkorn/farro/emmer expanding.
Einkorn and its cousin farro are hulled wheat. They were two of
the first types of wheat to be domesticated about 11 millennia
Oats
ago in Georgia, Armenia, and Turkey. Its name, however, is
from the German meaning first grain. Farro/emmer is known Oats are a popular grain in regions where they thrive, including
by different names depending on the region, with the most Russia, Canada, North America, Finland, Poland, and the
common ones being farro and emmer. Nutritionally, einkorn United Kingdom. In countries that were once part of the British
is somewhat higher in protein than modern wheat or farro, but Commonwealth, porridge was standard breakfast fare with
all have lysine as their limiting amino acid. Einkorn contains steel-cut or pin-milled oats being preferred for their chewy
somewhat higher levels of phosphorus, potassium, iron, and texture and nutty flavor. While this is still a breakfast option,
FOOD GRAINS AND THE CONSUMER | Cultural Differences in Processing and Consumption 39
changing habits have caused a decrease in the intake of por- with game. It is often mixed with other rice and can be used
ridge and an increase in oats in beverages, as additions to any way that rice may be.
yogurt and as the basis of ready-to-eat cereals and breakfast
and energy bars. For other parts in the British Isles and most of
Corn/Maize
Scandinavia, oats are found in many foods such as oatcakes,
biscuits, crackers, and even soup thickeners. In the United Corn, also known as maize, is a staple in parts of Central
States, most groats are steamed and flattened into rolled oats America, Africa, India, and China. It can be used alone or in
(old-fashioned or regular oats, quick oats, and instant oats). combination with other foods and is the most widely grown
The higher the degree of flattening and steaming, the quicker crop in the Americas. The domestication of corn occurred over
the cooking time and softness of the end oats. In Germany, 9000 years ago in Mexico. Corn was carried from the Americas
Switzerland, and other parts of Europe, muesli-type cereals, to Europe and Africa by early explorers and traders, so it
based on uncooked, rolled oats with a combination of nuts, became important in the cuisines of many countries through-
fruits, and other grains, are popular. out the world.
Part of oats popularity is due to their nutrition and satiety Maize contains a small amount of provitamin A (which can
properties. They contain more than 20 unique polyphenols contribute up to ten times the content of other grains) and is
called avenanthramides. They have high antioxidant, anti- high in antioxidants and carotenoids associated with eye
inflammatory, and anti-itching activities. They are high in health (lutein and zeaxanthin). Its protein quality is poor and
b-glucans. These fibers have been proven to lower serum cho- niacin tightly bound, so it is associated with pellagra in mar-
lesterol and help maintain a healthy blood glucose level. A ginal diets based on corn. Nixtamalization, a process used in
number of countries have health claims that tout these benefits. Latin America, releases the niacin and improves amino acid
availability in order to prevent pellagra. This process of soaking
the grain in lime (calcium oxide) gives it a flavor that charac-
terizes the Latin kitchen. Traditional dishes mixing corn with
Rice
beans and rice in many cultures provide amino acids to make a
Rices roots date back to South Asia. It has been dispersed and complete protein. Flour from the nixtamalized corn is masa
cultivated in a wide variety of habitats and consumed by harina, which is made into tortillas, enchiladas, and many
humans for centuries. It is grown on every continent except other dishes.
Antarctica. Rice provides about half the calories for up to half In the United States, corn was originally soaked in lye to
of the worlds population, especially in South and Southeast- remove the pericarp. It was then dried to make hominy and
ern Asia, Japan, Latin America, the southern part of the United hominy grits. Hominy and corn meal are popular throughout
States, and the Philippines. Rice has a role in religious and the southern and southwestern United States with foods such
ceremonial occasions and medicine in many cultures. It is so as corn pone and hush puppies. Cheese and grits are popular in
important in certain regions that saying Hi. How are you? is the southeastern United States and spicier corn dishes and
literally asking Hi. Have you eaten rice today? hominy lamb stews (posole) are popular in the Southwest.
Among various parts of the world, the type of rice desired Corn-based breads and hot cakes are used throughout the
can be very different. Rice is classified by texture, color, and United States and can be made of corn alone or have wheat
size: long-, medium-, or short-grain varieties. Short-grain vari- added to them. The method and ingredients vary across the
eties can be referred to as sticky because of their higher amy- country. In Italy and Spain, corn is used for dishes similar to
lopectin content (starch), while brown rice (WG rice) is known polenta. Ugali in Kenya and mealie porridge in South Africa are
for its higher levels of vitamins and minerals, and colored rices just a few of the corn dishes that are also important in Africa.
are prized for their variety and antioxidant capacity, the latter
being due to their high anthocyanin content.
Barley
Among the diverse rice uses throughout the world are Chi-
nese stir-fries, Spanish paellas, Japanese sushi, Indian Barleys use as human food is small despite its being the grain
biryani, Thai salads, Middle Eastern pilafs, Italian risottos, with the fourth highest production in the world. The amount
Mexican beans and rice, Senegalese yassa, Norwegian rice of milling required to make the barley edible varies by whether
pudding, American gumbos, and so many more. the grain is hulled or hulless (naked). Like chaff of the wheat,
Wild rice is not actually a kind of rice. Instead, it is a semi- the outermost layers are not digestible and must be removed by
aquatic grass historically grown in lakes, tidal rivers, and bays. processing. For hulless varieties, the outer hull is loosely
Native Americans traditionally harvested the wild rice in north- attached so very little polishing is required for its removal.
ern lakes and processed it by parching the green seeds in large Whether hulled or hulless, most barley is polished (pearled)
pots over open fires and then packing them away in birch-bark to some degree, varying from regular to medium to fine to baby
baskets. Commercially, wild rice farmers cure the rice until the pearled barley. On average, pearled barley is about one-third
chlorophyll dissipates, and then while the kernel is still in the lower in micronutrients than unpearled. Pot barley (also called
hull, it is dried. The inedible hull is removed mechanically, scotch barley) has the hull polished off, but the polishing
leaving the kernel exposed. All these processes give wild rice its process is least invasive.
signature nutty, smoky flavor. It is slightly higher in protein Barleys nutritional contribution is similar to wheat with a
than most other WGs and is a good source of fiber, folate, few differences. It is highest in fiber of all the WGs, containing
magnesium, phosphorus, manganese, zinc, vitamin B6, and on average 17 g of fiber/100 g, with a range from 12 to
niacin. Native Americans mixed it with berries and used it 40 g/100 g. The fiber in barley is b-glucans, and unlike in
40 FOOD GRAINS AND THE CONSUMER | Cultural Differences in Processing and Consumption
additional texture and flavor. Some amaranth applications are determines the presence and bioavailability of micronutrients
listed as follows: and macronutrients. In most developed countries where
overweight is a problem, consumption of WGs may be
South American countries: sold on the streets (like
advantageous, whereas refined grains remain popular and
popcorn)
may benefit areas where calories are in short supply. With
India, Mexico, Nepal, and Peru: traditional ingredient for
increasing globalization, grains and foods specific to one
breakfast porridge
region are spreading across the globe and enriching the diets
Mexico: dulce de alegria, a sweet cookie-confection dessert
of many both nutritionally and culturally. The resurgence of
plant-based and gluten-free diets has made traditional and
Buckwheat alternative cereal grains important dietary contributors.
The variety of ways in which cereal grains are processed and See also: Carbohydrates: Cereals: Chemistry and Physicochemistry
consumed around the world is both interesting and important of Non-starchy Polysaccharides; Food Grains and the Consumer:
in terms of providing calories and contributing to nutrients to Consumer Trends in Grain Consumption; Processing of Grains:
the diet. The extent of grain refinement and cultural practices Starch: Uses of Native Starch.
42 FOOD GRAINS AND THE CONSUMER | Cultural Differences in Processing and Consumption
Further Reading Shewry PR (2007) Improving the protein content and composition of cereal grain.
Journal of Cereal Science 46(3): 239250.
Abdel-Aal E and Rabalski I (2008) Effect of baking on nutritional properties of starch in
organic spelt whole grain products. Food Chemistry 111(1): 150156.
Jones JM (2012) Glycemic index: The state of the science, Part 1 The measure and its
variability. Nutrition Today 47(5): 207213.
Molina J, Sikora M, Garud N, et al. (2011) Molecular evidence for a single evolutionary Relevant Websites
origin of domesticated rice. Proceedings of the National Academy of Sciences
108(20): 8351. http://www.grdc.com.au/
Mosley BS, Cleves MA, Siega-Riz AM, et al. (2009) Neural tube defects and maternal http://www.homebaking.org/hbamembers/
folate intake among pregnancies conceived after folic acid fortification in the United http://hsc.csu.edu.au/food_technology/issues/
States. American Journal of Epidemiology 169(1): 917. http://www.ifst.org/learninghome/helpforteachers/lessonplantopics/cereals/
Schroeder N, Gallaher DD, Arndt EA, and Marquart L (2009) Influence of whole grain http://www.kswheat.com/
barley, whole grain wheat, and refined rice-based foods on short-term satiety http://wbc.agr.mt.gov/
and energy intake. Appetite 53(3): 363369. http://dx.doi.org/10.1016/j. http://www.wheatfoods.org/
appet.2009.07.019. http://wholegrainscouncil.org/whole-grains-101/
Fortification of Grain-Based Foods
CM Rosell, Institute of Agrochemistry and Food Technology (IATA-CSIC), Paterna, Valencia, Spain
2016 Elsevier Ltd. All rights reserved.
Topic Highlights Since fortification is a broader term, it will be used here while
referring to the addition of nutrients.
Fortification is a way of supplementing the diet through the
intake of some nutrients to combat deficiencies.
Foods selected as vehicles for fortification must meet cri- Criteria for Food Fortification
teria of availability and price.
Cereals are the best carriers for food fortification, mainly in Minerals and vitamins may be added to foods only for the
the case of vitamins and minerals. purpose of (1) restoration of nutrients lost during processing,
Chemical compounds must be carefully selected for ensur- (2) nutritional equivalence of substitute foods, (3)
ing micronutrient bioavailability after processing. fortification, and (4) ensuring the appropriate nutrient com-
Fortification is legislated in different countries. position for a special purpose food.
Some considerations should be taken into account prior to
addition of any nutrient to foods:
Learning Objective
The nutrient added should be present at a level that will not
result in either an excessive or an insignificant intake, con-
Understand the meaning of fortification and the role of
sidering the amount provided from other dietary sources.
cereals and cereal-based foods in fortification.
The maximum safe amount to be added should be set
considering the levels established by generally accepted
scientific data and bearing in mind the diverse degree of
Introduction
sensitivity shown by different consumer groups.
Fortification can be defined as the addition of one or more Nutrients should not be added to fresh products including
meat, fish, fruits, and vegetables, neither to beverages con-
vitamins and/or minerals to a food, regardless of their usual
taining more than 1.2% by volume of alcohol.
content in the food. Additions are carried out based on gener-
ally accepted scientific knowledge of the role of vitamins and The nutrient added should be biologically available from
the food and should not interfere with the metabolism of
minerals in attaining good health. Foods are fortified for the
any other nutrient.
following reasons:
The nutrient added should be sufficiently stable during
to prevent or correct a demonstrated deficiency of one or packaging, storage, distribution, and end use and should
more vitamins and/or minerals in the population or spe- not affect both the sensory characteristics and the shelf life
cific population groups; and of the food.
to improve the nutritional status of the population and Technology and processing facilities should be available to
dietary intake of vitamins or minerals due to changes in permit the addition of the nutrient and also methods of
dietary habits. measuring and controlling the levels of nutrient added.
health. An uncontrolled food fortification program could Table 1 Vitamins and minerals that may be added to foods
result in excessive intake of certain nutrients, thus creating
nutrient imbalances, and in consequence representing a risk Vitamins Minerals
to the health of consumers. It must be had in mind that while Vitamin A Calcium
deficiencies provoke diseases, the intake of too much of a Vitamin D Magnesium
vitamin or mineral also may cause health problems. In fact, Vitamin E Iron
too much vitamin A can cause birth defects, too much folate Vitamin K Copper
can hide symptoms of a vitamin B12 deficiency which can lead Vitamin B1 Iodine
to damage of the nervous system, and too much calcium can Vitamin B2 Zinc
induce kidney problems. Niacin Manganese
In developing countries, food fortification is one approach Pantothenic acid Sodium
Vitamin B6 Potassium
toward solving nutrient deficiencies, and so far the implemen-
Folic acid Selenium
tation of these programs has been very successful in correcting
Vitamin B12 Chromium
nutritional deficiencies of large segments of the population Biotin Molybdenum
within a very short period. In these countries, care should be Vitamin C Fluoride
taken that the food selected as a medium for the nutrient Chloride
addition is stable and routinely consumed by the population Phosphorus
at risk, and that the amount of nutrient added is sufficient to
correct the possible deficiency.
However, in any case fortification programs should include
adequate Quality Assurance and Quality Control (QA/QC) at
production level, regulatory monitoring of the nutrient content Once the food or carrier to be fortified is selected, the
of fortified foods, as well as assessment methods to document nutrient deficiencies affecting the population must be deter-
the nutritional and health effectiveness when implementing mined. There are only certain vitamins and minerals that may
fortification strategies. be added to foods (Table 1). Nevertheless, the main micronu-
trients recognized as being of concern to the health of the
population are iron, folic acid, B-vitamins, vitamin A, and
Cereal-Based Products as Vehicles of Minerals zinc, although in developed countries calcium also is
and Vitamins considered.
In general, the population of all the rice-consuming coun-
One of the most critical decisions concerning food fortification tries have vitamin A deficiency, which is associated with cor-
is the selection of the appropriate food to act as a carrier of the neal lesions that can lead to partial or total blindness, and with
nutrient. The basic criteria for identifying the food products for reduced resistance to infectious diseases. This would, in conse-
fortification include selecting a food that is regularly consumed quence, lead to an increase in morbidity and mortality.
by the target population group, affordability, and availability Another deficiency associated with people of rice-consuming
all through the year. countries is anemia due to iron deficiency. This has been linked
Cereals and cereal-based products constitute a major com- to reduced resistance to infection and also affects the cognitive
ponent of diet around the world. Cereals have been used as the development and physiological functions in children and, in
basis of human diet from the ancient times and their implica- severe cases, causes maternal deaths. Iodine constitutes the
tion in the human diet is still very significant. third major deficiency in rice-consuming countries. This min-
Globally, cereals account for 48% of total energy intake and eral is necessary for proper fetal development and also for
they are a cheap source of proteins compared to animal pro- normal physical and mental activities in adults.
teins. Although their relative contribution to human diet In developed countries, iron supplementation has been a
greatly depends on the production and the cultural differences, measure adopted to reduce the prevalence of iron-deficiency
the popularity of cereals, mainly rice and wheat, in the diet anemia. B-vitamins (thiamine, riboflavin, and niacin) and
worldwide make them the most appropriate vehicles for forti- calcium are also supplemented in developed countries to
fication strategies. Additionally, it must be remarked that ensure the adequate intake of those nutrients. Lately, some
cereals are not directly consumed as grains; processing is gen- studies confirming the role of folates in congenital malforma-
erally required to transform the grain prior to its consumption, tions (neuronal tube defect) and the development of chronic
with the subsequent reduction in nutrients and micronutrients diseases like Alzheimers in elderly people have recommended
that are mainly concentrated in the outer layers and germ. In the supplementation of folic acid in cereal-based foods. How-
the conversion of cereal grains to white flour, 20 or more ever, the effectiveness of this fortification program in the
ingredients (such as thiamin, riboflavin, niacin, pyridoxine, decrease of homocysteine concentration and its relationship
folate, panthothenate, biotin, vitamin E, calcium, copper, with a reduction in the incidence of cardiovascular disease is
iron, potassium, magnesium, manganese, phosphorus, zinc, still under research.
chromium, fluorine, molybdenum, and selenium) are lost to Selenium is an essential micronutrient for humans and may
the extent of 5090% during the milling process. Cereal forti- reduce the risk of degenerative diseases, including cancer, but
fication methods have been developed to restore the nutrients its deficiency has been detected in at least a billion people
that have been removed during milling and to improve the worldwide, although wheat is a major source of dietary sele-
nutrient intake level of specific population. nium in humans.
FOOD GRAINS AND THE CONSUMER | Fortification of Grain-Based Foods 45
Methods for Cereal Fortification nutrient supplementation in this method can be washed off if
the grains are rinsed before cooking.
Cereal fortification methods are mainly addressed to restore the The grain enrichment method involves the addition of a pow-
nutrients that have been removed during milling and to improve dered mixture of vitamins and minerals and a subsequent coating
the nutrient intake level of specific population. In addition, some of the grain with a water-insoluble substance. Usually, grains
efforts have focused on enhancing the nutritional quality of the enriched by this method contain a high nutrient concentration,
crops because half of the world population suffers from malnu- and they are mixed in a proportion of 0.5% with normal milled
trition with respect to iron, zinc, and selenium. rice to finally obtain an enriched product that meets the required
standard levels. Different variations of these methods have been
developed through the years. For instance, the powdered mixture
can be sprayed onto the rice and then coated with ethanol,
Biofortification
isopropanol, palmitic acid, or cellulose derivatives. This method
Biofortification, the increase of micronutrients in the edible has been effective for enriching rice with niacin, thiamin, pyri-
parts of the plants, can be achieved by either mineral fertiliza- doxine, vitamin A, vitamin E, folic acid, iron, and zinc by adding
tion or plant breeding. Biofortification by mineral fertilization alternative layers of nutrients and coatings.
is a common practice in some countries that applied selenium- Synthetic rice obtained by extrusion of rice flour to a rice
containing fertilizers as a short-term solution for improving kernel shape in the presence of vitamins and minerals also
selenium content of wheat. covers the fortification objective. The fortified simulated grains
Biofortification through plant breeding consists of the are then mixed with normal milled rice, although the con-
development of micronutrient-enhanced crop varieties sumer often notices and removes those grains before cooking.
through conventional breeding. Nevertheless, the implemen- This is obviated by dyeing them with a distinct color, so that
tation of this type of biofortification requires the identification the consumer clearly knows that it should be present.
of genetic resources to determine the interaction of genotype
and environment, to define the desired level of micronutrient
increase, and finally to assess the cost-effectiveness. So far, it Flour and Bakery Products
seems a sustainable and cost-effective approach for reaching
biofortification in zinc and iron. Fortification of flour is usually carried out in mills, where the
nutrient mixture is blended with the flour. In this practice, the
definition of the nutrients and the levels to be added must be
based on the nutritional needs and deficiencies of the popula-
Rice Fortification
tion, the common consumption of the fortifiable flour, the
Rice requires special methods for fortification as it is com- sensory and physical aspects, and economic cost. The addition
monly consumed as a whole grain and is extensively washed of B-vitamins, iron, and calcium is a common practice in some
prior to cooking or strained after cooking. There are different developed countries. However, the nutrient added must have
methods for rice fortification; the earliest one was the produc- good stability during storage, and in defining the levels to be
tion of parboiled rice. In the parboiling process, rice is steamed added must be considered the extraction rate during milling. In
under pressure to gelatinize the starch within the kernel; this fact, some recommendations have been established for wheat
process allows the transference of nutrients from the bran layer flour fortification (Table 2).
to the starchy endosperm. Fortification can even take place during the breadmaking
Other methods currently used are the powder and grain process. Different yeast strains have been suggested for obtain-
enrichment methods. A powder preblended with mixtures of ing enriched baked goods. A selenium-enriched yeast has been
vitamins and minerals is added to the rice. For white parboiled used for increasing the content of selenium in bread, obtaining
rice, the blend is added immediately after milling because the wheat bread rolls that can provide 25% of the recommended
heat and moisture content of the grain surface at that moment daily allowance in the form of selenomethionine, which is the
facilitates the adherence of the powder mixture. However, the best form of selenium for human consumption.
Table 2 Average levels of nutrients to consider adding to fortified wheat flour based on extraction, fortificant compound, and estimated per capita
flour availability
Level of nutrient to be added (ppm) by estimated average per capita wheat flour availability
(g day1)
Source: WHO, FAO, UNICEF, GAIN, MI, and FFI (2009) Recommendations on wheat and maize flour fortification. Meeting Report: Interim Consensus Statement. Geneva: World Health
Organization. http://www.who.int/nutrition/publications/micronutrients/wheat_maize_fort.pdf (accessed 10.12.13).
46 FOOD GRAINS AND THE CONSUMER | Fortification of Grain-Based Foods
Three different requirements should be considered when they are poorly absorbed and their absorption is highly depen-
selecting the source of nutrients: the chemical compound (1) dent on the meal served (phytates, calcium, and polyphenols
should be stable during packaging, storage, distribution, and decrease the bioavailability of iron). Ferrous sulfate is preferred
end use; (2) should not modify the sensory characteristics for enriching wheat flour because of its high bioavailability but
(appearance, texture, taste, and aroma); and (3) should ensure it often leads to unpleasant colors and flavors due to reactions
the bioavailability of the nutrient. Different chemical com- with other components of the food matrix. Some of these
pounds are used as a source of vitamins (Table 3) and minerals interactions with other food components can be avoided by
(Table 4) to be added to cereals. coating the iron form with hydrogenated oils or ethyl cellulose.
Nutrient stability is very important in the case of vitamin Iron fortification of flour is generally regulated in most coun-
fortification. Vitamins are sensitive to heat, oxidizing and tries by specifying a minimum iron level in the flour resulting
reducing agents, light, and other physical and chemical stress. from the sum of the added and native iron. Nevertheless,
For instance, vitamins are stable when added to flour but the Hurrell et al. reviewed the effectiveness of the iron fortification
combination of high humidity and temperature adversely programs in 78 countries and concluded that they are likely to
affects the stability of the vitamin A. This vitamin is labile to be ineffective. Legislation needs to be updated in many coun-
high temperature and unstable in the presence of oxygen and tries so that flour is fortified with adequate levels of the recom-
light, due to its susceptibility to oxidation. Therefore, a protec- mended iron compounds to meet population requirements.
tion and appropriate conditions of storage should be used in
order to reduce the vitamin A loss. Encapsulation in a more
hydrophilic coat is a common practice in order to obtain a
more water-dispersible product. Legislation Concerning Cereal Fortification
The supplementation of B-vitamins to flour is affected by
low stability of these compounds during storage. Hence, higher Although cereal fortification can be an effective means of
effectiveness is obtained when B-vitamins are added at the improving micronutrient status, there are some barriers to the
bakery rather than at the mill. general implementation. These include concerns related to a
The bioavailability of added nutrients, particularly iron, proliferation of the fortified foods if the fortification is volun-
varies widely depending on the iron compound used for forti- tarily practiced by manufacturers, which would lead to a simul-
fication and the physical and chemical properties of the food taneous replacement of the nonfortified foods in the diet. The
to be fortified. The selection of the iron form is based on success of fortification programs always depends on good con-
organoleptic considerations, bioavailability, cost, and safety. trol, so they should be set up, regulated, and enforced by the
Iron powders are extensively used in flour fortification, but national governments.
FOOD GRAINS AND THE CONSUMER | Fortification of Grain-Based Foods 47
Table 4 Compounds that may be added to cereals as a source of There are currently 14 countries with legislation or regula-
minerals tions that mandate the fortification of wheat flour, whereas in
countries such as Finland, New Zealand, and Norway law pro-
Minerals Chemical compound
hibits the addition of nutrients to wheat flour. In the United
Calcium Calcium carbonate States, the Food and Drug Administration approves the supple-
Calcium chloride mentation of four B-vitamins (thiamine, niacin, riboflavin, and
Calcium salts of citric acid folic acid) and iron to the wheat flour, for obtaining enriched
Calcium gluconate flour, although this practice is regulated by Federal law. The US
Calcium lactate government adopted a mandatory regulation concerning folic
Calcium hydroxide acid fortification. In the United Kingdom, it is mandatory for
Copper Cupric carbonate the restoration of all the nutrients removed with the bran in all
Cupric citrate
types of flours, with the exception of whole meal. From 1997,
Cupric gluconate
the Organizacion Panamericana de la Salud (OPS) is support-
Cupric sulfate
Iodine Sodium and potassium iodide ing the supplementation of iron to wheat flour due to its low
Sodium and potassium iodate cost and extensive consumption. Some countries from South
Iron Ferrous carbonate and Central America have implemented this measure for
Ferrous citrate reducing the incidence of iron-deficient anemia. The levels of
Ferric ammonium citrate nutrients to be added are also regulated by national
Ferrous gluconate governments.
Ferrous fumarate At the international level, GAINs (Global Alliance for
Ferrous lactate Improved Nutrition) National Fortification Program, which
Ferric diphosphate
began in 2003, has expanded to support 19 countries with
Elemental iron (carbonyl electrolytic hydrogen
high levels of vitamin and mineral deficiencies. A broad coali-
reduced)
Hydroxocobalamin tion of governments, businesses, international organizations,
Magnesium Magnesium acetate and civil society partners, known as National Fortification
Magnesium carbonate Alliances, support GAINs projects. Those projects aim to for-
Magnesium chloride tify foods and condiments including wheat and maize flour,
Magnesium salts of citric acid sugar, vegetable oil, milk, soy sauce, and fish sauce. Vitamins
Magnesium oxide and minerals used to fortify foods include vitamin A, vitamin
Magnesium sulfate D, iron, zinc, folic acid (B9), thiamin (B1), riboflavin (B2),
Manganese Manganese carbonate niacin (B3), pyridoxine (B6), and cobalamin (B12).
Manganese sulfate
Manganese chloride
Manganese citrate
Phosphorus Calcium salts of orthophosphoric acid
Zinc Zinc acetate
Zinc chloride Future Prospects
Zinc oxide
Zinc sulfate Currently, in the developed countries the term functional
Zinc carbonate foods or nutraceuticals seems to be old-fashioned, and
the cereal-based products surely may be one of the best
carriers of different compounds related to physiological
functions. Therefore, the near future in nutrition could be
The addition of nutrients is generally practiced by manu- the use of cereal-based products as carriers of o-fatty acids
facturers either voluntarily or because it is compulsory under and probiotic compounds such as fructo-oligosaccharides,
national rules. In general, vitamins and minerals may be added inulin, etc.
in conformity with the legislation of the country in which the
product is being sold, which must regulate the list of foods to
which vitamins and minerals may be added, the vitamins and
minerals permitted, and the levels at which they may be added. Exercises for Revision
The Economic and Social Department of FAO has elabo-
rated a review about the legislation status concerning rice What requirements should the nutrients to be used in
(Table 5) and wheat flour (Table 6) fortification. fortification meet?
Rice fortification is voluntary in the United States; however, When and where would food fortification be recommended?
most of the rice sold is fortified, and lately folic acid is also Which cereals would be the selection for food fortification?
being added to the enriched rice. In Canada, fortification of When using wheat white flour as a fortification carrier,
rice is also voluntary, and when labeled as enriched, it contains which chemical compounds would be advisable to be
vitamin B1, vitamin B3, and iron. Moreover, in Canada, the added? Cross reference to wheat milling chapters.
addition of vitamins B5 and B6 and folic acid is optional. Which cereal-based foods are usually fortified?
Fortification of rice in Philippines has a long tradition, and Considering iron deficiencies and consumption intake, jus-
there is a mandatory regulation for the addition of vitamin B1. tify where would be recommended iron food fortification.
48 FOOD GRAINS AND THE CONSUMER | Fortification of Grain-Based Foods
Angola No Vitamin A, I, Fe
Australia Prohibited
Canada Voluntary B1, B3
Costa Rica Voluntary No I, Fe
Cuba Voluntary No Fe, vitamins B1, C, and A
El Salvador No Vitamin A, I, Fe
Ethiopia No Vitamin A, I, Fe
Finland Prohibited Restoration allowed
Gambia No I, Fe, Ca, folic acid, vitamin B2
Haiti No
Hungary Voluntary 1 serving must contain 1/3 of RDA
Mauritania No I, Fe, folic acid
Morocco Voluntary No Vitamin A, I, Fe
New Zealand Prohibited In process of reviewing the legislation
Norway Prohibited
Peru Voluntary No
Philippines Voluntary Legislation for B1, B3, Fe Vitamins A, B1, and B2, I, Fe
Sweden May be allowed with special permission
Tanzania No Fe, I, folic acid, vitamin A
Turkey Voluntary In accordance with Codex Fe, I, Ca, vitamins B2, B6, C, and D
United Kingdom Voluntary Must be safe
Uruguay No Fe, I, vitamin D
United States Voluntary B1, B2, B3, folic acid
Vietnam Voluntary No Vitamin A, I, Fe
Country Law status B-vitamins Vitamin E Folic acid Iron Zinc Magnesium Calcium
Topic Highlights a way to feed the rapidly rising population against a back-
ground of significant pressure on land, water, and other valu-
The purpose of genetically modified (GM) grains is to able resources. At the other end of the spectrum are those who
improve yield, increase resistance to plant pests and diseases, think that the use of this technology is playing God and have
help plants adapt to stressful environmental conditions, dubbed GM foods frankenfoods.
confer resistance to certain herbicides, reduce spoilage mech- GM grains refer to those that contain a gene introduced by
anisms, and improve the nutrient profile of a crop. recombinant DNA methods to confer a desired trait. The abil-
GM grains may help address nutritional deficiencies in the ity to pick a specific gene for a desired trait is an outgrowth of
developing world. standard breeding practices that have been used for centuries
The scientific community agrees that GM foods present no where plants or animals with desired traits were bred together
greater safety risk than conventionally grown crops. in hopes of perpetuating the trait. For many grains, this would
Despite prevailing scientific evidence, consumers remain be a 10-year process. The use of genetic modification, also
wary. called biotechnology, has been used since the 1970s. It can
Awareness of GM foods is increasing among consumers, as shorten the time required to achieve the same outcome as
is skepticism too. standard breeding process and can be much more precise
European, Mexican, and some African consumers are the than traditional hybridization. Traits have been introduced to
most outspoken against GM foods, while consumers in the grain foods to improve yield, to increase resistance to plant
United States are the most open to the idea. pests and diseases, to help adapt to stressful environmental
Consumer perception is dependent on understanding of conditions such as drought or extremes of temperature, to
safety, perceived ratio of risks to benefits, religious/moral confer resistance to certain herbicides, to reduce spoilage, and
values, intended use of GM foods, scientific literacy, trust in to improve the nutrient profile of a crop.
regulatory organizations, price, and social economic status. Since the introduction of commercially available biotech
Yield gains through GM technology are critical for the crops in 1996, the global use of GM crops have sustained a
developing world to meet growing population, but mone- growth rate of 6% or 10 million hectares per year. Thus,
tary benefits often occur in developed countries. between 1996 and 2012, there has been nearly a 100-fold
Maize, soy, and rice are commercially grown and sold increase in acreage. Farmers in North America grow the most
worldwide, but GM wheat has not been commercialized. GM crops with Brazil in second place. GM soybeans, maize,
Labeling is mandatory on all GM foods and foods contain- cotton, and rapeseed were planted on 170.3 million hectares
ing GM ingredients in 21 EU countries, Australia, Brazil, worldwide in 2012, according to figures from the International
Russia, China, and Japan. Individual states in the United Service for the Acquisition of Agri-biotech Applications
States are considering GM labeling. (ISAAA). In developing countries, biotech grains accounted
for 52% of all biotech crops planted in 2012 worldwide. This
is up from 50% 1 year earlier. In contrast, Europe grew < 0.1%
Learning Objectives of the worlds biotech grains.
Scientific bodies around the world continue to evaluate
To understand the purpose of GM grains and their applica- evolving data and continue to aver that there are no health
tion to the greater majority of the world. risks in consuming foods including grains that are produced
To recognize common factors that influence consumer per- by biotechnology. The US National Academy of Sciences
ception of GM grains. concluded no adverse health effects attributed to genetic
To compare and contrast consumer perception of GM engineering have been documented in the human
foods/grains across varying regions of the world and to population. Similarly, the International Council for Science,
observe differences in the perceptions of the scientific and which includes 111 national academies of science and 29
consumer communities. scientific unions, concluded that there is no evidence of any
To understand the movement for labeling of GM foods: ill effects from the consumption of foods containing GM
Where is labeling already mandated, who are its propo- ingredients. The World Health Organization also issued a
nents/opponents, and what is the fundamental argument? response stating, No effects on human health have been
shown as a result of the consumption of such foods by the
general population in the countries where they have been
Introduction approved. In 2010, a European Commission review of 50
studies on the safety of biotech crops found no scientific
Genetically modified (GM) foods and grains create varied reac- evidence associating GMOs (Genetically Modified Organisms)
tions around the world. Some see the use of this technology as with higher risks for the environment or for food and feed
safety than conventional plants and organisms. The American convinced that the technology could increase yields for devel-
Medical Associations Council on Science and Public Health oping countries.
stated that Bioengineered foods have been consumed for close Variability in consumer perception is influenced by demo-
to 20 years, and during that time, no overt consequences on graphics of the consumer. Older consumers are more likely to
human health have been reported and/or substantiated in the accept the use of biotechnology that affords health benefits,
peer-reviewed literature. not as likely to accept it for benefits of increased freshness or
Despite acceptance by farmers and diverse expert scientific flavor. Those with more education are more likely to try GM
panels around the world, consumer attitudes to GM foods and crops. Women are more resistant to the use of biotechnology
grains vary markedly by geographic region. Various degrees of than men.
activism, prevailing public opinion, and the extent of eco- Canadians are also divided regarding the concept of GM
nomic development strongly influence attitudes toward GM foods. Less than a third of Canadian consumers say they
foods. However, two overarching factors seem to influence oppose the use of GM in food. This is a change in attitude
public opinion of GM grains worldwide: (1) the riskbenefit with more respondents in 2011 stating that they strongly
perception related to the personal and family situations and support or somewhat support the use of GM in food produc-
needs of the country and (2) ambivalence or fear about the tion than in 2001.
safety of GM foods, often resulting from either a lack of infor- Regulation in various countries often reflects public opin-
mation or misinformation generated by some inaccurate ion. Health Canada regulates GM foods and requires that
media. Often, media reports fuel fears of some untoward con- developers show that the seeds and plants have no untoward
sequence that may not be accurately predicted. In terms of risk effect on the environment or human health. Receiving
and benefit, some feel that the consumer bears a dispropor- approval from Health Canada can take 10 years and be very
tionate share of the risk while large companies reap the mon- costly. However, once approved, Canada does not require
etary benefit. In terms of adopting biotech crops, the farmers labeling on all GM foods, unless there are any health or safety
are as divided as public opinion. Subsistence farmers worry concerns such as a food containing an allergen.
about planting biotech crops because they fear they may be The United States currently only has voluntary labeling
unable to feed their families or earn a living. In contrast, other guidelines for foods produced by biotechnology. However,
farmers are lured by the promise of fewer inputs of water, the Food and Drug Administration (FDA) outlines certain
fertilizer, and pesticides and thus lower costs and higher yields. circumstances where labeling is required, including foods
The view is strongly influenced by region. that have significantly altered nutritional makeup, allergens,
or toxic levels of an ingredient as a result of GM. Regulatory
policies on GM foods are enforced by the FDA, Environmental
Protection Agency (EPA), and USDA.
Perceptions of GM Grains by Region The federal law regarding GM voluntary labeling does not
reflect the results of several polls indicating that over 90% of
GM Foods in the United States and Canada
US consumers want GM foods to be labeled. This is reflected in
Farmers in North America have dramatically increased their bills on GM labeling being introduced at the state level.
planting of GM grains between 1996 and 2013. This has been Approximately half of US states have pending legislations
driven by farmers expectations of higher crop yields and/or requiring labeling of GM foods to some extent. Maine and
lower production costs, a reduction in time devoted to crop Connecticut have passed laws mandating labeling of all GM
care and management, and other benefits including farm sus- foods, and many states are likely to follow suit. Those in favor
tainability. Reduced input, lower use of pesticides and herbi- of labeling argue that they have a right to know what is in their
cides, less acreage and tilling required, and fewer crop diseases food, some for ethical or religious reasons and some for health
are among some advantages cited. As a result, currently 93% of or environmental reasons. Much of the argument for labeling
the soybeans and 80% of the corn, according to the United is simply transparency. Americans claim a right to have imme-
States Department of Agriculture (USDA), grown in the United diate access to information on their foods content, so they can
States are GM. (Despite allegations to the contrary, no GM make a personal decision.
wheat is grown commercially in any part of the world.) Organic foods cannot contain GM ingredients, so some
Consumer acceptance of GM food in the United States has consumers used the organic label as a proxy for a label saying
increased slowly but steadily and somewhat parallels the GMO-free. Many manufacturers and sellers of organic foods
increase in number of hectares of biotech crops. It also reflects oppose the use of GM products. This meets the wishes of many
a less disparaging reaction to the technology by the press and organic consumers, but lack of GM labeling supports their own
the public. This is reflected in the International Food Informa- economic interests.
tion Councils 2012 survey on Consumer Perceptions of Food The calls for legislation on labeling of GM foods have
Technology, which shows that 74% of consumers have, at dismayed those in agribusiness as well as some scientists and
minimum, heard a little about biotechnology and as many consumers. Considering that 6070% of processed foods in
as 38% of US consumers are somewhat or very favorable the United States contain at least one GM ingredient, experts
toward plant biotechnology. 44% asserted they were worry that the reaction of consumers to such labels will cause
somewhat or very favorable regarding the use of biotechnol- them to reject use of GM products and seeds, turning back
ogy in grains specifically. Overall, consumers were likely to be gains made in yields along with profits. Food economists
more accepting biotechnology if the benefits of the technology suggest that not only it is these large corporations that will be
were clearly communicated and especially if consumers were burdened with the negative consequences of labeling but also
52 FOOD GRAINS AND THE CONSUMER | Genetically Modified Grains and the Consumer
consumers would likely see increased prices and changed avail- 2013 surveys carried out by the UK government showed that
ability. Groups who do not favor labeling often reference only 21% of the UK public support GM foods and less than
scientific literature, arguing that there is no significant differ- half of UK farmers believe GM technology is a good innova-
ence in the safety of GM foods and non-GM foods. They also tion. Over 40% of people stated that they were completely
note that manufacturers are required to alert consumers of any against the government promoting GM technology.
material difference in GM foods that could affect safety such as European acceptance of GM food, while limited, may be
the presence of allergens or potential toxins or decreased nutri- increasing. A poll of European consumers conducted between
tional quality. Those who oppose GM labeling state that con- 2002 and 2005 showed an increase of 6% in positive attitude
sumers wishing to avoid GM ingredients have the option to toward GMOs. 80% of EU consumers did not intentionally
purchase foods labeled organic, which precludes their use. discriminate against GM foods when shopping. Thus, Euro-
pean consumers stance on GM food may not be affecting
purchasing behavior. Recent research indicates that acceptance
GM Foods in Europe
of GM food is much improved if the consumer benefits such as
Strong resistance to GM foods pervades Europe. This occurs produced with fewer sprays and higher levels of a needed
despite numerous decisions regarding the safety of GM foods component such as omega-3 fats are clearly communicated
by bodies such as the European Commission (EC). However, to the consumer.
regulators dealing with GM foods in Europe invoke the pre-
cautionary principle, which states that if there are any unre-
Asia-Pacific Nations-Developed Countries
solved concerns, they will not accept the food process or
additive in the food supply. This principle aims at minimizing In Australia, consumer acceptance of GM foods has fallen since
any perceived or theoretical threats. It fails to consider any 2007, following a steep rise in consumer acceptance between
potential benefits where the science is indeterminate. It also 2005 and 2007. However, the acceptance level among the
is problematic as safety cannot be proven, only lack of safety. general public is still at 67%. Approval ratings of biotechnol-
Consumer resistance to GM foods and belief in the precau- ogy in food and drink have consistently remained lower than
tionary principle have led at least 21 European Union (EU) approval ratings of biotechnology in medicines, vaccines, and
countries to require labeling of GM foods or foods with GM stem cells, as well as for the purpose of creating pest-resistant
components. plants. Awareness of biotechnology in food is at a commend-
The overall sentiment in Europe is that the benefits must able 82%, and a solid two-thirds of the Australian public thinks
outweigh the risks. Furthermore, consumers and experts hold the use of biotechnology in food crops could be useful. Less
wholly different viewpoints toward GM foods, specifically than half of the population finds the application risky, but
regarding safety, ethical tradeoffs, benefits, and international GM-containing foods must be labeled.
trade. What might be seen as a benefit to experts is often viewed A study of 2000 New Zealand consumers sought to deter-
as a risk to consumers. When GM foods were first introduced mine whether consumers would purchase GM foods. Slightly
into the European market, consumers did not perceive that more than 30% of the over 700 who returned the survey said
there was a benefit of these products in that they were not that they would not purchase any of the GM foods listed on the
cheaper, did not seem tastier, and did not offer other added survey, which included bread from GM wheat and maize
value. The higher yields per hectare did not reduce prices for altered to resist insects. However, not all information from
consumers due to a general increase in prices for fuel and other New Zealand shows attitudes toward it to be negative, espe-
inputs that affect food prices. cially when the science is applied sensibly and consumers see it
Food scares in Europe in the 1990s eroded confidence in involves no great risks. Further biotechnology would be accept-
food regulatory bodies and the food supply. This influenced able in New Zealand if (1) the consumer is given a choice, (2)
discussions regarding the acceptability of GM foods. Con- the benefits hold greater weight than the risks, (3) the benefits
sumers continue to question aspects of consumer health, envi- are shared by the consumer as well as agribusiness, (4) the
ronmental risks that may occur over time, and economic effects practice is environmentally friendly, and (5) the technology is
on small European farmers. There are also fears of allergenicity safe, practical, and cost-effective. Concern and reasonable
and the development of microbial resistance to GM traits just doubt certainly exist in New Zealand regarding GM foods,
as has occurred with pesticides and herbicides. but not to an extent that completely stifles acceptance.
Farmers are wary of GM food as most of the GM food is Japan and Brazil, Russia, and China mandate labeling of all
imported, so there is an economic concern as well. Countries GMO-containing foods. While Japanese resistance to GM is not
who produce GM products have contested the use of the as strong as that in Europe, Japanese consumers continue to
precautionary principle as a trade barrier. Yet organizations remain unwilling to accept GM products. This was recently
such as the World Trade Organization are reluctant to impinge highlighted after finding of GMO wheat in Oregon. This
on the authority of various member states. revived the issue and Japans Ministry of Agriculture, Forestry
Despite over 25 years of use and no environmental or other and Fisheries opted for temporary import suspension and
problems, European consumers claim they have insufficient called for urgent greater GM inspection on imports to Japan.
and inconclusive information. They state that their confidence
could be improved if more definitive scientific information be
Asia-Pacific Nations-Developing Countries
made more readily available to the public.
Acceptance of GM foods varies among the countries with In Malaysia, almost 90% of consumers surveyed were aware of
some of the strongest resistance found in the United Kingdom. GM foods. However, the fraction of Malaysian consumers who
FOOD GRAINS AND THE CONSUMER | Genetically Modified Grains and the Consumer 53
accept GM foods may not be as high. The study revealed large it was completely acceptable, 42% deemed it relatively
concerns over food safety and perceived riskbenefit ratio. acceptable, 21% pleaded neutral, 5% were relatively opposed,
Other factors affecting consumer opinion were media influ- and only 1% were strongly opposed. The acceptance rate of
ence and lack of information. folate-enhanced rice was found to be > 60% in a study of
Filipinos, in general, trust the international agencies nearly 1000 consumers. Again, results were most strongly
responsible for conducting studies and disseminating informa- affected by perceived riskbenefit ratio.
tion on topics such as GMOs. This may be true due to the work
of the International Rice Research Institute (IRRI) and the work
Africa
of the International Maize and Wheat Improvement Center
(CIMMYT). Citizens are less confident in the government Attitudes regarding GMOs in many countries in Africa are
than university scientists. Overall, Filipinos believe the nutri- influenced by those in the rest of the world. Some developed
tive qualities of GM grains could prove beneficial, but they nations and world food organizations encourage the use of GM
remain guarded about safety. A study conducted in the Philip- crops for the purpose of alleviating hunger and tackling food
pines analyzed the attitudes of 423 Filipinos, who worked in a insecurity. Not all Africans are convinced that GMOs would be
variety of careers. The majority were middle-aged, educated a helpful strategy in addressing food insecurity. In fact, many
males with an allegiance to Roman Catholicism. Attitudes are distrustful despite a backdrop of decreased agricultural
were affected by education level and religious affiliation; productivity throughout most of the continent. The Center
those with higher education held more of a positive view on for Strategic and International Studies (CSIS) noted that
GM foods, while the strongly religious had moral concerns despite 70% of Africans working in agriculture, Africa imports
about GM technology. Many Filipinos across various disci- a quarter of its food.
plines claimed insufficient information on GMOs. Those who Across Africa as a whole, the mood toward GM foods can be
received information from the scientific community were more summarized as ambivalent, with some nations more resistant
likely to have positive opinions of the technology than those and others more welcoming. Tanzania, Uganda, Malawi, Mali,
who received information from popular media. Even without Zimbabwe, Nigeria, and Ghana have at least begun to consider
easy access to literature on biotechnology, Filipinos are less the possibility of growing GM plants, and many are conducting
likely to have extreme attitudes toward GM food. However, an research and field studies.
anti-GMO group of 400 interfered in 2013 with a GM trial on Awareness and acceptance vary dramatically across Africa.
Golden Rice, a variety of rice intended to decrease the preva- For example, the awareness level for GMOs in Ghana has been
lence of vitamin A deficiency. calculated at around 60%, meaning more than half of the
population of Ghana has heard to some extent of GM foods.
However, increased awareness does not assuage fear; rather, it
India
may lead to increased uncertainty. Part of the uncertainty stems
In India, most consumers are aware of GM crops, some to a from the perception that the safety of GMOs is still in question.
greater extent than others. While a whopping 96% of Indian Another part stems web posts and media stories that may
consumers deem biotechnology to be beneficial and worth inaccurately portray or overstate the risks and fail to tout the
further exploration, < 33% currently support the implementa- benefits and the numerous consensus statements averring their
tion of GM crops and hardly any want GMOs in their own food safety.
supply. Although awareness is high and an overwhelming Similar trends have been observed in other developing
majority views them as a promising prospect for the future, nations, even ones in the throes of a hunger crisis. Tens of
consumers do not necessarily trust the use of GMOs in their thousands of tonnes of corn donated by the United States was
food currently. They are more open to their use in medicine. rejected by Zambias president, who called it genetically mod-
The Supreme Court in India has just called for an indefinite ified poison. One stated rationale noted that since Europe
moratorium on field trials for new GM food crops. Just next objects to its use, it must not be safe. While a large number of
door, Bangladesh is allowing the planting of GM staples. Zambians are not welcoming to the idea of GMO, an increas-
ing fraction is warming (ever so slowly) to the possibility of
someday growing GMOs. However, someday sits far off on
China
the horizon as existing legislation all but bans the use of GM
Chinese consumers remain divided over the use of GM foods seeds and proponents have yet to unify and work toward
but are more accepting than consumers in Japan. A study acceptance.
published in 2010 by Zhang et al. recognized Chinese con- A larger portion of Kenyans compared to those from other
sumers as belonging to one of four categories: neutral (risks nations in Africa are more accepting regarding the use of GMOs
posed by GMOs are equal to benefits), relaxed (benefits of partially due to wider acceptance by prominent political and
GMOs outweigh the risks), skeptical (risks outweigh the ben- scientific leaders in Kenya. Greater acceptance and use of GM
efits), and uninterested (no perceived risks or benefits). Social crops are aided by two factors. First, the British left behind a
status and income level strongly influenced the particular cat- relatively advanced research center giving Kenyans in direct
egory. Although variability in degree of acceptance exists across contact with the latest scientific data on GMOs. Second, com-
China, results of the Zhang study suggest that more people mercial interests are exceptionally strong in Kenya, and they
accept biotechnology than object to it. When asked to rate promote the development and use of GM crops.
nutrition-improved GM rice on a scale from completely If Kenya is one of the more progressive nations regarding
acceptable to strongly opposed, 25% of 1000 surveyed said GMO use, South Africa is the most progressive, and GM crops
54 FOOD GRAINS AND THE CONSUMER | Genetically Modified Grains and the Consumer
have been grown in South Africa since 1997. As with Kenya, GM-produced golden rice adds vitamin A to help mitigate this
South Africas educational system is built on deep scientific tragic public health problem.
roots. Local scientists affirming GMOs safety and benefits No matter what the region, it is important that GM foods
help promote consumer acceptance of biotechnology. South and technologies address needs of local and subsistence
Africa, Egypt, Sudan, and Burkina Faso are the four nations in farmers, as well as respect cultural, dietary, and religious
Africa that sell GM crops commercially. importance of certain crops. In some cases, GM versions of
familiar crops may have greater acceptance than an unfamiliar
crop. While this is true in all regions, it may be especially true
Latin America for those in developing nations.
Judicious use of GM crops in developing nations means
Widespread production of GM crops in Brazil and Argentina
that efforts must be directed to ensure that the use of this
means that approximately one-third of the worlds acres
technology does not exacerbate the economic divide. Care
planted in GM crops occur in South America. GM maize
must be exercised to ensure that the benefits of GM crops
leads the production, but GM versions of cotton, soybean,
help address the problem of food security and improve the
and canola are also grown in South America.
environment in the region and that economic or other risks are
The widespread production of GM crops does not mean
minimized.
South American consumers readily accept GM technology.
Interestingly, the countries with high production Brazil and
Argentina also have high resistance. Arguments against GMs
use include a lack of human resources and appropriate equip-
Conclusion
ment in the region, concerns about viability of small farmers,
disputes about seed ownership, and fears that regulation sur-
The practice of biotechnology began in the 1970s, and grains
rounding GMOs is inadequate.
such as maize were some of the first commercially available
Peru and Colombia, in particular, are actively protesting
GM foods. Biotechnology has been employed with grains to
GMOs and resisting the efforts of multinational seed compa-
heighten the ability to withstand stress and resist pests, disease,
nies. Countries such as Mexico have actually banned the grow-
and herbicides; enhance the nutrient profile, freshness, or
ing of GM crops. In 1998, it became illegal to grow GM maize
flavor of the plant; and reduce spoilage and improve yield.
after the seeds were introduced into Mexico either through
Most crops available up until now have increased benefits to
accidental cross contamination or through illegal intent. How-
farmers and this indirectly can help contain food costs for
ever, the law has been loosely enforced. Reiteration of the ban
consumers. Soon, foods with increased flavor and nutrient
in 2013 by a judge may change this. Vocal leaders emphasize
profiles will be on the market.
corns essentiality and identity as Mexicos national crop and
Perception and acceptance of GM foods vary. The scientific
describe GM corn as a threat to their sovereignty and identity.
community concludes that GM foods, including GM grains, are
Mexico faces opposition from such bodies as the United
safe and pose no more risks than do conventionally grown
Nations Food and Agriculture Organization, which argue in
foods. Consumers remain wary despite the large body of sci-
favor of GMOs to eliminate hunger. Thus, across Latin Amer-
entific evidence and numerous scientific and government bod-
ica, mass scale production of GMO crops occurs while there are
ies deeming them safe and a possible way to address future
bans and widespread resistance, sometimes within the bound-
food needs. Attitudes toward GM foods strongly influence
aries of the same country.
perception of benefit and safety. Such attitudes are influenced
by media, legislation, science literacy, trust in regulatory orga-
nizations, price, religion and morality, and social economic
GM Foods in Developing Countries: A Beacon of Hope?
status.
In most developing countries, GM grains promise many agro- It is exceptionally difficult to capture an entire populations
nomic benefits. With the worldwide population expected to sentiment in one study, let alone a few sentences. Scientific
double by 2050, food security is of great concern. Biotechnol- literature shows a lack of consensus, even within a region. It is
ogy can be one of the most sustainable ways for ensuring even more difficult to mention or represent every countrys
adequate agricultural yields and value-added and nutritionally perspective. Despite limitations, this article divulges consumer
enhanced products. Statements such as those in the journal perception as it was most commonly portrayed in the scientific
GMO Biosafety Research dub the technology a beacon of hope. literature. Overall, awareness of GM foods has increased sig-
Biotechnology protects food security via three primary nificantly worldwide. In the developing world, GM grains offer
methods: increasing yields on existing land and optimizing the prospect of closing nutritional gaps, but still meet signifi-
other limited resources, advancing food quality and safety, cant resistance.
and potentially bettering the economic status of local farmers. Many in the developing nations remain wary of biotech-
GM grains can offer numerous nutritional advantages, such nology. Wariness is increased because most biotechnology
as fortification to address nutrient gaps or extreme deficiencies initiatives emanate from multinational companies. This fact
in developing countries. For example, WHO estimates that 250 means that not only there may not be adequate understanding
million preschool children are vitamin A-deficient and an of the consumers in a region but also there is worry about
estimated 250 000 to 500 000 vitamin A-deficient children motives of the multination and about who will reap economic
become blind every year, half of them dying within 12 months. gains. Acceptance of biotechnology may increase when goals of
FOOD GRAINS AND THE CONSUMER | Genetically Modified Grains and the Consumer 55
developed countries align with those of developing countries, Exercises for Readers to Explore the Topic Further
and the benefits are made reciprocal and equal. In some
regions, the technology is slowly gaining and more hectares To what extent do countries have access to scientific litera-
are being planted. ture concerning biotechnology and how does it affect their
The developed world is divided over the use of GM foods. overall consumer perception?
The United States, parts of South America, and South Africa How well do GM grains address nutritional deficits in the
currently have large areas devoted to GM-crop production. developing world?
Australia and New Zealand place great weight on the benefits How does labeling affect consumer perception of GM
and who reaps the benefits. In most regions, GM approval foods?
continues to be higher in medicine than food, but GM food
approval still is relatively high in these regions. The most
vehement opposition comes from parts of Europe. See also: Food Grains and the Consumer: Labelling of Grain-
Where GM crops are allowed, many jurisdictions require Based Foods.
labeling and regulation of crops produced by biotechnology.
Mandated labeling on all GM foods and foods containing
GM ingredients is currently enforced in several countries in Further Reading
Europe, Japan, Australia, Brazil, Russia, and China. In coun-
tries where no labeling of GM foods is required, labeling has American Medical Association, AMA (2013) Report of the Council on Science and
become a heated issue. Many consumers claim they have the Public Health: Labeling of Bioengineered Foods. http://www.ama-assn.org/ama
(accessed 01.14.).
right to know what is in their food, and the government has
American Council for Science and Health (2014) The co-existence of organic and GMO
an obligation to make such information readily accessible. foods. http://acsh.org/2014/04/co-existence-organic-gmo-foods/ (accessed 4/2014).
Those opposed to labeling of GM foods cite consensus state- Center for Food Safety (2013) US Polls on GE Food Labeling. http://www.
ments of government agencies and health organizations stating centerforfoodsafety.org/issues/976/ge-food-labeling/us-polls-on-ge-food-labeling
that food grown from GM seeds is not materially different (accessed 01.14.).
DeSteur H, Gellynck X, Storozhenko S, et al. (2010) Willingness-to-accept and
from food produced conventionally; therefore, there is nothing purchase genetically modified rice with high folate content in Shanxi Province,
to disclose. The US consumer population is largely in favor of China. Appetite 54(1): 118125.
labeling, but federal labeling guidelines are voluntary. The EuropaBio (2013) Abc stresses overwhelming weight of evidence on safety of GM
global status of labeling on GM foods may change in the crops. http://www.europabio.org/news/abc-stresses-overwhelming-weight-
evidence-safety-gm-crops (accessed 01.14.).
coming years, as consumers continue to demand transparency
European Commission (2006) Eurobarometer 66: Public Opinion in the European
from their governments. It may be interesting to observe how Union. http://ec.europa.eu/public_opinion/archives/eb/eb66/eb66_highlights_en.
consumer perception and purchasing behaviors of GM foods pdf (accessed 01.14.).
are altered in response to increased transparency. Hart J and Herrmann R (2009) Do they always say no? German consumers and second-
Encouragement of GM crops as a way to address the needs generation GM foods. Agricultural Economics 40(5): 551560.
Herath D (2001) Evolving Consumer Acceptance of Biotechnology Applications in
of the world population in 2050 by humanitarians such as Canada: Evidence from the Public Opinion Surveys in 2001 (fifth wave) and 2011
those working for the Gates Foundation could have a dramatic (seventh wave). http://ageconsearch.umn.edu/bitstream/150525/2/Herath8_1AAEA.
effect on attitudes and acceptance of GM foods in many parts pdf.
of the world. Further, open and transparent communication International Council for Science, ICSU (2003) New Genetics, Food, and Agriculture
Scientific Discoveries Societal Dilemmas. http://www.icsu.org/publications/
on the benefits of GM foods and the availability of foods with
reports-and-reviews/new-genetics-food-and-agriculture-scientific-discoveries-
specific consumer benefits may help to address the concerns of societal-dilemas-2003/ (accessed 01.14.).
many consumers and improve acceptance. International Food Information Council (2012) Consumer Perceptions of Food
Technology Survey. http://www.foodinsight.org/Content/5438/FINAL%20Executive
%20Summary%205-8-12.pdf.
International Service for the Administration of Agri-Biotech Applications, ISAAA (2013)
ISAAA Brief 442012: Press Release. http://www.isaaa.org/resources/publications/
Exercises for Revision briefs/44/pressrelease/ (accessed 01.14.).
Kaye-Blake W, Saunders C, and Fairweather J (2007) GM food and New Zealand
In one continent, take a sampling of countries from various Consumers. http://www.biotechlearn.org.nz/themes/
sectors and show in detail how these countries vary in their new_zealand_views_on_biotech/gm_food_and_new_zealand_consumers
(accessed 01.14.).
acceptance and regulation of GMO. Myhr AI (2007) The precautionary principle in GMO regulations. In: Traavik T and
Take a country or region and define the major concerns Lim LC (eds.) Biosafety First Norway: Tapir Academic Publishers (accessed 01.14.).
regarding acceptance of GMO. http://genok.no/wp-content/uploads/2013/04/Chapter-29.pdf.
Take a country or region and suggest communication The National Academy of Sciences, NAS (2004) Safety of Genetically Engineered Foods:
Approaches to Assessing Unintended Health Effects. http://dels.nas.edu/Report/
strategies that would be convincing for the acceptance of
Safety-Genetically-Engineered-Foods-Approaches/10977?bname (accessed
GMO. 01.14.).
What additional scientific information is needed for a cer- Nielsen T (2012) How do concerns about pesticides impact consumer willingness to
tain population to accept GMO? buy genetically modified french fries in Germany? Results from a purchasing
experiment. 2012 International European Forum, 1317 February, 2012. Austria:
What world events or program backing has the potential to
Innsbruck-Igls.
impact GMO acceptance? Paulsen T GM food fight: Why the Gates Foundation wants to make rice golden. http://
What problems has golden rice faced on its journey and www.humanosphere.org/basics/2013/09/gm-food-fight-why-the-gates-foundation-
how could such things be avoided in the future? wants-to-make-rice-golden/.
56 FOOD GRAINS AND THE CONSUMER | Genetically Modified Grains and the Consumer
Torres CS, Suva MM, Carpio LB, and Dagli WB (2006) Public Understanding and Relevant Websites
Perception of the Attitude Towards Agricultural Biotechnology in the Philippines.
ISAAA. SEARCA and CDC-UPLB. http://www.isaaa.org/programs/supportprojects/ http://www.agbioworld.org/ AgBio World, Peer Reviewed Publications on the Safety of
(accessed 01.14.). GM Foods.
USDA, Economic Research Service (2013) Adoption of Genetically Engineered Crops in http://dels.nas.edu/ National Academy of Sciences, Division on Earth and Life
the U.S. www.ers.usda.gov/data-products/adoption-of-genetically-engineered- Studies. National Research Council (U.S.).
crops-in-the-us/recent-trends-in-ge-adoption.aspx#.U0FCRMf95lk (accessed http://edis.ifas.ufl.edu/fe934 Evans, EA Ballen, FH. A synopsis of US consumer
01.14). perception of genetically modified (biotech) crops. University of Florida Extension.
Vinter R (2013) Livestock 2013: Survey shows consumer resistance to GMs. Farmers http://www.ers.usda.gov/data-products/adoption-of-genetically-engineered-crops-
Weekly. http://www.fwi.co.uk/articles/03/07/2013/139866/livestock-2013-survey- inthe-us/ Fernandez-Cornejo J, 2011 U.S. Economic Research Service, Adoption
shows-consumer-resistance-to.htm (accessed 01.14.). of Genetically Engineered Crops in the U.S.
World Health Organization, WHO (2014) 20 questions on genetically modified foods. http://www.gmo-compass.org/eng/agri_biotechnology/gmo_planting/ GMO
http://www.who.int/foodsafety/publications/biotech/20questions/en/ (accessed Compass.
01.14.). http://www.icsu.org/ International Council for Science.
Zhang X, Huang J, Qiu H, and Huang Z (2010) A consumer segmentation study http://www.isaaa.org/ International Service for the Acquisition of Agri-biotec.
with regards to genetically modified food in urban China. Food Policy 35(5):
456462.
Labelling of Grain-Based Foods
JM Jones, St. Catherine University, St. Paul, MN, USA
2016 Elsevier Ltd. All rights reserved.
Topic Highlights Coded or overt dates and lot numbers are required both to
track shelf life and to allow for product recalls. The manufac-
Basic label requirements are similar across jurisdictions turer contact information enables consumers to make queries
with the ingredient statement and contact information and to state complaints and facilitate recalls, if necessary. The
being critically important. net weight allows price and value comparisons of competing
Label terminology used to describe grain processing, products and ensures the consumer receives a fair measure.
milling, and grain fractions is understood by industry, but Some jurisdictions may require the country of origin labeling,
not by consumers. Further, it is not standardized across warning statements about allergens or ingredients of concern,
jurisdictions and is confusing to consumers. and advisory statements on safe handling of the product.
Nutrition labeling of grain-based foods is mandatory in Nutrition labeling, usually with a prescribed format, is
many countries. Whether required or voluntary, the format mandatory in many jurisdictions, while others allow or
is specified but formats vary across jurisdictions. encourage it voluntarily. Statements and visuals on the package
Regulation for labeling of organic and genetically modified and advertising must not be misleading. Foods for special
grains and other label descriptors such as fresh and natu- dietary or medical use such as cereals or bars that have func-
ral vary across jurisdictions. tional ingredients that reduce blood sugar may have more
Procedures and rules regarding the approval and use of extensive nutrition labeling requirements. Nutrient-content,
nutrient-content claims, structurefunction claims, and structurefunction, and health claims are allowed in some
health claims in grain-based foods depend on the jurisdictions, but specific claims and rules regarding them
jurisdiction. vary across jurisdictions.
Issues regarding the labeling of whole-grain foods are com- GBFs also carry label descriptors about the size, shape, and
plicated due to varying recommendations for whole-grain processing of the grain. In many cases, the terminology is
intake across regions, issues about the amount of dietary familiar only to those in the cereal industry. Thus, consumers
fiber or detractor nutrients that are in the whole-grain food, are confused about definitions surrounding grain fractions,
and the use of wet or dry weight for calculation of the whole whole grain (WG), and especially the definition of a whole-
grain. grain food (WGF). This article will discuss these and other
issues specific to GBF.
Learning Objectives
Label Descriptors for GBF
To be able to outline and define various requirements and
terms associated with labeling grain-based foods. Label descriptors for GBF can be those that are more general
To compare various label descriptors, nutrition labels, and such as fresh and those that communicate aspects regarding
types of nutritional claims used on grain-based foods in the product, form, and level and type of processing specific to
various jurisdictions. grain such as the method and degree of milling, the particle
To delineate issues associated with labeling of whole grains size, and other processing aspects. While these industry terms
and whole-grain foods within and across jurisdictions. associated with GBF are understood by industry, they can leave
consumers confused. Terms such as medium pearled or de-
germed are not always as clear or as easy to operationalize as
Overview terms for other common foods, such as canned sliced peaches
in syrup, where most of these terms need little further clarifi-
Grain-based food (GBF) labels allow consumers to select and cation. Nonetheless, this information is needed for selecting
purchase products with desired characteristics, ingredients, and the grain form for the best culinary use and for choosing
nutritional profiles. Labels act as both marketing tools and desired nutritional characteristics.
transmitter of legally required information. They must name Terms such as fresh, natural, homemade, artisan, tra-
and accurately describe the food and list the ingredients in ditional, ancient, and stone-ground are popular with mar-
descending order by weight. The ingredient statement is critical keters and consumers alike but have come under scrutiny by
for most consumers as it delineates what is in the product and various regulatory bodies. For the consumer, such terms give
gives the consumer an idea of the relative amounts of the products a goodness halo that may or may not be justified, so
various components in a product, since they must be listed in some jurisdictions have prescribed ways such terms can be
order by weight from the most to the least prevalent. In some used. In areas where there is no legal definition, the expecta-
countries, percent labeling is required, so that a precise amount tion is that such terms are not misleading. Terms such as
of a specific ingredient is declared. Interested consumers can natural or fresh may or may not be defined depending on
carefully scrutinize the ingredient statement for components or the jurisdiction. In some jurisdictions, the term fresh for bread
additives that they wish to include or avoid in their diets. is defined to mean prepared at the venue and sold there
shortly after baking. Preprepared dough off-site and baked on most countries have regulated formats. The nutrition label in
the premises cannot be labeled as fresh in some regions. It is most countries requires a declaration of energy in calories or
easy to understand why terms are confusing because the term joules, total fat, saturated fat, trans fat, cholesterol, sodium,
fresh for pasta denotes that the pasta is not dried. total carbohydrates, dietary fiber (DF), sugars, and protein.
Designations such as light, low-calorie, very low sodium, Rules about saturated fat, trans fat, and calories from fat vary
high in, or free from may be strictly defined or not allowed. across jurisdictions. Similarly, the list of micronutrients
For example, in the United States, rice cakes with under 40 cal required and allowed varies as does the percent daily value
might be labeled as a naturally low-calorie food and cannot (DV) standard. In most jurisdictions, labeling becomes com-
be labeled as low-calorie rice cakes implying that they are pulsory if a nutrition claim regarding a specific nutrient is on
lower in calories than other rice cakes. In certain jurisdictions, the label or in advertising. Nutrition labeling is harmonized
a GBF formulated to reduce an attribute such as calories, throughout the European Union and some regions; however,
sodium, or fat must be reduced by a specific percent and this is often not the case.
must be labeled and compared to either the original food or DF values in GBF may vary depending on which DF defini-
the average of foods in the category. tion and analytic method have been officially accepted. The
Claims that a food is high in x may need to follow rules for acceptance of Codex Alimentarius and aligned DF definitions
nutrient-content claims and may not be allowed for compo- and methods by Europe, Canada, and many other jurisdictions
nents that are not nutrients such as WG or antioxidants. In has helped harmonize fiber labeling across many jurisdictions.
some jurisdictions, free from labeling is specified in that foods Energy values may also vary depending on what calorie value is
that have never had component x cannot be labeled as free assigned to certain fibers in various regions.
from x. Statements about gluten-free must follow rules of the In most countries, nutrition labels are based on 100 g of
jurisdiction. The CODEX regulations, followed in a number of food, but some countries use dual labeling. Nutrient values are
areas, require < 20 ppm gluten, but a few regions also allow listed per 100 g of food and also for a serving size appropriate
low gluten with higher levels of gluten. Allergen labeling may to the food and the country. In the United States, the nutrient
be required, and so GBF would have the required disclosures of values are based only on government-mandated serving sizes
wheat, nuts, milk, eggs, and other common allergens. called reference amounts commonly consumed (RACCs).
Organic labeling for GBF aids consumers wishing to purchase The system using 100 g portions has the advantage of allow-
products that are produced without additives, pesticides (other ing easy comparison of products and enables consumers to
than those allowed by organic rules), and chemical fertilizers easily calculate the percent of various macronutrients. How-
and did not undergo genetic modification (GM) or irradiation. ever, some consumers cannot envision how 100 g of a specific
Some regions require official certification at all stages of the product a serving can be eaten. The following examples illus-
process in order to label products as organic. Table 1 shows a trate this confusion. A slice of bread is usually 30 g so labeling
gradation in organic that is allowed in the United States with per 100 g would be the nutrient value for three slices of bread.
products where 95% or more of the product is organic can bear Ready-to-eat cereals vary dramatically so a 30 g serving of a
an official seal. If a product has < 70% organic ingredients, it dense granola is 1/3c, whereas it is two cups of a puffed grain
cannot use organic on the main package label, but individual cereal. Unfortunately, many consumers are not able to use the
ingredients listed as organic on the ingredient statement. nutrition label effectively as they are unable to apply the listed
If GM grains are allowed, many countries require their values to the amount they have eaten.
labeling. In the United States, labeling of GM grains is not The nutrition label must list the nutrients in the package as
required because the USFDA views that there is neither a mate- sold, regardless of how the food is commonly ingested. Thus,
rial difference nor a safety or environmental issue associated some products have two values the one required for the food
with these products. However, deeply held beliefs by some as packaged and the one that gives nutrition information of the
consumers have prompted a number of state legislators to product as prepared or ingested. Thus, cereal labels often give
sponsor bills requiring GM labeling. To date, all such initia- values for the cereal as purchased and as consumed with 120 g
tives have failed. (c) of low-fat milk. A similar type of dual labeling may be on
packaged foods such as cake or pasta mixes, which have an
additional column giving nutrient values for the food prepared
The Nutrition Label according to package directions. Dual-language (or multilan-
guage) labeling of GBF enables their sale in countries such as
The nutrition labeling of packaged foods, including GBF, is
Canada with more than one national language or in a region
mandatory in many countries of the world. Where voluntary,
such as Southeast Asia.
100% organic Must contain only organically produced Labelling of the Various Grain Components
ingredients
Organic At least 95% of the products ingredients Grains have been processed since before biblical times to
must be organic remove the outer inedible husk, and in many cases, outer layers
Made with organic Must contain at least 70% organic may be removed. Grains may then be broken into particles of
ingredients ingredients various sizes, sifted into various streams, and kept separate or
Some organic Products contain <70% organic ingredients
recombined, and the various fractions may be processed fur-
ingredients
ther. The specific terminology used to describe grain
FOOD GRAINS AND THE CONSUMER | Labelling of Grain-Based Foods 59
components from various stages of milling and processing is Wholegrain is defined as intact and/or processed (e.g., de-hulled,
on the label, but consumers do not know if a meal is different cleaned, ground, cracked, flaked or the like) grains, where the frac-
from a grit and may think that degermed is a sterilization, and tions endosperm, bran and germ are present in the same proportion
not a milling, process. as found in the least processed traditional forms of the edible grain
kernel of the same species.
In many parts the world, GBFs are most commonly made
from refined or enriched grains. These foods have mostly the
endosperm of the grain, or in the case of enriched, the endo- All three definitions do not require that the grain remain as
sperm has some nutrients lost in milling added back. Current an intact kernel, but rather that all of its components be pre-
nutrition research suggests that WGs are advantageous in terms sent. However, a few countries using the terminology require
of more DF and reduced risk of chronic disease. Thus, some that whole defines the intact kernel. Each definition varies
consumers are trying to replace refined and enriched grain slightly in terms of both consumer friendliness and allowable
foods with WGFs, which makes the understanding of GBF loss of outer layers. The WGC definition is more consumer-
label terms so important. friendly than the AACCI definition that uses the botanical term
Enriched and Refined Grains: If the various parts of a kernel caryopsis. The HG definition allows some removal of a small
have been removed, it is a refined grain. If some vitamins and amount of grain such as that occurs in traditional processes
minerals lost in milling are added back, the grain is enriched. such as making bulgur or nixtimalizing corn or those that
Various food authorities decide if enrichment is allowed and, if make the grain healthier. Examples include the scoring of the
so, the specific nutrients added and the amount allowed. If outer kernel to increase availability of nutrients or nixtimaliz-
enriched flour is used in packaged food products, the nutrients ing to reduce mycotoxins in the outer layers. Some countries
added not only must be reflected on the nutrition facts panel equate whole-wheat or wholemeal flour; others allow the
but also may be reflected on the ingredient statement. The removal of some of the outer layers and still call it whole-
ingredient statement might read Ingredients: Sugar, Enriched wheat flour. This does create confusion.
Bleached Flour (wheat flour, niacin, iron, thiamine mononi- Definitions of WG help processors formulate food and help
trate, riboflavin, folic acid). . .. consumers describe WGFs that are all WG such as porridge oats
Grains may also be fortified to address nutritional inade- (oatmeal) or WG flours. However, the consumer has great diffi-
quacies in the region. Thus, the nutrient levels of fortified culty determining which foods are WGF or determining the num-
products may be higher than enriched products. Often, 25% ber of servings of WG to ingest in products that are partly WG.
of the DV is added, but some products have more. These Each current strategy for finding WGFs in the marketplace
additions are reflected on both the ingredient statement and has a disadvantage. One strategy suggests that a food is WG, if
the nutrition facts panel. If fortification levels in some GBF the first ingredient in the ingredient statement is WG. While
meet the daily requirement, for example, 100% of the DV, the this may correctly identify a WGF, it may not if the food has
product must be labeled as a multivitamin and mineral sup- many ingredients. This is explained by the following examples.
plement GBF. The intake of these fortified cereal products The ingredient statement from a bread mix is as follows:
increases intakes of nutrients of concern. In the case of folic enriched flour, whole-wheat flour, brown rice flour, rolled
acid fortification of grains, there has been a reduction of oats, nonfat dry milk, yeast, and salt. This could be a bread
neural tube defects, and certain birth defects have been mix with 90% enriched flour and 10% other ingredients mak-
decreased by 5070% in various countries. Yet some fear ing it not a WG bread mix. On the other hand, the ingredient
that such a policy could put some groups at risk for excess statement could reflect 35% refined flour, 30% whole-wheat
folic acid intake, even though most data do not indicate that flour, 30% brown rice flour, 3% rolled oats, and 2% other
this might be a problem. ingredients. This bread mix would clearly be a WGF even
WG was first defined by the American Association of Cereal though refined flour was the first ingredient. A 30 g serving of
Chemists (now AACCI) in 1999 as follows: granola could have 5 g of whole oats; 4 g each of peanuts,
almonds, pecans, and walnuts; and 5 g of raisin paste, honey,
and salt. The granola, while contributing WG and many nutri-
Whole grains shall consist of the intact, ground, cracked or flaked
caryopsis, whose principal anatomical components the starchy
tious ingredients, may not give a dietarily significant amount of
endosperm, germ and bran are present in the same relative pro- WG. Some suggest that combining the ingredient statement
portions as they exist in the intact caryopsis. with the products DF levels could identify WGFs, but this
approach works only if the formulation contains high-fiber
WGs have subsequently been defined by other organiza- WGs. Low-fiber WGs such as brown rice would have a low-
tions. The Whole Grains Council (WGC) created a more fiber level on the nutrition panel.
consumer-friendly definition. It reads as follows: Achieving international agreement on the amount of WG
that should be in a product to call it a WG serving is very
Whole grains or foods made from them contain all the essential difficult. There is no international agreement regarding the
parts and naturally-occurring nutrients of the entire grain seed. If the recommended daily intake of WG. Some regions make no
grain has been processed (e.g., cracked, crushed, rolled, extruded, recommendations. For those that do, recommended intakes
lightly pearled and/or cooked), the food product should deliver vary from 48 to 115 g day 1. This wide range makes agreement
approximately the same rich balance of nutrients that is found in
the original grain seed.
on the amount of WG in a product to call it a WGF-difficult.
Some jurisdictions allow manufacturers to label statements
The European HEALTHGRAIN (HG) workshop (2008) of fact, which means statements such as YXZ product gives xx
defined WG as follows: grams of WG per serving. The difficulty is that most consumers
60 FOOD GRAINS AND THE CONSUMER | Labelling of Grain-Based Foods
do not know the amount of WG recommended per day, so such It should be noted that it is merely a characterization and
labeling could easily be misinterpreted. Similarly labeled ban- does not speak of the nutritional attributes such as DF or
ners such as made with whole grain, whole-grain goodness, detractors like sugar or fat in the product. For some consumers,
and multigrain and even high-fiber make some consumers any designation of a food as WG is misinterpreted and creates a
think that the food meets one of the recommended WG servings. halo effect causing consumers to think such foods have more
The WGC stamp has high consumer recognition, is easy to DF, fewer calories, and detractor ingredients than they do.
understand, and is used internationally. The WGC has a stamp Further, this characterization is only voluntary and the net
on the food label that indicates the product has 8 g of more of result is that there can be many package statements claiming
WG per labeled serving or RACC. The stamp has two disadvan- WG content and consumer confusion.
tages. First, it causes some consumers to deduce that only those
products bearing the stamp are WG. This may occur because
consumers are unaware that some companies may not use the Nutrient Content, Health, and StructureFunction
stamp because of its cost. Its second disadvantage is that the Claims
stamp gives an unfair advantage to products with low moisture
or large serving sizes. This creates a scenario where a dough Nutrient-content claims are allowed in some jurisdictions pro-
made into a 25 g bun would not have the needed 8 g of WG to vided that they are factual and use terminology in ways spec-
receive the stamp, but the same dough in a 55 g bagel would. ified in the regulations. In many cases, a serving of a food
Since a clear WG characterization was suggested as being carrying a nutrient-content claim must provide a certain min-
important to the goal of eating more WG, AACCI and HG tried imum percentage of the DV, often 10% in a normal serving.
to respond by putting forward a way to identify WGFs. The For example, Acme cereal is a significant source of folic acid.
following questions were laboriously considered in the WGF Structurefunction claims are popular in some countries.
determination: Japan was the first to enfranchise these and allow a number
such claims. Such claims are also allowed in North America,
1. Should the WG characterization apply only to products that
Europe, Australia, and parts of Asia. Structurefunction claims
are in the grain group?
may point out the association with health but must not claim
2. Should it be based on a serving of food or be proportional
to cure a disease. Claims such as Acme cereal is a significant
and therefore serving size-neutral?
source of vitamin A, which has been associated with skin and
3. Should it be based on a wet (as served) or dry weight basis?
eye health, and Bran fiber in Acme cereal helps maintain
(Use of wet weight favors dry products and penalizes
bowel regularity are allowed in some jurisdictions. As with
cooked oatmeal, bread, and soup with pasta or barley.)
structurefunction claims, the amount of nutrient needed to
4. Should it apply only to the grain portion of the food? If so,
make a nutrient-content claim is often the same. Some coun-
should there be a minimum level or grain, even if all the
tries prohibit any claims if the food also contains the so-called
grain is WG?
detractor nutrients, such as saturated fat. If a claim is allowed,
5. Should it indicate that the food would deliver a certain
the detractor nutrient must also be prominently labeled in the
amount of DF or not have a certain level of detractor
same place as the positive claim. In order to have claims on a
nutrients such as sugar, salt, fat, or saturated fat?
food that diagnose, mitigate, treat, cure, or prevent a disease or
6. Should it apply to the entire meal in a frozen or other
specific class of diseases, officially approved health claims are
dinner where the side of grain would meet the requirements
used. Health claims are accepted only after significant scientific
but would fail to do so when all the food entree and other
agreement by the regulatory authorities in the region. The only
meal components are factored into the calculation?
exception is that of classical nutrient deficiency diseases.
7. Should half the grain in the product be WG?
Table 2 gives a sampling of health claims for GBF.
The AACCI Whole Grain Working Group put forward a WG
characterization that suggests a proportionality of 8 g of WG
per 30 g of food. This addresses problems created by small and Health Claims Regarding Blood Glucose
large serving sizes, but leaves some unanswered questions such
as WG pasta in a soup or side in a ready-to-eat meal. The 8 g/ The Swedish Nutrition Foundation (SNF) allows the following
30 g does not specify that the product must have half of the claim about blood glucose: oat bran -glucan reduces blood
grain be WG. Rather, it places a WG minimum, which allows glucose and insulin response.
formulators of GBF to increase the other grain fractions such as EFSA allows a claim on a lowered glucose response if 14 g of
bran, germ, or aleurone, which technically are not WG but resistant starch is substituted for 14 g of available starch.
carry many nutritional attributes of WG. Some are concerned
that the characterization could allow products to have higher
levels of refined grain component than WG. This could be true Summary
but the intent of using the 8 g per 30 g is that with the recom-
mendation of six 30 g servings of GBF needed for most calorie Labeling of GBF informs consumers when making grain pur-
levels, a person would get a minimum of 48 g of WG, an chases, enables industry to comply with rules regarding GBF
amount thought to be dietarily important. and grain components, and helps regulators ensure consumer
FOOD GRAINS AND THE CONSUMER | Labelling of Grain-Based Foods 61
Table 2 Health claims for grain-based foods in various jurisdictions protection and fair marketing of GBF. Understanding label
terms for any food is challenging, but terms for GBF, which
Fiber-containing grain products, fruits, and vegetables and cancer
describe grain forms, particle size, degree of milling and spe-
Sample claim: Low-fat diets rich in fiber such as Acme Bread may reduce
the risk of some types of cancer, a disease associated with many cific milling and processing treatments of various grain types,
factors and fractions, add to consumers confusion. This is com-
Requirements: GBF must meet criteria for low fat and, without pounded because terms vary across regions and among grains
fortification, be a good source of dietary fiber for a similar process. Terms such as enriched and refined are
Fruits, vegetables, and grain products that contain fiber, particularly applied to grains and not to most other foods. General label
soluble fiber, and the risk of coronary heart disease terms such as fresh and organic apply to GBF, but these may
Sample claim: Diets low in saturated fat and cholesterol and rich in fruits, or may not be defined in a jurisdiction. If defined, definition
vegetables, and grain products that contain some types of dietary fiber, even within a jurisdiction may vary among products with
particularly soluble fiber, may reduce the risk of heart disease, a
fresh on bread meaning something different than fresh on
disease associated with many factors
pasta. Some use organic labeling as a way to avoid chemicals
Requirements: GBF must meet criteria for low saturated fat, low fat,
and low cholesterol. They must contain, without fortification, at least and GMFs. Nutritionally labeled products are found in many
0.6 g of soluble fiber per reference amount, and the soluble fiber regions and help consumers meet individualized nutritional
content must be listed needs, although many consumers are confused by many terms
Folic acid and neural tube defects on the nutrition and grain label. GBF can carry a number of
Sample claim: Healthful diets with adequate folic acid may reduce a nutrient-content, structurefunction, and health claims. Each
womans risk of having a child with a brain or spinal cord birth defect region has specific rules regarding their approval and use.
Requirements: Enriched GBF must meet or exceed criteria for good Various health promotion agencies are suggesting the inges-
source of folic acid, that is, at least 40 mg of folic acid per serving (at tion of more WG. However, the ability of consumers to make
least 10% of the daily value)
informed choices as to what is a WGF is difficult. Even experts
Dietary soluble fiber, such as that found in whole oats, barley, oat bran,
disagree on the best and fairest strategy for such labeling. Vari-
and Oatrim or psyllium seed husk and coronary heart disease and
cholesterol lowering ous strategies have been proposed for the identification of label-
Sample claim: Diets low in saturated fat and cholesterol that include 3 g ing WGFs, but each has some drawbacks. Most recently, AACCI
of soluble fiber from whole oats per day may reduce the risk of heart has put forward a WG characterization, which is a proportional
disease. One serving of this whole oat product provides xx g of this minimum of 8 g or WG/30 g of product. Labeling of GBF is
soluble fiber important and complicated. It varies across regions due to non-
Requirements: Foods must meet criteria for low saturated fat, low standardized use of terms, varying nutritional requirements,
cholesterol, and low fat. Foods that contain whole oats must contain differing regulations by food authorities, and varying degrees
at least 0.75 g of soluble fiber per serving. Foods that contain psyllium of consumer understanding, needs, and perceptions.
seed husk must contain at least 1.7 g of soluble fiber per serving
Typical foods: GBF made with whole or rolled oats, oat bran, or whole oat
flour; barley; psyllium seed husk; and isolated or modified b-glucans
Exercises for Revision
from barley and oats
Note: The European Food Safety Authority (EFSA) allows the claim Oat
beta-glucan has been shown to lower/reduce blood cholesterol. Blood Take a country or region and compare the nutrition label
cholesterol lowering may reduce the risk of heart disease. The Swedish and general labeling requirements for GBF.
Code of Practice allows Soluble fiber from oat bran may help reduce Compare the various terms used to describe the milling
cholesterol. The United Kingdom allows The inclusion of oat bran can process of various grains such as pearling, polishing, pin
help reduce cholesterol as part of a diet low in saturated fat and a milling, and rolling and resultant particulate fractions such
healthy lifestyle. The Netherlands Nutrition Centre (Voedingscentrum) as kernels, groats, grits, meals, and flours.
allows One daily portion of ACME oat bran-based bread reduces
cholesterol within 3 weeks and helps maintain healthy arteries and
Compare health claim approval and use for GBF in various
countries.
blood vessels. And Switzerland BAG Oat bran favorably impacts
cholesterol levels Compare the labeling of WGs and WGFs across
jurisdictions.
Whole grain and heart disease
Diets rich in whole-grain foods and other plant foods and low in total fat, Compare the rules for using the terms natural, fresh, and
saturated fat, and cholesterol may reduce the risk of heart disease and organic across jurisdictions.
some cancers What contributions do enriched, refined, fortified, and
Dietary fiber content of at least 1.7 g per 35 g serving WGs make to the diet of a particular country or
Typical foods: Whole-grain breads and cereals (USFDA claim) subpopulation?
WG claims not been approved in many other jurisdictions because of
inadequate characterization of the ingredients and rulings of inadequate
evidence Exercises for Readers to Explore the Topic Further
Vitamins B12, B6, and folic acid and reduced risk of heart disease
Sample claim: Foods containing vitamin B12, as well as B6 and folic acid,
may reduce the risk of coronary heart disease
Label marketing statements, grain and processing
terminology, and nutrition labeling are confusing to con-
Typical foods. Fortified GBF may make such claims in the United States
sumers. Suggest some strategies to diminish confusion.
62 FOOD GRAINS AND THE CONSUMER | Labelling of Grain-Based Foods
What activities and labeling initiatives can be undertaken to Barr SI, DiFrancesco L, and Fulgoni 3rd VL 3rd (2013) Consumption of breakfast and
help consumers identify WGFs and increase their WG the type of breakfast consumed are positively associated with nutrient intakes and
adequacy of Canadian adults. Journal of Nutrition 143(1): 8692.
intake?
Berry RJ, Bailey L, Mulinare J, Bower C, and Folic Acid Working Group (2010)
Percent ingredient labeling is used in some countries for all Fortification of flour with folic acid. Food and Nutrition Bulletin 31(Supplement 1):
foods including GBF. Does this improve consumer under- S22S35.
standing of what is in the food? FDA Wants Consumers to Truly Get the Whole Grain (2014). http://usgovinfo.about.
com/od/consumerawareness/a/wholegrain.htm.
Grieger JA and Cobiac L (2012) Comparison of dietary intakes according to
breakfast choice in Australian boys. European Journal of Clinical Nutrition 66(6):
See also: Food Grains and the Consumer: Fortification of Grain- 667672.
Based Foods; Genetically Modified Grains and the Consumer; Grains Hamner HC, Tinker SC, Berry RJ, and Mulinare J (2013) Modeling fortification of corn
and Health: Misinformation and Misconceptions; Scientific masa flour with folic acid: The potential impact on exceeding the tolerable upper
Organizations Related to Grains: Research Organizations of the intake level for folic acid, NHANES 20012008. Food and Nutrition Research
57, 19146.
World: Global Trends and the Commercial Sector; Scientific Societies Holmes BA, Kaffa N, Campbell K, and Sanders TA (2012) The contribution of breakfast
Associated with Grain Science; The Basics: Grain: Morphology of cereals to the nutritional intake of the materially deprived UK population. European
Internal Structure; The Cereal Grains: Barley: An Overview of a Journal of Clinical Nutrition 66(1): 1017.
Versatile Cereal Grain with Many Food and Feed Uses; Maize: Yang Q, Cogswell ME, Hamner HC, et al. (2010) Folic acid source, usual intake, and
folate and vitamin B-12 status in US adults: National Health and Nutrition
Overview; Oats: Overview; Rice: Overview; Wheat: An Overview of the
Examination Survey (NHANES) 20032006. American Journal of Clinical Nutrition
Grain That Provides Our Daily Bread. 91(1): 6472.
Relevant Websites
Further Reading
http://wholegrainscouncil.org/whole-grains-101/definition-of-whole-grains
AACCIs Whole Grains Working Group Unveils New Whole Grain Products HealthGrain Forum. Definition and analysis of whole grain.
Characterization (2013). http://www.aaccnet.org/about/newsreleases/Pages/ http://wholegrainscouncil.org/whole-grains-101/definition-of-whole-grains Whole
WholeGrainProductCharacterization.aspx. Grains Council. Definition of whole grains.
Grains and Health
R Korczak, University of Minnesota, St. Paul, MN, USA
D Hauge and B Maschoff, Grains for Health Foundation, St. Louis Park, MN, USA
L Marquart, University of Minnesota, St. Paul, MN, USA; Grains for Health Foundation, St. Louis Park, MN, USA
P Jacques, Tufts University, Boston, MA, USA
R Lindberg, Minneapolis Heart Institute Foundation, Minneapolis, MN, USA
R Menon, The General Mills Bell Institute of Health and Nutrition, Minneapolis, MN, USA
2016 Elsevier Ltd. All rights reserved.
Moving from Traditional Research and Analytic foods supply, which interfaces with a community of clients
Thinking to Include Systems Approaches who frequent the food environment, such as restaurants
whereby these healthier grain-based foods are served and
Traditional nutrition research, from a biological and physio- potentially consumed.
logical dimension, has been primarily explored through Systems thinking is one approach to view problems as
in vitro, animal, observational, and experimental approaches parts of an overall system, rather than reacting to a specific
where reductionism is considered the gold standard. Analytic part, outcome, or event. It is about understanding and seeing
thinking has been the mainstay for scientific investigation and through the complexity and managing interdependency of
remains an important aspect of our current scientific practice. sectors, disciplines, and individual contributions. The process
These traditional approaches were extremely effective in encourages understanding of the big picture by sensing its
addressing nutrient deficiencies in the late nineteenth and large-scale patterns and responding to them. We might benefit
early twentieth centuries, up until the 1950s and 60s when by carefully crafting a process for defining problems and only
chronic diseases became a major issue of concern for food then designing collective solutions. Ultimately, this reflects
and nutrition researchers. design thinking in bringing more healthful ingredients,
As our society moves into a new millennium, innovative foods, and products into the kitchen and/or marketplace that
approaches are necessary to address the complex and dynamic consumers demand, by taking into account the influences of
health-related issues facing scientists through food technology, the overall food system, food environment, and community. A
nutrition and health, and community involvement. Unfortu- systems approach will require multicultural representation to
nately, we are routinely taught throughout our academic come around a specific initiative to prioritize, focus, and lever-
careers and professional lives on how to solve problems, but age the strengths of individuals throughout the various sectors
rarely taught how to identify problems. According to Russell and diverse disciplines of study.
Ackoff, We fail more often not because we fail to solve the
problems we face, but because we fail to face the right
problem. For example, how will we address the complexity of
what and how to deliver healthy grain-based foods that con- Partnerships to Achieve a Common Goal around
sumers find acceptable and will include in their daily diet? One Healthier Grain-Based Foods
solution, as recommended by the 2010 Dietary Guidelines for
Americans Committee, is for the sectors of government, industry, According to Rowe et al., PPPs can create a collaborative envi-
and nonprofits (including both large and small businesses) to ronment that maximize cross-disciplinary expertise among
come together in order to develop a system that would allow for government, academic, and industry researchers to help solve
health-promoting dietary patterns. At minimum, public health complex research problems. Leveraging the proficiencies and
officials are being asked to consider not only the overall roles of competencies of a broad range of stakeholder experts could
each player within the food system and how they work together maximize available expertise and collectively achieve the best
to create the food environment but also the significance of research outcomes while extending and enhancing limited
community and citizen representation in identifying and filling financial and other resources. Figure 1 addresses 12 proposed
the gaps, such as research projects that help meet the diverse and principles of PPPs, offered in no strict chronological order
complex demands of the public. because each partnership has unique characteristics.
For example, one way to fulfill the gap(s) and meet the
demands of the public is to develop a grain-based food that
is practical, healthier, affordable, and desirable (PHAD) such
that consumers will eat it in a restaurant setting. Yet the food It Starts by Defining Shared Value and Vision
must be practical for supply chain stakeholders along with
chefs and clientele in restaurants. A grain-based food might be Our vision is to bring together the various sectors, including
described as healthier, such that whole grain (WG) wheat is government, industry, nonprofit, academia, and activist
incrementally or gradually increased if the food/product groups, to begin articulating the needs, desires, and demands
meets PHAD criteria and restaurant clientele actually eat it. for targeting research dollars. The acquisition of research dol-
Affordability throughout the supply chain and in the food lars will help fill a void, which increases our ability as a society
preparation and restaurant setting can be a limiting factor. to cost-effectively deliver healthier grain-based foods into the
Overall desirability of these grain foods by supply chain marketplace. This is a vision of operation whereby sharing of
stakeholders, restaurant chefs, staff, and clientele along with intimate information, goals, and objectives makes it challeng-
community norms needs to be taken into account. ing to establish a common shared value leading to a common
In order to increase consumer intake of these grain-based purpose, which drives this network of individuals toward a
foods with higher WG content, it will require the development common outcome and benefit for business and society.
of tools, methods and approaches, systems, and necessary
infrastructure to successfully develop and deliver these foods
into the marketplace. The successful outcome of this effort
Vision
might be underpinned via research initiatives that focus on
increasing WG ingredients in grain-based foods for restaurant Increase the availability of grain-based foods that are PHAD
service. Ideally, these inherent research problems and solutions with the intention of making the healthy choice the easy and
are conceived and tested within the context of a complex grain natural choice for citizens.
FOOD GRAINS AND THE CONSUMER | Grains and Health 65
Figure 1 PPPs should do the following: Alignment among stakeholders through gauged,
1. Have a clearly defined and achievable goal to improve the health of coordinated, and prioritized research efforts that bridge
the public the gaps between research and consumer solutions
2. Articulate a clear statement of work, rules, and partner roles, A sustainable infrastructure guided by industry profitability
responsibilities, and accountability, to build in trust, transparency, while gradually improving the availability and consumer
and mutual respect as core operating principles acknowledging acceptance of healthier food products
there may be deal breakers precluding the formation of an effective
partnership in the first place Targeted, prioritized research efforts that allow scientists to
address the food attribute gaps related to quality, cost,
3. Ensure that objectives will meet stakeholder partners needs, with a
clearly defined baseline to monitor progress and measure success convenience, and efficacy, translating science into food
4. Considering the importance of balance, ensure that all members Research that ranges from fundamental to product applica-
possess appropriate levels of bargaining power tion of functional foods and health, including enhanced
5. Minimize conflict of interest by recruiting a sufficient number of delivery of bioactives
partners to mitigate influence by any single member and to broaden Research that focuses on the successful development,
private sector perspectives and expertise delivery, and enhanced consumer intake of healthier food
6. Engage partners who agree upon specific and fundable research ingredients (this knowledge can provide insight on what
question(s) to be addressed by the partnership innovation opportunities are feasible)
7. Enlist partners who are committed to the long-term goals as well as
to the sharing of funding and research data
8. Along with government and the private sector, include academics
and other members of civil society as partners Who Will Guide This Process?
9. Select objective scientific measurements capable of providing
common ground for both public and private sector research goals Assemble the right minds
10. Adopt research questions and methodologies established by The PPP will be composed of an expanded network of
partners with no vested financial interest in them, ideally in the working partners across all sectors and disciplines of the
precompetitive space food supply, closely linked to consumer groups.
11. Be flexible and ensure ongoing transparent communications We are asking stakeholders to be a part of a PPP, which
12. Consider a third-party convener to ensure equality at the table, will be formed to identify and to convene these partici-
clarify rules, establish operational guidelines, and specify funding pants and to solidify the project scope and the overall
arrangements approach.
Why Participate?
Values PPP members will be uniquely positioned to
Values are integral in the roles that individuals play through steer the fundamental research path from the ground floor,
the group process and dynamics, clarity of objectives, mutual before it becomes available to the greater grain-based food
trust, cooperation, and expected outcomes. industry/community;
view firsthand the output of this fundamental and transla-
tional research, which can feed research and development
Why Is There a Need for a PPP?
efforts to reach targeted customer segments such as schools,
Food and ingredient industries are uniquely positioned with restaurants, home, and retail;
the technical expertise to make gradual product formulation leverage the resulting innovation and new processing
changes that are well accepted by consumers. These gradual technologies.
product formulation changes will not only improve the prod-
uct nutritional profile but also move consumers toward dietary
recommendations: How Will This PPP Function?
How can we fund research initiatives that guide food for- 1. Key representatives, from across the food supply, will con-
mulation and product development to make it easier for vene to identify and prioritize research issues and gaps.
consumers to eat in a way that supports dietary guidance? 2. Meaningful collaboration will occur through application of
Is there a research approach that simultaneously encour- collective knowledge and targeted research.
ages new ingredient and innovative technology develop- 3. Mutually beneficial collective solutions for business and for
ment, supports a profitable industry business model, and society will result, ultimately positively influencing public
delivers products with enhanced nutritional value that health outcomes.
appeal to consumers?
What Are the First Steps? What Makes This PPP Approach Unique?
New research is needed to support system-wide changes and to This will be an innovative and a gradual shift in the way we
assure continuity from science (theory) to consumers (prac- think about our food supply resulting in a shared and coordi-
tice) throughout the food delivery system. This integrated and nated approach to research with the following governing
multifaceted process would deliver the following: guidelines:
66 FOOD GRAINS AND THE CONSUMER | Grains and Health
Open and safe environment to share ideas Follow-up from the meeting will be posted on the Grains for
Time to identify research gaps that limit availability of Health Foundations website (Whole Grains Summit 2012
healthier foods in the marketplace Events Page: http://grainsforhealth.org/news-and-events/
Supporting infrastructure to implement the changes events/2012-whole-grain-summit).
Freedom to innovate and to conduct the targeted research Further discussions with representatives from industry, gov-
necessary to transform science into food ernment, grain organizations, and academic institutions that
proposed research initiatives/projects include
What Will Success Look Like? I. standardized approaches and methods to WG research,
II. WG impact on the microbiome project,
Short term to midterm III. community WG farm-to-fork intervention in New Ulm, MN.
Coordinated and comprehensive plan that identifies research
gaps through an integrated and gradual approach so that basic,
applied, and translational research can be targeted toward the
development and delivery of healthier grain-based foods that
Standard Protocol for WG Intervention Studies
of Intermediate Metabolic End Points (Proposed
deliver the attributes of a food product that are appealing to
Project I)
consumers.
Purpose
Long term The main objective is to develop a standardized protocol
Sustainable infrastructure in place that supports a coalition
whereby future intervention studies of WG and surrogate end
across the food supply chain, developing a unified strategy
points would contain very specific research methods. Identify-
that leads to increased availability of grain-based foods that
ing the appropriate methods of WG intervention studies would
meet dietary guidance and resulting in a model that meets both
allow for easier comparisons across studies of similar WG and
business and societal needs.
of studies that investigate different types of WG. Conducting a
meta-analysis would then be more easily performed to ade-
quately address the role of WG in prevention of chronic
Overview of Foods Research and Public Health disease.
OVERVIEW FOR
ERNM
GOV
EN
T
PR
N- O
NO
FIT
PARTNERSHIPS
ME WGS
DE
ACA RY
DUST
IN
SCIENTIFIC FOOD
FOOD
INPUT SUPPLY CHAIN ENVIRONMENT
ATTRIBUTES
GROWIN G
P R O D U C TI O N
MANUFACTURING
CO NSUMER
CONSUMER BENEFIT
PROJECT SCOPE
DIET
Genetics Gut
Lifestyle Microbiome HEALTH
Life stage
Other
How does a whole grain diet How does the gut microbiota
influence the gut microbiota? influence health?
Figure 3 The possible scope of a collaborative research project to assess the impact of whole grain foods on the gut microbial communities.
FOOD GRAINS AND THE CONSUMER | Grains and Health 69
(CBPR) approaches will be utilized to increase WG availability, Yes, this has potential for significant public health impact.
sustainability, and consumption of AFH foods by increasing Investigating the current barriers to a sustainable WG supply
the number of WG offerings and percent WG content in grain- chain consistently delivering healthier WG options to AFH
based foods found in restaurants and grocery store eateries. settings benefits the consumer. By allowing chefs, manu-
This supply chain intervention will start in New Ulm, facturers, researchers, and policy makers to understand the
Minnesota, as part of GrainUP!, a grassroots WG consumer barriers, challenges, and opportunities to increasing WG
awareness campaign that encompasses the crucial industry- consumption AFH, steps can be taken to make the WG choice
facing supply chain work to create a road map for increasing the easy choice. This supports the Dietary Guidelines for
WG menu offerings AFH. Americans recommendation of eating at least three servings
The proposed study will test the following hypothesis: of WG a day.
increasing the WG food offerings by 2550% (depending 2. Does it have implications across many different food and/or
upon the AFH setting and food category) and gradually intro- beverage categories?
ducing an increase in the WG content of popular grain-based Grains are present in most AFH food categories, from
items in New Ulms AFH food environment (restaurants and appetizers to dessert. In addition, their appearance usually
grocery store eateries). The hypothesis is that this will increase includes other food groups, such as fruits and vegetables,
AFH WG consumption from an average of one-third serving meats and legumes, and dairy. Grains can serve as the
per day to 1 serving per day, thus contributing to the Dietary perfect vehicle for healthier foods AFH and provide an
Guidelines for Americans overall recommended three daily opportunity to address portion sizes and added sugars
servings of WG. We aim to address availability, consumption, and fats.
and sustainability at a systems level. 3. Is it precompetitive in nature and what does this mean for the
food industry?
Why This proposal provides the opportunity for both ingre-
dient and consumer packaged goods manufacturers to
For almost a decade, the Dietary Guidelines for Americans have
develop and pilot products and supply chain strategies, in
recommended that Americans consume at least three ounce
a precompetitive, collaborative environment. This applied
equivalents (servings) of WG foods daily for optimal health.
research model holds the potential to streamline processes,
WG foods improve heart health and weight management and
reduce costs, and ensure technology transfer that can be
lower risk of certain cancers. Yet in spite of health benefits and
applied to other cities and communities.
Dietary Guidelines for Americans recommendations, the average
American consumes only 0.8 servings per day. Low WG con-
sumption may be linked to the food environment and consump-
tion of foods outside of the home. Today, the average American Who
spends 42% of his or her food budget on AFH foods, which This proposal involves the resources and expertise of the Hearts
constitutes about 35% of daily calories. These foods tend to be Beat Back Project (Minneapolis Heart Institute Foundation,
lower in nutritional quality often high in calories, fat, sodium, Allina Health, and the community of New Ulm, MN), food
and cholesterol and offer less than one-third ounce of WG per industry, academics at the University of Minnesota, and project
day. Eating AFH as little as one meal per week can lead to management from the Grains for Health Foundation. The
approximately two extra pounds of weight each year for adults. Hearts Beat Back Project has previously received a USDA
Modifying the food environment where adults frequently eat is a Farmers Market Promotion Program grant, so has relationships
promising approach to bringing about sustainable improve- with farmers/producers, and is currently finishing a CDC Com-
ments in nutrition behaviors at a population level so that when munity Transformation grant (expanding their work to the rest
consumers are faced with making food choices, the healthier of Brown County, MN).
choice WG foods is the easier and preferred choice.
Recent research examining the relative healthfulness of
restaurant menus has shown that of 34 restaurants assessed
in New Ulm, fewer than 26% offered a WG bread option. Where
According to Rebecca Lindberg, researcher of Hearts Beat The intervention will support current work of the Hearts Beat
Back: The Heart of New Ulm Project, feedback from restaurant Back Project, a 10-year population-based demonstration pro-
owners and chefs involved in trying to improve the healthful- ject designed to deliver interventions at individual, institu-
ness of their menu offerings has indicated that the WG category tional, and community levels that reduce the rate of heart
is one of the most difficult menu items to change (oral com- disease risk factors in adult residents of New Ulm. Minneapolis
munication, 17 March 2014). Recent restaurant and grocery restaurants will be the next intervention site(s), once a sustain-
store eatery assessments demonstrated that increasing AFH WG able supply chain approach and model is established, with
menu choices remains difficult to influence when compared to eventual rollout to other nationwide cities and communities.
other healthful restaurant practices.
Phase 1: Environmental scan for assessments and establishment Public and private partnerships combine what type of sec-
of AFH community forums tors and disciplines?
Phase 2: Gradual implementation of intervention using the
CBPR approach
Phase 3: Social marketing and promotion Exercises for Readers to Explore the Topic Further
Phase 4: Final assessments and evaluation of likelihood
intervention outcomes will be sustained Identify the challenges and opportunities (gaps) related to
the substitution of WG wheat flour (50% whole wheat
flour) in a restaurant pizza. Track the pizza (1) through
How the supply chain from field to fork and (2) in the food
Typically, the last mile of the AFH food supply chain is where environment, such as in a restaurant, and explore the char-
many of the complications with WG foods reaching the con- acteristics of the community patrons who frequent this
sumer occur. Given this, intervention will occur at various local restaurant.
points along the food supply chain (miller, baker, food How might a flour milling or packaged foods company use
manufacturer, and AFH eating environment) to address WG this information related to the gaps in the food supply,
food quality, affordability, and desirability. An ecological food environment, and community patrons?
framework and the CBPR approach will be used to assess, What type of training programs might colleges, universities,
implement, promote, and measure a gradual increase in WG and industry consider in training of students and young
offerings and percent WG content of current grain offerings. professionals to understand more about the food system,
WG foods of varying refined grain/WG content will gradually food environment, and surrounding community who pur-
be introduced in these establishments, along with technologies chase and use their products?
to help reduce rancidity and enable tracking of shelf life of WG
flours to ensure the delivery of a fresh, safe, nutritious, and
palatable final product. The goal is that the increased consumer
Further Reading
acceptance and consumption of WG foods in AFH setting will
become permanent and sustainable. Ackoff RL (1981) Creating the Corporate Future: Plan or Be Planned for. New York:
Wiley.
Andersson A, Tengblad S, Karlstrom B, Kamal-Eldin A, Landberg R, Basu S, et al.
(2007) Whole grain foods do not affect insulin sensitivity or markers of lipid
Conclusions peroxidation and inflammation in healthy, moderately overweight subjects. Journal
of Nutrition 137: 14011407.
New research is needed to support system-wide changes and to Beauman C, Cannon G, Elmadfa I, Glasauer P, Hoffmann I, Keller M, et al. (2005) The
principles, definition and dimensions of the new nutrition science. Public Health
assure continuity from science (theory) to consumers (prac- Nutrition 8(6A): 695698. http://dx.doi.org/10.1079/PHN2005820.
tice) throughout the food delivery system. Public and private Bodinham CL, Hitchen KL, Youngman PJ, Frost G, and Robertson MD (2011) Short-
research is one approach to explore ways of identifying term effects of whole-grain wheat on appetite and of intake in healthy adults: A pilot
research initiatives that might help fill gaps in the food supply study. British Journal of Nutrition 106: 327330.
Ferruzzi M, Jonnalagadda S, Liu S, Marquart L, McKeown N, Reicks M, et al. (2014)
chain, food environment, and community settings to more
Developing a standard definition of whole grain foods for dietary recommendations:
successfully deliver healthier grain-based foods into the mar- Summary report of a multidisciplinary expert round table discussion. Advances in
ketplace that consumers demand and readily consume. Nutrition 5(2): 164176.
A process is necessary to examine the issues, problems, and Gharajedaghi J (2011) Systems Thinking: Managing Chaos and Complexity: A Platform
gaps that exist relative to the development, delivery, and con- for Designing Business Architecture, 3rd edn. New York: Elsevier.
Giacco R, Clemente G, Ciriano D, Luongo D, Viscovo D, Patti L, et al. (2010) Effects of
sumer intake of healthier grain-based foods. Each research the regular consumption of wholemeal wheat foods on cardiovascular risk factors in
project has merit as it relates to enhancing the development, healthy people. Nutrition, Metabolism, and Cardiovascular Diseases 20: 186194.
delivery, and consumption of grain-based foods made with The Giessen declaration. Public Health Nutrition (2005) 8(6A): 783786.
WG ingredients. Each project is complex in its own way and Grains for Health Foundation, Whole Grains Summit 2012: http://grainsforhealth.org/
news-and-events/events/2012-whole-grain-summit.
affects different aspects of the food system and food environ-
Hassel CA (2014) Reconsidering nutrition science: Critical reflection with a cultural
ment along with community input and influence. We began to lens. Nutrition Journal 13: 42. http://dx.doi.org/10.1186/1475-2891-13-42.
address these research initiatives during a PPP meeting held on Jacobs Jr. DR Jr., Meyer KA, Kushi LH, and Folsom AR (1998) Whole grain intake may
34 December 2014, in Washington, DC. The outcomes of this reduce the risk of ischemic heart disease death in postmenopausal women: The Iowa
meeting will be discussed in another communication venue. Womens Health Study. American Journal of Clinical Nutrition 68: 248257.
Jacques P, Hauge D, Voth K, Hermann M, Maschoff B, and Marquart L (2013)
Overcoming the challenges of translating the US Dietary Guidelines into healthier
grain-based foods. Nutrition Today 48(6): 254259.
Lankinen M, Schwab U, Kolehmainen M, Paananen J, Poutanen K, Mykkanen H, et al.
Exercises for Revision (2011) Whole grain products, fish and bilberries alter glucose and lipid metabolism
in a randomized, controlled trial: The Sysdimet Study. PLoS One 6: e22646.
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Food Grains: Intolerance, Allergy and Diseases
Contents
Cereal Allergens
Nutrition: Beriberi, A Deficiency Related to Grains
Celiac Disease
The Gluten-Free Diet
Cereal Allergens
AS Tatham, Cardiff Metropolitan University, Cardiff, UK
2016 Elsevier Ltd. All rights reserved.
their highly conserved sequence regions, structures, and func- binding to the a-, g-, and o-type gliadins and low-molecular-
tions and account for the cross-reactivity of allergens that are weight subunits of glutenin.
found within the cereals. The prolamins (defined as aqueous Detailed studies of sequentially extracted wheat flour, into
alcohol-soluble seed storage proteins) of wheat, barley, and salt-soluble (albumins and globulins) and alcohol-soluble gli-
rye, and to some extent oats, are related; those of maize, millet, adins and glutenins, indicate that patients sera recognize mul-
sorghum, and rice are in part related but differ significantly tiple allergens.
from wheat, barley, rye, and oats. Other protein families Other ingredients are used as flour/bakery additives, often
include the pathogenesis-related (PR) proteins, which are starch-cleaving enzymes, and some of these have been found to
involved in the defense systems of plants against a host of be allergenic. Fungal a-amylase from Aspergillus oryzae is added
different pathogens, and also albumins and globulins that to digest starch and is allergenic, as are cellulases and xylanases
can be involved in storage functions and metabolic proteins. from Aspergillus niger, added to flour to digest fiber. Soy flour,
Understanding these relationships can be critical in under- added to wheat flour as a source of lipoxygenase to whiten
standing cross-reactivity in cereal allergy and intolerance. flours, is responsible for food and inhalational allergies. Soy-
Allergic reactions to wheat can be divided into inhalational bean allergy has been implicated in bakers asthma.
allergy (bakers asthma), ingestion (WDEIA), and also wheat
allergy and contact dermatitis, caused by skin contact with
wheat proteins. In bakers asthma and WDEIA, the allergens Wheat-Dependent Exercise-Induced Anaphylaxis
are well characterized, whereas the allergens involved in wheat
allergy and contact dermatitis appear to be related to a range of Food-dependent exercise-induced anaphylaxis (FDEIA) is
proteins that vary between individuals. defined as when food intake, followed by exercise, results in
the display of allergic symptoms (from urticaria to anaphylac-
tic shock). When food intake and exercise are taken indepen-
Bakers Asthma dently of each other, no symptoms are observed. Ingestion of
the causative food and exercise within 23 h (moderate to
Bakers asthma is one of the most common forms of occupa- strenuous) are required for the allergic response to occur.
tional asthma, caused by the inhalation of wheat and cereal Wheat is reported, along with crustaceans, to be one of the
flours and of dusts and other substances commonly found in foods most commonly reported to be associated with FDEIA.
bakeries and flour mills. In bakery and flour-handling workers, The incidence of FDEIA is difficult to evaluate but estimated to
studies indicate 510% suffer from asthma and 1020% account for 515% of all anaphylactic cases; of this, one-third
from rhinitis, making it one of the most common occupational to one-half are estimated to arise from FDEIA. In a Japanese
allergies. study, the incidence was found to be 0.017% and 0.031% for
Studies have shown that multiple allergens are present in cases of FDEIA and EIA, respectively.
cereal flours that are recognized by IgE from patients sera. One The major allergens associated with WDEIA have been
group, the a-amylase inhibitors, contains the major allergens, identified as the o5-gliadins (from the B genome of wheat)
with other allergens (including the lipid-transfer proteins and the high-molecular-weight (HMW) subunits of wheat. In
(LTP), peroxidase, thioredoxin, and serine proteinase inhibi- one study, 80% of patients had IgE reacting to o5-gliadins,
tors) showing the strongest IgE reactivates in wheat flour; the the remainder to HMW subunits. The g-35 and g-70 secalins of
prolamins, thaumatin-like proteins, and acyl-CoA are also rye and the g-3 hordeins of barley cross-react with wheat
implicated. In addition, a range of flour improvers/additives allergic sera, but no cross-reactivity with the avenins of oats
contribute to the allergenic potential of cereal flours. has been reported.
The a-amylase inhibitors consist of 1216 kDa polypep- In wheat, the epitopes (amino acid sequences that cause the
tides with 45 intrachain disulfide bonds, which are tightly allergic response) are well characterized. Four immuno-
folded, making them thermally stable and difficult to digest. dominant epitopes in the o5-gliadins of wheat have been
They are encoded by multigene families in wheat, barley, and identified (QQIPQQQ, QQFPQQQ, QQSPEQQ, and
rye, giving rise to marked heterogeneity in their composition. QQSPQQQ). Mutational analysis, using synthetic peptides,
In addition, they can occur as monomers, homodimers, and indicated that Gln at positions 1, 5, 6, and 7 and Pro at
tetrameric aggregates. They are active against the a-amylase of position 4 of the heptapeptides were critical for IgE binding.
insects and in the cereals are active in plant defense. Homology Omega gliadins, derived from the A and D genomes of wheat,
between the proteins found in wheat, barley, and rye accounts do not contain these epitopes and are not implicated in
for the cross-reactivity between the flours from these cereals. WDEIA. Three IgE-binding epitopes, in the repetitive domains
They are panallergens, being present in the flours of other of the HMW subunits of wheat, have been identified, QQPGQ,
cereals. QQPGQGQQ, and QQSGQGQ, using synthetic peptides.
In addition, LTPs constitute a family of panallergens; their Aspirin increases the levels of gliadin-derived peptides cir-
main function is in plant defense against phytopathogens culating in the bloodstream and upregulates the allergic
(fungi and bacteria); they are classified as PR proteins. They response to specific foods, including wheat. Wheatexercise
have been identified as the major allergens in many fruits in and wheataspirin challenges increased the levels of gliadin-
the Rosaceae family (e.g., peach, plums, and apples). In bakers derived peptides in the blood, indicating that both exercise and
asthma, they are major allergens. aspirin facilitate allergen absorption from the intestine.
In addition to the water/salt-soluble allergens, which con- The specificity of the association between o5-gliadin and
stitute the major classes of allergens in bakers asthma, the WDEIA has led to the development of diagnostic tests measur-
prolamins are also implicated. Patients sera have shown ing the levels of IgE directed against recombinant o5-gliadins.
74 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Cereal Allergens
Wheat allergy caused by the ingestion of wheat proteins can Maize allergy has been reported in countries where maize
give rise to a range of symptoms including urticaria, atopic forms a staple in the diet, such as Africa, Central and South
dermatitis, and anaphylaxis. Wheat allergy in children, similar America, and southern Europe. In the United States, the prev-
to egg and milk allergies, is outgrown by the majority of alence of allergy to maize and maize-derived sweeteners is low
children by the age of 35. In children, the most common (estimated at 0.016% of the population); the incidence else-
symptoms are atopic dermatitis. In adults, wheat allergy is where is thought to be higher. Maize allergy is linked to occu-
less frequently observed. pational asthma and contact dermatitis through handling
Studies have shown that the IgE fractions for both adults maize flour and food allergy by maize ingestion.
and children react with gliadin (a-, g-, and o-types) and glute- The major allergens are LTPs and a-amylase inhibitors; the
nin fractions, albumins, and globulins, the latter two groups LTPs show cross-reactivity with LTPs from barley and rice;
being strongly correlated with atopic dermatitis. Similar stud- minor allergens include thioredoxins, g-zein (an alcohol-
ies have shown differences in the pattern of IgE recognition soluble seed storage protein), and trypsin inhibitors.
related to age and symptoms. Patients who showed anaphy-
lactic reactions, similar to those with WDEIA, had IgE to the
o5-gliadins.
Patients with this form of allergy often show IgE to multiple Allergies to Minor Cereals and Pseudocereals
allergens, and diagnosis, other than by food challenge, is prob-
lematic. Wheat-specific IgE concentrations do not appear to be Food and inhalational allergies have been reported for millets
good indicators of the outcomes of provocation (food chal- and sorghum and for the pseudocereals buckwheat, amaranth,
lenge) tests, so even with a low or negative outcome, wheat and quinoa. These crops are recommended for inclusion in
allergy cannot be excluded without a challenge test, if clinical gluten-free diets for celiac sufferers.
symptoms suggest wheat allergy.
Diagnosis of Cereal Allergy suggest that wheat gluten and wheat starch are found in more
than 30% of packaged supermarket foods.
If a cereal allergy is suspected, a number of tests can be per- Allergy medications control the symptoms and reactions to
formed for an accurate diagnosis. allergens but do not cure the allergy. Antihistamines are the
Skin prick tests (SPTs) involve applying allergens, in solu- most common form of allergy medication. During an allergic
tion, subcutaneously. In an allergic individual, these will pro- reaction, the immune system releases histamine, which starts
voke mast cells to release histamine and produce inflammation the cascade effect of allergy symptoms, including swelling or
within about 15 min. Blood tests measure IgE levels in vitro in edema, a drop in blood pressure, and itching. Antihistamines
blood serum. These tests measure sensitization, and not nec- work by blocking the action of histamine, and they work best if
essarily a clinically relevant allergy. The most reliable tests taken before an allergic reaction has started, but they can block
involve food challenges, which are used for immediate wheat the further release of histamine after a reaction has started.
allergy detection. Although diagnostic in vitro IgE and SPT Emollient lotions and creams, used for eczema and dry skin,
assays are known to be unsatisfactory, they are extensively may contain antibiotics to reduce the risk of infection and
used. The CAP system has a predictive value for wheat of steroids to reduce inflammation. Adrenaline is used as an
< 75%. The recent use of recombinant o5-gliadin for the diag- emergency treatment for anaphylactic shock, where sudden
nosis of WDEIA patients was reported to be around 80%, and high levels of histamine released during the allergic reaction
children with high concentrations of IgE specific to o5-gliadin cause difficulty in breathing. Adrenaline alleviates the symp-
had a higher risk of failing a wheat challenge. The tests may, in toms of the reaction and can be delivered by an autoinjector
some part, be unsatisfactory due to the allergen mixtures that before further emergency treatment.
are used. The albumins and globulins are relatively easy to
extract from wheat flour; the prolamins are less so, due to
their poor solubility in aqueous media and possible presenta- Application of Genetic Modification
tion in the IgE-binding assay.
The use of genetic engineering to downregulate gene expres-
sion has been demonstrated in a number of cereal crops,
particularly in relation to reducing the levels of specific pro-
Safe Amounts of Wheat Proteins for Patients with teins that contain celiac disease-activating epitopes. These
Wheat Allergy approaches have the potential to reduce allergens in cereals.
Hypoallergenic rice has been developed with downregulation
There is an inherent difficulty in determining the minimum
of the gene family encoding a-amylase inhibitors, which are
eliciting doses that will cause an allergic reaction. Individuals
major food allergens in Japan. These approaches reduced the
will differ in their sensitivity to different allergens, and often,
levels of the inhibitors down to about 20% of their levels in
multiple allergens are present; a combination of these may be
control plants. Wheat has been transformed to silence the o5-
required to elicit a response. In children with wheat allergy,
gliadins (active in WDEIA), resulting in complete or substan-
810 g flour has been shown to induce symptoms, whereas in
tially reduced levels relative to untransformed controls, with
another study, it required < 2 g; children generally have a lower
the retention of technological properties. These approaches
tolerance than adults due to their smaller body mass. WDEIA
could be used to reduce allergenicity by introducing stable
appears to be triggered by larger amounts of flour (60 g), which
heritable changes in the cereal genome.
may be related to the allergen being present in small amounts,
but there are few reported studies. There are ethical issues with
feeding cereal flours to individuals with wheat allergy (as you are
inducing a clinical response), but further study is required. Conclusions
related to labeling, greater confidence in the foods that they Further Reading
consume.
Altenbach SB and Allen PV (2011) Transformation of the US bread wheat Butte 86 and
silencing of omega-5 gliadin genes. GM Crops 2: 6673.
Gilissen LJWJ, van der Meer IM, and Smulders MJM (2014) Reducing the incidence of
allergy and intolerance to cereals. Journal of Cereal Science 59: 337353.
Exercises for Revision Hischenhuber C, Crevel R, Jarry B, et al. (2006) Review article: Safe amounts of gluten
for patients with wheat allergy or coeliac disease. Alimentary Pharmacology and
What are the major types of allergy associated with wheat? Therapeutics 23: 559575.
Inomata N (2009) Wheat allergy. Current Opinion in Allergy and Clinical Immunology
Why are patterns of cereal allergy changing worldwide?
9: 238243.
What are the major classes of proteins in cereal grains?
Matsuo H, Kohno K, Niihara H, and Morita E (2005) Specific IgE determination to
What are the main types of wheat allergy and their causative epitope peptides of omega-5 gliadin and high molecular weight glutenin subunit is
allergens? a useful tool for diagnosis of wheat-dependent exercise-induced anaphylaxis.
Journal of Immunology 175: 81168122.
Pastorello EA, Farioli L, Pravettoni V, et al. (2009) Maize food allergy: Lipid transfer
proteins, endochitinases, and alpha zein precursor are relevant maize allergens in
double-blind placebo-controlled maize-challenge-positive patients. Analytical and
Exercises for Readers to Explore the Topic Further
Bioanalytical Chemistry 395: 93102.
Salcedo G, Quirce S, and Diaz-Perales A (2011) Wheat allergens associated with bakers
Which groups of proteins account for cross-reactivity asthma. Journal of Investigational Allergology and Clinical Immunology 21: 8192.
between botanical species? Sheffer AL and Austen FA (1980) Exercise-induced anaphylaxis (1980). Journal of
Allergy and Clinical Immunology 66: 106111.
What are the potential uses of GM technology in reducing
Tada Y, Nakase M, Adachi T, et al. (1996) Reduction of 1416 kDa allergenic proteins in
cereal (and general) food allergies? transgenic rice by antisense gene. FEBS Letters 391: 341345.
What are pseudocereals? How can they contribute to the Tatham AS and Shewry PR (2008) Allergens in wheat and related cereals. Clinical and
diet of cereal allergy and intolerance sufferers? Experimental Allergy 38: 17121726.
Nutrition: Beriberi, A Deficiency Related to Grains
KJ Carpenter, University of California, Berkeley, CA, USA
2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition, volume 2, pp. 318325, 2004, Elsevier Ltd.
Topic Highlights struggling to understand this disease that led to the discovery
of the class of micronutrients that now is known as vitamins.
Beriberi, a name meaning great weakness is due to B-vitamin It is a complicated story with many false leads, but it pro-
(thiamin) deficiency, commonly associated with milled rice vides a valuable lesson in how science does eventually advance,
consumption. and the dangers of over-simplification of the problems of the
White rice alone, absent a diverse diet providing varied real world. It also shows how it finally proved possible, as a
sources of vitamins, is severely deficient in thiamin. result of collaboration between industry and academia, to
There is no easy, convenient food source that provides high enrich highly milled grain products so that they still pro-
thiamin. Parboiling of rice is beneficial, as is enriching rice vided a good contribution of vitamins and minerals.
with synthetic vitamins.
Occurrence in Asia
Learning Objectives The first Western physicians allowed to work in Japan in the
1870s were surprised to discover the existence of a serious
To understand the etiology of beriberi and the history of its disease previously unknown to them and second only to
dicovery. smallpox in its ravages. In Japan, it was known as kakke,
To understand the effect of grain processing on thiamin but was soon recognized as being identical to the disease
levels and hence on propensity to induce beriberi. known in South-east Asia as beriberi, a native name now
To understand the effects of other dietary habits, e.g. exces- universally adopted, which may originally have meant great
sive alcohol consumption, on thiamin deficiency. weakness. Characteristically, it began with a feeling of weak-
ness in the legs and a loss of feeling in the feet. Then, in many
but not all cases, the legs and then the trunk would swell with
retained water. Finally, the heart would be affected so that the
Introduction
subject gasped for breath, and would die from heart failure.
Older records from both Japan and China showed that it
Grains have been the major source of both energy and protein
had been known for some centuries, although it had been the
for the worlds population for millenia. However, it is now
opinion of two eighteenth century Japanese physicians that the
realized that, when the diet lacks a variety of other foods, the
disease had become worse after 1750. The early records also
grains are also needed to supply a considerable number of
indicated that it was largely a disease of the wet summer
other essential micronutrients if those eating them are to
months and could attack even the well-off.
remain healthy. But the processing of grains to remove the
bran and germ, in addition to its favorable effects of increasing
shelf life by removing much of the oils that can become rancid, Infection or Malnutrition?
and providing foods with a smoother feel by removing most of
the fiber, does also cause the loss of a large proportion of their Those most at risk were men in the newly modernized Japanese
other nutrients. army and navy, and also prisoners. As these were all people
It now appears that the emergence of the disease pellagra living together in large groups, and with the excitement in this
amongst communities in the southern states of the USA from period at other diseases being traced to the transmission of
around 1910 is explained by the introduction of machinery pathogenic bacteria, this seemed a likely cause for beriberi also.
that allowed the replacement of stone-ground whole corn Yet, it was difficult, on this basis, to explain the frequent
meal with de-germed meal, and that this, combined with observation that a naval ship would leave its base with all its
white wheaten flour and little in the way of lean meat, eggs, crew in good health, yet, after a month or more in isolation at
and milk, resulted in a diet deficient in the vitamin niacin. sea, the disease would sweep through the crew.
Until this came to be understood in the 1930s, several thou- Kanehiro Takaki, a surgeon on the naval staff, was directed
sand people died and even more suffered a miserable and in 1878 to work on the problem. He knew that the ships had
humiliating disease. been built in Britain and that they followed the general prac-
The following article summarizes the way in which another tices of the British navy where there was no beriberi. The only
disease, beriberi, that was causing huge suffering in the Far difference that caught his attention was in the rations issued to
East, particularly in the earlier period 18701910, came to be the men: the Japanese issues contained less protein and did not
understood as the consequence in most cases of people having meet the high standard in force at that time in Europe. He
polished white rice as their staple food with little in the way of therefore persuaded his superiors to permit a trial of modified
supplementary foods. It was the work of the pioneers rations with a proportion of the rice being replaced by meat,
condensed milk, vegetables, and barley. The change was a A long series of feeding trials confirmed that birds fed on
complete success, and it was found that even just the use of white rice would become sick with leg weakness, whereas those
barley in place of one half of the rice staple was enough to given supplements of rice polishings (still present in feed-grade
prevent the disease, which Takaki now believed to have rice) remained healthy. This was only an animal disease, but a
resulted from a deficiency of protein in the earlier rations. survey by the medical inspector of prisons in Java showed that
The Japanese army, perhaps as the result of interservice prisoners who had been receiving white rice as their staple
rivalry, did not follow the navy and in the short Russo-Japanese issue were susceptible to beriberi, whereas those receiving
war of 190405, some 100 000 of their soldiers had to be brown rice were not.
invalided home from Manchuria suffering from beriberi.
Isolation of Thiamin
The Analysis of Foods There is no evidence that cooked white rice has any posi-
tively harmful qualities, but if it is the major item in a diet that
Thiamin can be oxidized to a highly fluorescent derivative, contains only small amounts of foods that are richer in thia-
thiochrome. This property is used to measure the thiamin min, so that the diet as a whole provides no more than about
contents of different foods, even at levels of less than 1 ppm. 0.25 mg per 1000 kcal, it is not surprising that beriberi should
The procedure is specific, and no other naturally occurring gradually develop.
compounds have been found that give the thiochrome reaction. The data in Figure 2 also explain the Japanese experience
However, thiamin can react with polyphenols and a compound that serious problems with beriberi in their navy in the late
present in garlic to give derivatives that are still biologically 1800s disappeared when one-half of their rice ration was
active (as will be referred to again) but do not give the thio- replaced by barley.
chrome reaction. The analytical procedure may therefore under- The same figure also shows the low thiamin content of
estimate the efficacy of a product. It is essential, therefore, to tapioca prepared from cassava roots. This explains the exis-
have a confirmatory bioassay before knowing for certain that tence of beriberi in Brazil at the same period among even
any kind of processing has caused significant loss of thiamin. well-off people whose favorite foods were tapioca and molas-
ses. Their preferred protein supplement was dried, salted cod,
which had to be soaked for several days to leach out most of
the salt, which also removed most of the vitamin.
Rice and Other Staples The very first reports of beriberi to reach Europe came from
Portuguese priests working in the Molucca Islands (at the
Figure 2 illustrates the thiamin levels in the worlds major Eastern end of Indonesia) in the 1500s. Their staple was the
staple foods, both when fully milled and when minimally locally produced sago meal, now realized to be almost pure
processed. In the case of grains, the latter means removal of starch, and the priests correctly attributed their weakness to a
the husk (or hull) but no more. It is clear that, for each grain, the lack of something in this food and asked to be provided by
full milling that removes both the bran and germ results in the their superiors with wheat flour.
loss of a major portion of the thiamin originally present. Beriberi was also a serious problem in early spring in iso-
White rice is not that much lower than white wheat flour in lated communities in Newfoundland in the early years of the
its content of the vitamin, but after the grains have been twentieth century. Their families, who would be cut off for
prepared for consumption, the difference is increased. White the winter, had to buy 6 months of supplies, with white flour
rice is normally washed several times, and this alone can as their staple. They were also apparently not familiar with
remove half the thiamin present, and boiling in excess water using yeast to leaven bread, but cooked their flour with baking
can again halve the level of remaining vitamin. In contrast, soda, and it is known that much of the thiamin present
white wheat flour is most commonly baked into bread with would be destroyed under the alkaline conditions during this
yeast as the raising agent, and this causes little loss of thiamin. procedure.
Figure 2 Representative analytical values for the thiamin content of different staple foods: (1) after husking only ( ) and (2) after full processing
( ), as explained in the text, and also for some staple root crops, etc. Sago meal is not shown, as it contains only an insignificant level of thiamin.
(Reproduced with permission from Carpenter KJ (2000) Beriberi, White Rice and Vitamin B. Berkeley, CA: University of California Press.)
80 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Nutrition: Beriberi, A Deficiency Related to Grains
The Improvement of Rice it with certain varieties of rice, and not with the bulk of the rice
favored in Japan.
Once the association of beriberi with white rice in Asia had A traditional method of processing rice common in Bengal
been established, attempts were made to replace it in some way is called parboiling. It had been found that if rice in the husk
with other foods. As already mentioned, barley was an eco- were to be steeped for a period in hot water and then allowed
nomic and well-accepted alternative in the diet of the Japanese to dry in the sun, the husks cracked off more easily on pound-
armed forces. ing, and there was less breakage of the grains. Broken grains
In the Philippines, a proposal was made to enforce the use had a lower commercial value.
of brown rice by the local military. This, of course, is rice from In Malaysia, where many immigrant groups were employed
which the husk has been removed but not the entire bran layer as laborers but living on their habitual diets, it was realized in
and germ (embryo and scutellum) (Figure 3). This was the 1910 that Bengalis were remarkably free from beriberi, and
traditional staple of villagers in South-east Asia who had no studies in a mental hospital showed that replacing ordinary
access to mechanical rice mills. They would pound their paddy white rice with polished rice prepared from parboiled grains
(i.e., rice still in the husk) in some kind of bowl and then relieved the inmates from the disease. After thiamin had
winnow the product so that the lighter husks blew away, and been identified, analyses showed that the initial soaking of
the grains fell in a pile. the grain caused thiamin to diffuse into the endosperm and
This procedure was time-consuming but created no prob- to remain there as the grains dried out.
lems when only enough was pounded for immediate use in the Again, this does not appear to be something that could be
next 24 h. However, it was repeatedly found that in the tropics, applied more widely. The traditional soaking and sun-drying
brown rice on storage would become infested with insects of leaves the rice with a characteristic musty flavor and slight
different kinds, and the oil in the bruised germ would become discoloration, which is acceptable only to people who have
rancid. Since large organizations, or an army on the move, grown up with it. The process can be modernized, with con-
needed large-scale supplies ready for cooking, brown rice did trolled autoclaving and vacuum-drying for so-called conver-
not provide a practicable staple. sion of the rice, but this makes it too expensive for mass
The early workers who discovered the association of beri- consumption in developing countries.
beri with white rice had assumed that the important micronu- The final procedure for the production of enriched rice is
trient was concentrated in the bran of the grain. However, it to fortify it with the synthetic vitamin. For a flour, such as white
was later realized that more was present in the germ area wheat flour, this is relatively simple, requiring only very careful
(Table 1). Japanese millers then attempted to modify their mixing of the traces of vitamin with a small quantity of flour,
machinery so as to remove the bran without removing the then the blending of the premix to larger batches. This is
germ from the grain. The so-called germ rice that they were familiar, and indeed compulsory practice, in the USA and
able to produce proved to be both palatable and an improved UK, together with other vitamins and trace minerals. As a
source of thiamin. However, millers were only able to produce powder, thiamin cannot be blended with grains of white rice.
However, methods have been developed of preparing vitamin-
rich granules with the size and appearance of rice grains, and
blending these in the ratio of one granule to 200 grains, so that
the mix has at least the thiamin content of brown rice. To
reduce loss of the vitamin during washing and cooking, the
granules are coated with a nontoxic resin at the final stage of
their production. This method of enrichment has been tested
in an area of the Philippines where beriberi was endemic and
has proved successful in greatly reducing its incidence. Unfor-
tunately, where widely separated villages each had a small
electric mill, there were practical problems in persuading
millers to pay for the premix when the product appeared
unchanged, but the price had to be a little higher.
Supplementing Foods
Table 1 Thiamin contributed by the different parts of a sample of dehusked rice grains
Thiamin
Dissected parts of rice grain Proportion by weight (%) Concentration in fraction (mg per 100 g) Contribution to 100 g grain (mg)
Reproduced with permission from Carpenter KJ (2000) Beriberi, White Rice and Vitamin B. Berkeley, CA: University of California Press.
Japanese believed that consumptions of most foods other than Heat-Stable Antithiamins?
rice during the summer months caused heat. Hence, food
It has been found that when the thiamin in a food comes into
consumption was almost entirely restricted to rice conse-
contact with polyphenols such as caffeic acid, it no longer gives
quently leading to an increase in beriberi during these months.
a fluorescent product in the usual thiochrome procedure for
the estimation of thiamin. This led some workers to suppose
that drinking large quantities of tea or coffee, or chewing betel
Is there an Antithiamin Problem? nut all sources of polyphenols might induce a condition of
thiamin deficiency. However, biological assays have indicated
There are many references in the literature to at least a suspi- that the vitamin is still fully available.
cion that certain foods and beverages may be responsible for When thiamin is incubated with garlic extracts, it undergoes
beriberi appearing in people whose intake of thiamin would a reaction with the allicin present in which the thiazole ring
otherwise be adequate. opens, and the sulfur atom in the ring links to the alkyl sulfide
to form a disulfide compound. This is not measured in the
thiochrome reaction, but in the body it is reduced to re-form
Thiaminases
the active vitamin. In fact, compounds of this type can be
Many species of fish contain enzymes in their viscera that split absorbed more efficiently by alcohol-damaged intestinal
thiamin molecules at the junction between its two ring struc- walls than ordinary thiamin. Thiamin tetra-hydrofurfuryl
tures. This was discovered when foxes, being reared for their fur disulfide in particular is approved for this purpose in some
and fed on a mix containing a large proportion of whole raw countries.
fish, developed a form of paralysis that responded to injections Although there is no confirmed evidence of naturally occur-
with thiamin. A similar condition was seen later in cats that ring heat-stable compounds that inactivate thiamin, chemists
had been fed on a canned food containing a large proportion have synthesized such materials. One, named oxythiamin,
of whole fish. It was believed that the thiamin in the mix had has the amino group attached to the pyrimidine group in
been largely destroyed after the mix had been prepared and was thiamin replaced by a hydroxy group. Giving it to animals
waiting to be autoclaved. results in the more rapid production of some of the signs of
These experiences led to investigations as to whether thiamin deficiency, although it differs from thiamin in being
humans could be similarly at risk, but it appears not. The unable to pass the bloodbrain barrier.
enzymes are not present in fish muscles (i.e., fillets), and
even where small fish are eaten whole, they are not ground
up with other items of diet before being cooked. Lastly, it was
found in animal studies that a subsequent meal with different Acute Deficiency in the West
constituents was not affected by thiaminases being present in
an earlier meal. With the discovery that autoclaving yeast powder would
Another source of thiaminases was found to be bracken, destroy the thiamin, whereas the other B-vitamins were
and their presence explained the condition known as stag- retained, it was possible to place volunteers on an artificial
gers that occurs in horses that have been feeding on bracken. diet essentially free of thiamin. To the surprise of investigators,
Cooked bracken, in which the thiaminase was inactivated, some subjects had lost appetite within 2 weeks and became
proved harmless to horses. The only known case of thiaminase nauseated and dizzy, with other mental symptoms, by 6 weeks,
poisoning in humans occurred in a group exploring the inte- but with no sign of peripheral nerve damage or cardiac abnor-
rior of Australia in the 1860s. Running out of provisions on mality, which are characteristic of classic beriberi.
their return journey, they lived on the sporocarps in the fronds Trials using pigeons and rats with very deficient diets also
of a particular fern that is now known to contain a high level of produced appetite loss and death before any sign of leg weak-
a particularly heat-resistant thiaminase. All four men devel- ness had developed. It appeared that in both humans and
oped leg weakness and lassitude; three died, and the survivor animals, acute deficiency of thiamin resulted in damage to
remained lame even after his safe return. the central nervous system. With slightly higher intakes, the
82 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Nutrition: Beriberi, A Deficiency Related to Grains
CNS had priority, whereas peripheral nerves gradually Exercises for Revision
degenerated.
In your community, what dietary habits are most likely to
result in insufficient thiamin consumption?
Deficiencies in Total Parenteral Nutrition What are the mandatory (and common voluntary) enrich-
ments of cereal grain products in your country? Do you
There are many reports of people recovering from surgery of think the regulations make sense in relation to current
the gastrointestinal tract who have developed acute thiamin typical dietary patterns?
deficiency. They had been fed intravenously with a solution
providing energy, amino acids, and minerals, but no vitamins.
This is adequate for a short period, but thiamin is the first
vitamin to become depleted.
Exercises for Readers to Explore the Topic Further
In a number of cases where this type of parenteral feeding
has continued for some weeks, a condition called Wernickes What methods are used to measure thiamin content? How
stable is thiamin after typical food processing operations?
encephalopathy has developed. Patients are confused and
have characteristic involuntary eye movements. Where patients Are there any potential genetic solutions to low thiamin
content in milled rice?
have died, autopsies have shown brain lesions analogous to
those found in acutely deficient animals. The same outcome
has been seen in subjects voluntarily fasting for long periods or See also: Food Grains and the Consumer: Consumer Trends in
being unable to take food because of persistent vomiting in Grain Consumption; Cultural Differences in Processing and
pregnancy. Consumption; Grain Composition and Analysis: The Composition
of Food Grains and Grain-Based Products; Non-wheat Foods: Rice:
Chinese Food Uses; The Basics: Grain and Plant Morphology of
Alcoholism Cereals and how characters can be used to identify varieties; Grain:
Morphology of Internal Structure.
One material that can be responsible for the production of
thiamin deficiency is ethanol (i.e., alcohol in everyday
speech). In developed countries where nearly everyone can
afford a well-balanced diet, most of those diagnosed as being Further Reading
thiamin-deficient are alcoholics. The continued ingestion of
high levels of alcoholic beverages has many undesirable effects. Carpenter KJ (2000) Beriberi, White Rice and Vitamin B. Berkeley, CA: University of
In the present context, two are relevant. First, the alcoholic California Press.
Cook CC, Hallwood PM, and Thomson AD (1998) B vitamin deficiency and
typically no longer bothers to eat a normal range of foods, neuropsychiatric syndromes in alcohol misuse. Alcohol and Alcoholism
partly because the beverages provide a large portion of their 33: 317336.
calorie needs and partly from nothing but their next drink Eijkman C (1929) Antineuritic vitamin and beriberi. In: Nobel Lectures: Physiology or
being of immediate interest. Second, the high level of alcohol Medicine, pp. 19221941. Amsterdam: Elsevier.
Hinton JJC (1948) The distribution of vitamin B1 in the rice grain. British Journal of
ingestion damages the intestinal wall so that thiamin is
Nutrition 2: 237241.
absorbed less efficiently, and the requirement for the vitamin Jansen BCP and Donath WF (1926) On the isolation of the anti-beriberi vitamin.
increases. Koninklijke Akademie von Wetenschappen, Amsterdam, Proceedings
Unfortunately, a small proportion of such victims develop 29: 13901400.
Wernickes encephalopathy, which may lead in turn to Sauberlich HE, Herman YF, Stevens CO, and Herman RH (1979) The thiamin
requirement of the adult human. American Journal of Clinical Nutrition
Korsakoffs psychosis. Such people, sometimes referred to as 32: 22372248.
suffering from the WernickeKorsakoff syndrome, are at pre- Shimazono N and Katsura E (1965) Review of Japanese Literature on Beriberi and
sent incurable and have to be maintained in a mental hospital Thiamine. Kyoto: Vitamin B Research Committee of Japan.
for the rest of their life. The cost of this to the state is such that Takaki K (1906) The preservation of health amongst the personnel of the Japanese army
and navy. Lancet i: 13691374.
some specialists have argued that it would actually be cheaper
Truswell AS and Apeagyei F (1982) Alcohol and cerebral thiamin deficiency.
to have all beer and wine fortified with thiamin as a preventive. In: Jelliffe EF and Jelliffe DB (eds.) Adverse Effects of Foods, pp. 253258.
There may be a genetic factor making some Western people New York: Plenum Press.
susceptible to the cerebral form of beriberi and the Wernick- Williams RR (1961) Toward the Conquest of Beriberi. Cambridge, MA: Harvard
eKorsakoff syndrome, since it was seen even in Western pris- University Press.
Williams RR and Cline JK (1936) Synthesis of vitamin B1. Journal of the American
oners of the Japanese in the Second World War who had white Chemical Society 58: 15041505.
rice but no alcohol, but apparently has not been seen in Asian Wuest HM (1962) The history of thiamine. Annals of the New York Academy of Sciences
subjects. 98: 385400.
Celiac Disease
P Koehler, H Wieser, and KA Scherf, Deutsche Forschungsanstalt fur Lebensmittelchemie, Leibniz Institut, Freising, Germany
2016 Elsevier Ltd. All rights reserved.
Definition
Symptoms and Pathology
CD, also known as celiac sprue or gluten-sensitive enteropathy,
may be defined as an inflammatory disease of the upper small A range of symptoms may be associated with CD, and these
intestine (duodenum and jejunum) in genetically susceptible can be divided into intestinal disorders and extraintestinal
individuals. CD is a permanent food intolerance induced by features caused by malabsorption of essential nutrients such
ingestion of storage proteins (gluten) from certain cereals as deficiencies of vitamins and minerals. In infants, classical
(wheat, rye, barley, and oats) that cause damage to the enter- symptoms appear after weaning and introduction of cereals
ocytes and, as a consequence, malabsorption of important into the diet: diarrhea, abdominal distension, failure to thrive,
nutrients. CD patients develop a flat jejunal mucosa with the vomiting, muscle wasting, and apathy. Older children tend to
have more varied symptoms. Besides diarrhea or constipation, CD is associated with histocompatibility complex class II
features such as anemia, loss of appetite, and short stature may alleles HLA-DQ2 and HLA-DQ8. Almost all CD patients
be predominant. Diarrhea is the main presenting feature of ( 97%) possess one of these HLA genes. The large majority
adults; in some patients, anemia, osteoporosis, abdominal of patients ( 9095%) are DQ2-positive, and the remainder
pain, loss of weight, and weakness may be found. A minor are DQ8-positive. Two isoforms DQ2.5 (predominant) and
number of patients present psychological or psychiatric symp- DQ2.2 (rare) were identified among DQ2 alleles. HLA-DQ
toms such as depression, epilepsy, autism, or schizophrenia. genes encode protein heterodimers with a- and b-chains,
Frequently, the disease may be clinically silent or masked by both responsible for the CD-specific binding of gluten peptides
associated diseases like type 1 diabetes, although the patients by intestinal antigen-presenting cells. The HLA-DQ2.5 geno-
exhibit the full mucosal lesion. They are discovered only when type is associated with a very high risk for developing CD,
intestinal studies are undertaken. Around 25% of CD patients followed by DQ8 (high) and DQ2.2 (low). However, about
develop a resistance to a strict GFD. This condition, called 30% of the healthy population carry DQ2 and will never
refractory celiac disease (RCD), is characterized by a constant develop CD. Thus, HLA-DQ genes alone do not explain the
inflammatory process in the intestinal mucosa accompanied development of CD, and additional factors such as non-HLA
by persistent diarrhea, abdominal pain, and weight loss. RCD genes and environmental factors need to be involved in disease
carries a significant burden of morbidity and mortality, mainly development. Further, CD-specific genome research, mainly
from malnutrition and lymphoma. focused on non-HLA genes, indicated little consensus, and
The small intestinal lesion of CD has a highly characteristic the contribution of all non-HLA genes, known up to now,
morphology, but it may vary from patient to patient depending has been estimated to account for less than 10%.
on the severity and extent of disease. Patients show a wide Different environmental factors have been considered to be
variation of mucosa appearance from a complete flat to a important for the development of CD. For example, infections
convoluted pattern. The parameters usually assessed for the by a variety of pathogens, an imbalanced intestinal microbiota,
measurement of intestinal mucosa are villous height, villous the amount and timing of gluten exposure, and breastfeeding
width, villous height to crypt depth ratio, mucosal thickness, during first gluten intake have been associated with developing
epithelial cell height, crypt mitotic activity, and number of CD in infants.
IELs. Electron microscopy demonstrates fewer microvillous
intramembrane particles and abnormal tight junctions (TJs)
between epithelial cells. In untreated CD, serological anti- Pathogenesis
bodies to gliadin, deamidated gliadin peptides, reticulin, endo-
mysium, and tissue transglutaminase (TG2) are present. The development of CD is induced by the oral intake of gluten
present in wheat, rye, barley, and oat products. A daily intake
of more than 20 mg of gluten is estimated to be not tolerable
for CD patients. This approximately correspond to one-
Epidemiology and Genetics hundredth of a slice of bread. In contrast to most other food
proteins, gluten proteins are highly resistant to complete pro-
CD is prevalent mostly in Europe and those countries to which teolytic digestion within the gastrointestinal tract due to their
Europeans have emigrated including North and South America high proline content. Thus, relatively large protein fragments
and Australia. However, CD is also known to occur in many (peptides) accumulate in the small intestine and reach the
other parts of the world. In the past, CD was considered a intestinal brush border (step 1 in Figure 1). However, the
comparatively uncommon disorder with prevalence rates of 1 poor digestibility of gluten proteins is not sufficient to cause
in 2000 to 1 in 1000. Several recent studies using serological CD, and there is no known difference between healthy indi-
screening followed by small intestinal biopsy have shown a viduals and CD patients in their (in)ability to digest these
much higher prevalence, and it is now estimated that CD may proteins. Changes in the mucosal permeability for gluten pep-
affect as many as 1 in 300 to 1 in 100 individuals. The iceberg is tides are supposed to be the first pathogenic event in the
a common model to describe the epidemiology of CD. The tip development of CD. Gluten peptides are bound to the intesti-
of the iceberg is formed by patients who have been diagnosed nal mucosa before passing through the enterocyte layer either
by conducting a biopsy demonstrating a flat mucosa. Below transcellularly or paracellularly (Figure 1, step 2). The most
the waterline, there is a big group of undiagnosed patients with important structures of the paracellular route are the epithelial
a flat mucosa, but with no or weak symptoms (undiagnosed pore-forming TJs. They are composed of a number of proteins
silent CD). At the bottom of the iceberg, there is a small section (e.g., zonulins, claudins, and occludin) that regulate paracel-
of patients with a normal mucosa but with the genetic predis- lular permeability. Zonulin 1, known to enhance permeability
position and increased levels of celiac-specific antibodies. and to be upregulated during the acute phase of CD, has been
These subjects may develop clinically overt CD later in life proposed to be one of the initial factors of the pathogenesis of
(undiagnosed potential CD). CD. When crossing the epithelium via the transcellular path-
The occurrence of CD among first-degree relatives of CD way, gluten peptides are differently degraded in the intracellu-
patients is fairly strong with a prevalence of 1015%. The lar compartments of the enterocytes of CD patients. The
concordance between dizygotic twins is reported to be in a transcellular permeability is much higher for intact CD-active
range of 1120% and that between monozygotic twins peptides compared with nonactive peptides.
7580%. CD can occur at any age, but in adults, the peak After passing through the epithelium, CD-active peptides
incidence is in the fifth decade. Females are more commonly arrive at the lamina propria and induce two different immune
affected than males with a suggested ratio of around 2:1. responses: the adaptive and the innate responses. Within the
FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Celiac Disease 85
I
1
E E 2 E E
II
IFN
5 TNF
T
MMP A B
III P T IgA
C B P
T IgG
4 B 6
T
B
Th 1
Th 2
adaptive mechanism, gluten peptides are specifically modified common to both conditions. It appears as patches of itchy
by TG2 (Figure 1, step 3) either by partial deamidation of red papules and blisters on the extensor surface and pressure
glutamine residues leading to negatively charged glutamic acid areas; healing up results in pigmentation. Histologically, there
residues or by cross-linking to TG2 by isopeptide bonds between is infiltration of the dermal papillae with inflammatory cells.
glutamine residues of the peptides and lysine residues of TG2. The dermal histology is similar to that of the small intestine in
Particularly, deamidated peptides are selectively bound to DQ2/ CD, but the abnormalities tend to be milder and patchier. In
DQ8 heterodimers on the surface of antigen-presenting cells many cases, a GFD results in a significant improvement.
(dendritic cells, macrophages, and B cells) and then presented Gluten ataxia is one of the neurological manifestations
to the T cell receptor of gluten-sensitive CD4 T cells. The stim- attributed to gluten intake. All these patients present gait
ulation of T cells leads to the secretion of proinflammatory T ataxia, and the majority have limb ataxia and evidence of
helper (Th-1) cytokines (Figure 1, step 5), in particular cerebellar atrophy. The current diagnostic recommendation
interferon-g, tumor necrosis factor, matrix metalloproteinase, is a serological screening for antigliadin, anti-TG2, and anti-
and nitric oxide, which lead to premature senescence of small TG6 antibodies. Patients positive for these antibodies should
intestinal enterocytes, shedding of these cells, and, in turn, the follow a GFD.
observed small intestinal enteropathy. Simultaneously, T cells Wheat-dependent exercise-induced anaphylaxis (WDEIA) is
provide help for the production of IgA and IgG antibodies to a severe IgE-mediated allergic reaction provoked by the com-
gluten proteins and autoantibodies to TG2 (Figure 1, step 6). bination of wheat ingestion with intense physical exercise over
In contrast, the rapid innate immune system is characterized the next few hours. Aspirin and ethanol, together with wheat-
by the massive increase in IELs, one of the hallmarks of CD, which containing food, are further cofactors for WDEIA. Peptides
can be observed very early before the onset of villous atrophy. Two derived from o5-gliadins and high-molecular-weight (HMW)-
subsets of IELs bearing the abT cell receptor or the gdT cell glutenin subunits have been identified to be critical for IgE
receptor are linked to CD innate immunity. Interleukin-15 binding.
(IL-15) has been considered a central player in this part of the Non-celiac gluten sensitivity (NCGS) is a gluten-mediated
gluten-induced immune response. IL-15 is produced by epithelial disorder characterized by gastrointestinal complaints including
and lamina propria cells in active CD, but not by T and B cells bloating, diarrhea, weight loss, and abdominal pain. The small
involved in the adaptive immune response. Recently, it has been intestine of patients is usually normal, and serum antibodies
suggested that amylase-trypsin inhibitors (ATIs) present in cereal typical for CD are absent. ATI or fermentable oligo-, di-, and
flours are coactivators of the innate immune system. How the monosaccharides and polyols may be implicated in NCGS, but
cross talk between innate and adaptive immunities is orchestrated to date, there is too little knowledge on responsible triggers of
is an outstanding question that will need to be addressed. this currently underrecognized and undertreated disorder.
Recent screening studies have shown an increased preva-
lence of CD in autoimmune disorders such as type 1 diabetes,
Associated Diseases thyroid disease, primary biliary cirrhosis, and Sjogrens syn-
drome. Moreover, IgA deficiency, chronic fibrosin alveolitis,
Dermatitis herpetiformis, a skin disorder, is associated with other interstitial lung diseases, and concomitant distal ulcera-
CD, because some degree of gluten-sensitive enteropathy is tive colitis have been reported in association with CD. In other
86 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Celiac Disease
instances, there are unexpected associations such as epilepsy wheat, rye, and barley. Starches of these cereals are permitted,
and various undefined neurological disorders. A small number if the gluten level is lower than 20 mg kg1 (Draft Revised
of patients with CD have exhibited symptoms of various psy- Codex Standard 2008). The toxicity of oats has been discussed
chiatric disorders, such as schizophrenia. A lymphoma of the controversially, but recent studies have provided strong evi-
small intestine is the classical malignancy associated with CD. dence that only a very small subgroup of CD patients cannot
The prevalence of carcinoma of the small intestine, esophagus, tolerate oats. The daily intake of gluten should not exceed
or pharynx, crypt hyperplasia, villous atrophy, and mucosal 20 mg. Since small amounts of gluten are hidden in many
hypertrophy also increases with CD. It has been reported that a foods, dietary counseling is absolutely necessary. In most
strict GFD helps safeguard against these malignancies. countries, celiac societies are an invaluable help in providing
dietary guidelines and food lists. The prognosis for patients
who are correctly treated is excellent, whereas failure to imple-
Diagnosis ment a strict diet may result in continuing symptoms and in
the two major complications: osteoporosis and malignancy.
The diagnosis of CD is complex and tends to require a high CD patients may consume gluten-free foods from two differ-
level of clinical suspicion. The diagnostic approaches have ent categories. Firstly, they are allowed to eat a wide range of
changed during the last decades, and recommendations for common products such as meat, fish, milk, fruits, and vegeta-
the procedures are still different. In summary, the diagnostic bles. In the case of composite foods, however, it is difficult to
scheme may consist of the following points: (1) clinical history recognize whether they are gluten-free or not. Labeling of gluten
and symptomatology, (2) serology, (3) small intestinal as an allergen is mandatory in the framework of CODEX Ali-
histology, (4) response to a GFD, and (5) HLA status. Classical mentarius and in the European Union. Secondly, patients con-
intestinal symptoms should prompt the primary doctor to sume dietetic food that is gluten-free according to the CODEX
initiate a serological test. Extraintestinal symptoms are more Alimentarius standard. The current standard and legislation in
difficult to interpret, and medical specialists (e.g., dermatolo- the United States, Canada, and the European Union set a gluten
gists, dentists, and neurologists) should be aware of possible level of 20 mg kg1 food as the threshold for a gluten-free claim.
CD. For initial testing, a quantitative test detecting IgA class In some countries, foods containing up to 100 mg gluten/kg,
antiendomysium, anti-TG2, and antideamidated gliadin pep- such as wheat starch, can be labeled as low in gluten. For gluten
tide antibodies from a blood sample is recommended. To analysis, extraction with 60% ethanol and an immunochemical
achieve the clinical demand for both maximal sensitivity and method for quantitative determination (enzyme-linked immu-
specificity, the combination of different tests is recommended. nosorbent assay (ELISA) with monoclonal or polyclonal anti-
In case of IgA deficiency, IgG class antibodies have to be used. bodies) are recommended. Dietetic gluten-free foods are mostly
The histological judgment of small intestinal mucosa is substitutes of products usually containing wheat, rye, and bar-
commonly regarded as the gold standard for the reliable diag- ley, such as bread, other baked products, pasta, and beer. Com-
nosis of CD. Biopsy specimens of active CD demonstrate a flat mon raw materials used for these products are flours from corn,
mucosa with the absence of intestinal villi, a cellular infiltrate sorghum, rice, buckwheat, quinoa, teff, and chestnut, and com-
of the lamina propria, and an increase in the number of IELs. mon thickening agents are locust bean gum, guaran gum, and
In the ESPGHAN guidelines revised in 2012, the omission of hydroxypropyl methyl cellulose.
the small intestinal biopsy is only tolerated in children and Owing to the enormous increase of knowledge about the
adolescents with distinct signs and symptoms suggestive for pathomechanism of CD, a number of novel strategies for the
CD and very high anti-TG2 antibody titers with levels exceed- prevention and treatment of CD have been developed. Usually,
ing ten times the upper limit of normal. CD-toxic peptides derived from digested gluten survive the
Genetic tests based on HLA-DQ2 and DQ8 alleles cannot normal digestive process. Supplementation with additional
be used to confirm CD, because they are common in the peptidases may degrade CD-toxic epitopes to avert an immune
general population (around 30%). In the case of doubtful response. Peptidases from bacteria, fungi, and germinating
diagnostic results, however, HLA-DQ genotyping can be used cereals have been proposed for oral therapy. Some of these
to rule out the existence of CD, because of its high negative approaches are already in phase 1 and 2 clinical trials. More-
predictive value. If a patient is negative for HLA-DQ2/8, CD over, inhibitors of zonulin, a key molecule for intestinal
can be excluded lifelong and no further investigations and permeability; inhibitors of TG2, the autoimmunogen of CD;
follow-up are needed. The final confirmation of CD is estab- inhibition of gluten peptide presentation by HLA-DQ2
lished by a clinical, serological, and histological response to a antagonists; and modulation or inhibition of proinflammatory
GFD. When keeping a strict GFD, patients show a rapid cytokines have been proposed as alternative therapies. How-
improvement of symptoms within weeks. Histological regen- ever, the risks, benefits, and costs of alternatives have to be
eration is slower and complete recovery can take months or carefully weighed, and the conditions and indications under
years. which such alternative therapies might be warranted have to be
accurately defined.
Therapy
Toxicity Testing
After the diagnosis of CD has been established, permanent
withdrawal of gluten from the diet is the current essential Numerous in vivo and in vitro methods have been used to
treatment. The GFD involves avoiding products containing identify CD toxicity and immunogenicity of different cereal
FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Celiac Disease 87
and noncereal materials, proteins, and peptides. The tests can Toxicity of Cereals
be differentiated into (1) in vivo challenge of patients, (2)
in vitro tests with tissues and cells of patients, and (3) animal The relationship between CD and the ingestion of wheat flour
models. Most investigators would agree that in vivo testing is was established in 1950. Soon after, a series of investigations
the gold standard for assessing CD toxicity of proteins or led to the conclusion that rye and barley were also harmful,
peptides. Early workers established toxicity in a series of feed- whereas corn, rice, buckwheat, and potatoes were not. There
ing tests based on the production of symptoms such as steat- has been disagreement about the toxicity of oats; the reason for
orrhea or on tests such as malabsorption of fat or xylose. that could have been that commercial oat flour is frequently
However, an important impediment was that CD patients contaminated with small amounts of wheat, rye, or barley.
differed widely in their sensitivity to gluten and, consequently, Most clinical studies indicate that pure oats do not activate
the optimal amount of gluten equivalent used to challenge CD and it is proposed that only a very small subgroup of CD
patients and the duration of challenge were uncertain. In any patients is sensitive to oats. Accordingly, CD is closely related
case, 10100 g of gluten equivalent was necessary for the test- to the taxonomy of cereals: only species found in the tribe
ing of each patient, and such large amounts were the most Triticeae within the grass family Poaceae are likely to trigger
crucial limiting factor for feeding tests of purified proteins or CD. All wheat species besides bread wheat such as durum
peptides. Direct instillation into the small intestine followed wheat, spelt, emmer, einkorn, and kamut have been suggested
by biopsy allows the reduction of required amounts to 1 g to be toxic for CD patients due to homology of their storage
equivalent of gluten. Biopsies are taken from patients in remis- proteins to wheat.
sion at the beginning and after several hours of testing.
Changes in enterocyte height, ratio of villous height to crypt
depth, and the number of IELs are considered to be reliable
parameters for toxicity assessment. Toxicity of Cereal Proteins
Because in vivo tests require large quantities of substances
and only a limited number of test patients are available, a series Immediately after wheat was established as the CD-activating
of in vitro tests has been developed. The organ culture of cereal, fractionation of the flour and in vivo testing led to the
human small intestine, which requires only milligram equiva- conclusion that gluten is toxic, whereas starch and albumins
lents of gluten, provides the most reliable in vitro approach. are not. Gluten is the rubbery protein mass that remains when
The intestinal tissue from patients with active CD is removed as wheat dough is washed with water to remove starch granules
part of the diagnostic procedure and incubated in a culture and other soluble constituents. Wheat gluten proteins can be
medium. The tissue shows improvement of enzyme activity divided according to their solubility in aqueous alcohols into
and morphology in the medium alone, but not in the presence the soluble gliadins and the insoluble glutenins. Toxicity tests
of CD-toxic substances. Apart from human material, tissue indicated that the gliadin fraction was the most toxic factor,
cultures with fetal rat and chicken intestine have been used. T whereas the effect of the glutenin fraction was described either
cell lines and clones raised against gluten are useful for per- nontoxic, weakly toxic, or as toxic as gliadins, but on very
forming a large number of experiments and readily demon- inadequate evidence. More recent studies on pure glutenin
strate the T cell response to stimuli. Gluten-sensitive T cells can subunits indicated that the glutenin fraction is also harmful
be isolated from intestinal biopsies cultured and stimulated for CD patients.
in vitro or can be found in the blood of treated CD patients after Flour protein fractions of rye and barley have not been
oral gluten challenge. A frequently used test is the T cell prolif- tested for celiac toxicity until now, but equivalent to the gliadin
eration assay performed by incubation of intestinal CD- fraction of wheat, the corresponding prolamin fractions of rye
sensitive T cells, the putative antigen ( 100200 mg ml1) (secalins) and barley (hordeins) are associated with CD.
treated with TG2, antigen-presenting cells (e.g., B cells), and Although the prolamin fractions of cereals are crude mixtures
tritiated thymidine. The proliferation of T cells determined by of different proteins, their amino acid compositions (Table 1)
scintillation measurements is a parameter for immunogenic show a close relationship to both taxonomy and CD toxicity.
effects. Additionally, the production of IFN-g or interleukins Toxic gliadin, secalin, and hordein fractions are characterized
can be measured. T cell tests have been used widely to compare by the highest contents of glutamine (3537 mol%) and pro-
the level of immunogenic effects. Generally, it should be men- line (1723 mol%). Both amino acids have been considered to
tioned that T cells do not necessarily reflect the small intestine be important for CD toxicity. The prolamins of rice, millet, and
in vivo, because they lack any connection to other cell types corn are lower in glutamine and proline, but rich in alanine
present in the intestine. In addition, T cells are frequently (914 mol%) and leucine (1219 mol%). Oats are in an inter-
different in their reaction to antigens; application of several mediate position, the glutamine content being similar to the
simultaneous T cell lines from different patients is therefore Triticeae prolamins and the values of proline and leucine in
needed. Moreover, immunogenicity does not always corre- agreement with those of other species.
spond to toxicity demonstrated by in vivo or organ culture Among the prolamin fractions, only wheat prolamin (glia-
tests. In any case, in vivo testing ultimately will be necessary din) has been investigated in detail. The digestion with pepsin
to evaluate conclusions on in vitro testing. Among animals, and trypsin alone or followed by pancreatin resulted in the
there are three models that spontaneously generate gluten- retention of toxicity, and a peptide mixture with a molecular
dependent diarrhea: the Irish setter dog, the rhesus macaque, mass less than 2000 was still toxic. Also, the breakdown of
and the horse. However, the animal model that reproduces all the disulfide bonds by oxidation or heating during the
important aspects of CD still awaits development. baking process did not abolish toxicity. In consequence, the
88 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Celiac Disease
Amino acid Wheat (gliadin) Rye (secalin) Barley (hordein) Oats (avenin) Rice (oryzin) Millet (kafirin) Maize (zein)
Table 2 Amino acid sequences of peptides tested for CD toxicity by instillation and organ culture tests
Table 3 Amino acid sequences of selected gluten peptides would be helpful to establish a more precise epidemiological
stimulatory for intestinal T cells of CD patients picture of CD. Little is known on the complex interplay
between genetic and environmental factors leading to CD
Protein Amino acid sequencea
development, and it will be a major area of research to under-
a2 LQLQPFPQPQLPYPQPQLPYPQPQLPYPQPQPF stand their respective contributions and develop possible pre-
a20 PFRPQQPYPQPQPQ vention strategies. Further studies are required to allow a better
g5 FLQPQQPFPQQPQQPYPQQPQQPFPQ diagnostic differentiation between CD, NCGS, and irritable
g5 QFSQPQQQFPQPQ bowel syndrome, because the symptoms and nonspecific find-
LMW 156 QQQQPPFSQQQQSPFSQQQQ ings may be similar in patients. While the role of gluten pep-
LMW 17 QQPPFSQQQQQPLPQ tides in the adaptive immune response is well characterized,
HMW 2 GQQGYYPTSPQQS much less is known regarding the innate immune response.
HMW 2 QGYYPTSPQQSG
Elucidating the mechanism of the innate immune response
Sec a2 QPFPQPQQPFPQSQ
will contribute to a better understanding of the CD pathome-
Sec a9 PQQPFPQPQQPFPQ
Hor a2 QQFPQPQQPFPQQP chanism. Gluten-sensitive T cell assays are used to compare the
Hor a9 PQQPFPQPQQPFRQ level of immunogenic effects, but immunogenicity measured
Av a9A QYQPYPEQQEPFVQ by T cell tests does not necessarily correspond to toxicity
Av a9B QYQPYPEQQQPFVQ revealed by in vivo or organ culture tests. This is the reason
why it would be of interest to test immunogenic peptides by
a
Glutamine (Q) residues targeted by TG2 are in bold. in vivo assays. The GFD will remain the preferred therapy of CD
Source: Wieser H, Koehler P, and Konitzer K (2014) Celiac Disease and Gluten:
in the future. Regarding alternative therapies, CD patients have
Multidisciplinary Challenges and Opportunities, 1st edn. London, Waltham, San Diego:
expressed the wish to have access to a vaccine against CD or to
Academic Press, with permission.
a pill that would occasionally allow them to eat gluten-
containing food. Such alternative therapies are still in the
wheat gliadins and glutenins, rye secalins, barley hordeins, and
early stages of development, and they will have to undergo
oat avenins. Some authors have subdivided CD-active peptides
safety and toxicology testing before they can enter the market.
into toxic peptides generating an innate immune response
and immunogenic peptides generating an adaptive immune
response. The discrepancy between the numbers of immuno-
Exercises for Revision
genic peptides identified by T cell tests and of toxic peptides
identified by in vivo or organ culture tests is extraordinary: 1014
peptides have been classified as immunogenic, 10 peptides as
What is the primary environmental trigger for CD and
which cereals are harmful?
toxic, and 5 peptides as both toxic and immunogenic. The
question of whether immunogenicity corresponds to toxicity
What other gluten-related disorders are there?
in any case remains to be answered.
What are the steps in the diagnostic scheme to identify CD
and what is the essential treatment once the diagnosis has
In summary, a common feature among toxic/immunogenic
been established?
peptides is the presence of multiple proline and glutamine
residues, which gives rise to four unique structural and func-
What methods can be used to measure CD activity?
tional properties:
What foods are allowed to be consumed by CD patients?
What are the characteristic properties of CD-toxic/
1. These peptides are exceptionally resistant to peptidolysis immunogenic peptides?
by gastric, pancreatic, and intestinal digestive peptidases
because of their high proline content. As a result, a high
intestinal concentration of potentially immunogenic pep- Exercises for Readers to Explore the Topic Further
tides is maintained following a gluten-containing diet.
2. Selected glutamine residues in these gluten peptides are CD patients need to adhere to a strict GFD. Compare the
deamidated or transamidated by TG2 under physiological characteristics of wheat bread to those of gluten-free breads.
conditions, leading to enhanced immunogenicity. The increased knowledge about the pathomechanism of
3. The proline-rich gluten peptides naturally adopt a left- CD has led to the development of novel treatment strate-
handed polyproline II helical conformation, which is the gies, such as oral peptidase therapy. Look at the different
preferred conformation of all bound major histocompati- proposed treatment options and identify which steps in the
bility complex class II ligands. pathomechanism are targeted.
4. Large peptides that contain multiple HLA-DQ-binding epi- Take a look at the database of CD-active gluten peptides of
topes have a greater T cell stimulatory activity than small the Food Allergy Research and Resource Program of the
peptides containing only one epitope. Department of Food Science and Technology, University
of NebraskaLincoln, the United States (www.allergenon-
line.org/celiacbrowse.shtml), and find the CD-active pep-
tides derived from alpha-gliadins.
Future Prospects
A standardization of criteria for prevalence determination and See also: Appendix 1: Nutrient-Composition Tables for Grains and
a differentiation into cases of active, silent, and potential CD for Grain-Based Products; Appendix 2: Lists of Standardized Testing
90 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Celiac Disease
Methods for the Analysis of Grain and Grain-Based Foods; Food Dicke WK, Weijers HA, and van de Kamer JH (1953) Coeliac disease. II. The presence in
Grains: Intolerance, Allergy and Diseases: Cereal Allergens; The wheat of a factor having a deleterious effect in cases of coeliac disease. Acta
Paediatrica 42: 3442.
Gluten-Free Diet; Grain Composition and Analysis: Standardized Husby S, Koletzko S, Korponay-Szabo IR, Mearin ML, Phillips A, Shamir R, et al. (2012)
Test Methods for Grains and Grain-Based Products; The Composition European Society for Pediatric Gastroenterology, Hepatology, and Nutrition
of Food Grains and Grain-Based Products; Units of Grain Science and guidelines for the diagnosis of coeliac disease. Journal of Pediatrics
Trade: Equivalence between the US, Chinese, and Metric Units; Gastroenterology and Nutrition 54: 136160.
Proteins: Protein Synthesis and Deposition; Proteomic Analysis Kasarda DD (1994) Toxic cereals in coeliac disease. In: Feighery C and OFarelly C
(eds.) Gastrointestinal Immunology and Gluten-Sensitive Disease, pp. 203220.
Relevant to Cereal Grains; The Protein Chemistry of Cereal Grains; The Dublin: Oak Tree Press.
Basics: Taxonomic Classification of Grain Species; The Grain Crops: Lindfors K, Rauhavirta T, Stenman S, Maeki M, and Kaukinen K (2012) In vitro models
An Overview; ; The Cereal Grains: An Overview of the Family of for gluten toxicity: Relevance for celiac disease pathogenesis and development of
Cereal Grains Prominent in World Agriculture; Barley: An Overview of a novel treatment options. Experimental Biology and Medicine 237: 119125.
Ludvigsson JF, Leffler DA, Bai JC, Biagi F, Fasano A, Green PH, et al. (2012) The Oslo
Versatile Cereal Grain with Many Food and Feed Uses; Oats: Overview; definitions for coeliac disease and related terms. Gut 62: 4352.
Wheat: An Overview of the Grain That Provides Our Daily Bread; Richey R, Howdle P, Shaw E, and Stokes T (2009) Recognition and assessment of
Wheat Processing: Gluten and Modified Gluten; The Gluten Proteins coeliac disease in children and adults: Summary of NICE guidance. British Medical
of the Wheat Grain in Relation to Flour Quality. Journal 338: b1684.
Sapone A, Bai JC, Ciacci C, Dolinsek J, Green PHR, Hadjivassiliou M, et al. (2012)
Spectrum of gluten-related disorders: Consensus on new nomenclature and
classification. BMC Medicine 10: 13.
Further Reading Schuppan D, Junker Y, and Barisani D (2009) Celiac disease: From pathogenesis to
novel therapies. Gastroenterology 137: 19121933.
Abadie V, Sollid LM, Barreiro LB, and Jabri B (2011) Integration of genetic and Sollid LM, Qiao S-W, Anderson RP, Gianfrani C, and Koning F (2012) Nomenclature
immunological insights into a model of celiac disease pathogenesis. Annual Review and listing of celiac disease relevant gluten T-cell epitopes restricted by HLA-DQ
of Immunology 29: 493525. molecules. Immunogenetics 64: 455460.
Arendt EK and Dal Bello F (2008) Gluten-Free Cereal Products and Beverages, 1st edn. Wieser H (2007) Chemistry of gluten proteins. Food Microbiology 24: 115119.
Burlington, San Diego, London: Academic Press. Wieser H and Koehler P (2008) The biochemical basis of celiac disease. Cereal
Catassi C, Fabiani E, Iacono G, DAgate C, Francavilla R, Biagi F, et al. (2007) A Chemistry 85: 113.
prospective, double-blind, placebo-controlled trial to establish a safe gluten Wieser H, Koehler P, and Konitzer K (2014) Celiac Disease and Gluten:
threshold for patients with celiac disease. American Journal of Clinical Nutrition Multidisciplinary Challenges and Opportunities, 1st edn. London, Waltham, San
85: 160166. Diego: Academic Press.
The Gluten-Free Diet
V Zevallos, Johannes Gutenberg Mainz, Mainz, Germany
I Herencia, Universidad Politecnica de Madrid, Madrid, Spain
2016 Elsevier Ltd. All rights reserved.
Figure 2 Villous architecture of the proximal intestine in a treated (a) and an untreated (b) CD patient, using scanning electron microscopy (left) and
histology (right). Reproduced from Gasbarrini G and Mangiola F (2014). Wheat-related disorders: a broad spectrum of evolving diseases.
United Eur. Gastroenterol. J. 2(4): 254262.
disorders. The pathogenesis of IBS, Crohns disease, and ulcer- nongluten fraction (amylase trypsin inhibitors, ATIs), seems to
ative colitis remains to be fully elucidated, but genetic predis- play a significant role in the onset of disease.
position, microbiome, and environmental/dietary triggers are RA increases the risk of atherosclerosis and cardiovascular
the most important factors involved in the onset of disease. disease; it has been shown that a gluten-free vegan diet
Furthermore, in vivo experiments using HLA-DQ8 mice sensi- decreased total cholesterol. It is antiatherogenic and signifi-
tized with gluten show that an altered barrier function could be cantly reduces disease activity and the levels of antibodies to
a critical step in facilitating exposure of the mucosal immune b-lactoglobulin. Type I DM is also affected by gluten and
system to dietary antigens that subsequently will trigger an having a GFD has beneficial effects on the preservation of B-
immune response. Thus, a GFD recommended to patients cell function. Wheat consumption also affects neurological
with IBD-like symptoms resulted in improved symptoms, sug- dysfunctions, although the precise mechanism on the nervous
gesting the potential role of a GFD as adjunctive therapy. system is still unclear. Thus, a GFD has been shown to improve
Nonceliac gluten/wheat sensitivity (NCGS) is a condition that and stabilize ataxia by preventing irreversible loss of Purkinje
significantly improves on a GFD, initially reported in the late cells in the cerebellum and improve neuronal sensory action
1970s; NCGS can be defined as a reaction to gluten-containing potential and headaches. Furthermore, there are reports indi-
grains without involvement of allergy or autoimmune cating that a GFD has a positive effect on symptom reversal of
mechanisms. It is associated with gastrointestinal (bloating, diar- multiple sclerosis.
rhea, abdominal pain, and nausea) and predominantly non- Dietary trends are an important cultural factor often asso-
gastrointestinal symptoms (headache, foggy mind, chronic ciated with health and disease; in recent years. Being on a GFD
fatigue, joint and muscle pain, numbness of leg and arms, and has become fashionable in most Western societies. In the
anemia). The prevalence of NCGS is between 1% and 5% and United States, it is considered the most trendy diet with 1.5
seems to be more common in females and young/middle-aged million citizens (without any underlying medical condition)
adults although the prevalence in children needs further consuming gluten-free products. This proportion is similar in
investigation. The pathogenic mechanism is still unclear, but other Western countries and is predicted to increase even more
the innate immune response to wheat, particularly part of the in the coming years. The popularity of gluten-free products is
94 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | The Gluten-Free Diet
partly motivated by the association with healthy eating, weight junctions, a paracellular pathway, epithelial transcytosis, and
loss, and a recent negative perception of wheat as an over- direct sampling by dendritic cells. Internalized gluten peptides
utilized staple of the Western diet. are then deamidated by the enzyme tTG II, increasing nega-
tively charged peptides that can be accommodated in binding
pockets of the heterodimeric HLA class II alleles encoding for
Gluten and Nongluten Wheat Proteins HLA-DQ2 and HLA-DQ8. This increase in antigenicity acceler-
ates the secretion of interferon-g (IFN-g) by gluten-specific
The storage proteins are the main immunogenic components CD4 T-helper cells, contributing to the expansion of
of wheat, barley, and rye kernels. These proteins are located in cytotoxic T cells and fibroblasts and the release of metallopro-
the endosperm of kernels and can be subdivided according to teases to degrade the extracellular matrix of enterocytes at the
their solubility in water, salt, alcohol, and alkali into albumins, lamina propria and also to start secretion of anti-tTG by gluten-
globulins, prolamins, and glutelins, respectively. For wheat, specific plasma cells into the lamina propria and then to the
the glutenprotein complex can readily be purified by making bloodstream.
a dough with wheat flour and water and kneading the dough The other major grouping of wheat-grain proteins is the
under running water. The starch and water-soluble proteins water-/salt-soluble proteins referred as nongluten wheat
wash out and a cohesive ball of gluten remains. The cohesive proteins, the most relevant in terms of innate inflammation
properties of wheat gluten are the reason why only wheat flour being the ATIs. Some of these proteins play a significant role as
is well suited to bread making. The storage proteins of barley a plant defense mechanism against pests and during the met-
and rye are similar to those of wheat, but they lack the dough- abolic process of seed development. Wheat ATIs were identi-
forming properties of wheat gluten. For dietary purposes, the fied as contaminating proteins within pepsintrypsin gliadin
storage proteins of wheat, barley, and rye can be considered as digests; they represent between 2% and 4% of total grain
gluten, being a mixture of both prolamins and glutelins. protein compared to 7080% for gluten. ATIs are resistant to
Prolamins are alcohol-soluble storage proteins character- proteolytic digestion due to their compact secondary structure
ized by their higher content of proline and glutamine, which that includes five intrachain disulfide bonds. ATI subfamilies
in combination could constitute between 30% and 70% of the can be monomeric or form noncovalently linked dimers or
total amino acid composition. The molecular weight of pro- tetramers. There are up to 17 ATI species of around 150-amino
lamins ranges from 10 to 100 kDa and prolamins are a signif- acid length with a variant primary structure and conserved
icant source of nitrogen, sulfur, and carbon for the developing secondary structure.
embryo and they are known as gliadins in wheat, hordeins in ATIs engage the toll-like receptor 4 (TLR4)-MD2-CD14
barley, and secalins in rye. complex and lead to upregulation of maturation markers and
According to their decreasing electrophoretic mobility, gli- release of proinflammatory cytokines in myeloid cells of celiac
adins can be divided into a, b, g, and o gliadins. Based on their patients. In vivo feeding experiments in mice showed that ATIs
N-terminal amino acid sequences, gliadins can be categorized increased intestinal and systemic release of cytokines and che-
into a, g, and o. There is another classification based on the mokines like IL-8, TNF-a, and CCL-2 within 212 h, and
amount of sulfur, o gliadins are sulfur-poor and a and g ex vivo celiac biopsy studies demonstrated an adjuvant effect
gliadins are sulfur-rich. The first prolamin to be fully of added ATIs for the gluten-specific T-cell response. Therefore,
sequenced was A-gliadin (an a-type) using direct protein ATIs appear to be the sought triggers of innate immunity in CD
sequencing; the remaining amino acids were sequenced using and NCGS.
cloned DNA, facilitating the development of synthetic pep-
tides. Other studies using different wheat, barley, and rye cul-
tivars found that purified prolamins from rye (g and o
Gluten-Containing and Gluten-Free Plants
secalins) and from barley (o hordeins and C-hordeins) could
be identified using chromatographic methods. Prolamins are Gluten-containing cereals are part of the Gramineae family,
also present in other plants but at a different proportion also known as Poaceae due to its association with the type
and with different amino acid sequences than in wheat, barley, genus Poa. This family includes 12 subfamilies, in which two
and rye. are more relevant to CD: Pooideae that includes wheat, rye,
The glutelin protein of wheat is termed glutenin. It consists and barley, which are considered toxic for celiac patients, and
of alcohol-soluble subunits that are cross-linked by interchain Panicoideae that includes maize, millet, and sorghum, which
disulfide bonds to form alcohol-insoluble polymeric glutenin. are considered suitable for the GFD. The food product corn
These subunits have been classified according to their electro- gluten is a misnomer; it is the protein washed from maize
phoretic mobility into high-molecular-weight (HMW) and (corn) grain during wet milling. It is not related to wheat
low-molecular-weight groups. The HMW glutenin subunits gluten.
make a major contribution to dough formation. Specific sub- The taxonomic plant classification proposed by Kasarda
units are identified according to the relevant genes, such as provided a simple guidance to separate safe from unsafe grains.
1Dy10, 1Dx5, 1Dx4, and 1Dy9. Rye and barley have equiva- Monocotyledonous plants, members of the Gramineae
lent glutelin fractions (HMW secalins and D-hordeins, respec- family, and the Triticeae tribe (wheat, barley, and rye) were
tively) but they are less suitable for dough formation. considered celiac-toxic, while dicotyledonous plants such as
Immunogenic gluten (prolamins and glutelins) peptides Chenopodiaceae and Amaranthaceae (quinoa and amaranth)
trigger an adaptive immune response in CD. These peptides and those from other tribes (maize, rice, and tef) are consid-
reach the lamina propria via a combination of defective tight ered safe for celiac patients.
FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | The Gluten-Free Diet 95
Early studies on oats, probably contaminated with gluten, contaminated with gluten (up to 3000 mg kg1). Although
indicated that oats were unsuitable, but recent feeding studies these products were not labeled as gluten-free, they still repre-
comparing a GFD with and without oats concluded that mod- sent a risk for celiac patients, particularly since clinical studies
erate amount of oats (5070 g per day) can be safely added to have suggested that 100 mg kg1 of gliadin or more can trigger
the GFD of celiac patient. Another study assessing the long- an inappropriate immune response leading to histological
term effects of large amount of oats (up to 100 g per day) damage of the intestinal mucosa in diagnosed patients.
found that histology, serology, and well-being were constant Similarly, it was found that the daily intake of gliadin at
throughout the study. However, there have also been reports 414 mg did not produce the same effects for most celiac
that activation of CD with morphological deterioration patients.
occurred after oats challenge. They described T-cell lines reac- The codex standard for gluten-free foods (Codex Stand 118,
tive to avenin (the storage protein of oats) from duodenal 1981) regulates the level of gluten in gluten-free foods and is
mucosa of five patients consuming oats as part of their GFD often reviewed by a special FAO/WHO committee. The last
that were positively stimulated by avenin peptides rich in review of Codex Alimentarius Commission (2008) indicates
proline and glutamine residues. Other reports indicated with- that the maximum gluten level for gluten-free products is
drawals from some large feeding studies, which added to the 20 mg kg1. In 2012, new legislation based on these guidelines
T-cell data, suggest that for a minor proportion of the celiac was issued by the European Commission, and the 20 mg kg1
population, oats are unsuitable. threshold was applied to all GF products, that is, naturally GF
Furthermore, gluten-containing plants (wheat, rye, barley, (maize, rice, and uncontaminated oats) and derived from
and their earlier ancestors) rich in ATIs have a strong innate wheat starch. Other organizations such as the Food and Drug
stimulatory capacity that contrasts with a rather minimal Administration have adopted similar a similar threshold level
innate stimulatory effect of ATIs from gluten-free plants. (20 ppm), and the Food Standards Australia New Zealand has
Thus, gluten-free plants are also ATI-free. The average amount chosen a more strict approach considering no detectable
of wheat flour consumed per adult/day is 250 g, indicating amount of gluten or as low as 35 ppm (www.standards.gov.
that around 1 g of ATIs could be enough to trigger a moderate au; www.coeliac.org.au).
innate immune response in the gut of people affected by CD or
NCGS, also having an effect on nongastrointestinal organs.
Nutritional Value of the GFD
The morphological effect of ATIs is discrete and characterized
by an increase in inflammatory cells in accordance to histolog- Restrictive diets such as a GFD tend to have an inherent risk of
ical changes observed in NCGS patients. Furthermore, nutritional deficiency. Previous studies mostly agreed that a
the effect is dose-dependent, and therefore, a reduction GFD is nutritionally inadequate, but other studies did not
(8090%) of daily ATI consumption may be sufficient to elim- reach the same conclusion (http://www.food.gov.uk/science/
inate the stimulatory effect, and therefore, a strict GFD does research/allergy-research/t07053), and in general, information
not need to be maintained. Over the last decades, an increased about the nutrition adequacy of a GFD is still rather limited.
prevalence of allergies and autoimmune diseases has been There is evidence that adults following a GFD have
observed, particularly in developing countries. This can be increased levels of cholesterol, altered homocysteine levels,
partly explained by a parallel increase in wheat (ATIs) con- and a potentially high risk of developing cardiovascular dis-
sumption since traditional (gluten-free) staples were replaced ease. Furthermore, another study showed that gluten-free
by more affordable alternatives (modern wheat). products, particularly breads and pasta, have low protein,
high saturated fat, sodium, and cholesterol compared to
gluten-containing products, reflecting the nutritional conse-
Legislation of Gluten-Free Products
quences (low protein/high fat) particularly in females.
The denomination gluten-free theoretically implies without This nutritional imbalance can be due to the ingredients
any amount of gluten; however, having such a diet will be used. For instance, wheat starch requires an industrial process
impractical and very difficult to implement since gluten is a that inevitably alters the macro- and micronutrient composi-
ubiquitous ingredient and therefore will be almost inevitably tion of wheat, a source of protein, iron, and vitamins (thiamin,
present in trace amounts within any diet. Furthermore, com- riboflavin, folate, and niacin). In addition, replacing tradi-
pliance to a GFD is mainly related to palatability and the tional gluten-free products for alternative naturally gluten-free
availability of gluten-free products. This difficulty was partly products such as Andean grains (quinoa and amaranth) can
addressed by the addition of wheat starch flour to the gluten- increase variety, palatability, and most importantly significant
free list, particularly in Scandinavia and the United Kingdom amounts of nutrients such as protein, fiber, iron, calcium,
where patients consumed traces up to 200 ppm without clini- vitamins, and polyunsaturated fatty acids.
cal evidence of exacerbation of the disease. However, there is Baking gluten-free products remains a challenge mainly
an individual level of tolerance to gluten and even small because the main ingredient, gluten, cannot be easily replaced.
amounts of gluten can be of relevance for a group of celiac Most of the gluten-free flours lack the viscoelastic properties of
patients. wheat dough when kneaded in water and usually require more
The other alternative to increase palatability and reduce the water to form a dough, which is difficult to handle. It is often
risk of nutritional imbalance is to include alternative naturally referred to as a batter. The main problem arises when the
gluten-free ingredients. However, there is a risk of gluten con- mixture of water, yeasts, gluten-free flours, and starches forms
tamination that could occur from harvest to packaging, for few coherent structures because the gases from fermentation
example, samples of millet, sorghum, and soy heavily are not entrapped, directly affecting quality parameters, having
96 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | The Gluten-Free Diet
reduced volume and crumbly structure, with poor texture, Investigate the methods used to determine the amount of
mouthfeel, and flavor. gluten in food products and ingredients. Start with the
However, the gluten-free baking industry through food encyclopedias articles on the standardization of methods
technologists and nutritionists maintains a continuous process for analyzing grains and Appendix 3, which lists testing
of quality development, particularly by the addition of novel methods.
ingredients and new technologies. Thus, the main goal is to
create gluten-free products with similar quality standards to the
gluten-containing counterparts that will ultimately help to
See also: Appendix 3: Grains, Foods, and Ingredients Suiting
improve the quality of life of people affected by wheat-related
Gluten-Free Diets for Celiac Disease; Food Grains and the
conditions.
Consumer: Consumer Trends in Grain Consumption; Grains and
Health: Misinformation and Misconceptions; Labelling of Grain-Based
Foods; Food Grains: Intolerance, Allergy and Diseases: Celiac
Conclusion and Future Developments Disease; Cereal Allergens; Proteins: The Protein Chemistry of Cereal
Grains; The Basics: Taxonomic Classification of Grain Species; The
Patients affected by wheat and related cereals benefit from a
Cereal Grains: Wheat: An Overview of the Grain That Provides Our
GFD. There is clear evidence that gluten proteins exacerbate
Daily Bread; Wheat-Based Foods: Cereals: Overview of Uses:
celiac disease. However, non-gluten proteins as activators of
Accent on Wheat Grain.
innate immunity might also play a significant role in CD and
other inflammatory conditions. Health care professionals sup-
port patients on a GFD by improving their psychological well-
being, reducing social restrictions, developing new products to
Further Reading
meet nutritional targets, and ultimately improving their quality
of life. The increased number of people benefited from a GFD Catassi C, et al. (1993) Dose dependent effects of protracted ingestion of small amounts
as well as a recent social trend indicates that the gluten-free of gliadin in coeliac disease children: A clinical and jejunal morphometric study. Gut
market is projected to grow significantly over the coming years. 34(11): 15151519.
Catassi C, et al. (1999) Why is coeliac disease endemic in the people of the Sahara?
Lancet 354(9179): 647648.
Catassi C, et al. (2013) Non-celiac gluten sensitivity: the new frontier of gluten related
Exercises for Revision disorders. Nutrients 5(10): 38393853.
Ciacci C, et al. (1999) Low plasma cholesterol: A correlate of nondiagnosed celiac disease
in adults with hypochromic anemia. Am. J. Gastroenterol. 94(7): 18881891.
Describe the most common symptoms of CD for children
Dicke WK, Weijers HA, and Van De Kamer JH (1953) Coeliac disease. II. The presence
and for adults. in wheat of a factor having a deleterious effect in cases of coeliac disease. Acta
What are the screening tests and the gold standard tests to Paediatr. 42(1): 3442.
diagnose CD? Dieterich W, et al. (1997) Identification of tissue transglutaminase as the autoantigen of
Outline the taxonomic groupings of grain species that are celiac disease. Nat. Med. 3(7): 797801.
Dupont FM, et al. (2011) Deciphering the complexities of the wheat flour proteome
celiac-toxic or not. Refer to the encyclopedias article on the
using quantitative two-dimensional electrophoresis, three proteases and tandem
taxonomy of grain-bearing plants. mass spectrometry. Proc. Natl. Acad. Sci. U. S. A. 9: 10.
Summarize the development of legislation to assist con- Ejderhamn J, Veress B, and Strandvik B (1988) The long-term effect of continual
sumers in choosing gluten-free foods, also considering the ingestion of wheat starch-containing gluten-free products in coeliac patients.
means of communication, mainly via product labeling. In: Kumar PJ (ed.) Coeliac Disease: One Hundred Years, pp. 294297. Leeds: Leeds
University Press.
Gasbarrini G and Mangiola F (2014) Wheat-related disorders: a broad spectrum of
evolving diseases. United Eur. Gastroenterol. J. 2(4): 254262.
Exercises for Readers to Explore the Topic Further Haas SV (1924) The value of banana in the treatment of celiac disease. Arch. Pediatr.
Adolesc. Med. 28(4): 421.
Hadjivassiliou M, et al. (2003) Dietary treatment of gluten ataxia. J. Neurol. Neurosurg.
Investigate the recorded evidence for oats being harmful or
Psychiatry 74(9): 12211224.
not in CD. Refer to recommendations in Appendix 3, and Hafstrom I, et al. (2001) A vegan diet free of gluten improves the signs and symptoms of
examine articles such as the following: rheumatoid arthritis: The effects on arthritis correlate with a reduction in antibodies
- Kemppainen, T.A. et al., 2008. Unkilned and large to food antigens. Rheumatology 40(10): 11751179.
amounts of oats in the coeliac disease diet: a randomized, Hager AS, et al. (2012) Investigation of product quality, sensory profile and
ultrastructure of breads made from a range of commercial gluten-free
controlled study. Scandinavian Journal of Gastroenterology, flours compared to their wheat counterparts. Eur. Food Res. Technol. 235(2):
43(9), pp. 10941101. 333344.
- Lundin, K.E.A. et al., 2003. Oats induced villous atrophy Herfarth HH, et al. (2014) Prevalence of a gluten-free diet and improvement of clinical
in coeliac disease. Gut, 52 (11), pp. 16491652. symptoms in patients with inflammatory bowel diseases. Inflamm. Bowel Dis. 14.
Junker Y, et al. (2012) Wheat amylase trypsin inhibitors drive intestinal inflammation via
In recent years, a GFD has become fashionable in many
activation of toll-like receptor 4. J. Exp. Med. 209(13): 23952408.
societies. Investigate the existence of this fashion in your Kasarda DD (1994) Toxic cereal grains in coeliac disease. In: Feighery C and
region. Determine the extent to which this trend, if appar- OFarrelly C (eds.) Proceedings of the 6th International Symposium on Coeliac
ent, has a cultural-fashion basis or if it may be associated Disease, pp. 203220. Dublin: Oak Tree Press.
with valid health and disease reasons. Kasarda DD, et al. (1984) Nucleic acid (cDNA) and amino acid sequences of alpha-type
gliadins from wheat (Triticum aestivum). Proc. Natl. Acad. Sci. U. S. A. 81(15):
In parallel, investigate the availability of gluten-free foods 47124716.
in your region, considering over-the-counter goods and Kemppainen TA, et al. (2008) Unkilned and large amounts of oats in the coeliac disease
packaged foods in the local shops. diet: a randomized, controlled study. Scand. J. Gastroenterol. 43(9): 10941101.
FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | The Gluten-Free Diet 97
Lee AR, et al. (2009) The effect of substituting alternative grains in the diet on the Peraaho M, et al. (2003) Wheat-starch-based gluten-free products in the treatment of
nutritional profile of the gluten-free diet. J. Hum. Nutr. Diet. 22(4): 359363. newly detected coeliac disease: prospective and randomized study. Aliment.
Lo W, et al. (2003) Changing presentation of adult celiac disease. Dig. Dis. Sci. 48(2): Pharmacol. Ther. 17(4): 587594.
395398. Schuppan D, Junker Y, and Barisani D (2009) Celiac disease: from pathogenesis to
Lundin KEA, et al. (2003) Oats induced villous atrophy in coeliac disease. Gut 52(11): novel therapies. Gastroenterology 137(6): 19121933.
16491652. Shewry PR and Tatham AS (1990) The prolamin storage proteins of cereal seeds:
Miranda J, et al. (2014) Nutritional differences between a gluten-free diet and a diet structure and evolution. Biochem. J. 267(1): 112.
containing equivalent products with gluten. Plant Foods Hum. Nutr. 69(2): Stern M (2000) Comparative evaluation of serologic tests for celiac disease: a
182187. European initiative toward standardization. J. Pediatr. Gastroenterol. Nutr. 31(5):
Molberg O, et al. (1998) Tissue transglutaminase selectively modifies gliadin peptides 513519.
that are recognized by gut-derived T cells in celiac disease. Nat. Med. 4(6): 713717. Strsrud S, et al. (2003) Adult coeliac patients do tolerate large amounts of oats. Eur. J.
Osborne TB (1907) The Protein of the Wheat Kernel. Washington, DC: Carnegie Institute Clin. Nutr. 57(1): 163169.
of Washington, 84. Zanini B, et al. (2013) Impact of gluten-free diet on cardiovascular risk factors. A
Pastore MR, et al. (2003) Six months of gluten-free diet do not influence autoantibody retrospective analysis in a large cohort of coeliac patients. Dig. Liver Dis. 45(10):
titers, but improve insulin secretion in subjects at high risk for type 1 diabetes. 810815.
J. Clin. Endocrinol. Metabol. 88(1): 162165. Zevallos VF, Herencia LI, and Ciclitira PJ (2014) Gastrointestinal effects of eating
Pengiran Tengah CDSNA, et al. (2004) Multiple sclerosis and occult gluten sensitivity. quinoa (Chenopodium quinoa Willd.) in celiac patients. Am. J. Gastroenterol.
Neurology 62(12): 23262327. 109(2): 270278.
Proteins
Contents
The Protein Chemistry of Cereal Grains
The Protein Chemistry of Dicotyledonous Grains
Protein Synthesis and Deposition
Nitrogen Metabolism
Nitrogen in Grain Production Systems
The Enzymes Associated with Food Grains and Their Functions in Processing
Proteomic Analysis Relevant to Cereal Grains
Thiolomics of the Gluten Protein Network of Wheat Dough
The importance of dietary protein from cereal grains rice, Worldwide, plant proteins contribute more than half of our
wheat, maize, and others. dietary supply of protein. Over 200 million tonnes of protein
The purification of wheat gluten from dough as an exper- are harvested each year in the form of cereal grains, based on FAO
iment in the kitchen and as a commercial product. statistics (http://faostat.fao.org). This protein harvest includes
The simple structure of proteins a linear polymer of that derived from the three major cereal grains, wheat, rice, and
amino acids produces the diversity of properties maize, and also less widely grown species: barley, rye, triticale,
of enzymes, hormones, hair and horn, silk and sinew, and sorghum. In some developing countries, a single cereal
and gluten together with the many other grain storage species may be the major or sole source of protein in the diet.
proteins. In many countries, cereal proteins enter the diet in various
Folding and cross-linking of the amino acid polymer gives forms, including wheaten breads and pastries, noodles, fried
three-dimensional structure that is needed for many pro- rice dishes, rye crispbreads, cookies, biscuits, pearl barley,
tein functions, such as enzymatic activity. pasta, corn tortillas, breakfast cereal, oatmeal porridge,
Osbornes solubility-based protein classes: water-soluble couscous, and many more foods. Most obviously, cereal grain
albumins, salt-soluble globulins, prolamins extractable proteins are eaten directly in these many processed forms. The
with 70% aqueous ethanol, and the residual glutelins. second route is indirect, in the form of animal products, such
Protein quantity is determined by various methods, espe- as meat, milk, and eggs, following the feeding of cereal grains
cially by near-infrared spectroscopy, based on total nitrogen to animals.
content. Whereas the protein part of the grain is important nutri-
Protein quality relates to its nutritional value and its con- tionally, it also serves functional purposes, relevant to the
tribution to functional uses. processing of the grain. Of these, wheat and rye are unique
The specific proteins are described for wheat, rye, triticale, because of the dough-forming properties of their storage pro-
barley, rice, maize, and sorghum. teins, known as gluten when formed into dough.
Early Concepts
The word protein was proposed by the French scientist Berze-
lius, writing to the German scientist Mulder, with the appro-
priate meaning of primary substance, as recently as 1838, long
after the acceptance of the terms for specific types of proteins,
such as gluten, zymom, and fibrin. Early concepts of protein
chemistry imagined it to have the complex formula
C20H31N5O6, meaning that 20 molecules of carbon (C) were
compounded with 31 molecules of hydrogen, five molecules
of nitrogen, and six molecules of oxygen. The dogma of those
early days also maintained (incorrectly) that this complex was
absorbed (unchanged) directly by the intestine. It is now
known that dietary proteins are hydrolyzed to the component
amino acids for absorption into the bloodstream.
Wheat proteins
HMW
glutenin subunits
gliadin
gliadin
LMW
glutenin subunits gliadin
gliadin
S-rich prolamins
Figure 2 Classification of the proteins of the wheat grain, showing the electrophoretic patterns for some of them.
groups as sulfur-rich, sulfur-poor, and high-molecular-weight There are about 20 different amino acids, each of them
(HMW) prolamins, representing the a-, b-, and g-gliadins having the combination of an acidic group (COOH) and a
plus the low-molecular-weight (LMW) glutenin subunits, the basic group (NH2) and each having a distinct side chain that
o-gliadins, and finally the HMW glutenin subunits, respectively, provides the functional differences between them all. The
in the old classification system. amino acids are joined together by peptide bonds to form a
polypeptide chain, by the following formula (for two amino
acids, R1 and R2):
So What Is a Protein?
R 1 COOH H2 N R 2 ! R 1 CO NH R2 H2 O
A protein molecule is simply a linear polymer of amino acids.
The structure of protein molecules also depends on the folding The sequence and specificity of the amino acids are spelled
and cross-linking of the polymers, giving a three-dimensional out exactly in the genetic code of the genes, thereby ensuring
structure that is especially important for functions such as that the protein will be made in the same manner each time
enzymatic activity. in the specific species and tissue. For this reason, the determi-
It is no wonder that these early protein chemists wrestled nation of grain protein composition is a valuable means of
with reaching this understanding, when we now see that this establishing the identity of a variety within a grain species.
relatively simple chemical structure can produce the great
diversity of properties that are found in proteins, such as
Many Different Cereal Grain Proteins
enzymes and epidermis, hair and horn, feathers and flagella,
silk and sinew, and even spiders web. In the past century, we have come a long way from Osbornes
The diversity depends on the selection and sequence of the concept of four classes of proteins to realize that within each of
amino acids plus the length of the protein (polypeptide) chain. his classes, there are a large number of proteins differing in
Further diversity is provided by cross-linking between polypep- charge and size; this great diversity and heterogeneity can
tide chains and from linking to accompanying molecules readily be shown by fractionation methods such as gel electro-
(lipids, carbohydrates, or metal ions). The resulting protein phoresis. Each of the horizontal lines (bands) in the electro-
molecules are large compared with many biological com- phoretic patterns in Figure 3 represents a separate protein
pounds, with molecular weights often in the range of component (more than one in many cases). The combination
10 000100 000 Da (1 Da being equivalent to the mass of a of gel electrophoresis and gel isoelectric focussing as used in
hydrogen atom). Some proteins are even larger, the largest proteomics, provides a two-dimensional display of spots
glutenin proteins of wheat gluten having sizes extending up (Figure 4), each being a distinct protein (polypeptide, mean-
to tens of millions of Daltons. ing polymer of amino acids).
PROTEINS | The Protein Chemistry of Cereal Grains 101
Figure 3 Lab-on-a-chip analysis of the composition of prolamin proteins extracted from grains of wheat varieties (L to R: Halberd, Janz, Frame, and
Yitpi) and of barley varieties (Excalibur, Sloop, Schooner, Flagship, Gairdner, and Baudin). Each protein extract was applied to a small lab chip for
microfluidic capillary electrophoresis. Each analysis took about 50 s. The elution patterns were manipulated to simulate gel electrophoresis patterns.
3 pH 10
kDa
130
10
Crude protein Crude fat Crude fiber
Figure 4 Proteome map of the polypeptides (reduced proteins)
extracted from mature wheat grain (cv Rosella). Fractionation involved Crude ash Starch Cellulose
isoelectric focussing between pH 3 (left) and pH 10 followed by SDS gel Figure 5 Chemical composition typical of a whole cereal grain.
electrophoresis in a gradient from 8% (top) to 18% polyacrylamide.
After in-gel digestion with trypsin, 315 protein spots (as numbered) were
excised and identified by MALDI-TOF and LCMS/MS. Adapted from
Mak, Y., Willows, R. D., Roberts, T. H., Wrigley, C. W., Peter Sharp, P.,
and Copeland, L., 2009. Germination of wheat: A functional proteomics the prolamin proteins extracted from grain samples of specific
analysis of the embryo. Cereal Chemistry 86 (3), 281289. varieties of wheat and of barley. Each of the two sets of
varieties has a distinct pattern of bands, while all the wheat
patterns differ from all the barley patterns. This type of anal-
The diversity of grain protein composition is such that ysis is thus useful for the identification of varieties in the
differences in composition can readily be demonstrated practical situation of grain delivery when identity must be
between closely related varieties of a grain species. This is established to define quality type and royalty payment for
evident in Figure 3 where each column of bands represents distinct varieties.
102 PROTEINS | The Protein Chemistry of Cereal Grains
Why Is Protein So Important in Grains? and the fiber components (Figure 5). The protein content of
cereal grains ranges from 8% to 15% of the grain weight (Table 1).
The protein component of the grain is critical to the grains role Even within a grain species, the actual protein concentration may
as a seed the beginning of a new plant. The grain protein has cover a range of values, depending on the genetic potential of a
the natural purpose to serve as a reserve of amino acids for the specific variety of that grain species, on the nutritional status of
embryo (germ) to draw upon during germination, while the the plant, and on climatic and other environmental factors.
roots and shoots are growing until the leaves can support Most of the protein is in the grains endosperm (the part
growth and development via photosynthesis. that becomes white flour after milling the cereal grain), the
The protein component of the grain is important nutrition- endosperm being the largest part of the grain by mass. How-
ally as a source of amino acids, especially the essential amino ever, the proportion of protein may be higher in some of the
acids that we cannot synthesize ourselves. The amino acids that non-endosperm tissues of the grain, especially in the germ (the
are essential include lysine, methionine, leucine, isoleucine, embryo, which will become the new plant after germination).
and several more. The barley grain is taken as an example of this in Figure 6.
On the other hand, grain protein is important for the grain Figure 7 shows the chemical composition of wheat grain, with
processing industry, because it is often the most important part of respect to the distribution of protein (and other components)
the grain in determining processing quality and nutritional value. in its various anatomical parts.
In grain composition, protein is next in abundance after starch The importance of protein content is seen in the valuation
of grain shipments. The single figure of the protein content of
a grain sample is an important specification in the marketing
Table 1 Approximate protein content of the more common cereal of most cereal grains. In general, grain of higher protein con-
species, as percentages of the as is weight of whole grain, except for tent commands a higher value. For wheat, a higher protein
rice (milled grain)
content indicates better baking quality, and a higher protein
Cereal species Protein content content for feed grains provides a richer supply of essential
amino acids (irrespective of the proportion of essential amino
Barley 912 acids in their storage protein).
Maize (corn) 812 However, malting barley is an exception to the rule that
Oats (as groats) 1215 high protein content is desirable, because of the need to have a
Rice (milled) 811 high starch content to provide a correspondingly high extrac-
Rye 1215
tion of fermentable sugars in the malting process. Nevertheless,
Wheata 916
even for malting barley, there is a minimum level of protein
a
Durum wheat, a distinct species from common wheat, generally requires a higher (8%), to ensure that there is an adequate supply of the hydro-
protein range, say, 1216%. lytic enzymes needed throughout the malting process.
Crude protein
Crude fat
Crude fiber
Crude ash
Nitrogen-free extract
Cellulose
12% Protein
2% Lipids
60% Starch
2% Red. sugars
6% Pentosans
% of grain 2% Cellulose
2% Ash
0 50 100
Embryo Scutellum Pericarp Aleurone Endosperm
Protein
Lipids
Starch
Red. sugars
Pentosans
Cellulose
Ash
Figure 7 Chemical composition of the wheat grain; distribution between anatomical parts. Red. Sugars means reducing sugars, such as glucose.
Table 2 Methods used to determine the protein content of cereal elements and determination of the proportion of the element
grain, especially wheat nitrogen. This figure is multiplied by 6.25 to obtain the protein
content or by a factor of 5.7 in the case of wheat. This proce-
Process Material assayed Assay method dure assumes that virtually all the nitrogen in the grain is in the
form of protein. A lower multiplication factor is used for wheat
Grain NIR
Mill Flour/meal NIR
because its protein contains a high proportion of the amino
# acid glutamine, giving its protein unusually high nitrogen
Wash (for wheat Gluten Weight (wet or dry) content (Figure 8). The total nitrogen content is used as an
only) absolute value in establishing correlations for the routine
# method of NIR spectroscopy to determine the protein content
Extract Protein in solution Biuret or dye-binding of whole grain or of milled products (Table 2).
reaction
#
Hydrolyze Amino acids Gas or liquid Protein Quality
chromatography
# The single figure of protein content, however, does not take
Alkaline digestion Amide N Titrate ammonia released into account the quality of the protein. This quality is indicated
#
for nutritional purposes by analysis of the levels of essential
Acid digestion Ammonia or Kjeldahl or Dumas method
amino acids after hydrolysis of the peptide bonds. For wheat,
nitrogen
however, the quality of the protein also relates to the dough
strength provided by the particular combination of gluten pro-
teins present, determined partly by the genotype (the variety)
and also by the growth conditions.
Analysis of Protein Content For the animal feed industry, the amino acid composition is
especially important, due to the need to ensure an adequate
Today, the routine determination of protein content of all supply of essential amino acids. The protein of cereal grains is a
grain species generally involves the use of near-infrared (NIR) good source of most of the essential amino acids, with the
spectroscopy, applied to whole grain, milled grain, or flour, exception of lysine and possibly tryptophan, which are lower
based on correlations to total nitrogen analyses (Table 2). In in the cereal grains than in most animal proteins. The amino
the case of wheat only, protein content can also be determined acid composition for wheat is shown in Figure 8, with respect
by washing out the gluten protein and weighing its wet mass. to the major classes of wheat proteins. The amino acid com-
Even though gluten protein is less than 80% of the total grain positions of whole grains of several cereals are compared in
protein, this approach can be used if appropriate correlations Table 3.
are established between the level of washed gluten and true
grain protein content.
Further possibilities include colorimetric methods (e.g., the Comparison of the Proteins of Cereal Species
biuret reaction) or the amide distillation procedure (Table 2).
The standard (true) method for protein determination The various cereal species differ in the composition of their
involves the complete digestion of the grain or flour to its grain proteins. Just as the various cereal species are related
104 PROTEINS | The Protein Chemistry of Cereal Grains
Albumin
Globulin
Prolamin
Glutenin
Ile Leu Lys Met Cys Phe Tyr Thr Trp Val Arg His Ala Asp Glu Gly Pro Ser NH3
Table 3 The amino acid compositions of whole grains of several the cereal endosperm have been given the specific names
cereals, expressed as grams per 16 g of nitrogen according to each cereal genus, as listed in Figure 9.
Alanine 3.6 6.0 7.5 4.0 4.5 The Proteins of Wheat, Rye, and Triticale
Arginine 4.6 8.3 4.2 4.7 6.3
Aspartic acid 4.9 10.3 6.3 5.7 7.7 Wheat and rye are closely related; triticale is a man-made
Cysteine 2.5 1.1 1.6 2.3 2.7 hybrid between wheat and rye. The genetic control of the
Glutamic acid 29.9 20.6 18.9 23.6 20.9 storage proteins of these cereals is detailed in the specific
Glycine 3.9 5.0 3.7 3.9 4.7 articles on wheat, rye, and triticale. Their protein compositions
Histidine 2.3 2.5 2.7 2.1 2.1 are thus similar to a limited extent, but the dough-forming
Isoleucine 3.3 3.8 3.7 3.6 3.8
quality of rye and triticale is much poorer than that of wheat.
Leucine 6.7 8.2 12.5 6.7 7.3
The popularity of rye-based foods is not so widespread as that
Lysine 2.9 3.8 2.7 3.5 3.7
Methionine 1.5 2.3 1.9 1.7 1.7 of foods made from wheat. Rye is especially popular in Eastern
Phenylalanine 4.5 5.2 4.9 5.1 5.0 European and Scandinavian countries. Baked products are
Proline 9.9 4.7 8.9 10.9 5.2 commonly crispbread and varieties of bread, including
Serine 4.6 5.4 5.0 4.0 4.7 pumpernickel.
Threonine 2.9 3.9 3.6 3.3 3.3 Whereas true rye bread is made from 100% rye, it is
Tryptophan 1.1 1.2 0.6 1.5 1.3 common for rye bread to be made from a blend of rye and
Tyrosine 3.0 3.5 3.8 3.1 3.0 wheat flours, the contribution of the wheat gluten being
Valine 4.4 5.5 4.8 5.0 5.1 needed to provide the baking quality that is lacking in the rye
flour. The poorer dough-forming characteristics of rye gluten
are largely due to the lower level of the glutelin-type protein,
combined with the more hydrophilic (water-loving) properties
of the rye secalins, compared with the corresponding classes of
genetically, these genetic similarities and distinctions are protein in wheat.
reflected directly in the amino acid sequences of their storage The water-soluble proteins of rye contain the albumin and
proteins. Both the relevant gene sequences and the correspond- globulin classes of protein, similar to those in wheat, but for
ing amino acid sequences also reflect the taxonomic relation- rye, there is generally a higher level of amylase enzymes than in
ships among the cereal species, as is illustrated in Figure 9. To a wheat because of the greater susceptibility of rye to preharvest
large extent, these differences explain how the storage proteins sprouting and the consequent production of starch-degrading
of the respective cereal grains make their distinct contributions enzymes.
with respect to their food uses. The essential amino acid composition of rye is slightly
The most obvious example is wheat, whose gluten-forming better than that of wheat, because of the lower content of
storage protein is unique among the cereals in suiting wheat prolamin-type proteins in rye (Figure 9). The properties of
flour for bread making. Furthermore, the proteins of the rice triticale are intermediate between those of rye and wheat,
grain contribute to its eating quality. The storage proteins of reflecting the genetic origins of triticale.
PROTEINS | The Protein Chemistry of Cereal Grains 105
Globulins Prolamins
Maize, oats, Wheat Oats Rice Maize Sorghum Rice Maize Oats Triticeae:
rice, barley, barley, wheat,
rye, wheat rye
Embryo,
aleurone Endosperm
Figure 9 Storage protein classes in cereals. Adapted from Shewry, P. R., Tatham, A. S., and Halford, N. G., 1999. The prolamins of the Triticeae. In:
Shewry, P. R. and Casey, R. (Ed.), Seed Proteins. Kluwer Academic, The Netherlands, pp. 3578.
Hordein class Loci Gene copy number Proteins present in the cv Carlsberg MW (kDa) Corresponding wheat prolamin
The Proteins of the Barley Grain cereal grains, still being deficient in lysine, methionine, and
threonine. Nevertheless, it is a popular source of protein and
The protein concentration of cultivated barley ranges from 9% energy in feedstuffs, in combination with other sources of
to 12%. For barley, the protein content is generally calculated energy. High-lysine barley genotypes have been developed,
as total nitrogen content multiplied by the factor of 6.25. The analogous to those in maize, the first of these being the
protein content of hull-less (naked) barley genotypes is gener- Hyproly line.
ally higher (even up to 18%) because of the absence of the The hordein polypeptides show considerable polymor-
outer hull layers (the glumes the palea and lemma), which phism, like the prolamins of wheat and rye. A large proportion
contain relatively little protein. of the barley prolamins are disulfide-linked (both intra- and
The production of the enzymes of germination is an impor- interchain linkages) with naturally occurring polymers of high
tant part of a good malting barley, but paradoxically, high molecular weight. Unlike the glutenin polymers of wheat,
protein content (e.g., over 11%) is not necessarily desirable these are not desirable for the processing of barley. Because
for the best malting quality. This is because the ultimate test of they generate problems in the malting process and down-
malting quality is the extent of extract produced during the stream, they are likely to produce haze in the final product
mashing process. The amount of extract determines how much (i.e., beer) if they have not been adequately hydrolyzed during
beer can be produced from a given amount of malt. Extract processing.
represents the amount of soluble fermentable sugars from the The hordein polypeptides are classified into groups (A, B, C,
hydrolysis of starch, resulting from the combination of very and D hordeins) according to their decreasing mobilities dur-
active amylases and high starch content. For this reason, a low ing SDS gel electrophoresis. The C hordeins have analogy to
protein content (i.e., less than 8%) is also undesirable, as it the omega-gliadins of wheat, having a general absence of the
may have the consequence of inadequate enzyme production. sulfur-containing amino acids. They are coded by genes (locus
Consumption for human food is a traditional use of barley, Hor 1) on the short arm of barley chromosome 1H, as indi-
but this now accounts for only 5% of total barley consumption cated in Table 4. The genes (Hor 2 and Hor 5) for the sulfur-rich
worldwide. Feed use predominates, amounting to 75% of B and gamma-hordeins are also coded on chromosome 1H.
barley use. The amino acid composition of barley does not The D hordeins, which occur as disulfide-linked polymers, are
provide a consistently better balance nutritionally than other analogous to the HMV subunits of wheat glutenin; like them,
106 PROTEINS | The Protein Chemistry of Cereal Grains
their genes (Hor 3) are located on the long arm of the corre- Like the prolamins from other cereals, rice prolamins are
sponding barley chromosome (1H). readily soluble only in alcohol/water mixtures. However, the
classical extraction procedure had not been useful in early
studies. More recently, these estimates of prolamin content
have been shown to be low. Using 55% propan-L-ol and reduc-
The Proteins of the Rice Grain ing agents, higher yields of prolamins (2025% of the total
protein) have been obtained. Rice prolamins are 1016 kDa in
The proteins of rice are a contrast to those of wheat and barley, molecular weight. Thus, their molecular size range is signifi-
being predominantly of the globulin class. As a protein source, cantly smaller than the prolamins of other cereals. They are
milled rice contains the lowest amount of protein (about 5%) highly variable between rice cultivars, based on their electro-
among the major cereals. Moreover, this protein is not easily phoretic and isoelectric focussing fingerprints.
digestible by humans and monogastric animals. However, According to the Osborne classification system, the least
compared with that of other cereal proteins, the overall soluble proteins of the rice grain are oligomeric globulins
amino acid composition of rice protein is significantly more representing up to 80% of the total protein fraction
balanced due to the relatively higher level of lysine content. (Figure 9). They have been difficult to study because of their
This unusual amino acid composition results because rice is general insolubility in all solvents except dilute alkali, due to
one of the few cultivated plants in which there are significant their high molecular weight and their heterogeneity. When
levels of globulins and prolamins. These are the two major analyzed by gel filtration chromatography, the purified globu-
classes of storage proteins in the seeds of higher plants. Unlike lin subunits were resolved into three subunits, linked by disul-
other cereals that accumulate prolamins as their primary nitro- fide bonds, which varied in stoichiometry depending on the
gen reserve, the major storage proteins in rice are the glutelins, report and on the rice variety used.
which are homologous at the primary sequence level to the 11S Despite their general insolubility, rice glutelins are homol-
globulin proteins (Figure 9), a class that is the dominant form ogous in structure to 11S globulins (Figure 9). The structural
of nitrogen deposition in legumes. Rice prolamin proteins relationship is evident when comparing the N-terminal amino
have a number of characteristics that are different from the acid sequences and using immunologic methods. The extent of
prolamins present in most other cereals. homology between the rice glutelin and legume 11S globulins
The most important characteristic of the deposition of is 3035%. Structural similarity is evident in the primary
nitrogen in the rice kernel is that there are three kinds of sequences of homology to a motif common in wheat, rye,
protein bodies in the rice endosperm, namely, large spherical, barley, and maize prolamins.
small spherical, and crystalline protein bodies. Each of these is
surrounded by a single continuous membrane. Although the
spherical protein bodies form within vacuoles, the proteins are
synthesized in the endoplasmic reticulum and in the Golgi The Proteins of Maize
apparatus and are then transported to the vacuoles via vesicles.
Removal of the husk (hull) from rough (paddy) rice yields the Maize (or corn), often classed in world trade as one of the
kernel. This is composed of the pericarp, seed coat, aleurone, coarse grains, is an important source of protein in the diets of
endosperm, and the germ and is known as brown rice. It has a many people in countries such as Africa and South America. In
protein content of 910%, with a significantly higher nutri- addition, maize is used extensively as a feed grain and in a
tional value than the most commonly utilized rice product range of industrial processes. The process of corn starch man-
white polished (milled) rice, whose protein content is 8%. ufacture produces the grain protein as a by-product. It is some-
The milling process for paddy rice results in 4055% white times referred to as corn gluten in the trade, but this name is
milled rice, together with three major by-products, namely, inappropriate, because maize protein has no relationship to
husks (20%), bran (10%), and brokens (1022%). These by- the gluten of wheat and is safe for consumption by those
products have protein contents of 3%, 17%, and 9%, respec- susceptible to celiac disease.
tively. The aleurone layer, the tissue with the highest level of Like the other cereal grains, maize protein is slightly defi-
proteins and nutritionally important minor components, is cient in lysine. This limits its utilization as an animal feed. This
removed during the rice milling process. nutritional imbalance is largely due to the low lysine content of
The subaleurone region of the rice grain plays an important zein, the major group of maize proteins (Figure 9). The
part because of its nutritional value. It is a globulin-rich layer, development of the opaque-2 and floury-2 mutations has
being several cell layers thick. Its lysine content is much higher led to the development of maize genotypes with higher levels
than that of the proteins located in the rice endosperm. Thus, of lysine, due to a lower content of zein and more of the
rice should be milled as lightly as possible to retain as much as nutritionally superior albumin type of protein. Although
possible the subaleurone layer. these new maize types offer nutritional advantages, they carry
The albumin fraction of rice is highly heterogeneous, con- a grain yield penalty.
taining biologically important components. It can be separated The zein protein of maize can be isolated by extraction of the
into four subfractions, based on the molecular size of its pro- crushed endosperm (the starchy part of the grain) with 70%
teins. More than 50 individual polypeptides were observed in aqueous ethanol, after breaking disulfide bonds. Maize zein has
the albumin fraction, using isoelectric focussing. As for cereal been further fractionated into subclasses of proteins, designated
grains in general, the albumins are mainly enzymes and alpha-, beta-, gamma-, and delta-zeins. The genes for their syn-
enzyme inhibitors. thesis have been mapped to maize chromosomes 6 and 7.
PROTEINS | The Protein Chemistry of Cereal Grains 107
The major storage proteins of sorghum, termed kafirins How did the early chemists views on protein chemistry
(Figure 9), are fractionated by a procedure similar to that for differ from our present knowledge? Why were they unaware
the zeins of maize. Alpha-kafirin, composing about 80% of the of the great heterogeneity of grain proteins?
overall kafirin content, can be further subdivided into two What are the aspects of protein composition that explain
proteins with molecular weights of about 23 000 and the great differences between the various proteins in their
25 000 Da. Three beta-kafirins and one gamma-kafirin have properties?
been identified, all of them having molecular weights in the Prepare a table of differences and similarities between the
range of 1628 kDa. Like maize, high-lysine mutants of sor- major cereal species with respect to their protein
ghum have been developed. compositions.
What are the nutritional deficiencies of cereal grain proteins
with respect to the essential amino acids? Describe the genetic
achievements that have partially overcome these deficiencies.
Future Prospects
Our current understanding of grain protein chemistry contrasts Exercises for Readers to Explore the Topic Further
strongly with the rudimentary knowledge of the early twentieth
century. That knowledge has progressed from the recognition Examine the taxonomic relationships between the major
of only a few distinct grain proteins to fractionating and char- cereal species, as provided in this encyclopedia. Compare
acterizing thousands of proteins from all the tissues of the the associations between the proteins of these grains in
cereal grain the endosperm, germ, scutellum, and bran. Figure 8. Look at the genus names to see the origins of
This knowledge permits more efficient selection for improved the grain-specific protein names.
processing quality in the breeding of new varieties. The knowl- Obtain the amino acid compositions of the grain proteins
edge is also being applied to testing for protein quality after the for the major cereals. What are the main differences? How
grain is harvested, so that grains of specific quality can be would these differences account for functional and nutri-
segregated for appropriate marketing and processing. tional differences between these grain species?
Knowledge at the chemical level is now complementing Contrast the differences in function for an enzyme (to
new genetic insights, so that breeders and molecular geneticists catalyze a specific reaction) versus a storage protein
can employ conventional and novel methods to create quality (a source of amino acids for the new plant). Why would
types that have previously not been possible varieties with both types of protein be produced under the same strict
new protein functionality. Thus, the future holds great promise genetic control?
for grains that will be processed more efficiently with nutri- Compare the various methods of determining the amount
tional advantages. For example, it has been possible to incor- of protein in a grain or flour sample (Table 2 and else-
porate purified gluten proteins into rice dough, thus to where). Which would you expect to be the most accurate?
monitor their effects on the mixing properties.
The protein content of the cereal grains has always been
See also: Genetics of Grains: Maize: Other Maize Mutants; Maize:
important for marketing and processing. The generally desir-
Quality Protein Maize; Grain Marketing and Grading: Identification
able trait of high-protein concentration has often been
of Varieties of Food Grains; Proteins: Protein Synthesis and
inversely proportional to grain yield. Improvements in genet-
Deposition; Proteomic Analysis Relevant to Cereal Grains; The
ics and agronomic practice have tended to reduce grain pro-
Enzymes Associated with Food Grains and Their Functions in
tein levels, because high yield has been a common goal for
Processing; The Basics: Taxonomic Classification of Grain Species;
farmers.
The Grain Crops: An Overview; The Cereal Grains: An Overview of
The coming decades will bring a new factor that will further
the Family of Cereal Grains Prominent in World Agriculture; Wheat
erode protein levels the rising concentrations of carbon
Processing: The Gluten Proteins of the Wheat Grain in Relation to
dioxide in the atmosphere. For whatever reason, it is inevitable
Flour Quality; Wheat-Based Foods: Cereals: Overview of Uses:
that CO2 levels will continue to rise, although the rate of
Accent on Wheat Grain.
increase is difficult to predict. Many experiments have been
conducted to simulate effects of future levels of CO2. They have
consistently indicated increased yields of grain and biomass,
together with decreased levels of grain protein content. The Further Reading
higher atmospheric concentration of CO2 acts as a carbon
Bailey HC (1941) A translation of Beccaris lecture Concerning Grain (1729). Cereal
fertilizer, with consequent increases in starch synthesis but
Chem. 18: 555.
little change in protein synthesis. There will thus be a need to Bushuk W (ed.) (2001) Rye: Production, Chemistry and Technology, second ed. St.
rethink the role of protein in cereal grains if it will gradually Paul, MN: American Association of Cereal Chemists.
become lower in concentration. Dendy DAV (ed.) (1995) Sorghum and Millets: Chemistry and Technology St. Paul, MN:
At the manufacturing level, there is increasing interest in pro- American Association of Cereal Chemists.
Hartley H (1951) Origin of the word protein. Nature 168: 244.
viding greater protein content for cereal-based snacks and break- Henry RJ and Kettlewell PS (1996) Cereal Grain Quality. London: Chapman and Hall.
fast foods. Cereal proteins are also undergoing texturization and Islam S, Ma W, Yan G, Bekes F, and Appels R (2011) Modifying processing and health
extrusion with the aim of creating synthetic meat products. attributes of wheat bread through changes in composition, genetics and breeding.
108 PROTEINS | The Protein Chemistry of Cereal Grains
In: Cauvain SP (ed.) Bread making. Improving quality, second ed. Boston, Shewry PR and Miflin BO (1985) Seed storage proteins. In: Pomeranz Y (ed.) Advances
New York: CRC Press. in Cereal Science and Technology vol. 7, pp. 183. St. Paul, MN: American
Juhasz A, Bekes F, and Wrigley CW (2014) Wheat proteins. In: Ustunol Z (ed.) Applied Association of Cereal Chemists.
Food Protein Chemistry, Chapter 11, pp. 219304. West Sussex, UK: Sikorski ZE (2001) The Chemical and Functional Properties of Food Proteins.
Wiley-Blackwell, John Wiley & Sons. Lancaster, PA: Technomic Publishing.
Juliano BO (ed.) (1985) Rice: Chemistry and Technology, second ed. St. Paul, MN: Watson SA and Ramsted PE (eds.) (1987) Corn: Chemistry and Technology St. Paul,
American Association of Cereal Chemists. MN: American Association of Cereal Chemists.
Kent NL (1975) Technology of Cereals, with Special Reference to Wheat, second ed. Webster FH (ed.) (1985) Oats: Chemistry and Technology. St. Paul, MN: American
Oxford, UK: Pergamon. Association of Cereal Chemists.
Lasztity R (1999) Cereal Chemistry. Budapest, Hungary: Akademiai Konyvkiado. Wrigley CW (2012) Proteins the basis of life. Teach. Sci. 58: 5659.
McGregor AW and Bhatty RS (eds.) (1993) Barley: Chemistry and Technology. St. Paul, Wrigley CW and Batey IL (eds.) (2010) Cereal Grains: Assessing and Managing Quality
MN: American Association of Cereal Chemists. Oxford, UK: Woodhead Publishing Ltd.
Mertz ET (ed.) (1992) Quality Protein Maize. St. Paul, MN: AACC International.
Pomeranz Y (ed.) (1988) Wheat: Chemistry and Technology, third ed. St. Paul, MN:
American Association of Cereal Chemists. Relevant Websites
Seneweera S, Fernando N, Panozzo J, Tausz M, Norton RM, Fitzgerald GJ, Myers S, and
Nicolas ME (2014) Intra-specific variation of wheat grain quality in response to http://www.aaccnet.org American Association of Cereal Chemists.
elevated [CO2] at two sowing times under rain-fed and irrigation treatments. http://www.campden.co.uk Campden & Chorleywood Food Research Association.
J. Cereal Sci. 59(2): 137144. http://www.icc.or.at International Association for Cereal Science and Technology.
Shewry PR and Casey R (eds.). (1999) Seed Proteins. The Netherlands: Kluwer http://www.usda.gov United States Department of Agriculture.
Academic. http://www.wheat.pw.usda.gov Grain genes.
The Protein Chemistry of Dicotyledonous Grains
PR Shewry, Rothamsted Research, Harpenden, UK; University of Reading, UK
2016 Elsevier Ltd. All rights reserved.
Topic Highlights globulins are called legumins after the family Leguminosae.
The individual types of 7S and 11S globulins in different crops
The major types of storage proteins present in seeds of are also often given specific trivial names, as listed in Table 1.
dicotyledonous (noncereal) crops are globulins and It has long been recognized that members of these two
albumins. protein groups also vary in their precise S20,w values, with
Two types of storage globulin are recognized with sedimen- legumin-like globulins often having values of 12 or more
tation coefficient S20,w values of 7 and 11. (e.g., the 13S lupin globulin) and vicilin-like globulins
7S and 11S globulins have related structures and belong to 8 (e.g., in peanut). Furthermore, not all 11S globulins are
the cupin superfamily of plant proteins. readily soluble in dilute saline, with notable exceptions being
2S albumin storage proteins are related to other types of 11S storage globulins of cereals, which are either insoluble in
small sulfur-rich proteins and belong to the prolamin saline (rice) or only soluble at higher concentrations
superfamily, which includes cereal seed storage proteins. (0.81.0 M) of NaCl (oats). Consequently, the term 11S is
used in this article to describe storage proteins with a
legumin-like structure and the term 7S to describe vicilin-like
Learning Objectives proteins.
Table 1 Protein contents and types of storage proteins in major Table 2 Polypeptide molecular masses (Mr 103) of 7S and 11S
legume, oilseed, and pseudocereal crops globulins present in species
-Helical -Helical
domain 4 domain 2
Domain 3
Domain 1
Figure 2 Structure of the 7S globulin canavalin from jack bean (Canavalia ensiformis) seeds. The protein is trimeric, with a single subunit
shown in lighter gray. Domains 1 and 3 are b-barrels and domains 2 and 4 a-helical. Reproduced with permission from Shewry, P.R., Jenkins, J.,
Beaudoin, F., Mills, E.N.C., 2003. The classification, functions and evolutionary relationships of plant proteins in relation to food allergens.
In: Mills, E.N.C., Shewry, P.R. (Eds.), Plant Food Allergens. Blackwell Science, Oxford, UK.
soybean) confirm this scheme and show remarkable conserva- contain a well-defined family of albumin storage proteins with
tion of structure. Each protein consists of three subunits S20,w values of 2. These 2S albumins are major storage pro-
arranged around a threefold axis (Figure 2), with each subunit teins in a number of dicotyledonous crops (e.g., oilseed rape/
consisting of two closely related structural modules and each canola and sunflower) and minor components in others (e.g.,
comprising an antiparallel b-barrel domain and an a-helical soybean). Although they have not yet been identified in some
domain. The subunits of 7S and 11S globulins have similar dicotyledonous crops, this may reflect lack of study rather than
structures, as shown by the overlay in Figure 3, and the mature true absence, and they may well prove to be ubiquitous in
hexameric 11S globulin protein is considered to be a dimer of dicotyledonous seeds. A preliminary report of the presence
trimers. of 2S albumin storage protein in seeds of one monocotyledon-
The similarity in the structures of the 7S and 11S globulins ous species (yucca) has not been confirmed by more detailed
is consistent with the observation that both proteins can exist studies, but they are known to be structurally related to several
in trimeric forms (the mature 7S and intermediate 11S globu- other families of low-molecular-mass sulfur-rich proteins
lins) and in hexameric forms (the mature 11S globulins and including nonspecific lipid transfer proteins and trypsin/a-
the mature 7S globulins under specific conditions). It is also amylase inhibitors of cereal grain.
consistent with the fact that the 7S and 11S proteins appear to A typical 2S albumin consists of two subunits of
be stored as a homogeneous mixture in protein bodies in seeds Mr 810 kDa and 45 kDa, which are associated by two
of legumes and other dicotyledonous species. disulfide bonds. However, these two subunits are initially syn-
A final point of interest is that both 11S and 7S globulins thesized as a single precursor polypeptide, and hence, by anal-
may be allergenic, notably in soybean and peanut. ogy with 7S and 11S storage globulins, they should perhaps be
called polypeptide chains. Nevertheless, the term subunit is
widely used and will be retained here.
Albumins
The proteolytic processing to release the 2S albumin sub-
Albumins are defined as proteins that are soluble in water. units may result in the loss of short peptide sequences from the
Consequently, albumin fractions extracted from most plant N- and C-termini of the proteins and from between the two
tissues comprise complex mixtures of largely unrelated com- subunits (linker peptides). Furthermore, this proteolysis only
ponents. However, the seeds of many dicotyledonous plants occurs after the proproteins are folded and four disulfide
112 PROTEINS | The Protein Chemistry of Dicotyledonous Grains
Figure 3 Overlay of the backbone structures of single subunits of proglycinin (11S globulin precursor) from soybean (black) and phaseolin
(7S globulin) of bean (gray). Reproduced with permission from Shewry, P.R., Jenkins, J., Beaudoin, F., Mills, E.N.C., 2003. The classification, functions
and evolutionary relationships of plant proteins in relation to food allergens. In: Mills, E.N.C., Shewry, P.R. (Eds.), Plant Food Allergens.
Blackwell Science, Oxford, UK.
Signal peptide
removed
cotranslationally
21 157
NH2 S A B C COOH
Small Large subunit
subunit
N-terminal C-terminal
prosequence Linker prosequence
peptide
Peptides removed by
proteolysis in vacuole
Figure 4 Schematic sequence of the precursor protein of napin (the 2S albumin of oilseed rape) indicating peptides removed by proteolysis and
the large and small subunits of the mature protein.
Storage Proteins of Legumes the 7S conglutin b, with slightly less conglutin a (11S globulin)
and 25% of the 2S albumin conglutin d. Conglutin a has a
Many leguminous species store 7S globulin, 11S globulin, and
mass of 336 kDa and readily dissociates to a trimeric form
2S albumin proteins, but the proportions of these vary and
(7.1S, Mr 170 kDa) at low ionic strength or high pH (8.8). It
exceptions do occur. For example, Phaseolus contains 7S but
comprises acidic chain varying in Mr (within and between
not 11S globulins. Additional types of storage protein also
species) from 25 to 63 kDa linked to basic chains of
occur in some species, including various lectins and protease
Mr 21 kDa. Conglutin a is unusual among 11S legumin-
inhibitors, which can have negative impacts on nutritional
type globulins in that it is glycosylated on a single site. Con-
quality.
glutin b comprises polypeptides that range in mass between
Soybean is the most widely studied legume species. It con-
and within species from 15 to 60 kDa.
tains major groups of 11S (glycinin) and 7S (b-conglycinin)
Little detailed work has been carried out on the other
globulins, in ratios ranging from 1:1 to 6:1 depending on the
legumes/pulses discussed in this article. Both chickpea and
genotype, with only small amounts of 2S albumins. In addi-
lentil contain 11S and 7S storage globulins (in a ratio
tion, a number of other proteins can each account for up
of 6:1 in chickpea). In addition, chickpea contains a major
to 5% of the total protein in some lines, notably Kunitz-
albumin of Mr 20 kDa, which comprises subunits with Mr 10
and BowmanBirk-type protease inhibitors, lectins, urease,
and 12 kDa.
and lipoxygenase. Sedimentation coefficients and masses for
Adzuki bean is rich in 7S globulins with major components
glycinin have been reported to range from 11.8S to 13.1S and
of Mr 55 kDa, but small amounts of 11S globulins and 2S
from 30 to 380 kDa and for b-conglycinin from 6.7S to 8.0S
albumins are also present.
and from 105 to 330 kDa.
Glycinin has a typical hexameric 11S globulin structure,
with subunits comprising basic chains of Mr 20 kDa linked Storage Proteins of Oilseeds
to acidic chains varying in mass from 10 to 40 kDa. Glycinin
The two most intensively studied oilseeds are sunflower and
is also unusual in that the hexamer can undergo reversible
oilseed rape (canola). Both contain predominantly 11S-type
dissociation into trimers, depending on the ionic strength.
globulins with some 2S albumins.
b-Conglycinin comprises various combinations of three
The 12S globulins (cruciferin) of oilseed rape account
major subunits a (Mr 80 kDa), a0 (Mr 80 kDa), and b
for 40% of the total seed proteins and have Mr of
(Mr 50 kDa) resulting in seven major forms. Both 7S and
30360 kDa. They have a typical hexameric structure with
12S globulins of soybean are also thought to be allergenic.
acidic chains of Mr 3040 kDa linked to basic chains of Mr
Pea seeds contain legumins (12S to 13S, Mr 330420 kDa)
20 kDa. The 2S albumins of oilseed rape (napins) also have a
and vicilins (7S to 8S, Mr 150190 kDa). The legumin subunits
typical structure. The major napins have masses of 14.5 kDa
consist of acidic chains of Mr 25 kDa and 40 kDa linked to
(subunit Mr of 9 and 4 kDa) but minor forms of
basic chains of Mr 20 kDa. The 7S vicilins comprise subunits
Mr 12.5 kDa also occur. Napin processing includes the loss
of Mr 50 kDa, but proteolysis at one or two sites and glyco-
of N-terminal, C-terminal, and linker peptides, as discussed
sylation at a single site may occur, resulting in polypeptide
earlier.
chains varying in Mr (1250 kDa). Larger subunits of
Sunflower, a member of the family Compositae, contains
Mr 70 kDa also occur, which form a second trimeric 7S pro-
12S globulins (helianthinin) and 2S albumins in a ratio
tein called convicilin. Pea seeds also contain two major albu-
of 2:1. The Mr 300 kDa helianthinin comprises acidic chains
mins, PA1 and PA2, of which the former may be a 2S storage
of Mr 3234 kDa (a and a0 chains) linked to basic chains of
albumin. PA1 accounts for 10% of the total seed protein but
Mr 2227 kDa (b chains). Sunflower 2S albumins comprise
half of the total sulfur, in the form of cysteine ( 11 mol.%).
1113 components with masses of 1018 kDa. All are single-
The dominant peanut proteins are arachin (13S14S, Mr
chain proteins (i.e., they lack a proteolytic cleavage site
330400 kDa) and conarachin (8S to 9S, Mr 140190 kDa).
between the putative large and small subunits), but most are
The subunits of arachin comprise acidic chains of
synthesized as pairs, two albumins being released from a single
Mr 1448 kDa linked to basic chains of Mr 21.4 kDa,
precursor protein by proteolysis. One sunflower albumin that
whereas conarachin comprises polypeptide chains ranging in
is synthesized from a separate precursor protein is called SFA8.
Mr from 23 to 75 kDa. It is probable that both large and
This contains 16 methionine and 8 cysteine residues in a
typical conarachin subunits are present, as in soybean and
mature protein of 103 residues (i.e., a total of 23 mol.% of
pea, with proteolytic processing contributing to the polymor-
sulfur-containing amino acids). Safflower, a related member of
phism. Both proteins also exhibit complex association/disso-
the family Compositae, also appears to contain major groups
ciation behavior, depending on the ionic strength. The major
of 12S globulin (called carmin) and 2S albumin storage
peanut allergen Ara h 1 is a glycosylated 7S globulin with
proteins, but little detailed work has been carried out.
Mr 63.5 kDa, whereas Ara h 3 is an 11S globulin.
Cottonseed is unusual among nonleguminous dicotyle-
Most of the knowledge of peanut albumins comes from the
donous species in containing 7S globulin storage proteins as
analysis of genomic clones and is related to their role as aller-
well as 11S globulins and 2S albumins, with most of the
gens (Ara h 2, 4, 6, and 7). They show high homology to 2S
knowledge being based on DNA-derived sequences with little
albumins from other legumes (soybean and lupin) and are
direct analysis of the seed proteins. Linseed (and linola) has
presumed to have a typical heterodimeric structure.
not been widely studied but the major seed protein is reported
The salt-soluble (i.e., albumin and globulin) proteins of
to be a 12.7S globulin of Mr 250300 kDa. SDS-PAGE
lupins are called conglutins. The major component is usually
showed six nonidentical subunits.
114 PROTEINS | The Protein Chemistry of Dicotyledonous Grains
Storage Proteins of Pseudocereals Compare the contents of albumin and globulin storage
proteins in the major types of dicotyledonous crops:
Pseudocereals are not true cereals but three dicotyledonous
legumes and pulses, oilseeds, and pseudocereals.
crops that produce small grain-like seeds. Consequently, their
seed proteins are similar to those of other dicotyledonous
What are the relationships of albumin and globulin storage
proteins to other grain proteins?
plants rather than related to cereal seed storage proteins.
Seeds of amaranth (Amaranthus spp., Amaranthaceae)
contain 13S legumin-type globulins (amaranthin) and 8S
Exercises for Readers to Explore the Topic Further
vicilin-type globulins (S20,w value of 8.6) (conamaranthin).
In addition, albumin proteins with S20,w values of 2 are
present, with SDS-PAGE showing major bands of Mr 9 kDa
How do the structures and compositions of storage proteins
determine the functional properties of the seeds for food
and 4 kDa. This is consistent with the presence of typical 2S
processing and their role in the human diet?
albumins. Two methionine-rich proteins with Mr 18 kDa
have also been reported. Quinoa (Chenopodium quinoa,
How can these structures and properties be manipulated
(by breeding or GM technology) to improve their quality
Chenopodiaceae) seeds also contain 11S globulins (chenopo-
for nutrition and processing and to reduce adverse reactions
din), of Mr 320 kDa, and 2S albumins, the latter being rich in
in humans (allergies and intolerances)?
cysteine (15.6 mol.%) but low in methionine.
In buckwheat (Fagopyrum esculentum, Polygonaceae), the
major 13S legumin-like globulin is located in the cotyledons See also: Oilseed and Legume Processing: Canola: Processing;
with smaller amounts of putative 8S globulins in the peri- Soybean: Soy Concentrates and Isolates; Proteins: Protein Synthesis
sperm. Methionine-rich ( 9 mol.%) 2S storage albumins are and Deposition; The Protein Chemistry of Cereal Grains; The Basics:
also present. Taxonomic Classification of Grain Species; The Legumes and
Pseudocereals: Amaranth: Overview; Beans: Overview; Buckwheat:
Overview; Chickpea: Overview; Lupin: Overview; Pea: Overview;
Conclusions Pseudocereals: Overview; Quinoa: Overview; The Oilseeds: Oilseeds:
Overview; Sunflower: Overview; Wheat Processing: The Gluten
The 2S albumins, 7S globulins, and 11S globulins are major Proteins of the Wheat Grain in Relation to Flour Quality; Pulses,
groups of storage proteins that are widely distributed in seeds Overview.
of dicotyledonous plants, although their relative amounts vary
and other types of storage proteins also occur in some species.
Further Reading
Consequently, they largely determine the nutritional quality
and processing properties of the seeds. A detailed knowledge of Bewley JD and Black M (1999) Seeds Physiology of Development and Germination.
their structures and properties is therefore essential in order to New York: Plenum.
elucidate structurefunction relationships. This will facilitate Black M and Bewley JD (2000) Seed Technology and its Biological Basis. Sheffield:
Academic Press.
the optimization of seed quality for traditional and novel
Boulter D and Croy RRD (1997) The structure and biosynthesis of legume seed storage
products with improvement using classical genetic or genetic proteins: a biological solution to the storage of nitrogen in seeds. Adv. Bot. Res.
engineering approaches. 27: 184.
Salunkhe DK, Chavan JK, Adsule RN, and Kadam SS (1992) World Oilseeds Chemistry,
Technology, and Utilisation. New York: Van Nostrand Reinhold.
Shewry PR (1995) Plant storage proteins. Biol. Rev. 70: 375426.
Exercises for Revision Shewry PR and Casey R (eds.) (1999) Seed Proteins. Dordrecht: Kluwer.
Shewry PR, Jenkins J, Beaudoin F, and Mills ENC (2003) The classification, functions
and evolutionary relationships of plant proteins in relation to food allergens.
Compare the structures and properties of albumin and In: Mills ENC and Shewry PR (eds.) Plant Food Allergens. Oxford: Blackwell
globulin storage proteins. Science.
Protein Synthesis and Deposition
PR Shewry, Rothamsted Research, Harpenden, UK; University of Reading, Reading, UK
P Tosi, University of Reading, Reading, UK
2016 Elsevier Ltd. All rights reserved.
Table 1 Summary of the characteristics of albumin, globulin, and prolamin storage proteins
Source: Shewry PR (2000) Seed proteins. In: Black M and Bewley JD (eds.) Seed Technology and Its Biological Basis, pp. 4284. Sheffield: Blackwell Science and Sheffield Academic
Press, with permission.
Storage protein
synthesis occurs on
ribosomes attached to ER
RER
Smooth
ER
Transport Protein
in body
vesicles formation
within RER
Golgi
apparatus
Protein
body
formation
Secretory
vesicle Within
Plasma vacuole
membrane
Figure 1 Schematic summary of storage protein synthesis, processing, trafficking, and deposition in the endomembrane system of the cell.
These vacuoles may subsequently divide to form protein bod- In rice and maize (and probably also in sorghum and mil-
ies. However, all three types of storage protein may also lets), prolamins accumulate directly within the lumen of the ER
undergo modification, either in the vacuole or during their to form a population of protein bodies surrounded by a mem-
passage through the ER and Golgi apparatus. brane of ER origin, which may remain studded with ribosomes.
The 2S albumins are synthesized as precursor proteins, In contrast, it appears that two separate pathways occur in wheat,
which undergo folding and disulfide bond formation in the barley, and rye, with some prolamins being trafficked through
ER lumen. The precursors are then proteolytically processed in the ER and Golgi apparatus and deposited in storage vacuoles (as
the vacuole to generate the mature two-subunit structure. This described earlier for albumin and globulin storage proteins),
processing involves cleavage adjacent to specific asparagine while others are retained in the ER as in maize and rice. It has
residues and the loss of one or more short peptide fragments been suggested that the polymeric prolamins are preferentially
(from the protein N-terminus and C-terminus and between the retained in the ER and monomeric forms transported to vacu-
two subunits). oles, but it seems likely that the division is not so clear-cut. It is
The 7S and 11S globulin subunit is also folded in the ER also probable that the relative amounts of prolamins deposited
lumen with the formation of a single intrachain disulfide bond in the two types of protein body vary with the rate of protein
in the latter (7S globulins lack cysteine residues and hence have synthesis and with the developmental state of the tissue. Thus, it
no disulfide bonds). Both types of subunits are then assembled can be envisaged that high levels of protein synthesis would lead
into trimeric structures stabilized by noncovalent forces. There to a high level of accumulation within the ER lumen and that this
has been considerable debate about the extent to which route would also be favored as the cells become distended with
globulin storage proteins are modified by the addition of deposits of starch and proteins. In wheat, the individual protein
oligosaccharides (glycosylation). It is now established that bodies collapse during the later stages of seed maturation form-
this rarely, if ever, occurs for 11S globulins but that some 7S ing a continuous matrix in the cells of the mature grain.
globulin subunits are modified by glycosylation of specific
asparagine residues (N-glycosylation). This occurs in several
stages, with the initial attachment of a high-mannose sugar What Determines the Destination of Storage Proteins
occurring cotranslationally (i.e., concurrent with folding and in the Endomembrane System?
disulfide bond formation). Subsequently, enzymes present in
the ER and Golgi apparatus may act to remove or add further As discussed earlier, trafficking through the endomembrane
sugar residues to give a more complex structure with final system is a highly regulated process with the destinations of
trimming occurring within the storage vacuole. Once within proteins being determined by their sequences and structures.
the vacuole, the 11S globulins are proteolytically cleaved at an The retention of storage proteins within the ER or their
internal asparagine residue to give acidic and basic chains that transport to the vacuole should, therefore, result from the pres-
remain associated by the single disulfide bond formed in the ence of specific signals. A number of proteins are permanently
ER. After this proteolytic processing, the intermediate dimers resident in the lumen of the ER, including PDI and BiP. In these
formed in the ER are able to assemble to form the mature cases, retention results from the presence of a specific ER reten-
hexameric structure. tion signal, which is a tetrapeptide located at the protein
The 7S globulin subunits may also be proteolytically pro- C-terminus. The two most widespread tetrapeptides are KDEL
cessed in the vacuole, but this depends on the species and the and HDEL, which correspond to the amino acids lysine or
precise protein subunit. For example, in bean (Phaseolus histidine, aspartic acid, glutamic acid, and leucine. There is no
vulgaris) and soybean (Glycine max), the 7S globulin subunits evidence that such a retention signal is present at the C-termini
do not undergo any proteolysis, while in pea (Pisum sativum), of prolamins that are retained in the ER, and a different mech-
some subunits are processed at one or two sites to give anism must therefore be sought. It has been suggested that BiP
polypeptide chains ranging in mass from 12 to 35 kDa, in plays a role in rice, by transiently binding to prolamins and
addition to unprocessed chains of mass 50 kDa. However, retaining them in the ER to allow assembly into protein bodies.
the trimeric structure is not affected by these processing events. However, this mechanism has not so far been shown to be
applicable to other cereals. An alternative explanation is that
retention results from the insolubility of the proteins in aqueous
environments (i.e., the ER lumen) and their propensity to form
Cereal Prolamins Are Deposited in Two aggregates under such conditions. It can be envisaged that such
Types of Protein Body aggregates would accumulate directly in the ER lumen and then
be budded off to form protein bodies.
Cereal prolamins do not undergo glycosylation or proteolytic Some proteins are targeted to the vacuole by the presence of
processing (except for cotranslational removal of the signal specific prosequences that are removed by posttranslational
sequence), but some subunits are assembled into high- proteolysis once the protein arrives in the vacuole. These pro-
molecular-mass polymers stabilized by interchain disulfide sequences may be at the protein N-terminus, as in the barley
bonds. In wheat, these glutenin polymers may have masses of cysteine proteinase aleurain and the sweet potato storage pro-
between 1 and 10 MDa, and it is not known how such complex tein sporamin (which is stored in tubers) or at the C-terminus
multisubunit structures are assembled. However, it is probable as in lectins; more rarely, a noncontiguous series of residues
that assembly and interchain disulfide bond formation occur form a tridimensional structure in the mature protein, which
rapidly after synthesis within the ER lumen. Subsequently, pro- acts as a vacuolar sorting signal. None of the 2S albumins, 7S
lamins are deposited in two discrete types of protein body. globulins, 11S globulins, or prolamins that are deposited in
PROTEINS | Protein Synthesis and Deposition 119
interact strongly with each other and with the monomeric storage tissue(s) and cease when these cells mature and desic-
gliadins by noncovalent forces, particularly hydrogen bonds. cate. The onset and duration of the phases vary widely between
This combination of covalent and noncovalent interactions, species and genotypes and are greatly affected by the environ-
together with the intrinsic structures of the individual subunits, mental conditions. In small grain cereals (wheat and barley)
results in the viscoelastic properties associated with wheat grown in the United Kingdom, storage protein synthesis is
dough and gluten. usually first detected in appreciable amounts at 1214 days
The limited information that is available indicates that after pollination and is most active over the next 23 weeks.
glutenin polymers have defined structures, with specific inter- However, synthesis may be initiated as early as 6 days in plants
actions between individual subunits. Similarly, it is clear that grown in hot and dry climates. In maize (corn) grown in North
some allelic forms of HMW subunits are associated with the America, storage protein synthesis occurs between 3 and 7
formation of larger polymers, leading to positive effects of weeks after pollination.
quality. However, the mechanisms responsible for defining Storage protein fractions are, of course, not single proteins
specific structures are not known. but mixtures of components, which may be encoded by several
Finally, the proportion of high-molecular-mass glutenin multigene families. In general, these genes show highly coor-
polymers increases dramatically during the desiccation phase of dinated patterns of expression, with only small differences in
grain development, suggesting that polymer formation is also their timing and relative levels of expression during develop-
driven by dehydration. This is consistent with the model that ment. For example, in sunflower, the expression of genes for
glutenin polymers are stabilized by strong arrays of hydrogen 11S globulin storage proteins (helianthinins) is detected
bonds, but rearrangements of disulfide bonds may also occur. slightly before that of genes for 2S albumins, but in wheat
and barley all groups of prolamin storage proteins show sim-
ilar expression patterns.
Regulation of Storage Protein Synthesis
Tissue Specificity Environmental Control
Storage protein synthesis in seeds is strictly regulated with Environment has a major impact on seed development, includ-
respect to timing, tissue specificity, and, to a lesser extent, nutri- ing storage protein synthesis, with higher temperatures
ent availability. In addition to the embryo, which is a diploid generally increasing the rate of development. In addition,
zygotic tissue that gives rise to the seedling, all seeds also contain specific heat-shock effects may occur when temperatures
an endosperm, which is triploid and is derived from a second exceed 35 C: these can result in detrimental effects on the
fertilization event. However, the size of the endosperm varies quality of wheat grown in Australia and some other parts of the
greatly, forming 8090% of the mature cereal grain but being world. The most dramatic effect of environment on storage
largely broken down during development in soybean and many protein synthesis is that of nutrition. It is well established
other seeds. Similarly, whereas some seeds such as quinoa and that the protein content of seeds is regulated by the availability
amaranth contain a perisperm (derived from the maternal of nitrogen, with storage protein acting as a sink for nitrogen,
nucellar tissue), this fails to develop and is absorbed in most which is not required by the plant for other purposes. For
species. In all seeds, at least one of these three tissues functions example, increasing the total nitrogen content of barley grain
as a storage tissue, and in some cases, more than one. The central from 1% to 3% is associated with an increase in prolamin
part of the endosperm is the major storage tissue in cereals, (hordein) storage proteins from 35% to 50% of the total
accumulating protein (prolamins with varying amounts of grain nitrogen. This effect can be exploited to grow grain with
11S-type globulins), and starch. However, 7S globulins and oil defined protein contents for specific end uses (e.g., low protein
are also stored in the outer cells of the endosperm (the aleurone for malting barley and high protein for bread-making wheat).
layer) and the single cotyledon (scutellum) of the embryo. The However, almost all storage proteins also contain sulfur in
cotyledons of the embryo form the major storage tissue in grain the form of the amino acids cysteine and methionine. Many
legumes (including soybean, peanut, lupin, and pea), species are able to adjust to variation in the relative availabil-
sunflower, and oilseed rape/canola, while proteins and oil are ities of nitrogen and sulfur by having specific types of seed
stored in the endosperm and embryo of quinoa with starch storage proteins with high and low contents of sulfur amino
being stored in the perisperm. acids. Thus, the 2S albumin storage proteins of oilseed rape
It should also be noted that the synthesis of seed storage and sunflower have higher contents of cysteine and methio-
proteins has never been detected in nonseed tissues of the nine than the globulin storage proteins, and the synthesis of
plant, whether by determination of gene expression (i.e., the former is therefore reduced under conditions of low sulfur
mRNA populations) or by protein accumulation. availability (Figure 3(a) and 3(b)). Similarly, restricted avail-
ability of sulfur favors the synthesis of the sulfur-poor C hor-
deins of barley (Figure 3(c)) and o-gliadins of wheat over
Temporal Regulation
other more sulfur-rich prolamins, which can have adverse
Seed development proceeds through a series of genetically consequences for grain quality in the latter species.
programmed stages, starting with cell division and differentia-
tion to establish the basic structure of the component organs
Regulation of Storage Protein Gene Expression
and tissues (i.e., the embryo and endosperm). The synthesis
and accumulation of storage products, including proteins, The regulation of seed storage protein genes usually occurs at
occur during the subsequent expansion of the cells of the the level of gene transcription, although some fine-tuning can
PROTEINS | Protein Synthesis and Deposition 121
Conclusions
3
It is clear that we still have much to learn about the precise
mechanisms of storage protein synthesis, trafficking, and depo-
sition, although the broad principles are now understood.
A detailed understanding of these mechanisms is important
as interactions (i.e., between proteins and with other grain
components) established during synthesis and deposition
may ultimately affect the functional properties of the proteins
(a) + (b) + (c) + in food products. Furthermore, it is becoming clear that the
synthesis, assembly, deposition, and interactions of seed pro-
Figure 3 The effect of sulfur on the total seed proteins of oilseed rape
teins may be affected by environmental factors that ultimately
(a) and sunflower (b) and on the prolamin (hordein) proteins of barley
impact on seed quality. Identifying the major sites of these
(c). Note the presence of S-rich low-molecular-mass proteins (probably
2S albumins) in oilseed rape and sunflower (bracket) and of S-poor C effects and the critical stages of seed development should
hordeins (2) in barley. Parts (a) and (b) are reproduced from Spencer D, allow them to be predicted and ultimately manipulated to
Rerie WG, Randall PJ, and Higgins TJV (1990). The regulation of pea give a higher level of stability.
seed storage protein genes by sulfur. Australian Journal of Plant It is also important to understand the mechanisms of seed
Physiology 17: 355363, with permission. Part (c) is reproduced from protein synthesis, trafficking, and deposition in order to facil-
Shewry PR, Franklin J, Parmar S, Smith SJ, and Miflin BJ (1983) The itate the application of genetic engineering to improve grain
effects of sulfur starvation on the amino acid and protein composition of quality. It is essential that any novel or mutant proteins
barley grain. Journal of Cereal Science 1: 2131, with permission. expressed in transgenic plants should be accepted by the
secretory system as authentic and transported to their correct
occur at the level of translation of the mRNA into proteins. destination. Any errors in folding or failure to contain the
Consequently, interest has focused on the identification of correct targeting information could lead to protein turnover
regulatory sequences present upstream of the gene sequences or accumulation in incorrect compartments (e.g., ER instead of
that encode storage proteins (called 50 upstream sequences) storage vacuoles in dicotyledonous seeds) with adverse effects
and of specific proteins that bind to these (transcription fac- on grain development, quality, and yield.
tors). This has led to the identification of short conserved
nucleotide sequences (often called boxes) that regulate expres-
sion of the genes. Exercises for Revision
Comparison of the 50 upstream sequences of a number of
genes encoding 11S storage globulins has led to the identifica-
tion of a 28 base pair (bp) legumin box. Deletion of a central
Summarize the compartments of the endomembrane sys-
tem of the plant cell in relation to the trafficking and
7-nucleotide motif from the center of this box resulted in a processing of seed storage protein.
drastic reduction in the gene expression in transgenic plants,
demonstrating its importance. Related boxes are also present
Discuss the origins of protein bodies in cereal and dicoty-
ledonous seeds.
in the 50 upstream sequences of 7S globulin and 2S albumin
genes. However, further studies have demonstrated the pres-
Discuss the pathways and mechanisms of processing and
deposition of different types of storage proteins: albumins,
ence of several additional upstream regulatory sequences that globulins, and prolamins.
may modulate storage globulin gene expression.
Similar studies have shown that many prolamin genes also
contain a conserved sequence of 30 bp located about 300 bp
upstream of the gene-coding region. This is called the Exercises for Readers to Explore the Topic Further
endosperm box or prolamin box and is unrelated in
sequence to the legumin box. The prolamin box actually Identify the structure/sequences that determine storage pro-
comprises two conserved sequences separated by a more vari- tein targeting and trafficking in the endomembrane system.
able region. One of these conserved sequences is thought to be Determine the organization and interactions of storage pro-
responsible for conferring endosperm specificity, while the teins within protein bodies.
other may play a role in the regulation of the gene. However, Determine the precise mechanism of storage protein gene
additional regulatory sequences are undoubtedly also present regulation in the context of grain development, the nutri-
including a short sequence (the E motif) in the a-zein genes of tional status of the plant, and abiotic stress.
122 PROTEINS | Protein Synthesis and Deposition
Shewry PR (2000) Seed proteins. In: Black M and Bewley JD (eds.) Seed Technology
See also: Proteins: The Protein Chemistry of Cereal Grains; The
and its Biological Basis, pp. 4284. Sheffield: Blackwell Science and Sheffield
Protein Chemistry of Dicotyledonous Grains; Wheat-Based Foods: Academic Press.
Cereals: Overview of Uses: Accent on Wheat Grain. Shewry PR (2003) Plant proteins. In: Thomas B, Murphy D, and Murray B (eds.)
Encyclopaedia of Applied Plant Science, pp. 889896. London: Elsevier
Science.
Shewry PR and Casey R (1999) Seed Proteins. Dordrecht: Kluwer Academic.
Further Reading Shewry PR and Halford NG (2002) Cereal seed storage proteins: Structures, properties
and role in grain utilization. Journal of Experimental Botany 53: 947958.
Bechtel D, Wilson JD, and Shewry PR (1991) Immunocytochemical localization of the Shewry PR, Franklin J, Parmar S, Smith SJ, and Miflin BJ (1983) The effects of sulphur
wheat storage protein triticin in developing endosperm tissue. Cereal Chemistry starvation on the amino acid and protein compositions of barley grain. Journal of
68: 573577. Cereal Science 1: 2131.
Bewley JD and Black M (1995) Seeds Physiology of Development and Germination, 2nd Shewry PR, Halford NG, Tatham AS, Popineau Y, Lafiandra DL, and Belton PS (2003)
edn. New York: Plenum Press. The high molecular weight subunits of glutenin and their role in determining
Buchanan BB, Gruissem W, and Jones RL (2000) Biochemistry and Molecular Biology wheat processing properties. Advances in Food and Nutrition Research
of Plants. Rockville, MD: American Society of Plant Physiologists. 45: 221302.
Larkins BA and Vasil IK (1997) Cellular and Molecular Biology of Plant Seed Spencer D, Rerie WG, Randall PJ, and Higgins TJV (1990) The regulation of pea seed
Development. Dordrecht: Kluwer Academic. storage protein genes by sulphur. Australian Journal of Plant Physiology
Lending CR, Chesnut RS, Shaw KL, and Larkins BA (1989) Immunolocalization of 17: 355363.
avenin and globulin storage proteins in developing endosperm of Avena sativa L. Tosi P (2012) Trafficking and deposition of prolamins in wheat. Journal of Cereal
Planta 178: 315324. Science 56: 8190.
Muntz K and Shutov AD (2002) Legumains and their functions in plants. Trends in Plant Yamagata H and Tanaka K (1986) The site of synthesis and accumulation of rice storage
Science 7: 340344. proteins. Plant and Cell Physiology 27: 135145.
Nitrogen Metabolism
CA Atkins, The University of Western Australia, Crawley, WA, Australia
2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition, volume 2, pp. 287292, 2004, Elsevier Ltd.
Topic Highlights stage we must infer the commonality of processes that are so
far only adequately documented for a few species.
Mature plant seeds contain nitrogen-based nutrient
reserves, particularly proteins.
Nitrogen is transported into grain or seed, where it is
metabolized into amino acids. Nitrogen Nutrition of Developing Seeds
Most nitrogen entering the seed is sued to synthesize stor-
age proteins, which are utilized by the germinating seedling Before considering seed nutrition it is useful to review the basic
(or enter into food processing). structural features of seeds and the consequences of this struc-
ture to solute import into and transfer within the component
tissues of a seed as it develops. The outer seedcoat tissues are
Learning Objectives maternal while the endosperm and embryo within this coat are
filial. There is no vascular connection between the two. Thus,
To understand the dynamics of nitrogen metabolism that whether stored reserves of the seed accumulate in the endo-
results in the storage of useful plant proteins in the mature sperm, the embryo, or associated tissues, in all cases solutes
grain. that supply C and N (assimilates) pass across an apoplastic
To understand the transportation of nitrogen-containing compartment. It follows that in receiving translocated solutes
solutes into the seed, in relation to the concept of sink before they pass to the embryo the seedcoat may regulate both
strength. the rate and nature of solute transfer.
To understand the metabolism of nitrogen within the seed Although developing seeds and their surrounding support-
and the formation of storage reserves. ing structures may have the capacity for photosynthesis during
early stages of development, this is limited. Consequently their
major source of C is assimilates translocated in phloem and
Introduction xylem. These are principally in phloem as sucrose, longer
raffinose-based oligosaccharides, or in some species cyclitols,
The success that flowering plants enjoy is in no small way a like mannitol and sorbitol. The same translocation streams
consequence of the effectiveness of their seed as a perennating supply N but typically phloem contains 1020 times more N
organ. The embryo is in a metabolically dormant state and is than xylem. However, even though both streams may carry a
accompanied by a reserve of nutrients that can be mobilized on wide array of amino acids, as well as other N-solutes (e.g.,
germination to support the activated meristems of the new proteins, nucleic acids, plant growth regulators), usually one
plant. The diversity of composition exhibited by mature seeds or two low molecular weight organic compounds predomi-
reflects variation in the size and nature of this stored reserve. nate. For many dicotyledonous species, including temperate
Some are high in carbohydrate, others in protein, and others in legumes, these are the amides, glutamine and asparagine or
oil. Yet all are viable and are able to convert the reserves into their parent acids, glutamate, and aspartate, especially in
translocatable solutes with a balance of C and N that supports cereals. In some species arginine, proline, or non-protein
the active root and shoot meristems before uptake from soil amino compounds like homoserine, ornithine, citrulline,
provides an alternative source of nutrients and before the canavanine, and putrescine predominate. In others, particu-
seedling is autotrophic for C through photosynthesis. The larly nodulated legumes of tropical origin, the ureides, allan-
reserves are fashioned from organic assimilates that supply C, toin, and allantoic acid, account for almost all the N in xylem
N, and S through the long distance translocation channels of and a significant component in phloem-N. There are some very
the vegetative structure of the plant. unusual N-solutes that translocate the bulk of N, for example
Metabolism of N has been studied in a restricted range of djenkolic acid (two cysteines bridged at their S groups by a
seed types, mostly those of cereals, legumes, and oilseeds of methylene) occurs at very significant levels in the xylem of
agricultural importance and more recently in the model species some Acacia spp., and no doubt the plant kingdom employs
Arabidopsis thaliana. Because of the high proportion of protein many other forms of translocated N that have yet to be discov-
stored in legume seeds these have received most attention in ered. Thus protein synthesis in sink organs does not receive a
this regard. However it is well to realize that plants produce an mixture of amino acids that can be simply used to form pro-
almost bewildering array of N-containing compounds, many tein, but rather a mixture in which just one or two solutes
of which are amino compounds that accumulate in seeds and provide the bulk of N that has to be metabolized to generate
are likely to impart unique nutritional, toxicological and phar- the appropriate proportions of the 20 protein amino acids. In
macological properties. Our understanding of this whole area their passage from sites of phloem unloading to the embryo,
in angiosperms is thus limited and fragmentary, and at this N-solutes may be transiently stored, for example in the
seedcoat, and are metabolized in highly specific ways within translocation stream passes through stem segments immedi-
the tissue compartments they traverse. ately before entering the inflorescence. Similar specific
There seems little doubt that almost all the N entering seeds is exchanges of amino acids have also been found for developing
in an organic form. Although the literature records quite high cowpea fruits/seeds. The precise tissue or cellular site where
levels of ammonium ion in both xylem and phloem, it is not these transfer mechanisms act and the molecular basis for their
likely that this is the norm and may, in many cases, be a conse- regulation is unknown but a role for transfer cells has been
quence of hydrolysis of the amide groups of glutamine and inferred.
asparagine during or subsequent to collection of vascular exu- The developing seed is supported on or within fruiting
dates. Xylem transports considerable quantities of nitrate from structures that not only bear the vasculature which brings the
the root system to the major transpiring surfaces of the shoot but translocation streams to the seed but which serve also as inter-
negligible quantities of this inorganic ion are transferred to mediate sites for metabolism and transient accumulation of
phloem. As a result, fruit tissues and especially developing seeds nutrients, including N. For example, in the period from anthe-
are unlikely to receive significant levels of nitrate and conse- sis to 11 days after anthesis, in cowpea fruits 21 mg N enters in
quently exhibit low or negligible capacity for its assimilation. phloem and 17 mg N in xylem. Of this N less than half
Early in legume fruit/seed development when a significant (15 mg) reaches the developing embryo and endosperm, the
proportion of translocated N is delivered in xylem, the C:N rest is sequestered into the pod wall (16 mg) and the testa
ratio of incoming solutes is more narrow than that required by (7 mg). Importantly, however, the incoming N-solutes are
the developing tissues, and excess N is transiently stored. The metabolized extensively before being transferred to the
endosperm in many dicotyledonous seeds accumulates a sig- embryo. At this early stage of development xylem-N is 70%
nificant proportion of this N as ammonium ion and levels ureide and phloem-N 20% ureide, but ureides account for
approaching 0.1 M have been recorded for periods of 34 less than 20% in the podwall and testa and no more than 2%
weeks in white lupin (Lupinus albus) seeds. Although the of soluble-N in the liquid endosperm or embryo. Later in
ammonium is promptly reassimilated when the developing development (1222 days after anthesis) phloem accounts
embryo expands to fill the endospermic apoplast, such high for 80% of N entering the fruit and, although the ureide level
levels are normally regarded as extremely toxic and mecha- is low, ureide-N accumulates preferentially in the seedcoat.
nisms that prevent this ammonium entering the embryo to Despite this enrichment literally no ureide is transferred to
any great extent must be functioning at this time. In pea the apoplast surrounding the embryo. Rather, ureide-N is con-
(Pisum sativum), the major forms of accumulated N in the verted to glutamine and this is the major source of N for
endosperm are homoserine, glutamine, and alanine, in cow- protein synthesis. Similar observations have been made for
pea (Vigna unguiculata) endosperm there is negligible ammo- soybean, another plant that translocates significant levels of
nium ion but 3055% of the N is histidine, while in chickpea ureide-N in xylem and to a lesser extent in phloem.
(Cicer arietinum) ammonium and citrulline predominate. The In pea a major N-solute entering the seedcoat in phloem is
significance of ammonium or other N solutes that accumulate homoserine and, although this may accumulate transiently in
to such high levels in endosperm when embryo development is endosperm, most is converted to threonine and glutamine in
characterized by cell division is unknown, but it is interesting the seedcoat during embryo expansion. Even in species that
to note that coconut endosperm (milk), used in pioneering have an asparagine-rich phloem stream reaching the seedcoats,
studies of plant cell and tissue culture, contains high concen- a significant proportion of asparagine-N is converted to
trations of N, including ammonium ion, along with plant glutamine-N during passage to the apoplast and embryo.
growth regulators. Whether this apparent preference for N as glutamine in the
A positive correlation has been established between rates of embryos nutrition reflects some metabolic requirement or the
ammonia emission (i.e., N loss) and ammonium concentra- specificity of transporters involved in transferring solutes
tion in leaves. There are no emission data available for devel- across tissue compartments of the seed has yet to be clearly
oping seeds/fruits, but given the high levels of ammonium ion resolved. In maize the endosperm of the developing kernel
that may accumulate in liquid endosperm it seems possible supplied assimilates through the pedicel and even though the
that transient periods of N loss occur due to its volatilization. It major phloem N-solute is aspartate this is largely metabolized
is perhaps not too surprising then that the highest rates of with its N being transferred to glutamine. The maize pedicel
ammonia emission in barley occur during grain filling. expresses a unique cytosolic form of glutamine synthetase,
The mix of solutes in phloem is not constant during repro- GSp1, which is developmentally regulated, increasing as storage
ductive growth, the C:N ratio changing to more closely match protein synthesis increases. Glutamine is the major amino acid
the C and N requirements of the developing seed. In white released from the pedicel to the apoplast surrounding the
lupin, sucrose levels in phloem decline progressively after endosperm and a transporter expressed in the basal endosperm
anthesis while the concentration of amino compounds transfer cells apparently enhances its uptake. Thus specificity of
increases, almost doubling over the period of grain filling. both metabolism and transport combine to ensure that a pre-
This pattern of change is consistent with a declining rate of ferred N-solute reaches the embryo tissues.
photosynthesis and increasing senescence of leaves, with
release of protein-N as amino compounds, during reproduc-
tive development. However, further studies with lupin have Transport of N-Solutes within Seeds
shown that in fact the phloem N level also increases to narrow
the C:N ratio as a consequence of solute specific mechanisms Rates at which solutes that supply C and N to the embryo are
that achieve an enrichment of phloem with asparagine as the transported are likely to contribute to determining the
PROTEINS | Nitrogen Metabolism 125
potential accumulation of assimilates or sink strength of seeds The Arabidopsis genome has revealed transporters for other
and consequently grain yield and harvest index. Studies with types of N-solute that are expressed and could have significant
four cultivars of bean (Phaseolus vulgaris) that differed in their roles in seed metabolism. These include oligopeptide trans-
rates of seed growth found that although sink size, as assessed porters, such as AtPTR2-B, that might be responsible for the
by surface area and volume of cotyledons, were major deter- movement of peptides released from storage proteins during
minates of growth rate, the rate of transfer of dry matter was germination to new meristems or in the transport of signal
also a contributing factor. Not surprisingly, maximal sucrose peptides from sites of synthesis to sites of action. The possible
fluxes into cotyledons were positively correlated with levels of importance of peptide transport in seed development was
expression of the sucrose/H symporter and accompanying highlighted in a study where antisense lines for AtPTR2-B
H-ATPase in the dermal cells facing the apoplast between were generated in Arabidopsis. The transgenic plants showed
the cotyledons and the seedcoat. Specifically in relation to normal silique development but formed fewer, larger seeds
N nutrition, activity of amino acid transporters has been sug- than controls. Specifically about half of the embryos were
gested to regulate the rate of seed protein synthesis. Although arrested at an early stage of their development, suggesting
most research into solute unloading from phloem and trans- peptide transport was essential for N nutrition at this time or
port within developing seeds has involved sucrose, some of the that transfer of some bioactive peptide, essential for embryo-
amino acid transporters expressed in seed tissues have been genesis, was limiting.
identified and their location studied in relation to likely Plants also express high affinity transporters specific for
function. purines and purine derivatives (PUPs) including bases, nucle-
In pea, two cDNA clones (PsAAP1 and 2) belonging to osides, purine alkaloids, and cytokinins. Northern analysis for
the AAP (amino acid permease) family of H/amino acid AtPUP1 in Arabidopsis has however shown only a low level of
co-transporters have been isolated from cotyledons. Like the expression in developing siliques compared to other tissues
sucrose transporter, PsAAP1 is highly expressed in special- and their functional significance in seed development or in
ized epidermal transfer cells abutting the seedcoat and the mobilization of N during germination remains to be
shows broad specificity, mediating transport of acidic, neu- established.
tral, and basic amino acids. Similar broad-spectrum mem- A novel superfamily of transporters has recently been
bers of the AAP family have been described for developing described for allantoin and other oxidized derivatives of het-
Arabidopsis and castor bean seeds. In Arabidopsis AtAAP1 is erocyclic N solutes in Arabidopsis. In view of the predomi-
localized to endosperm and cotyledons while AtAAP2 is nance of ureide-N in translocation streams serving developing
restricted to the vasculature of siliques and the funiculus. fruits in certain tropical legume species, but with the meta-
Thus, AtAAP2 is thought to mediate exchange of amino bolic requirement that ureide-N is not transferred from the
acids from xylem to phloem. Such an exchange has been maternal tissue to the embryo, it will be interesting to see
identified as a specific asparagine transfer in the upper stem whether or not fruit and seed tissues express these trans-
segments and raceme of lupin to enrich phloem with N, porters. Enzymes of purine oxidation (urate oxidase and
narrowing the C:N ratio to match more closely seed allantoinase) have been demonstrated at elevated levels dur-
demands for these commodities. ing germination of a number of seeds, and it has been
The broad substrate specificity found for most plant amino inferred that purines are released due to intense nucleic acid
acid transporters does not conflict with the idea that translo- hydrolysis. If indeed allantoin provides a supplementary
cated N is characterized by just one or two amino acids. Most source of translocated N to the newly formed root and
of the amino acid transporters from plants have been charac- shoot apices then the novel allantoin transporters may be
terized by yeast mutant complementation or functional analy- important in phloem loading at this time.
sis in Xenopus oocyte assays and specificity in vivo may be quite
different. Indeed, differences in affinity for amino acid species
have been described. Furthermore, specific transporters are Primary N Metabolism
likely to be found when more extensive genomic data is avail-
able for a wider range of plants. The primary pathways of N metabolism in seeds are the reac-
As noted above seed tissues produce and may transiently tions that utilize incoming N-solutes to form the 20 protein
accumulate significant amounts of ammonium ion. It appears amino acids. As noted above N-solutes translocated to tissues
that this is a feature of their metabolism throughout devel- surrounding a developing seed or to the embryo itself are
opment, raising the question of transport mechanisms for characterized by one or two predominant compounds.
the ion within seed tissues. Preliminary data indicates that Among these asparagine and glutamine appear to be the
genes homologous to the high affinity ammonium trans- most common. Although some incoming asparagine is incor-
porter family (AMT), AtAMT1 from Arabidopsis, are not porated directly into seed protein the majority is metabolized
expressed in the seedcoat and cotyledons of developing (in white lupin more than 80% over 13 weeks of develop-
lupin seeds. However, the recently described high affinity ment). Supplying 15N (amide)-labeled asparagine to develop-
AMT2 transporter gene is expressed in Arabidopsis siliques ing lupin fruits in phloem resulted in label being recovered in
and this may be the transporter that functions in seeds. In 15 amino acids following hydrolysis of seed protein. In addi-
view of the widely diverse levels of ammonium that form in tion to asparagine, glutamine, aspartate, and glutamate, those
seed tissues at different stages of development it seems likely labeled included arginine and histidine, those derived from
that other transporters/channels with quite different kinetic pyruvate (alanine, leucine and valine) and 3-phosphoglycerate
features are also involved. (glycine and serine), those of the aspartate family (lysine and
126 PROTEINS | Nitrogen Metabolism
isoleucine) as well as aromatic amino acids (tyrosine and mapping population with the mutant Ask2 was a good candi-
phenylalanine). Clearly seeds express all the major pathways date gene for a quantitative trait locus (QTL) affecting free
for synthesis of protein amino acids. amino acid content in the harvested grain. Dihydropicolinate
There are two potential pathways releasing asparagine-N to synthase (DHDPS) converts the product of AK (aspartate 4
amino acid synthesis. Hydrolysis of the amide-group by aspar- semialdehyde) to an intermediate in lysine synthesis and is
aginase produces ammonia and aspartate, the latter providing also subject to feedback inhibition by lysine during seed
an amino donor to the many aminotransferases (AT) that have development.
been detected in seeds. The second pathway utilizes an Specific features of C metabolism within the developing
asparagine-AT that transfers the amino group to glyoxylate seed are required to accommodate some aspects of amino
forming glycine and 2-oxosuccinamic acid. The acid is reduced acid synthesis. Phosphoenolpyruvate carboxykinase (PEPCK)
to 2-hydroxysuccinamate and deamidated to ammonia and is present in a wide range of developing seeds. In grape seeds,
malate. Although both potential pathways are expressed in PEPCK is expressed in the inner layer of the seedcoat, in the
developing pea fruits, asparagine-AT is largely confined to chalaza and in cells at the boundary of the storage tissue;
pod tissue and is very low in developing seeds. Asparaginase expression coinciding with maximum deposition of storage
on the other hand is expressed at high levels in seedcoats early reserves. Feeding asparagine to developing grape seeds caused
in development, transiently in endosperm and in filling coty- a strong induction of PEPCK indicating that the enzyme func-
ledons of legume seeds. The same could be said for predomi- tions in the metabolism of N-solutes after unloading from
nant non-amino acid solutes like the ureides. Although there phloem. These data are consistent with an anaplerotic role
are a number of enzymic routes for the degradation of allan- for PEPCK in supplying acetyl-CoA when the TCA cycle is
toin that are expressed in different plants, differing even involved in the interconversion of C skeletons for 4C and 5C
between cultivars, the N is released as ammonia. Thus reassi- amino acids.
milation of ammonia is a major activity of seed tissues whether Because seeds are formed within enclosing structures that
in the seedcoat, endosperm, or the components of the embryo. limit gas exchange the concentrations of respired CO2 within
Not surprisingly, both glutamine synthetase (GS) and gluta- the structures may build up to quite high levels and refixation
mate synthase (GOGAT) activities are expressed in seed tissues, of some of this C is achieved through RubisCO (Ribulose bis P
providing a high affinity utilization mechanism for ammonia carboxylase/oxygenase) and especially PEP carboxylase. In
that is likely to function in reassimilation throughout seed lupin embryos some of this refixed C is incorporated in sugars
development. Seeds express glutamate dehydrogenase (GDH) and TCA cycle intermediates (malate, isocitrate, and citrate),
activity and this may also participate in reassimilation. While but as much as one-third provides C for a range of amino acids,
the role of GDH has been the subject of considerable debate principally alanine, aspartate, and serine, but also glutamate,
and conflicting opinion the consensus view that has emerged glutamine, asparagine, glycine, valine, leucine, and isoleucine.
for most tissues appears to be one of oxidation of glutamate In Vicia faba PEP carboxylase is up-regulated in cotyledons
rather than synthesis. However, the very high concentrations of during grain filling. Again this suggests an anaplerotic role for
ammonium ion that form transiently in the endosperm of the provision of 5C skeletons to the TCA cycle functioning in
some seeds would be kinetically favorable for GDH to func- amino acid synthesis in an intensely respiring tissue.
tion, for a short while at least, in its assimilation.
Among the protein amino acids that are synthesized largely Synthesis of Stored N Reserves in Seeds
in situ to meet the needs of seed storage protein accumulation
are those of the aspartate family; namely lysine, threonine, The major dietary sources of plant protein are seeds of cereals
isoleucine, and methionine. Not surprisingly expression of and legumes. Legumes and some oilseeds typically contain
the first committed step in the aspartate pathway, catalyzed 3050% dry weight as protein while cereals are lower, 10%.
by a bifunctional aspartate kinase (AK)-homoserine dehydro- The storage proteins constitute up to 90% of the seeds protein
genase (HSDH) in Arabidopsis, is coincident with storage pro- at maturity and as a consequence it is this group that largely
tein synthesis in the embryo. Cereals are deficient in lysine and determines the nutritional and processing qualities of grain.
in some, threonine and tryptophan, while legume seeds are Features of the synthesis, localization, and properties of the
poor sources of S-amino acids, particularly methionine. As a major storage proteins have been reviewed extensively and,
consequence there has been considerable research into regula- more recently, details of the regulatory mechanisms that deter-
tion of the pathway for these essential amino acids. There mine their synthesis in response to changes in plant nutrition
have also been numerous attempts to exploit this knowledge and environmental conditions are being revealed.
to enhance the nutritional and/or processing qualities of both While most of the incoming N is used for protein synthesis,
cereal and pulse grains. AK activity is subject to feedback inhi- seeds may also accumulate quite a significant pool of free
bition by pathway products, specifically lysine or threonine. protein amino acids that persists to maturation and constitutes
However, in the high lysine opaque-2 mutants of maize very a store at maturity. The amides glutamine and asparagine are
significant levels of the aspartate family accumulate as a pool of common constituents of this soluble-N pool but other amino
free amino acids in the mature endosperm. Accumulation acids may predominate in some species.
results in part from lysine degradation being compromized, One of these soluble reserves is arginine. In soybean argi-
because lysine-ketoglutarate reductase activity is suppressed, nine is synthesized in developing cotyledons as well as being
but other studies have identified changes in sensitivity of the supplied from the seedcoat to the embryo through the apo-
monofunctional AK (Ask2) to feedback regulation by lysine as plast. Its accumulation to more than 60% of the free amino
a major genetic determinant of free amino acid content. In a acid pool apparently occurs because arginase is not expressed
PROTEINS | Nitrogen Metabolism 127
in the embryo. The accumulation of lysine and other amino reassimilation in regulating the germination process. Such
acids of the aspartate family in maize endosperm was noted markers offer the potential to breed for complex traits related
above and doubtless there are numerous other examples that to seed N metabolism in grain improvement program.
occur in nature.
Seeds accumulate significant quantities of non-protein
amino acids. Some of these, such as homoserine, are interme- Future Prospects
diates in the synthesis of the 20 protein amino acids while
others are accumulated as end products of metabolism. Despite the wealth of knowledge about the composition of
Some, such as the arginine analogue canavanine, are highly seeds and the nature of their stored reserves, factors that deter-
toxic to insects and other potential herbivores apparently serv- mine which reserves are formed and stored and the roles of
ing to protect the developing or mature grain. Canavanine can environmental cues in these processes have yet to be estab-
reach levels that are 6% of the dry weight in the seeds of lished. Clearly there is tissue-specific and temporal expression
Canavalia ensiformis and so represent a significant stored of genes that encode biosynthetic enzymes but the nature of
reserve of N. Mature groundnut (Arachis hypogea) seeds contain the mechanisms that integrate metabolism so that stored
high levels of g-methylene-glutamine that is mobilized to the N-reserves are generated to maximize the available resources
developing seedling following germination but is not metabo- of C, N, and S need to be resolved. Although functional geno-
lized to any great extent, apparently accumulating again in the mic or proteomic analyses have been applied to developing
next generation of seed. seed tissue they have yet to reveal new information about more
There are hundreds of nonprotein amino and imino acids detailed aspects of the pathways and regulation of N metabo-
formed by plants with the overwhelming majority of these lism and transport. As genomic analysis is extended to a wider
accumulated in seeds. Many of these occur in sufficient range of plants this information will become available provid-
amounts that they could be construed as a stored reserve but ing new tools for both understanding and manipulating grain
the physiological and biochemical studies to confirm this role development. Significant changes to grain composition
have yet to be undertaken. through selection from germplasm collections, conventional
breeding and the exploitation of direct genetic engineering will
be an ongoing area for research as novel, more nutritious and
Mobilization of N Reserves in Germinating Seeds health promoting food ingredients are sought. Because plants
engage in a bewildering diversity of N metabolic end products,
While the N reserves in seeds are principally in the form of many of which accumulate in seeds, it is important that critical
storage proteins, as germination proceeds N from nucleic acids studies of N metabolism are extended beyond the handful of
or metabolic proteins that might have been formed before species for which information is currently available.
dehydration or subsequently following imbibition is also
available. The possibility that allantoin is formed from nucleic
acid catabolism has been indicated earlier. Additionally seeds Exercises for Revision
mature with what can be a significant soluble N pool and this
appears to have a specialized role in the first few hours/days of Draw up a flow chart showing the movement of nitrogen
germination when the newly activated meristems of the from the farmers field to the final grain-based food
embryo begin to grow rapidly. product.
Arginine has been recognized as a common constituent that Investigate further the concepts of source and sink in rela-
fulfils this role. Its accumulation as the seed matures results tion to carbon and nitrogen metabolism in cereals. How
from suppression of an enzyme, arginase, which would nor- useful are these concepts?
mally catabolize the amino acid. However, within a day after
imbibition arginase transcript level and activity increase
sharply in the cotyledons to release urea and, as a result of Exercises for Readers to Explore the Topic Further
urease activity, ammonium ion for reassimilation and translo-
cation to the new meristems. It is likely that the translocated Investigate the current understanding of nitrogen metabo-
N-solute at this time is glutamine. Thus arginine is a compact lism in relation to endosperm protein formation in a
form of stored N (C:N ratio of 1.5) that provides an almost selected cereal grain.
instant N source, and is no doubt used in this role in a wide What techniques are available to study nitrogen fluxes in
range of species (e.g., pumpkin and pea seeds). Whether other cereal plants?
soluble N-compounds that accumulate in maturing seeds func- What are the energetic limitations to increased protein
tion as initial sources of N for germination is not known. content in cereals?
Ammonia released from oxidation of free and mobilized
amino acids can account for 50% of the N made available
during germination of Arabidopsis seeds indicating, as during See also: Carbohydrates: Carbohydrate Metabolism; Starch:
seed development, a central role for GS/GOGAT in reassimila- Chemistry; Proteins: Protein Synthesis and Deposition; The Enzymes
tion. QTLs for enhanced germination efficiency in maize have Associated with Food Grains and Their Functions in Processing; The
been identified that are co-localized with two of the GS1 Protein Chemistry of Dicotyledonous Grains; The Basics: Grain:
(cytosolic forms of glutamine synthetase) structural genes, Morphology of Internal Structure; Wheat Processing: The Gluten
gln3 and gln4, emphasizing the importance of ammonia Proteins of the Wheat Grain in Relation to Flour Quality.
128 PROTEINS | Nitrogen Metabolism
Further Reading Lea PJ and Ireland RJ (1999) Nitrogen metabolism in higher plants. In: Singh BK (ed.)
Plant Amino Acids, pp. 147. NY: Marcel Dekker.
Editorial (1995) The plant cell special review issue on plant biochemistry. Plant Cell Murray DR (1987) Nutritive role of seedcoats in developing legume seeds. American
7: 7951111. Journal of Botany 74: 11221137.
Atkins CA (2000) Biochemical aspects of assimilate transfers along the phloem path: Rosenthal GA (1982) Plant Nonprotein and Imino Acids. NY: Academic Press.
N-solutes in lupins. Australian Journal of Plant Physiology 27: 531537.
Azevedo RA, Arruda P, Turner WL, and Lea PJ (1997) Review Article Number 121: The
biosynthesis and metabolism of the aspartate derived amino acids in higher plants.
Phytochemistry 46: 395419.
Bewley JD and Black M (1985) Seeds. Physiology of Development and Germination. Relevant Websites
NY: Plenum Press.
Kozlowski TT (ed.) (1972) Seed Biology, vol I. Importance Development and mpiz-koeln.mpg.de http://www.mpiz-koeln.mpg.de.
Germination. NY: Academic Press. dijon.inra.fr http://www.dijon.inra.fr.
Nitrogen in Grain Production Systems
XM Zhou, McGill University, Ste-Anne-de-Bellevue, QC, Canada
BL Ma, Agriculture and Agri-Food Canada, Ottawa, ON, Canada
DL Smith, McGill University, Ste-Anne-de-Bellevue, QC, Canada
2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition, volume 2, pp. 277286, 2004, Elsevier Ltd.
Topic Highlights most common inorganic forms include dinitrogen (N2) gas,
nitric acid (HNO3), nitrate (NO
3 ), nitrite (NO2 ), ammonium
The nitrogen cycle refers to the dynamic flow of nitogen (NH4 ), gaseous nitrous oxide (N2O), gaseous nitric oxide
from the atmosphere to plants and animals and its (NO), gaseous nitrogen dioxide (NO2), and gaseous ammonia
recycling. (NH3). The most common N-containing biomolecules belong
Nitrogen is fixed through nitrogen (atmospheric fixation), to one of the following three chemical families: proteins
biological fixation, and industrial fixation through the (including enzymes) and related compounds (including pep-
Haber-Bosch process. tides and amino acids), nucleotides (largely RNA and DNA),
Nitrate and ammonium are taken up and assimilated by and porphyins (e.g., cytochromes and chlorophyll).
plants, and produce amino acids and thence the storage Although N constitutes only 17% (w/w) of plant dry
proteins of the grain. matter, it is the element most frequently limiting the plant
Inappropriate use of fertilizers may result in nitrate run-off growth. Sufficient nitrogen is fundamental for crop growth,
to surface water, and consequent environmental health development, and yield. Improvement in N use-efficiency is
risks. thus the key to produce safe and high-quality food and feed in
an environmentally responsible way.
Learning Objectives
To understand the fixation of nitrogen from the atmo- The Nitrogen Cycle
sphere and its processing into storage proteins of the grain
or seed. Nitrogen is a dynamic element and is recycled continually by
To understand the importance of crop management prac- plants and animals. It can be transformed from one oxidation
tices in reducing environmental harm caused by nitrogen state to another through processes that comprise the N cycle
runoff. (Figure 1). The term cycle is really a misnomer as the path-
ways of N in the biosphere are actually a simple web. Nitrogen
moves among oxidation states, with ammonia (NH3) being
Introduction the most reduced and nitrate (NO 3 ) the most oxidized. The
tremendous chemical stability of dinitrogen (N2) gas makes
Nitrogen (N) is the largest component of the Earths atmo- the conversion of this compound into ammonia (NH3) or
sphere, comprising 78% (by volume) of the air we breathe. oxides of N (NO 3 ) energetically expensive and the rate-
It is usually present in the atmosphere as a dinitrogen gas (N2), limiting step in this part of the cycle. This causes N2 to
which is colorless, odorless, tasteless, and relatively inert at accumulate in the atmosphere. Thus, cycling of nitrogen by
room temperature. various biological entities determines a major part of the com-
Nitrogen was first identified by the chemist and physician position of the Earths atmosphere.
Daniel Rutherford in 1772. It is now known to be present in a It is a paradox that while 78% of the Earths atmosphere is
vast range of materials encountered in our day-to-day exis- N2 gas, and plant stems, leaves, flowers, and fruits are
tence; foods, fertilizers, poisons, and explosives are a few gen- immersed in this gas, biologically available N is generally the
eral examples. Some specific examples are: nitrogen gas is used most limiting nutrient element for crop production. A solution
as a blanketing medium during the production of electronic to this problem evolved in the exquisitely adaptable prokary-
components, as an agent for annealing stainless steel, in bev- otes. A limited number of microorganisms are able to react N2
erage processing, as a refrigerant for food protection and pres- with electrons to form NH3, which they then incorporate into
ervation, and as a major fertilizer in crop production. various biomolecules. When these microorganisms die, N is
Atmospheric nitrogen is also responsible for the orange-red, mineralized to simple forms (NH3, NO 3 ) through decompo-
blue-green, blue-violet, and deep violet colors of the aurora. sition by other microorganisms. In these forms, N is available
More importantly, N is a component of a wide range of to higher plants. Nitrogen is taken up by crop plants, which
biological compounds, being fourth in abundance after become food for animals, including human beings, and energy
carbon (C), hydrogen (H), and oxygen (O). sources for bacteria or fungi; N is then cycled back to the soil
Nitrogen exists in organic and inorganic forms in the atmo- when dead cells and tissues of organisms (plant debris for
sphere, biosphere, hydrosphere, and geosphere. It can be pre- example) or N-containing waste products (such as animal
sent in gas, liquid (dissolved in water), or solid phases. The urine and feces) are broken down by soil microbes.
Removal N
from harvest
Nitrification
Soluble and exchangeable
ammonium NH4+
Micelle fixation
Nitrites NO2 Nitrification
In natural ecosystems, N changes from one form to the production systems is taken up by the plants, with the remain-
other, but most of it remains within the nearly closed system der being unusable or lost. Globally, various forms of N are in a
in a dynamic balance. In agricultural systems, humans often dynamic balance.
add chemical N, fixed through industrial processes, a signifi- Soil microbes play a wide range of roles in the N cycle.
cant portion of which is eventually removed from the field in Because of this, these processes are dependent on temperature,
harvested material. Food, fiber, and livestock are now often moisture, and the quality and quantity of soil organic matter
produced primarily in one location and transported to other available to them. It is microbes that break down most of the
regions of the world for consumption, making crop produc- organic material, leading to mineralization of organic N.
tion fields into nutrient flow-through systems. Nitrogen input Microbial populations in the soil can take up large amounts
into agriculture production systems is largely from chemical of N released from decomposing plant tissues, as long as there
fertilizers, or N-fixing crops (including those grown as green is sufficient reduced carbon available, leading to immobiliza-
manures). There are other minor sources, such as biosolid tion of N. The conversion of ammonium (NH 4 ) to nitrite
waste products, which are sometimes added to croplands. (NO
2 ) and to nitrate (NO3 ) is referred to as nitrification. All
A small amount of N comes with precipitation, sometimes forms of N in the soil (except N2 gas) can be converted to NO 3
associated with lightning discharges, which reacts the N2 and by soil microorganisms, under aerobic conditions when soil
O2 in the atmosphere. There is some N-cycling within agricul- temperatures are above freezing.
tural systems, with livestock manures constituting the major The amount of nitrification is controlled by the supply of
component of this process. NH 4 which is, in turn, controlled by decomposition rates,
Nitrogen outputs from crop production systems include plant and microbial growth, and soil pH, or by N-fertilizer
removal of harvested grains and forage biomass from produc- addition rates. Nitrate can be converted back into N2, or into
tion systems and losses of various N forms through runoff nitrous oxide (N2O), through denitrification. This process
(surface water), leaching (to ground water), denitrification often takes place under anaerobic (generally water saturated)
(production of gaseous N2O), and volatilization (generally soil conditions. In this process, soil NO 3 can be reduced to N2
NH3). Because of leaching and gaseous losses, and aspects of gas through a series of intermediate steps [NO
3 ! NO2 ! NO
plant N physiology, only 50% of the N applied to crop (gaseous nitric oxide) ! N2O] and finally to N2 (gas). The
PROTEINS | Nitrogen in Grain Production Systems 131
intermediate and final products of denitrification, are released (N released as decomposition of plant and animal residues).
into the atmosphere. Denitrification may lead to the loss of The potentially available N (9099%) in the soil is in organic
45% of soil NO 3 each day when soil remains water saturated. forms, the rest being in inorganic (or mineral) forms. The
Microbes play a key role in most aspects of the N cycle, gener- organic forms are not immediately available for plant use and
ally extracting energy by converting forms of N out of equilib- have to be decomposed by soil microorganisms through min-
rium with the oxidation state of their environment into forms eralization. In addition, a very small amount of organic N may
that are in equilibrium, and extracting a reasonable portion of exist in relatively small soluble compounds, such as urea or
the energy liberated through this process. This causes N to amino-sugars, which may be slightly available to plants.
move among forms that are biologically available or unavail- Symbiotic (largely legume) N-fixation and mineral fertilizer
able, mobile or immobile, oxidized or reduced, gaseous or N are the major sources of N for grain production systems, with
solid. animal and green manures, crop residues, composts, sludges,
soil micro- and macro-biota being the other possible sources.
Again, organic N must be mineralized before it is available to
Nitrogen Sources plants. Sodium nitrate (NaNO3) and potassium nitrate
(KNO3) are formed by the decomposition of organic matter.
Nitrogen exists in inorganic or organic forms in our environ- In certain dry areas of the world, these saltpeters are found in
ment. Inorganic forms include NO
3 , NO2 , NH3, and N2 quantity and are used as fertilizers. Other inorganic N com-
whereas organic N is found in a wide range of biomolecules. pounds are ammonia (NH3), nitric acid (HNO3), the N oxides
Protein is one of major classes of biomolecules and generally (NO, NO2, NO3, N2O), cyanides (CN), etc. However, inor-
the largest repository of organic N in the biosphere. RuBisCo, ganic N in soil generally comprises only a small portion of
the plant protein responsible for the uptake of CO2 by C3 soil N. Hence, despite N being one of the most abundant
plants, is the most abundant protein on the planet. The other elements on the biosphere, N deficiency is probably the most
two most abundant classes of biocompounds, carbohydrates, common nutritional problem affecting crop plant production
and lipids, generally have little N associated with them. Over worldwide.
9099% of N in the surface (or tillage layer; 025 cm) layer of
most soils is in organic forms. Although there is an abundant
potential supply of N in the Earths atmosphere (N2 gas), the Nitrogen Availability to Crop Plants
unique geometry of the triple bond between the two N atoms
results in a very stable compound that is very energetically The availability of N sources is a fundamental requirement for
expensive to break. Thus, this form of N cannot be used directly metabolism as it allows N assimilation into amino acids (for
by most organisms. Whenever dinitrogen is converted into which photosynthetically produced carbon compounds are
other forms, a large input of energy is involved. The major also required) and their availability for protein synthesis. An
sources of this energy are: (1) lightning in the case of atmo- adequate supply of N allows leaf growth and photosynthesis.
spheric fixation, which accounts for 58% of total N fixed and Inorganic (or mineral) forms of N, such as NH
4 and NO3 ,
potentially available to the biosphere. This process results in comprise the majority of plant-available N. Other forms of N
the combination of oxygen from the atmosphere with dinitro- must be converted to one of these compounds by either natural
gen, causing the formation of N oxides. These compounds or artificial means before they can be utilized by plants. When
dissolve in rain water and are carried to Earth; (2) reduced N is first released from organic matter, it is in the form of
carbon and adenosine triphosphate (ATP) in the case of bio- gaseous NH3, which is volatile and can be lost to the atmo-
logical N fixation, conducted by certain prokaryotes alone or in sphere. When NH3 dissolves in water, it acquires a proton and
symbiotic relationships with plants and sometimes animals; becomes ammonium (NH 4 ), which carries a positive charge
and (3) fossil fuels for industrial fixation. and is attracted by the soil clay colloids and soil organic matter,
Industrial fixation relies on the HaberBosh process, devel- which carry negative charges. Once attached to the soil matrix,
oped during the First World War for the production of explo- NH 4 becomes part of the cation-exchange process whereby
sives. This process uses a catalyst, high pressure, and high plants exchange a hydrogen ion (H) for one of the positively
temperature (600 C), to combine N from the atmosphere charged molecules in the soil. Only a small portion of the soil
with hydrogen (usually derived from natural gas or petroleum) NH
4 exists in the soil solution; this NH4 is available to crop
to produce ammonia (NH3). Ammonia can be used directly as plants. On the other hand, NO3 carries a negative charge, and
fertilizer, reacted with CO2 to form urea ((NH2)2CO), and/or so is not bound by clay particles. Thus, NO 3 molecules are free
oxidized to form nitrate (NO 3 ) which can be salted with other to move with the soil water and are more readily available to
compounds (such as ammonium, NH 4 ) to form other types of crop plants than NH 4 . However, in coarse textured or water-
N fertilizers (e.g., ammonium nitrate). The large requirement saturated soils, NO 3 is much more prone to leaching into
for energy makes N fertilizer easily the largest component of groundwater than NH
4 ; once in ground water NO3 becomes
fossil fuel consumption in crop production, being close to 50% a potentially serious pollutant.
of the total for crops such as maize.
The bulk of the soil N in noncropland systems comes
from fixation by symbiotic bacteria (largely genera associated Nitrogen Fixation
with legumes and collectively referred to as rhizobia and spe-
cies of Frankia, usually symbiotic with woody plants), fixation A symbiotic relationship between leguminous plants and
by free-living bacteria and mineralization of organic matter N-fixing bacteria allows for plant-based biological N fixation.
132 PROTEINS | Nitrogen in Grain Production Systems
The major conversion of N2 into NH3 (a biologically available Table 1 Annual N inputs and outputs of typical agricultural land
form) is the biological N-fixation process, carried out by single-
celled prokaryotes. This process requires a great deal of energy, N (kg ha1)
supplied as ATP, which is derived from the breakdown of Inputs
organic matter, or through photosynthesis (e.g., cyanobacteria). Plant residues 10100
It is estimated that biological N fixation accounts for the annual Biological N fixation 10250
production of as much as 250 109 kg of ammonia, which is Natural fertilization through precipitation 050
twice the amount produced by the HaberBosch process. Chemical fertilizer 0400
Biological N fixation can be described as the following Outputs
equation: Volatilization of NH3 060
Dinitrification 1070
N2 8H 8e 16ATP 2NH3 H2 Leaching 1070
16ADP 16Pi Crop harvest 10150
often the limiting factor in plant growth. The absorption of often the form of N preferentially taken up by the plant.
NO 3 by roots is not only determined by the N demand of Many forest species take up primarily NH 4 , since forest soils
the plant, but also by its availability in the soil environment. are relatively acidic, inhibiting the development of popula-
Nitrate absorbed via roots is either reduced in situ into tions of bacteria that cause nitrification. The average NH 4
ammonium, imported into vacuoles (by high- and low-affinity concentration in well-aerated soil is 10100 times lower
transport systems) for storage, or transported via the xylem to than NO 3 , rarely exceeding 0.12 ppm. However, absorption
the shoot where it can either be metabolized or stored as a of NH 4 by plant roots can occur at very high rates, due to the
reserve. A low-affinity root NO 3 uptake system may play a presence of transport systems in the root plasma membrane
greater role in nitrate uptake. In some cases, a high-affinity with high substrate affinities. Like NO
3 , NH4 uptake by roots
system has been shown to be the major mode of nitrate uptake is diurnally regulated, with maximum uptake during the light
through uptake competition studies and membrane voltage period. Uptake is stimulated by sugar supply, implying that
change studies. The low affinity transport system (LATS) the greater uptake during the day is related to the availability
(possibly a carrier system or an anion channel), and the of photosynthate.
inducible, high affinity transport system (HATS) are regulated
by cellular energy supply, intracellular nitrate levels, and a
proton electrochemical gradient. The latter is regarded as an
H/anion cotransport carrier system that produces transient Assimilation of Nitrogen
plasma membrane depolarization upon addition of nitrate.
The depolarization is counteracted by the plasma membrane Both nitrate and ammonium are taken up and assimilated by
H-ATPase. The plasma membrane proton ATPase is induced plants. In general, NO 3 is the predominant form of inorganic
by nitrate. In addition to the nitrate uptake system, plants N taken up by grain crop plant roots (except paddy rice).
have an inducible nitrate efflux system, requiring both RNA Under most circumstances, absorbed nitrate is reduced to
and protein synthesis. The efflux system, however, has a much ammonium in roots and/or leaves, and is then utilized in the
slower rate than the uptake system. synthesis of amino acids and proteins. Whatever the source of
In the agricultural soils of tropical and temperate regions ammonium in plants, this ion is first converted into amino
(except paddy rice fields), nitrate accounts for 7090% of soil acids, the building blocks of protein.
mineral N. Nitrate in the soil solution is 10 times more mobile Nitrate is assimilated into organic compounds by GS/
than NH
4 . Two reserves of soil NH4 exist: a small portion of GOGAT (GS glutamine synthetase/glutamate synthase;
NH 4 is in the soil solution, and a large portion of NH 4 is GOGAT glutamate oxidase, glutamine amino tranferase).
adsorbed on soil colloids. Only water soluble NH 4 is available The GS/GOGAT pathway is considered the major pathway for
for plant uptake. Depending on the plant species, the bulk of incorporating reduced N into organic molecules. This pathway
the nitrate entering the plant is reduced to ammonia and recycles glutamate; amino groups are eventually transferred to
assimilated into organic compounds in the roots or in the other amino acids and utilized for protein synthesis and also
leaves. Plants can take up either NO
3 , NH4 , or mixtures of for synthesis of other key biomolecules such as RNA, DNA, and
the two. In most cases, crop plants perform better when the porphyrins.
134 PROTEINS | Nitrogen in Grain Production Systems
Ammonia assimilation
Proteins Proteins
Seeds
Mitochondrion Chloroplast
AA
Albumins NH1
Globulins GS2 NIR
NH4+ NO2
Prolamins NH1
(cereal) Vacuole
NH4+
NH4+
NH+4 HNO3
Leaf
Stem
NO3 NO2
Phloem
Nitrate reduction
(cytoplasm)
NR
HNO3
NO3 NO2
NH+4
Plasma membrane
Nitrate reduction
AA GS2 Plastid (cytosol)
AA NIR
+
NH4 NO2
GS1
Vacuole NR
NH+4 NH+4 NH+4 NO3 HNO3
NH+4 NO3
GS and GOGAT are located in the cytosol (for a review of incorporation where glutamate dehydrogenase is involved:
the cytosol, see www.worldreference.com) and the chloro- a-ketoglutarate NH3 NADH ! glutamate NAD; (2) GS-
plast (for a review of chloroplast, see www.worldreference. GOGAT cycle: GS catalyses the ATP-dependent conversion of
com). Their activity in particular plant tissues is closely linked glutamine utilizing ammonia as substrate and is represented by
to specific primary N assimilation, NH3 recycling during pho- two protein groups, plastid (GS1) and cytosolic (GS2) isoen-
torespiration, or general N remobilization. There are two path- zymes; GS1 is located exclusively in chlorophyllous tissues while
ways for incorporation of N into amino acids: (1) Direct GS2 is located predominantly in roots, nodules, and floral
PROTEINS | Nitrogen in Grain Production Systems 135
organs. GOGAT catalyzes the conversion of glutamine and using ferredoxin (Fd). Nitrite reductase is located in plastids
a-ketoglutarate to glutamate: glutamine a-ketoglutarate and possesses FeS and heme cofactors. Subsequently, the
NADP H H ! 2 glutamate NADP. The assimilation of NH 4 is incorporated into carbon compounds: amides
NH3 may be by either glutamate dehydrogenase or GS/GOGAT; (amino acids) and ureides.
the first of these requires quite high concentrations and the Nitrate reduction occurs mainly within the chloroplasts of
bulk of uptake is thought to be through GS/GOGAT. green leaves, but can also occur within plastids of roots. Induc-
Following the incorporation of NH3 into glutamine, N can tion and the rate of nitrate uptake depend on the external
be transferred directly to the same position in asparagine by nitrate concentration, light, pH, temperature, the concentra-
asparagine synthetase. Amino transferase can transfer the tion of other ions, anaerobic conditions, metabolic inhibitors,
amino group to aspartate, which is converted to asparagine. It and biological genotypes. Since nitrate reductase contains Mo,
has been suggested that the reaction catalyzed by Fd-GOGAT is Mo deficiency can cause N deficiency.
the key regulatory element controlling N partitioning and
redistribution during plant growth and development. To
date, research has focussed on modifying ammonia assimila-
tion in transgenic plants and this has demonstrated that the Nitrogen Storage
amplification or shifting of ammonium assimilation in a par-
ticular organ or tissue has strong effects on plant growth and Seed storage proteins comprise 70% (up to 100% in some
development. Thus, there appears to be great potential for developing countries) of total intake of dietary protein by
improving N-use efficiency in crop plants. humans. It also provides the major protein source in the diet
for nonruminant livestock. The amino acids in the leaves,
produced from assimilated NO 3 with energy and carbohy-
drates produced by photosynthesis, are transported to the
embryo and the cotyledons via mass flux in phloem vessels
Reduction of Nitrate and Nitrite of the leaf. The main transport forms of amino acids are:
glutamine, asparagine, serine, alanine, glutamate, and aspar-
Once NO 3 enters the root symplast, it may follow five tate. As a result, storage proteins are the products of the
pathways: secretary pathway, which resides within the endomembrane
system of the cell. In general, N assimilates available to
1. efflux back to the apoplast [for a review of apoplast see
developing grains are mainly used for the synthesis of pro-
www.worldreference.com];
teins. These proteins then are accumulated and stored in
2. reduction of NO
3 to NO2 , then NH4 by nitrate reductase
specialized tissues such as the endosperm of cereals, other
(NR) using the reduced form of nicotinamide adenine
endospermic monocotyledons, or the cotyledons of the
dinucleotide (NADH) and by nitrite reductase (NIR) using
embryo for the pulse legumes and other nonendospermic
ferredoxin (Fd) in root cells, leading to the production of
dicotyledons.
amino acids;
Grains of cereal crops usually contain 719% protein while
3. accumulation into vacuoles of root cells, involving trans-
the seeds of legume crops contain up to 40% protein, in both
port across the tonoplast [for a review of tonoplast see
cases on a dry matter basis. Tuber crops are rich in soluble
www.worldreference.com];
carbohydrates and low in protein. For example, the crude
4. secretion into xylem vessels for long distance transport to
protein concentration in sweet potato is twice that of potato
the shoots; and
(12.1% versus 5.4%). Thus, when tuber crops are used as
5. reduction of NO 3 in the leaves, often using excess reductant
human food or animal feed, they are a good source of energy,
in upper, CO2 limited, leaves.
but require protein supplementation.
High concentrations of nitrate can be found in vacuoles, indi- Almost all storage proteins in the seeds of major crops can
cating that nitrate not only acts as a nutrient but also partici- be grouped into three categories according to their functions:
pates in osmotic maintenance. The first step in the nitrate (1) globulins (soluble in dilute saline solutions), the most
assimilation pathway is the reduction of nitrate to nitrite, common storage proteins, are present in all angiosperm
catalyzed by assimilatory NR. When reduction occurs in leaves, seeds; (2) albumins (soluble in water); (3) prolamins (soluble
the nitrite formed by NR activity is then translocated to the in dilute alcohol) and restricted to the seeds of Gramineae
chloroplast, where it is further reduced to NH 4 by NIR. During (grass) family. This protein represents the main storage protein
the nitrate reduction process, NR activity appears to be the rate- in cereals such as wheat (gliadins), barley (hordeins), oat
limiting step in the conversion of NO
3 to NH3/NH4 , because (globulins), rice (glutelins), rye (secalins), and maize (zeins).
it is (1) the first enzyme in the pathway NO3 of assimilation, Both albumins and globulins have biological functions. This is
(2) substrate inducible, (3) relatively unstable, and (4) its particularly true of enzymes during seed germination. In most
activity relative to other enzymes in the NO 3 assimilation cases, storage proteins have no biological activity and only act
pathway is low and its Km (related to binding affinity) for as a source of N, sulfur, and carbon skeletons for the develop-
NO 3 is high. Further, there is evidence that when nitrate is the ing seed. Thus, storage proteins are not as evolutionarily con-
dominant N source, nitrate reductase activity (NRA) is the strained as those of other proteins, such as enzymes, although
limiting factor to the growth of many plants. NR has therefore there is a requirement that proteins should be efficiently syn-
been intensively studied in order to understand its catalytic thesized, packaged, and stored, and then remobilized during
efficiency and regulation. Nitrite is reduced to NH 4 by NIR the process of seed germination.
136 PROTEINS | Nitrogen in Grain Production Systems
Interactions between Nitrogen and Carbon Metabolism is not toxic, it can easily be reduced, internally, to the toxic
compound nitrite. Reduction to nitrite in vegetables can occur
The products of photosynthesis are the building blocks of postharvest as well as after ingestion, in saliva and in the
agricultural production. From a production point of view, the gastrointestinal tract. One of the symptoms of nitrite toxicity
efficiency of photosynthesis is mainly determined by the avail- is methemoglobinaemia, in which NO 2 binds tightly to hemo-
ability of light, CO2, water, heat (temperature), and key ele- globin reducing the ability of the blood to carry oxygen and
ments in the soil, with N being the most likely to be limiting. leading to respiratory dysfunction. Chronic nitrite toxicity may
Precipitation is outside management control in crop produc- lead to the formation of carcinogenic nitrosamines. The fre-
tion systems, although irrigation may provide reliable water quency of gastric cancer could be reduced by avoiding high
where water resources are available. Nitrogen application, intake of nitrate.
including aspects such as quantity, timing, and method is According to the International Fertilizer Association,
part of most crop management regimes. Grain yield is largely annual world consumption of N fertilizer is equivalent
a function of accumulation of photosynthetic assimilates. The to 83 million tonnes (Mt). North America accounts for 15%
interactions between carbon dioxide (CO2) and nitrate assim- of this total world consumption. Nitrogen that is not taken up
ilation and their dynamics are key elements of crop produc- by plants may lead to gaseous loss to the atmosphere, or an
tion. As a result, an adequate supply of N, its assimilation into increased level of nitrate in the soil. This nitrate is vulnerable to
amino acids (for which photosynthetically produced carbon run off in surface water or leaching into groundwater, causing
compounds are required), and their availability for protein significant environmental pollution. High concentrations of
synthesis, are essential for metabolism. NO 3 in drinking water due to surface and ground water con-
Nitrate is reduced to NH3 by NR and NIR using electrons tamination can also lead to methemoglobinaemia.
from photosynthetic electron transport. The NH3 is then con- Nitrous oxide (N2O) is a greenhouse gas with a heat-
verted into amino acids by the GS/GOGAT reaction (the main trapping capacity that is 310 times greater than that of CO2.
pathway by which plants convert ammonia into amino acids), Since the last century N2O emission has been increasing 0.3%
and the carbon-skeletons are provided as organic acids year1. Emissions of N2O from cultivated lands are estimated
derived from the tricarboxylic acid cycle. Carbohydrates for at 3.03.5 Mt N2O-N year1. The bulk of current N2O emis-
organic acid synthesis, ultimately from photosynthetic CO2 sions are caused mainly by the application of fertilizer N and
assimilation, and ATP for the GS/GOGAT reaction are pro- biomass burning. When soils become anaerobic, or sometimes
duced by photosynthesis and respiration. Thus, there is close during the conversion of NH
4 to NO3 , N2O is produced.
interaction in the very earliest phases of N and carbon metab- N2O escapes the soil as a gas and accumulates in the atmo-
olism, both using light energy, with some 10% of the electron sphere where it is both an agent of stratospheric ozone
flux in photosynthesizing leaves used for N reduction. destruction and a very potent greenhouse gas. High levels of
During the last 46 decades grain yields of major field soil organic matter and nitrate combined with low oxygen
crops have increased 2.5 fold, largely due to an increase in levels promote rapid denitrification. Increasing emissions
the production and storage of carbon in grains by efficient of greenhouse gases are likely to accelerate global climate
crop varieties and increased use of N fertilizer. Although large change. Average global surface temperature is expected to
variations in grain protein concentrations exist among varie- rise 0.62.5 C (33.136.5 F) in the next fifty years, and
ties within a species, an increase in yield has generally 1.45.8 C (34.542.4 F) in the next century.
resulted in a slight to severe decrease in the protein/starch Reducing the contamination of drinking water associated
or oil ratio. For example, in the case of maize, the increase in with excess application of inorganic fertilizer and manure in
the genetic-based yield potential after 1967 was, on average, agriculture production is a high priority. There is an urgent
accompanied by a decrease in the concentration of protein need to develop new crop production systems that maximize N
( 1.5%) and an increase in that of starch ( 2%). In the case use efficiency and are less hazardous to the environment.
of soybean, genetic selection leading to a seed yield increase However, the reduced application of fertilizers conflicts with
of 100 kg ha1, increased seed oil yield by 1 kg ha1, and the need for greater crop yields to feed rising world popula-
decreased protein yield by 23 kg ha1. For wheat, an tions. Crop yields would drop to a very low level if N-fertilizer
increase in yield of 2 kg ha1 led to a decrease in the applications were to cease. Many countries that use substantial
protein concentration of more than 2% of the dry weight. amounts of fertilizers and export agricultural produce would
Plant geneticists and breeders continually make efforts to barely be able to feed their own population in the absence of
break this relationship, hoping for increases in yield without N-fertilizer applications. To overcome the potentially danger-
decreases in quality. ous social problems of food shortage, environmental degrada-
tion and pollution, agricultural yields per unit land area must
be increased at the same time as dependence on applied fertil-
izers is decreased. Agriculture uses N fertilizer very inefficiently
Nitrogen and Environmental Health (even in developed countries), in part because other environ-
mental conditions are often limiting. Thus, understanding the
Nitrate accumulation in plants occurs when the speed of process of N uptake and assimilation, and mechanisms associ-
nitrate uptake exceeds the rate of reduction and subsequent ated with increased N-use efficiency by crops would be a ben-
assimilation into amino acid and proteins for growth. A high efit not only in the context of environmental concerns but also
nitrate accumulation in edible parts of plants consumed as crop quality. Consequently, selection of superior genotypes
vegetables is a potential health hazard. Although nitrate itself with efficient N-use and development of sustainable
PROTEINS | Nitrogen in Grain Production Systems 137
production systems for specific regions will solve or at least Exercises for Revision
reduce the negative impact of fertilizer N use on the
environment. Compare nitrogen use in leguminous and in cereal grain
crops.
Explain the environmental effects of large-scale nitrogen
Conclusions and Prospects use in crop agriculture.
Table 1 Composition of cell walls in cereal endosperm of arabinofuranosyl residues to the xylan backbone. There have
been suggestions that xylan synthesis might also involve
Cereal Source of walls Major polysaccharides enzymes of the cellulose synthase or cellulose synthase-like
Barley Aleurone 71% Arabinoxylan enzyme families, given the structural similarities between cel-
26% (1,3;1,4)-b-Glucan lulose and the xylan backbone of arabinoxylans. As the
2% Cellulose genomes of more cereal species are sequenced, it is clear that
2% Glucomannan most genes involved in cell wall polysaccharide synthesis are
Starchy endosperm 75% (1,3;1,4)-b-Glucan members of multigene families.
20% Arabinoxylan
2% Cellulose
2% Glucomannan Enzymic Hydrolysis of Wall Polysaccharides
Wheat Aleurone 65% Arabinoxylan in Germinated Grain
29% (1,3;1,4)-b-Glucan
Most of the enzymes involved in (1,3;1,4)-b-glucan and arabi-
2% Glucomannan
2% Cellulose noxylan degradation in the endosperm of germinated grain
Starchy endosperm 70% Arabinoxylan have been characterized in detail, particularly in barley,
20% (1,3;1,4)-b-Glucan wheat, and maize. In each case, a battery of hydrolytic enzymes
7% Glucomannan is required to completely depolymerize the polysaccharides.
4% Cellulose The enzymes responsible are generally synthesized de novo in
Rice Starchy endosperm 27% Arabinoxylan the aleurone layer or the scutellum and subsequently secreted
20% (1,3;1,4)-b-Glucan into the starchy endosperm. Walls of the starchy endosperm
28% Cellulose are completely degraded during this process, but a thin inner
3% Pectin
layer of the aleurone cell walls appears to resist degradation,
15% Xyloglucan/mannans
presumably to maintain the structural integrity of the aleurone
cells while enzyme secretion is in progress.
The (1,3;1,4)-b-glucans of endosperm cell walls are depo-
The fertilized endosperm mother cell divides for 70 h to lymerized by (1,3;1,4)-b-glucan endohydrolases, of which
form up to 2000 free, individual nuclei in the cytoplasm of there are one or two isoforms in most cereal species. The
the central cell. Walls subsequently grow between the free (1,3;1,4)-b-glucan endohydrolases release low-molecular-
nuclei. After a lag phase, further expansion of the cellular mass (1,3;1,4)-b-oligoglucoside products that can be further
endosperm occurs through meristematic activity of peripheral hydrolyzed by exohydrolases such as b-glucan glucohydrolases
cells that eventually differentiate to form the aleurone. In and b-glucosidases. The gene families that encode these
barley, the major components of the starchy endosperm enzymes are generally larger, with five or more members.
walls, (1,3;1,4)-b-glucan and arabinoxylan, accumulate A second, distinct class of (1,3;1,4)-b-glucan endohydrolase
steadily between 13 and 40 days post anthesis (dpa). has been reported in germinated barley and in maize coleop-
Despite the importance of cell wall biosynthesis as a deter- tiles. The enzymes release (1,3;1,4)-b-glucans with degrees of
minant of final cereal grain quality, polysaccharide synthases polymerization of 60100, but the enzymes have not been
involved in wall synthesis have proved very difficult to purify characterized in detail. Their action patterns are consistent
by conventional biochemical procedures, mainly because they with a cellulase, but there remains some doubt as to their
are membrane-bound enzymes and can be rapidly inactivated precise role in (1,3;1,4)-b-glucan depolymerization in germi-
in vitro. A key breakthrough in this area occurred when cellu- nated grain.
lose synthase genes (CesA) were identified in higher plants; Arabinoxylan depolymerization in germinated cereal grains
subsequently, multiple CesA genes and multiple groups of also requires the concerted action of a range of different hydro-
cellulose synthase-like (Cs1) genes have been identified. It lytic enzymes. (1,4)-b-Xylan endohydrolases hydrolyze glyco-
has now been demonstrated that the CslF, CslH, and CslJ sidic linkages in the (1,4)-b-xylan backbone of the
genes encode (1,3;1,4)-b-glucan synthases and that the CslF polysaccharide and probably require a short, unsubstituted
group is probably the most important in cereal grain develop- region of the (1,4)-b-xylan chain for activity. Genes and
ment. Indeed, mutations of the CslF6 gene result in grain with cDNAs encoding barley (1,4)-b-xylan endohydrolases have
little or no (1,3;1,4)-b-glucan, and when the CslF6 gene is been isolated and used to define the expression patterns and
overexpressed in developing barley grain, (1,3;1,4)-b-glucan chromosome locations of the genes. There is another class of
levels increased significantly, as did dietary fiber content of enzymes that can hydrolyze polymeric arabinoxylans from
the grain. Nevertheless, it is becoming increasingly apparent cereal cell walls. These enzymes hydrolytically remove single
that other proteins and enzymes are also required for (1,3;1,4)- a-arabinofuranosyl substituents from the (1,4)-b-xylan back-
b-glucan synthesis and regulation. bone of the polysaccharide and have been designated arabi-
Identification of the xylan synthase genes in cereals has noxylan arabinofuranohydrolases (AXAH). They are probably
been particularly challenging, and although genes encoding responsible for the removal of arabinofuranosyl residues prior
enzymes from the glycosyltransferase (GT) GT43 and GT47 to (1,4)-b-xylan endohydrolase action in germinated grain but
families have been implicated in the synthesis of the (1,4)-b- might also be involved in the modification of arabinoxylan
xylan backbone of cereal grain arabinoxylans, precise details of fine structure during wall deposition, maturation, or expan-
the roles of these enzymes are now yet available. Enzymes sion. Despite these possibilities, there has not been any defin-
encoded by the GT61 family appear to mediate the addition itive description of the role of the AXAHs in cereal grain
140 PROTEINS | The Enzymes Associated with Food Grains and Their Functions in Processing
development or germination. Complete depolymerization of effect in long fermentation processes, which applies to most
oligoarabinoxylosides released from cell wall arabinoxylans by types of western-style pan breads and leads to sticky dough that
(1,4)-b-xylan endohydrolase requires the action of a- is harder to handle, a sticky crumb, dark-colored crust and
arabinofuranosidases and b-xylosidases, both of which have loaves with large holes that are difficult to slice and may jam
been detected in germinated cereal grains. Enzymes with dual a- slicing machines. During cooking or baking, the starch
arabinofuranosidase/b-xylosidase activity have been described becomes increasingly susceptible to attack by a-amylase, and
and again these hydrolases are encoded by multigene families. even small amounts of enzyme can cause dramatic differences
in starch viscosity and product texture. Flat breads, chapatis,
and sourdough breads are more tolerant to a-amylase, whereas
Enzymes in Grains That Affect Processing, End-Product Japanese-style (udon) noodles are very sensitive. Irrespective
Appearance, Nutritional Value, or Storage of Grain of the end product, appearance and quality deteriorate as a-
amylase activity increases.
This section will focus primarily on wheat that is a staple food High levels of a-amylase activity in ripe wheat grain can
in many parts of the world and unique in terms of the myriad arise via a number of mechanisms. First, the low-pI a-amylase
of end products that are consumed. that is synthesized in the green (chlorophyllous) pericarp of
immature grains may be retained rather than being gradually
degraded as the grain matures (Figure 1).
a-Amylases in Developing and Ripe Wheat Grains
Frost damage to partially developed grains and conditions
Starch constitutes up to 80% of the material in the endosperm of of high humidity and low light intensity during ripening have
cereal grains and is stored in a highly crystalline form in granules been associated with this abnormal retention of low-pI a-amy-
that are laid down during development and ripening. a-Amylase lase. Second, some bread and durum wheat genotypes contain
is a critical enzyme in remobilization of the endosperm since it a genetic defect referred to as late-maturity a-amylase (LMA) or
initiates hydrolysis of the starch granules, reducing the large, prematurity a-amylase (PMAA) that may be result in the syn-
complex molecules of starch to smaller units as already thesis of high-pI a-amylase in the aleurone tissue during the
described. Wheat and barley grains can synthesize two groups mid to later stages of grain development (Figure 1). Induction
of a-amylases, or a-amylase isoenzymes, that differ in genetic of high-pI a-amylase synthesis in LMA-prone varieties may be
control, temporal pattern of synthesis, and physical and chem- constitutive, in tall wheat varieties and in semidwarf varieties
ical properties. Perhaps, the most important group in grains are grown under mild conditions free from moisture stress, or
the high-pI (germination or malt) a-amylases that are products require a cool temperature shock in semidwarf varieties ripen-
of the a-Amy1 genes that are normally only synthesized in the ing under warm to hot conditions. LMA, unlike germination or
grains during the early stages of germination. The second group, sprouting, is not dependent on rain and does not appear to
low-pI (pericarp or green) a-amylases, are products of the a- involve changes in, or signals from, the embryo. Finally,
Amy2 genes. They are present in the pericarp of immature grains, adverse weather conditions (rain accompanied by high humid-
at low levels in mature grains, and are also synthesized during ity) at maturity and prior to harvest may trigger grains to
the later stages of germination. In comparison with other cereal commence germination while still in the spike. This phenom-
enzymes, the a-amylases are relatively thermostable and remain enon, known as preharvest sprouting (Figure 2), is relatively
active at temperatures above 60 C where the starch is gelati- common in many parts of the world.
nized and becomes extremely sensitive to enzymic hydrolysis. Preharvest sprouting-resistant varieties in which the grain is
Under ideal growth conditions, ripe wheat grains contain a still dormant at harvest ripeness have been developed to combat
small amount of low-pI a-amylase and no high-pI a-amylase, this problem. The sequence of events that occurs during sprout-
and following milling, the flour can be used to produce a vast ing follows very closely those observed in isolated grain that is
array of processed foods including pan breads, flat breads, germinated under more controlled conditions. a-Amylase syn-
noodles, cakes, and biscuits as well as industrial products. For thesis begins adjacent to the embryo or germ end of the grain
leavened products such as western-style pan breads, some a- and progresses in a wave toward the distal end. The system has
amylase is required to produce the fermentable sugars needed been very widely studied by plant biochemists endeavoring to
by the yeast. This a-amylase, often microbial in origin, is understand the intricacies of control of enzyme synthesis and
normally added in measured amount during mixing particu- the role of plant hormones. Initially, only high-pI isoenzymes
larly in the highly automated bread-making plants that require are synthesized; however, as germination progresses, the low-pI
strict adherence to specifications to maintain quality and effi- groups are also produced. While LMA and sprouting are readily
cient production. Under nonideal growing conditions, or as a distinguished both in genetic control and in the pattern of
result of adverse genotype environment interactions, variable enzyme distribution within the grain, the a-amylase isoenzymes
amounts of either, or both, groups of a-amylases may be involved are similar, as are the effects on the quality of end
retained or synthesized prior to harvest and as a consequence products. In contrast to LMA, where only high-pI a-amylase
the grain may be unsuitable for processing. Wheat with unac- appears to be synthesized, sprouted grains may contain elevated
ceptably high levels of a-amylase is commonly downgraded in levels of a range of hydrolytic enzymes including endoproteases,
quality, often to be restricted to animal feed, and attracts lipases, and cell wall-degrading enzymes.
substantial financial dockages. While some of the a-amylase
is removed with the bran and germ during milling, at progres-
Endoproteases in Developing and Ripe Wheat Grains
sively higher levels of activity, more of the enzyme will have
been transported into the endosperm and will be recovered in Endoproteases attack the internal peptide bonds of proteins
the flour. Excess levels of a-amylase in wheat have a disastrous and may make them more accessible to exoproteases that
PROTEINS | The Enzymes Associated with Food Grains and Their Functions in Processing 141
50
45
40
35
-Amylase activity
30
25
20
15
10
0
0 5 10 15 20 25 30 35 40 45 50
Days after anthesis
Sprouted Nonsprouted
70 Figure 2 Sprouted (left) and nonsprouted (right) spikes of wheat.
60
critical to the production of high-quality end products such as
bread and noodles.
50
-Amylase activity
protein. The starting point for synthesis of flavonoids is phe- The orthodiphenols are oxidized further by PPO to s-benzo-
nylalanine and the enzyme phenylalanine ammonia lyase, quinones, which in turn polymerize nonenzymatically to mel-
which leads into a biosynthetic pathway that produces a vast anins (brown pigments). Fortunately, most of the PPO in
array of structures including flavones, cinnamic acids and lig- wheat grains is removed with the bran during milling, but
nins, anthocyanins, and tannins. Xanthophylls and flavonoids sufficient enzyme is carried through in the flour of many
accumulate during the early to middle stages of grain develop- wheat cultivars to present problems with the color stability of
ment, depending on the activity of the enzymes in the respec- products that are not cooked immediately. While the endoge-
tive biosynthetic pathways. In the case of xanthophylls, the nous substrate of PPO in grains has not been identified, the
levels decline markedly as the grain ripens. Lutein may also role of PPO in darkening of WSN, YAN, and durum pasta is
be converted to mono- and di-fatty acid esters during storage of now clear. Addition of exogenous substrates such as tyrosine,
ripe grain or flour. The concentrations of xanthophylls and catechol, and L-DOPA (L-3,4-dihydroxyphenylalanine)
flavonoids in flour are a product not only of the balance of enhances noodle darkening, whereas darkening is greatly
synthesis and degradation during grain development but also reduced in the presence of specific PPO inhibitors or in culti-
of the milling process, in particular the efficiency of separation vars selected for low or zero grain PPO activity (Figure 3).
of starchy endosperm from other grain tissues such as the germ Variation in PPO activity explains most of the genetic differ-
and seed coat. ence in noodle darkening between cultivars, but is only respon-
sible for a portion of the total darkening that occurs. Physical
effects and other unidentified oxidation reactions also contrib-
Lipoxygenase
ute to the reduction in noodle brightness over time.
Lipoxygenases (LOX) catalyze the oxidation of polyunsatu-
rated fatty acids such as linoleic and linolenic acid to hydro-
peroxides. Fatty acid radicals produced during the intermediate
Peroxidases and Hydrogen Peroxide Synthesis and Catalysis
steps in the reaction can initiate oxidative degradation of pig-
ments such as lutein and b-carotene. As a result of this co- Peroxidases are oxidoreductases widely distributed in higher
oxidation, lutein and lutein ester pigments may decline plants and have been shown to oxidize many compounds,
during dough mixing in preparation for the production of including phenolics, in the presence of hydrogen peroxide.
WSN and durum pasta, leading to poor color stability. Loss Oxidation of phenolic compounds by these enzymes can lead
of pigment can be reduced by the addition of compounds such to the formation of brown pigments similar to those formed by
as a-tocopherol that inhibit LOX activity or by using bread and PPO, and consequently, they have also been implicated in
durum cultivars that have been selected for zero LOX activity. browning or darkening of noodles and pasta. In the case of
In addition to its effects on color, LOX has been associated with peroxidase, the situation is complicated by the presence of a
the formation of compounds with undesirable odors and fla- number of forms with different substrate specificities that are
vors in a range of products. Loss of lutein and carotene is also localized in embryo, endosperm, and seed coat tissues of wheat
undesirable due to the role these compounds play in the health grains and by the interaction with systems that either generate
of our eyes, in iron uptake in the intestinal tract, and as general H2O2 (superoxide dismutase and NADH oxidation) or catalyze
scavengers of peroxy radicals that reduce oxidative damage to (catalase) its destruction.
tissues. In addition to its possible role in product darkening, per-
oxidase has been implicated in the development of symptoms
of a grain quality defect known as black point. Both wheat
Polyphenol Oxidase and barley are prone to this defect, which manifests as a dark
Polyphenol oxidases (PPO) catalyze the oxidation of phenolic stain in the seed coat or husk, respectively, at the embryo end
compounds to produce brown pigments on the cut or dam- of the grain and is associated with warm, humid conditions
aged surfaces of fruits and vegetables and can cause major during ripening (Figure 4). Unfortunately, this pigmented
economic losses. PPO located in the seed coat of ripe wheat tissue fragments during milling and black specks are carried
grains is able to hydroxylate certain phenols in the s-position through into wheat flour or semolina. While it has no effect on
adjacent to the existing OH group to form orthodiphenols. processing properties or texture, the specks are visible in
Figure 3 Yellow alkaline noodle sheets: from left to right control (2% sodium carbonate), 10 mM tyrosine (polyphenol oxidase substrate),
and 10 mM tyrosine plus 10 mM tropolone (specific polyphenol oxidase inhibitor).
PROTEINS | The Enzymes Associated with Food Grains and Their Functions in Processing 143
noodle and pasta products and high levels of black point result durum wheat that lack the D-genome Wheat Genomics or in a
in downgrading and substantial economic losses. small number of bread wheat cultivars that contain a non-
The enzymes discussed in this section all have specific functional version of the gene. Since adequate lutein intake is
functions within the tissues of developing grains, although in associated with a reduction in the incidence of glaucoma and
some instances the exact role has not been identified unequiv- macular degeneration, a major cause of blindness in elderly
ocally. Their physiological functions are quite distinct from people, this modification may have implications for absorp-
their effects on processing and end-product appearance. tion, bioavailability, and attempts to improve eye health
While in many cases it is possible, or at least conceivable, to through dietary supplementation.
develop germplasm with extremes of variability for levels of Enzymes have been implicated in the deterioration in qual-
these enzymes, some caution is required in case such dramatic ity of rice during storage. This deterioration may be linked to
changes have adverse effects on plant growth, disease resis- lipid peroxidation, peroxidative changes in polyunsaturated
tance, or grain yield. fatty acids that in turn are governed by the balance of produc-
tion and dissipation of reactive oxygen species. Peroxidases are
key scavengers of reactive oxygen species and recent studies
have indicated that the activity of this enzyme declines during
Enzymes that Are Active in Ripe, Quiescent Grains storage of rice.
Despite the low moisture content of ripe cereal grains and the
death of the seed coat and starchy endosperm tissues during
grain ripening, it is clear that respiration and some other Endogenous Enzyme Inhibitors
enzyme-catalyzed reactions continue at a slow rate during stor-
age of harvested grain. Some of these activities result in the Cereal and legume grains contain inhibitors against a range
modification of the structure of grain constituents and secondary of enzymes such as proteinase, a-amylase, endoxylanase, and
metabolites, while others may impact on the storage quality or limit dextrinase. While some inhibitors are only effective
the capacity to maintain freshness and palatability. In addition, against exogenous enzymes of microbial or insect origin,
ripe grains contain a range of enzymes such as exopeptidases and others are capable of inhibiting endogenous grain enzymes.
b-amylase that are sequestered in active or inactive form. These The physiological function of these inhibitors is not certain but
enzymes can spring into action once endo-acting enzymes, it has been suggested that they may be involved in plant
synthesized during germination or sprouting, catalyze the initial defense mechanisms, protecting the stored reserves in the
depolymerization of protein and starch. seed from invading microbes and insects, and/or play a regu-
At maturity and depending on the genotype and growing latory role in plant development and metabolism. A role in the
conditions, wheat grains contain varying levels of xanthophyll plant defense mechanisms seems plausible given the effective-
(hydroxy carotene) present primarily as free lutein. During ness of many endogenous inhibitors against hydrolytic
storage of wheat grain and wheat flour, and again depending enzymes from microbial and insect sources. Support for a
on genotype as well as temperature and humidity, considerable regulatory role is less plausible and requires unequivocal evi-
proportions of the free lutein (up to 60%) may be converted to dence that the inhibitors reduce the activity of particular
mono- and di-fatty acid esters. A specific GDSL-lipase (the endogenous grain enzymes at physiological concentrations
prefix GDSL is added to distinguish this group of lipases from and that the inhibitors are located in the same tissue as the
common lipases and refers to specific sequence of amino acids target enzyme. Most of the proteinase inhibitors that have been
at the active site of this group of lipases), coded by a gene characterized to date are inhibitors of a particular class of
located on chromosome 7D, appears to be responsible for protein hydrolases, the serine proteinases. As yet, there has
esterification possibly acting as a trans-esterase to transfer lino- been no conclusive demonstration of serine proteinases of
leic and palmitic acids from membrane phospholipids to the the chymotrypsin family or their genes in plants. In the case
hydroxyl groups of lutein. The lipase is synthesized during the of the bifunctional a-amylase/subtilisin inhibitor that is com-
early to middle stages of grain development but does not mon in barley grains, there is clear evidence that it can signif-
catalyze esterification of lutein until the ripe grain is stored icantly reduce the activity of endogenous high-pI a-amylase
under warm, dry conditions. Esterification does not occur in extracted from germinated or malted grain and ameliorate
144 PROTEINS | The Enzymes Associated with Food Grains and Their Functions in Processing
some of the deleterious effects of high-pI a-amylase on loaf genomics technologies, coupled with rapid functional analysis
volume and structure. However, recent studies of gene expres- systems, are now available to researchers. These new technol-
sion have indicated that the inhibitor is primarily located in ogies can be confidently expected to identify more genes and
the maternal seed coat tissues of barley grains rather than reveal potential strategies for further manipulation of enzyme
endosperm. Similarly, whereas proteins of the serpin super- and grain constituent levels relevant to the grains industries in
family are localized in both vegetative and endosperm tissues the near future.
of barley grains, they only inhibit specific mammalian serine
proteinases. Limit dextrinase inhibitors isolated from barley
inhibit the activity of endogenous limit dextrinase and appear
Exercises for Revision
to be ineffective against a number of other enzymes capable of
hydrolyzing a-1,6-linkages in dextrins. In theory at least, these What enzymes are important in the development and
breakdown of endosperm cell walls?
inhibitors could interfere with the malting process although
a role in commercial malting has not been demonstrated. What enzymes are important during germination?
When are the different a-amylase families synthesized in
Two groups of endoxylanase inhibitors (XIP-type and TAXI-
wheat grains?
type) have been isolated from cereal grains that are effective
against fungal endoxylanases and both bacterial and fungal Which enzymes present in ripe wheat grains affect end-
product quality?
endoxylanases, respectively. Microbial endoxylanases are
commonly used in bread making to improve loaf volume. What methods are used to overcome the deleterious effects
of some enzymes on the quality of wheat-based end
Irrespective of possible roles in plant defense and regula-
products?
tion, significant numbers of inhibitors are active either against
endogenous enzymes that are important in processing or What is black point and does it affect end-product quality?
against exogenous enzymes, usually of fungal or bacterial ori-
gin, that are added as ingredients to improve processing and
quality of end products. In these instances, the inhibitor and Exercises for Readers to Explore the Topic Further
the enzyme are brought into contact as a result of milling or
deliberate physical admixture and there may be important Fully characterize all the gene families involved in endo-
implications for industrial applications. sperm (aleurone and starchy endosperm) cell wall synthesis
and degradation, develop a better understanding of their
respective roles and interactions, and explore ways of mod-
Future Prospects ifying their activity to improve products and processing.
Genetic and molecular characterization of late-maturity a-
Many cereal products and processes depend on, or are amylase in wheat to assist in the development of varieties
adversely affected by, specific enzymes either that are present that maintain low a-amylase activity and to provide new
in grains at maturity or that develop as the result of interactions insights into the processes involved in the control of grain
between grains and the environment. As more of these development.
enzymes are recognized, and their impact quantified, many Utilize genetic and molecular techniques to optimize the
opportunities will be presented to improve products and to grain content of enzymes and constituents that impact on
develop new and novel products and processes. In many cases, end-product quality and grain storage without compromis-
there is already sufficient genetic variation within existing ing productivity.
germplasm or near relatives for the selection of lines with
enzyme levels more appropriate to the particular end-use pro-
See also: Barley, Rice and Maize Processing: Barley: Malting;
cess, while in other instances, it may be possible via mutation,
Carbohydrates: Cereals: Chemistry and Physicochemistry of Non-
genetic engineering, or modification of milling techniques to
starchy Polysaccharides; Starch: Synthesis; Fats: Lipid Chemistry;
extend variation beyond that currently available. Genetic engi-
Genetics of Grains: Wheat Genetics and Genomics; Proteins:
neering, in particular, opens up the possibility of introducing
Protein Synthesis and Deposition; The Basics: Grain: Morphology of
genes encoding novel enzymes into cereal grains, overexpres-
Internal Structure; The Cereal Grains: An Overview of the Family of
sing genes controlling synthesis of existing enzymes, or synthe-
Cereal Grains Prominent in World Agriculture; Wheat Processing:
sizing enzymes in tissues such as the starchy endosperm where
The Gluten Proteins of the Wheat Grain in Relation to Flour Quality;
they are currently absent. As an example, identification of the
Wheat-Based Foods: Noodles: Asian Wheat Flour Noodles.
genes should reveal potential strategies for the manipulation of
cell wall (1 ! 3,1 ! 4)-b-D-glucan and arabinoxylan levels, for
the enhancement of cereal quality and productivity. The 3-D
structure of one of the barley (1,3;1,4)-b-glucan endohydro- Further Reading
lases has been solved and the structural information used to Asenstorfer RE, Appelbee MJ, and Mares DJ (2009) Physical-chemical analysis of non-
rationally design mutated forms of the enzyme with enhanced polyphenol oxidase (non-PPO) darkening in yellow alkaline noodles. Journal of
thermostability. The increased thermostability of this barley Agricultural and Food Chemistry 57: 55565562.
(1,3;1,4)-b-glucan endohydrolase could have commercial Chen TF and Chen CL (2003) Analysing the freshness of intact rice grains by colour
determination of peroxidase activity. Journal of the Science of Food and Agriculture
potential through its ability to overcome filtration problems 83: 12141218.
and other difficulties encountered in the malting and brewing Dominguez F and Cejudo FJ (1996) Characterization of the endoproteases appearing
industries. Increasingly sophisticated and high-throughput during wheat grain development. Plant Physiology 112: 12111217.
PROTEINS | The Enzymes Associated with Food Grains and Their Functions in Processing 145
Fincher GB (1989) Molecular and cellular biology associated with endosperm Howitt C and Pogson BJ (2006) Carotenoid accumulation and function in seeds and
mobilization in germinating cereal grains. Annual Review of Plant Physiology and non-green tissues. Plant, Cell and Environment 29: 435445.
Plant Molecular Biology 40: 305346. Mares D and Mrva K (2008) Late maturity a-amylase: low falling number in the absence
Fincher GB and Stone BA (1986) Cell walls and their components in cereal grain of preharvest sprouting. Journal of Cereal Science 47: 617.
technology. Advances in Cereal Science and Technology 8: 207295. Shirley BW (1998) Flavonoids in seeds and grains: Physiological function,
Fuerst EP, Anderson JV, and Morris CF (2006) Delineating the role of polyphenol agronomic importance and the genetics of biosynthesis. Seed Science Research
oxidase in the darkening of alkaline wheat noodles. Journal of Agricultural and Food 8: 415422.
Chemistry 54: 23782384. Stewart RJ, Varghese JN, Garrett TPJ, Hoj PB, and Fincher GB (2001) Mutant barley
Garcia-Olmedo F, Salcedo G, Sanchez-Monge R, Royo J, and Carbonero P (1987) Plant (1 ! 3,1 ! 4)-b-glucan endohydrolases with enhanced thermostability. Protein
proteinaceous inhibitors of proteinases and a-amylases. Oxford Surveys of Plant Engineering 14: 245253.
Molecular and Cell Biology 4: 275334. Williamson PW (1997) Black point of wheat: In vitro production of symptoms, enzymes
Hidalgo A and Brandolini A (2012) Lipoxygenase activity in wholemeal flours from involved and association with Alternaria alternate. Australian Journal of Agricultural
Triticum monococcum, Triticum turgidum and Triticum aestivum. Food Chemistry Research 48: 1319.
131: 14991503.
Proteomic Analysis Relevant to Cereal Grains
A Juhasz, Centre for Agricultural Research of the Hungarian Academy of Sciences, Martonvasar, Hungary
R Haraszi, Campden BRI, Gloucestershire, UK
F Bekes, FBFD PTY LTD, Beecroft, NSW, Australia
DJ Skylas, Grain Growers Limited, North Ryde, NSW, Australia
CW Wrigley, The University of Queensland, Brisbane, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
Metabolites
Sugars, nucleotides, amino acids, lipids
Environment
detection techniques (e.g., NaCl ionization in the ESI of a mass gel electrophoresis, liquid chromatography, and mass spec-
spectrometer suppresses the peptide ionization), they need to trometry and subsequent protein identification (Figure 2).
be removed at a proper time of analysis. Starch and complex
carbohydrates can also seriously reduce protein extraction,
Gel-Based Methods
which might be a primary problem in proteomic analyses of
maize or rice seeds, however. The ideal extraction method Gel-based proteomics is still the main approach used in grain
should be compatible with downstream protein analysis by proteomics. For decades, one-dimensional gel electrophoresis
148 PROTEINS | Proteomic Analysis Relevant to Cereal Grains
(1-D GE) was the most popular method in protein chemistry including Coomassie or silver staining, but the more common
due to its ability to determine and compare the molecular and routinely used stains now are fluorescent dyes, such as
weights of proteins. It was used in different approaches, such SYPRO Ruby protein stain or CyDyes (Cy2, Cy3, and Cy5).
as to compare and determine the number and size of seed Two-dimensional Fluorescence Difference Gel Electrophoresis
storage proteins in wheat. It is also used in immunodetection (2-D DIGE) was developed to overcome problems related to
or Western blotting. Today, it is most often used as the second moderate gel-to-gel reproducibility observed in 2-D gel electro-
dimension in two-dimensional gel electrophoresis. Two- phoresis. In this approach, two protein samples and an internal
dimensional gel electrophoresis (2-D GE), introduced in standard are labeled prior the gel electrophoresis with one of
1975, is the most widely used gel-based technique to separate the three size- and charge-matched, spectrally resolvable fluo-
proteins. The first dimension involves isoelectric focusing rescent dyes. The three samples are then mixed and uploaded
(IEF), in which proteins are fractionated based on their isoelec- onto a single IPG strip followed by SDS-PAGE. The simul-
tric points (pI) across a specific pH range using commercially taneous separation of multiple fluorescent-labeled samples,
available immobilized pH gradient (IPG) gel strips. The together with an internal standard, results in fewer gels needing
second-dimension of fractionation resolves the proteins on to be analyzed in a significantly shorter time frame. Using the
the basis of their molecular mass, using sodium dodecyl sulfate internal standard, the gel-to-gel variation can be eliminated as
polyacrylamide gel electrophoresis (SDS-PAGE). The SDS no technical replicates are needed to confirm differences in
detergent binds to the denatured proteins, overriding any protein abundance.
intrinsic charge differences, to provide a uniform charge-to-
mass ratio, allowing proteins to resolve according to their
respective sizes. Due to its excellent resolving power, the tech- Gel-Free Methods: LC-MS and LC-MS/MS Methods
nique is suitable to separate over 1000 proteins on a single gel. In proteomics, high-throughput protein identification is usu-
The 2-D GE proteomic map in Figure 3 represents the pro- ally done by mass spectrometry (MS) or coupling it with more
teins of immature wheat grain flour. It shows the polypeptides sensitive methods. Shotgun or bottom-up proteomic
extracted from the immature endosperm of the soft wheat approaches are targeting the analysis of a peptide mixture
variety Wyuna, the endosperm being the floury part of the that is a derivative of the target proteome. Proteolytic enzymes
grain, excluding the germ, embryo, and bran layers. The pro- or specific chemicals can be used to digest proteins before
teome map of Wyuna appears in two parts, reflecting two chromatographic separation that is followed by peptide ioni-
different pH ranges used in IEF. Protein extracts were fraction- zation and detection. The general use of trypsin digestion failed
ated in both acidic (pH 47) and basic ranges (pH 611). The in the case of gluten, which contains low level of arginine and
proteomic pattern is thus a map of pI, in the horizontal dimen- lysine. Replacement by chymotrypsin was first shown to effec-
sion increasing from left to right, and of molecular size, in the tively produce peptides suitable for chromatography. A novel
vertical dimension. SDS-PAGE provides fractionation on the combination of direct enzymatic digestionLC-MS/MS (ESI
general basis of the size of the protein molecules, from largest ion trap) method was developed for the semiquantification
(top) to smallest (bottom). of gluten traces in food samples, which applies a two-step
After separation, the proteins in the gels must be visualized. multienzymatic digestion. Recent studies are more and more
There are various stains available for protein visualization, acknowledging the odd nature of cereal proteins and
1
4
5
2 6
Figure 3 The acidic and basic proteome maps of immature (17 days postanthesis) wheat grain endosperm of the soft Wyuna variety. Protein spots
were characterized by N-terminal amino acid sequencing and identified from protein-sequence database interrogation.
PROTEINS | Proteomic Analysis Relevant to Cereal Grains 149
demonstrating the need to use unconventional proteomic The relatively novel aspect of proteomic analysis is the
approaches for proteins like gluten. subsequent stages of postseparation microcharacterization of
Top-down proteomics involves the gas-phase ionization of the individual protein spots. The ability to characterize and
intact proteins and subsequent high-resolution mass measure- identify proteins in a high-throughput manner is a critical
ment of intact protein ions followed by their direct fragmenta- factor in proteome projects. Two widely used protein-
tion inside the mass spectrometer without prior digestion. Top- characterization methods are N-terminal amino acid sequenc-
down approaches enable a more complete characterization of ing and MS, with the latter being the preferred method of
protein isoforms and any PTMs than the bottom-up approaches choice used in proteome projects.
but are not exempt from limitations (e.g., high purity, HPLC N-terminal amino acid sequencing has been a popular
separation, and high molecular weight (>50 kDa)). New meth- method for characterizing proteins in the past. The most com-
odological advances with ultra high performance liquid chro- mon form of this method is the traditional Edman degradation
matography (UHPLC) and mass spectrometers with ion chemistry, which can be used to sequence proteins from gels,
mobility are opening up new ways for top-down approaches or it can be carried out on proteins electroblotted from the
in the proteomics field. proteome gel to a polyvinylidene difluoride membrane. In
Matrix-assisted laser desorption/ionization time of flight either case, Edman chemistry involves the reaction of phenyl
(MALDI-TOF) can be used for indirect quantification of cereal isothiocyanate with the free amino group of the protein; deriv-
proteins like gliadins even in processed foods and the direct atives are cleaved and then converted to phenylthiohydan-
identification and exact molecular mass determination of cereal toins, which differ in their amino acid side chain, allowing
proteins without prior separation. MALDI-MSPMF (peptide them to be characterized by retention time using reversed-
mass fingerprinting) is a combination, which involves enzy- phased high-performance liquid chromatography. This is not
matic digestion of proteins resulting in peptide fingerprints. the preferred method in high-throughput large-scale proteome
Sensitivity and resolution of LC-MS methods (e.g., MALDI- projects, due to the relatively low number of samples that can
TOF and ESI-Q) can be increased with the application of be processed in parallel (generally only three or four samples),
coupled (tandem MS) techniques like ESI-QqQ and ESI-Q- its low sensitivity, and the time and cost for analysis.
TOF, where selected ionized peptides are fragmented before A gel spot can be directly ionized by MALDI and detected
entering the detector of the mass spectrometer. The monitoring with a TOF or can be destained, reduced, enzymatically
of specific mass transitions as SRM (single reaction monitor- digested, and then extracted (in gel digestion) in order to
ing) or multiple reaction monitoring (MRM) provides a unique have a peptide mixture, which then can be analyzed with LC-
identification and quantification tool for the analysis of com- MS or LC-MS/MS for protein composition.
plex mixtures including overlapping peptide characteristics. Peptides and proteins are identified from the recorded and
processed mass spectra with a peptide matching algorithm and
against a protein database. Besides experimental conditions
Identification and Characterization of Protein Gel Spots including the chromatographic separation, the precision of
This procedure of 2-D GE has been available for some decades, peptide identification is highly depending on the quality of
providing proteomic maps such as in Figure 3. Six spots are the mass spectra and the database.
numbered as examples of polypeptide identification. Table 1 Peptides with completely different amino acid sequences
shows the results from excising these six spots from these gels and functionality may coelute during LC separation, have
for identification by N-terminal sequencing. As a result of identical or very similar masses, and may fragment similarly.
matching these sequences to databases, the identities could This is particularly true for proteins belonging to a protein
be determined for five of these six spots. family like alpha-gliadins and low molecular weight (LMW)
Table 1 N-terminal amino acid sequence characterization of immature wheat endosperm proteins, taking the six numbered spots in Figure 3 as
examples
1 XATFTLPD Superoxide dismutase 87.5% in 8 aa Wheat N-terminal sequence starts at residue 22 of O82571 with residues
(EC 1.15.1.1) 121 possibly being a signal sequence or truncated. Also
matches to P93606 and Q96185
2 SGPWMCYV a-Amylase inhibitor 100% in 7 aa Wheat P10846 (fragment). Also matches other a-amylase inhibitors
WDAI-3 P01084 and P01085
3 MKLIAAYL 60S acidic ribosomal 100% in 8 aa Wheat Subunits P1 and P2 exist as dimers at the large ribosomal subunit
protein P2 (ribosomal P05390
protein A)
4, 5 NMQVDPSG g-Gliadin 100% in 8 aa Wheat N-terminal sequence starts at residue 20 of P21292 as signal
sequence of precursor is residues 119
6 DILRSDQP No close matches
Protein sequences were used to interrogate SWISS-PROT and TrEMBL databases via FASTA algorithm.
150 PROTEINS | Proteomic Analysis Relevant to Cereal Grains
glutenin subunits. Additionally, the protein content in a gel that is, the many different characteristics of the plant and of the
spot is usually so little that without sufficient sensitivity and grain produced from it. There are different subdivisions of
resolution, the detection and identification of their peptides proteomics that use different approaches and targets to answer
are not possible. different questions related to protein identification, expres-
Generally, marker peptides are used for the quantitative sion, function, interactions, etc. These different types are sum-
detection of proteins. The first step is to do discovery work in marized on Figure 4.
order to map the most unique and detectable peptides using a
high-resolution mass spectrometer like an ESI-Q-TOF or a
software tool such as Skyline or the ProPepper database,
Proteome Mining
which provides the starting point in the marker selection pro-
cess. After selecting a potential pool of peptides, the next step is The water-soluble albumins and salt-soluble globulins of
the experimental optimization of the quantification method wheat grain comprise mainly metabolic and protective
using, for example, an ESI-QqQ and synthetic peptides in order proteins, while at least some of the insoluble structural pro-
to tune up the mass spectrometer for the accurate detection of teins may be extracted with dilute acid or alkali in the glutenin
specific predefined mass transitions of the targeted peptides. fraction. Albumins and globulins, each accounting for 10%
The size of a database due to false-positive hits and the of total flour proteins, are known to be soluble enzymes and
accuracy of entries (proper annotations) are further critical proteins soluble in polar solution, respectively. More than
factors in the relation of peptide matches to the protein source. 2000 protein spots can be extracted from flour with salt
This is relevant in the discovery phase and especially complex solution.
for the seed proteome, where the plant genomes are not fully The structure of the wheat grain can be categorized simply
sequenced yet. In targeted shotgun proteomics, the design of into three distinct fractions, the outer layers, the germ, and the
an SRM for protein quantification needs to fulfill the criteria endosperm. The endosperm plays the most important role
that the amount of peptides and that of proteins are equimolar, from a human consumption point of view and is the fraction
which is not the case for, for example, gluten, where the high studied in most detail, identifying a large number of proteins
level of homology and the multiple prevalence of sequences in the developing grain.
limit the establishment of quantitative relationships between a The proteins located in the wheat endosperm determine the
detected peptide, its individual protein source, its protein characteristics of elasticity, extensibility, and gas-holding abil-
family, and its plant origin. ity of the wheat dough. Particularly important are the proteins
within the gluten complex, the low- and high-molecular-
weight glutenin subunits, proteins, and the diverse group of
The Potential Value of Proteomics to Grain Science gliadins. This group of proteins shows extensive polymor-
phism across wheat genotypes. Wheat also contains smaller
Using proteomic analysis, we can identify the individual pro- amounts of other storage proteins such as globulins and
teins that are responsible for specific aspects of the phenotype, triticins and proteins such as amylase and protease inhibitors
Post-
Proteome translational
mining proteomics
Proteomics
Figure 4 Proteomic approaches used in grain science. Adapted from Juhasz J, Moolhuijzen P, Bellgard M, Appels R, and Bekes F (2013) Wheat grain
proteomics for the food industry. In: Toldra F and Nollet LML (eds.) Proteomics in Foods. Principles and Applications. Food Microbiology and Food
Safety, vol. 2, pp. 341379. New York: Springer.
PROTEINS | Proteomic Analysis Relevant to Cereal Grains 151
that may protect against insects and fungi. A range of enzymes Numerous studies have been carried for the identification
exist in very small amounts. and mapping of wheat proteins. Studies, for example, on ker-
Amphiphilic proteins are detergent-soluble membrane pro- nel responses to heat stress have enabled identification of
teins and together with gluten proteins play an important role many of the enzymes in kernel of wheat. However, the major-
in wheat quality. Some of them, like puroindolines, are lipid- ity of soluble proteins still need to be identified and mapped.
binding proteins and are strongly linked to dough-foaming Proteomic research activities have been reviewed describing
properties and to fine crumb texture. applications of proteomics in wheat research: dough quality,
The proteomic investigation of the wheat amyloplast can be starch properties, puroindoline, chromosomal location of cod-
carried out on whole amyloplasts or purified amyloplast mem- ing genes, effects of growth environment, insect damage to
branes, identifying the total number of 171 and 108 proteins, wheat grains, and genotype identification. The review of Juhasz
respectively. The largest class of proteins with a known func- el al. provides a table listing 38 application areas in 113 pub-
tion (24%) was found to be starch synthesis and storage lications, using the different techniques of proteomics in both
enzymes. Affinity electrophoresis using glucan (cornstarch or plant biology and food science.
amylopectin) as affinity substrate has been utilized successfully The development of techniques to improve extraction and
in the identification of enzyme-type proteins, especially the separation and to identify proteins and peptides is facilitating
proteins related to starch biosynthetic and hydrolyzing functional proteomics, together with the analysis of sub-
enzymes and their inhibitors. proteomes from small amounts of starting material, such as
Achievements of proteomic research on bran and germ seed. The combination of proteomics with structural and func-
tissues are described by Juhasz et al. tional analysis is increasingly applied to target subsets of pro-
teins. These next-generation proteomic studies differentiate
between special subproteomes and functional subproteomes
Functional Proteomics and contribute to understanding the processes controlling
cereal grain development, nutritional and processing
Functional proteomics aims to characterize how changes in
characteristics, and the changes caused by biotic and abiotic
protein localization, expression, and interaction are affected
stresses. The importance of abiotic stresses caused by global
in the different developmental stages or by different environ-
climate change has particular significance in the future of
mental cues. These processes include germination and seedling
agriculture and food industries. Advances in crop proteomics
development, tillering, stem elongation, head emergence,
provide the basis for interpreting changes in the grain as a
flowering, seed development, and dormancy. Functional pro-
result of climatic factors including photosynthetic stress, air
teomics is suitable to determine the dynamics of how proteins
pollutants, thermal stress including heat and cold, osmotic
are related into complex networks and how their expression
stress (drought, salt, and flooding), and metal stress to help
influenced by coexpressing proteins or other biomolecules
to eliminate or to reduce the negative effects of these factors on
present in the tissue. The approaches used are based on quan-
the food production chain
titative analyses of the proteome, combining approaches such
as analytical chemistry, biochemistry, MS, and bioinformatics.
For instance, large-scale proteomic experiments enable us to Posttranslational Proteomics
understand the key players and mechanisms involved in grain
Polypeptides produced during translation often undergo dif-
filling and seed maturation (i.e., the expression of cereal grain
ferent posttranslational alterations by modifications of amino
proteins) in order to understand how environmental stresses
acid side chains through the addition of a wide range of
affect wheat quality. These mechanisms determine regulatory
molecules, mostly carbohydrates, lipids, phosphate, nitrate,
principles of transcriptional activation and protein turnover in
or methyl groups. These modifications are covalent processing
response to abiotic or biotic stresses as well as to differences in
events that can modulate the activity, localization, turnover,
developmental processes in the different cereal species or
and interaction of many seed proteins. The resulted modifica-
cultivars.
tions, such as glycosylation, phosphorylation, methylation,
ubiquitination, and deamidation, of the proteins are often
associated with changes in the protein functions. PTM results
Expression Proteomics
in changes in the molecular weight, charge, and hydrophobic-
Quality attributes of a flour sample are defined by the allelic ity of proteins. The modified subproteomes can be studied at
composition of gluten proteins (the high molecular weight both protein and peptide levels using different gel-based anal-
(HMW) and low molecular weight (LMW) glutenin subunits ysis techniques, as well as MS. Glycosylation and phosphory-
together with gliadins) as well as their relative expression lation are two common forms of modifications affecting about
levels. Proteomic analysis, used to monitor the expression of 40% and 60% of the seed proteins. Glycosylation events are
gluten proteins during grain development, specially the glute- characteristic on all secreted and membrane proteins, while
nin subunits, provides useful information about polymer phosphorylation is often linked to proteins related to enzyme
formation and helps in relating protein composition to quality metabolism, stress responses, and growth. Storage proteins of
attributes. Nongluten proteins encompass smaller percentages the cereal seeds are rich in amino acids like proline or gluta-
of the seed protein content; however, they play a crucial role mine. Deamidation of glutamine-containing proteins into glu-
in different metabolic processes during grain development, tamic acid results in higher water-binding capacity. Polymer
ripening, and desiccation, and they can have a relevant role formation of storage proteins in wheat gluten may result from
in defense mechanisms against biotic and abiotic stresses. PTM events, by which sulfur-containing amino acids are
152 PROTEINS | Proteomic Analysis Relevant to Cereal Grains
bound to each other via disulfide bonds, so that a stabile combined with Western blotting may be used to overcome
chemical interaction is formed. These abilities of cereal and this challenge. The power of proteomics used for the identifi-
especially wheat grain proteins are important in the structure cation and characterization of protein components in the grain
function relationship analysis of wheat gluten for food prod- is greatly magnified when combined with other postgenomic
ucts and also for nonfood uses. techniques such as metabolomics. Such multidisciplinary ana-
lyses, including tools of proteomic, metabolomic, and bioin-
formatic analyses, can provide direct evidence of the effects of
Computational and Structural Proteomics
protein expression.
Structural proteomics deals with the complex information hid- The next major challenge toward this aim is to identify the
den in the amino acid sequence of a protein. It determines how constituents of the cereal grain proteome in order to under-
the order and characteristics of the constituent amino acids stand the cereal genome. Of particular importance will be the
influence the secondary and tertiary structure of a protein deciphering of protein alterations between normal and stressed
and how these features determine the ability of these proteins conditions to permit the identification and prioritization of
to function as interacting partners in protein complexes. The agronomically important targets. Proteome analysis aims to
major challenge of structural proteomics is to assign structure identify changes not only in protein expression but also in
data determined by x-ray or NMR crystallography or in silico PTMs, proteinprotein interactions, cellular and subcellular
modeled to protein sequences obtained from genome and distribution, and temporal patterns of expression.
transcriptome sequencing projects by the use of high- The power of proteomics as a tool for the discovery of
throughput characterization of the 3-D structure. Structural biological pathways and processes related environmental
proteomic studies of cereal grains can be used to analyze changes is now well established. Indeed, proteomics has
enzyme complexes present in the seed, to identify receptor already uncovered many potential targets for specialized qual-
functions by the analysis of different proteinligand interac- ity purposes or different biotic and abiotic stress conditions.
tions, to predict how pathogens (including viruses, fungi, or The current era of proteomics is now beginning to investigate
insects) interact with the host plant, or to identify and predict how this technology can serve the breeders, the food industry,
allergen components present in the seed. Structural proteomics and the consumer for high-throughput identification and
strongly builds on computational biology utilizing the poten- prognostic applications.
tial of sequence data analysis and 3-D modeling.
Computational proteomics is a term generally describing a
broad range of computational tools, including algorithms, bio- Exercises for Revision
informatics techniques, and data acquisition models to analyze
protein or peptide related sequence data. The collection and Develop an extended definition of the proteomics concept,
evaluation of data from protein databases, combined with gel- including mention of the genome, growth conditions, tis-
based and MS-based or immunoanalytical computational ana- sue identity, stage of maturity, and functional relationships.
lyses, can deliver outputs in drug discovery, food-safety analyses, Explain the relative advantages and disadvantages of gel-
and even the development of personalized medicine. based and gel-free technologies for proteome analysis.
Which is better suited for polypeptide identification and
for eventually elucidating the functions of the many
Improving on Current Proteome Technology polypeptides?
Several distinct versions of proteomics are described:
The successful completion of wheat genome and genome- expression, structural, functional, posttranslational
sequencing projects of related grain species will lead us to proteomics, etc. Explain the differences between these
better understand mechanisms involved in the stages of plant concepts and their successive roles in understanding the
development, such as seed development, maturation, seed sequence of events from genes to mature grain.
dormancy, and germination. Additional insights will come Describe some practical applications of proteomics in grain
from using genome sequences combined with genotype-level science.
information, plus additional achievements in protein separa- Construct a diagram (and explanation) showing relation-
tion and identification techniques. Results can be expected in ships between genomics, various forms of proteomics, and
genotype-level complex information about what features are other -omics. Describe their relevance to grain science.
responsible for specialized end-use quality attributes, for sus- Consider thiolomics and zymomics, as examples (but not
tainable quality under different environmental conditions, and economics). The following reference provides an introduc-
for the complexity of wheat-allergen proteins. To study these tion to zymomics: Kosar-Hashemi B, Irwin JA, Higgins J,
mechanisms, projects need to involve multidisciplinary Rahman S, and Morell MM (2006) Isolation, identification,
approaches, such as genomics, proteomics, metabolomics, and characterization of starch-interacting proteins by 2-D
and bioinformatics. affinity electrophoresis. Electrophoresis 27: 18321839.
Currently, proteomics receives criticisms related to its low
reproducibility, evidenced by gel-to-gel variations between lab-
oratories. By the standardization of extraction and separation Exercises for Readers to Explore the Topic Further
protocols, these issues will be improved. Identification of
weakly expressed molecules via MS remains problematic, It is stated that the results of proteomic analysis of grain
although techniques such as protein and peptide chips proteins have wide-ranging implications for research on
PROTEINS | Proteomic Analysis Relevant to Cereal Grains 153
structurefunction relationships for grain proteins. Find dimensional electrophoresis, three proteases and tandem mass spectrometry.
out about some examples of proteomics being used to Proteome Science 9: 10. http://dx.doi.org/10.1186/1477-5956-9-10.
Finnie C, Sultan A, and Grasser KD (2011) From protein catalogues towards targeted
demonstrate how specific proteins influence significant
proteomics approaches in cereal grains. Phytochemistry 72: 11451153.
aspects of grain development and utilization. Gao L, Wang A, Li X, et al. (2009) Wheat quality related differential expressions of
Develop some diagnostic applications of proteomic analy- albumins and globulins revealed by two-dimensional difference gel electrophoresis
sis, based on the knowledge that specific polypeptides dis- (2-D DIGE). Journal of Proteomics 73: 279296.
tinguish one cereal variety from others (variety Gao L, Ma W, Chen J, et al. (2010) Characterization and comparative analysis of wheat
high molecular weight glutenin subunits by SDS-PAGE, RP-HPLC, HPCE, and
identification), or one grain tissue from another (milling) MALDI-TOF-MS. Journal of Agricultural and Food Chemistry 58: 27772786.
or one grain genus from another, or grain tissue from that Hashiguchi A, Ahsan N, and Komatsu S (2010) Proteomics application of crops in the
of a predator. context of climatic changes. Food Research International 43: 18031813.
Investigate applications of proteomics in plant breeding to Jorrin-Novo JV, Komatsu S, Weckwerth W, and Wienkoop S (eds.) (2014) Plant
Proteomics Methods and Protocols. In: Methods in Molecular Biology, vol. 1072.
achieve aims such as improved pathogen resistance and
Totowa, NJ: Humana Press.
better grain quality. Juhasz J, Moolhuijzen P, Bellgard M, Appels R, and Bekes F (2013) Wheat grain
Construct a diagram (and explanation) showing relation- proteomics for the food industry. In: Toldra F and Nollet LML (eds.) Proteomics in
ships between genomics, various forms of proteomics, and Foods. Principles and Applications. Food Microbiology and Food Safety,
other -omics. Describe their relevance to grain science. 2: pp. 341379. New York: Springer.
Kav NNV, Srivastava S, Yajima W, and Sharma N (2007) Application of proteomics to
Consider thiolomics and zymomics, as examples (but not investigate plant-microbe interactions. Current Proteomics 4: 2843.
economics). The following reference provides an introduc- Manjasetty BA, Bussow K, Panjikar S, and Turnbull AP (2012) Current methods in
tion to zymomics: Kosar-Hashemi B, Irwin JA, Higgins J, structural proteomics and its applications in biological sciences. 3 Biotech 2(2):
Rahman S, and Morell MM (2006) Isolation, identification, 89113.
Martnez-Maqueda D, Hernandez-Ledesma B, Amigo L, Miralles B, and Gomez-Ruiz J
and characterization of starch-interacting proteins by 2-D
(2013) Extraction/fractionation techniques for proteins and peptides and protein
affinity electrophoresis. Electrophoresis 27: 18321839. digestion. In: Toldra F and Nollet LML (eds.) Proteomics in Foods. Principles and
Applications. Food Microbiology and Food Safety, vol. 2, pp. 2150. New York:
Springer.
See also: Breeding of Grains: Wheat Breeding: Exploiting and McLaffety FW, Breuker K, Jin M, et al. (2007) Top-down Ms, a powerful complement to
the high capabilities of proteolysis proteomics. FEBS Journal 274: 62566268.
Fixing Genetic Variation by Selection and Evaluation; Genetics of
Miernyk JA and Hajduch B (2011) Seed proteomics. Journal of Proteomics
Grains: Development of Genetically Modified Grains; Genomics of 74: 389400.
Food Grains; Wheat Genetics and Genomics; Grain Marketing and Pappini D, Hojrup P, and Bleasby A (1993) Rapid identification of proteins by peptide-
Grading: Identification of Varieties of Food Grains; Proteins: Protein mass fingerprinting. Current Biology 3: 327332.
Synthesis and Deposition; The Protein Chemistry of Cereal Grains; Sealey-Voyksner JA, Khosla C, Voyksner RD, and Jorgenson JW (2010) Novel aspects
of quantitation of immunogenic wheat gluten peptides by liquid chromatography-
Thiolomics of the Gluten Protein Network of Wheat Dough; The mass spectrometry/mass spectrometry. Journal of Chromatography A
Basics: Grain and Plant Morphology of Cereals and how characters 1217: 41674183. http://dx.doi.org/10.1016/j.chroma. 2010.01.067.
can be used to identify varieties; Grain: Morphology of Internal Shewry PR, DOvidio R, Lafiandra D, Jenkins JA, Mills NF, and Bekes F (2009) Wheat
Structure; Wheat Processing: The Gluten Proteins of the Wheat grain proteins. In: Khan K and Shewry PR (eds.) Wheat Chemistry and Technology,
pp. 223298. St Paul, MN: AACC Press.
Grain in Relation to Flour Quality.
Skylas DJ, Mackintosh JA, Cordwell ST, et al. (2000) Proteome approach to the
characterisation of protein composition in the developing and mature wheat-grain
endosperm. Journal of Cereal Science 32: 169188.
Skylas DJ, Copeland L, Rathmell W, and Wrigley CW (2001) The wheat-grain proteome
as a basis for more efficient cultivar identification. Proteomics 1: 15421546.
Further Reading Skylas DJ, Van Dyk D, and Wrigley CW (2005) Proteomics of wheat grain. Journal of
Cereal Science 41: 165179.
Branlard G and Bancel E (2007) Protein extraction from cereal seeds. In: Thiellement H, Tasleem-Tahir A, Nadaud I, Girousse C, Martre P, Marion D, and Branlard G (2011)
Zivy M, and Damerval C (eds.) Plant Proteomics Methods and Protocols. Methods Proteomic analysis of peripheral layers during wheat grain development.
in Molecular Biology, vol. 355, pp. 1527. Totowa, NJ: Humana Press. Proteomics 11: 371379.
Camafeita E, Sols J, Alfonso P, Lopez JA, Sorell L, and Mendez E (1998) Selective
identification by matrix-assisted laser desorption/ionization time-of-flight mass
spectrometry of different types of gluten in foods made with cereal mixtures. Journal
of Chromatography A 823: 299306. Relevant Websites
Colinge J and Bennett KL (2007) Introduction to computational proteomics. PLoS
Computational Biology 3(7): e114. http://dx.doi.org/10.1371/journal. http://expasy.org Proteomics tools.
pcbi.0030114. http://propepper.net ProPepper is a protein, peptide and epitope analysis suitable for
Cramer R and Westermeier R (eds.) (2012) Difference Gel Electrophoresis (DIGE). cereal prolamin analysis.
Methods and Protocols. In: Methods in Molecular Biology, vol. 854. Totowa, NJ: www.uniprot.org Uniprot is a comprehensive database and analysis tool used protein
Humana Press. science.
Dupont FM, Vensel WH, Tanaka CK, Hurkman WJ, and Altenbach SB (2011) http://wiley-vch.de Proteomics is a scientific journal specializing in articles on
Deciphering the complexities of the wheat flour proteome using quantitative two- proteomics.
Thiolomics of the Gluten Protein Network of Wheat Dough
F Bonomi and S Iametti, University of Milan, Milan, Italy
2016 Elsevier Ltd. All rights reserved.
related to structural issues in these proteins. These include (1) side chains, and of thioldisulfide exchange reactions.
the role of noncovalent (mostly hydrophobic) interactions Gliadins also interact with the flour lipids or with added
in network formation; (2) the availability of thiols and disul- lipids, and there is a general consensus on their role in deter-
fide to exchange reactions, as determined by overall and mining dough extensibility. Gliadins have molecular masses
local protein flexibility, in turn related itself to protein solv- ranging from 30 to 45 kDa and had been subdivided into a/b,
ability and to the intensity of physical deformation; and g, and o fractions, according to their mobility in polyacryl-
(3) the possible role of low-molecular-weight thiols and amide gel electrophoresis (PAGE) at acid pH. Alpha- and
of low-abundance thiol-rich proteins in facilitating thiol b-gliadins are currently considered a single gliadin (a/b
disulfide exchange reactions. type) and contain six Cys residues in their C-terminal
Structural studies on proteins that cannot be solubilized domain. Eight Cys residues are found in the C-terminal
without resorting to the use of denaturants (alcohols or deter- domain of g-gliadins, whereas no cysteine is present in
gents) are intrinsically complicated. Methods capable of eval- o-gliadins.
uating protein surface hydrophobicity in systems made up of The glutenin fraction in wheat consists of protein aggregates
water-insoluble proteins have been developed and applied to from about 60 000 to more than 10 millions Da. Glutenin sub-
protein characterization in cereal-based starting materials and units have been divided into high-molecular-weight glutenins
products without resorting to protein separation that necessarily (HMW-GS) and low-molecular-weight glutenins (LMW-GS).
uses solvent systems that affect protein structure and interactions HMW-GS represent 510% of gluten proteins, but dough
or to standard proteomics or peptidomics approaches, in properties are reportedly strongly influenced by their relative
which a combination of separative techniques is used to clarify abundance. The bread-making quality of wheat genotypes
the whole pattern of proteins and peptides and to quantitate appears to be determined by the expression of specific HMW-
their relative abundance. Protein solubility in the absence/pres- GS genes. The N- and C-terminal domains in HMW-GS contain
ence of chaotropes and in the absence/presence of disulfide cysteine residues that form inter- and intrachain disulfide
reductants, such as dithiothreitol (DTT) and 2-mercaptoethanol bonds, whereas the central domain comprises repeated
(2-ME, also known as beta-mercaptoethanol), has been used to sequences that may promote intermolecular hydrogen bonding.
study hydrophobic and covalent interactions among proteins In particular, the x-type subunit includes four cysteines; three at
and their impact on the rheological properties of dough. These the N-terminal and one at the C-terminal. The y-type subunit
properties are reportedly affected by agents that block the has seven cysteines; five in the N-terminal domain, one within
reactivity of thiols in wheat proteins, thus preventing many of the repetitive domain, and one close to the C-terminus. These
the permanent structural changes that occur during further proteins promote the formation of larger glutenin polymers by
processing. forming intermolecular disulfides with other glutenins.
The thiolomics approach described in what follows aims LMW-GS are classified into B (4251 kDa), C (3040 kDa),
at combining information about the amount of thiols present and D types (5570 kDa), the B type being the most abundant.
(and accessible under native or denaturing conditions) in indi- D-type LMW-GS correspond to a o-gliadin-like protein with an
vidual protein components in a complex system such as grain additional cysteine that allows interchain disulfide bond for-
proteins. This approach offers the advantage of combining mation in the glutenin polymer. C-type LMW-GS have a struc-
information about thiols on individual proteins along with ture similar to a/b- and g-gliadin but contain an extra unpaired
their location on specific components (which may be identi- cysteine residue thought to be involved in an interchain disul-
fied by further proteomics analysis), thus contributing to a fide. The presence of an odd number of cysteines makes the
more comprehensive assessment of the molecular determi- C- and D-type LMW-GS behave as terminators of the growing
nants of grain performance. glutenin polymer chains, with a presumed negative effect on
flour quality.
B-type LMW-GS are subdivided in LMW-s, LMW-m, and
Protein Thiols in Cereal Systems LMW-i types, on the basis of the first amino acid in the mature
polypeptide being Ser, Met, or Ile, LMW-s-type subunits being
Wheat proteins will form the basis of the discussion here, as the most abundant. The sequences of LMW-s and LMW-m
the occurrence and relevance of thiols in small molecules (the consist of a short N-terminal region including the first cysteine,
so-called low-MW thiols that include free and protein-bound a glutamine-rich repetitive domain, and a C-terminal region
glutathione and free cysteine itself) and in the water-soluble containing seven cysteines. Mature LMW-m and LMW-s con-
albumin/globulin components of flours will be discussed tain eight cysteines, of which the first and seventh are involved
elsewhere. It has to be noted that some of the processes used in intermolecular disulfide bonds, whereas the others partici-
to convert some cereals into foods (including fermentation, pate in intramolecular bonds. LMW-i contain the typical
as in the case of sorghum) result in a significant decrease of eight cysteine residues, all in the C-terminal domain and all
the total content of reactive cysteines that appear to be pref- involved in interchain disulfide bonds. The ability of LMW to
erentially metabolized by the involved microorganisms. form a variable number of intermolecular disulfide bonds
These effects on protein thiols are not as evident in other defines their role as chain extenders or chain terminators.
pretreatments of cereal and noncereal grains (e.g., germina- B-type LMW-GS allow the extension of glutenin polymers
tion and/or malting). (chain extenders). LMW-GS of the C type and D type behave
As summarized in Figure 1, wheat gliadins are monomeric as chain terminators, in what they block further extension
proteins with a relevant role in the development of the gluten of the disulfide network, as they lack the additional free
intermolecular network through the formation of hydrogen cysteine residues required for formation of further interchain
bonds, of hydrophobic bonds between nonpolar amino acid linkages.
156 PROTEINS | Thiolomics of the Gluten Protein Network of Wheat Dough
C1 C5
C1 C7
Gliadins
HMW
N-TER REP C-TER x-type
1 2 3
C C C C4
SH SH
LMW C 1
C7
SH SH
Glutenins
Figure 1 Location of cysteine residues and of disulfide bonds in wheat gliadins and glutelins.
Thiol Accessibility as a Protein Structural Probe colored thionitrobenzoate anion upon reacting with an SH
group. This method can be applied in principle regardless
Accessibility of thiols to common colorimetric or fluorescent of the solubility of the proteins being investigated, but it is not
reagents has been studied as a function of added denaturing suitable for permanent (i.e., covalent) labeling of thiols, in
agents (urea, thiourea, guanidinium ions, and detergents) and view of the lability of the resulting mixed disulfide.
of reaction time to assess overall and local rigidity of proteins Other reagents can form much more stable bonds with
in either the starting material or the finished products. This protein thiols, generating species that may resist the chemically
approach could to some extent be combined with solubility severe conditions used for resolving individual components in
and compositional data, in an attempt to gather a unifying
vision of individual molecular features that contribute to the O OH
O
properties of these systems.
The approach presented here relies on the observation that N+
O O
thiols buried within the structure of a protein (or a protein S
aggregate) often become available to suitable reagents only HO S
O
upon protein denaturation by physical or chemical agents. +
N
Accessibility of the target thiol(s) is thus determined by steric
O
factors, in particular when bulky reagents are used. From a
methodological standpoint, readily accessible thiols may be O
NO2 NO2 NO2
SH + S OH S +
assessed in the absence of suitable protein denaturants, R HO S
R S
OH
HS
OH
O2N O O O
whereas the addition of denaturing agents should permit the
detection of all free thiols in the investigated proteins. Figure 2 Structure of 5,50 -dithiobis-(2-nitrobenzoic acid) (DTNB)
Simple spectrophotometric assays rely on using 5,50 -dithio- (Ellmans reagent) and its reaction with thiols, leading to the formation of
bis-(2-nitrobenzoic acid) (DTNB, see Figure 2) that releases a the yellow-colored (A412 13 700) thionitrobenzoate anion.
PROTEINS | Thiolomics of the Gluten Protein Network of Wheat Dough 157
protein mixtures. There are three fundamental types of chemical additional thiols buried in the hydrophobic regions of proteins
reactions that can be used for this purpose (see Figure 2), or at hydrophobic interfaces among proteins should become
involving (1) mercuric derivatives, (2) N-ethylmaleimide available to the labeling agent if denaturants were added prior
derivatives, (3) iodinated compounds, such as monoiodobimane to IAF. As stated earlier, measurements based on thiol accessi-
or iodoacetamide derivatives. Selected examples of compounds bility may be carried out regardless of whether the involved
in each of these classes of reactivity are presented in Figure 3. proteins are soluble or insoluble (as is the case for most of
wheat proteins) in the conditions used for labeling.
As indicated by electrophoretic separation of IAF-labeled
wheat proteins in the presence of sodium dodecyl sulfate
Mapping the Pattern of Thiol Exposure in Individual (SDS-PAGE; see an example in Figure 5), treatment with IAF
Cereal Proteins (Thiolomics) in the presence of urea gave the expected increase in the inten-
sity and number of fluorescent bands, as a consequence of
The possibility of covalently labeling any exposed thiol prior to increased thiol exposure in the presence of the chaotrope.
separation of individual polypeptides has been exploited to The SDS-PAGE tracings in Figure 5 also allow to appreciate
assess the pattern of thiol exposure also in the insoluble com- some specificity of the labeling procedure in wheat flour and in
ponents of wheat flours and semolina. By introducing dena- semolina and a different sensitivity of otherwise similar pro-
turing and disulfide-reducing steps at appropriate points in the teins in each of the samples to the presence of urea during the
thiol-labeling procedure, it is possible to use 2-D electropho- labeling step. In the case of soft wheat, labeling in the presence
resis techniques to obtain a visually informative thiolomics of urea apparently modifies only the intensity of the fluores-
map of the conditional accessibility of thiols in individual cence associated with all protein bands, and no additional
proteins in the materials under investigation. These maps pro- bands were present in the sample labeled in the presence of
vide information as for the type of proteins where thiols are urea. In the case of durum wheat semolina, the fluorescence
available and on the relative abundance of proteins and reac- intensity of specific protein bands was sensibly increased when
tive thiols on them (Figure 4). labeling was carried out in the presence of urea, and the effect
In the original procedure, thiols in wheat flour or semolina of the denaturant on the accessibility of thiols was most evi-
proteins were covalently labeled by 5-iodoacetamid- dent in the largest polypeptides. This may be taken as an
ofluorescein (IAF) in the absence or in the presence of 4 M indication of the presence of multiple thiols on the involved
urea, allowing to evaluate thiols accessibility before and after proteins and of changes in their accessibility to the reagent in
protein unfolding and as related to the dissociation of non- the presence/absence of urea.
covalently linked protein complexes. Proteins labeled under Given the possible ambiguities associated with identifica-
either condition were then separated by SDS-PAGE and/or by tion of specific components from SDS-PAGE gels, 2-D electro-
2-D electrophoresis. Visual inspection and quantitative com- phoretic maps were also obtained, similar to those used in
parison of the maps for proteins and for accessible thiols in classic proteomics studies. In this technique, proteins are
different and variously treated starting materials were then made soluble by unfolding in urea and reduction with DTT
carried out to identify specific proteins or protein classes and are separated first according to their isoelectric points (pI)
whose thiol functions may specifically contribute to specific in an immobilized pH gradient. The second run is performed
traits of the material. in an orthogonal direction with respect to the first separation
Only readily accessible protein thiols should be labeled in step and separates them by molecular mass. A comparison of
flour samples treated with IAF in the absence of urea, whereas protein-stained 2-DE maps obtained in the presence and
absence of treatment with the fluorescent thiol probe confirms
O that covalent binding of the probe may introduce some alter-
Iodoacetamide
I ations in the pI of labeled proteins, although the impact of this
NH2
substitution on the acidic proteins that are predominant in
wheat does not impair analysis of the resulting maps.
O O Typical 2-D maps of the IAF-labeled polypeptides in various
H3C
N
CH3 Monobromobimane wheat samples are shown in Figure 6. These maps can be easily
N analyzed by standard image analysis techniques that may be
H3C CH2Br used to convert positive and negative images (the original
fluorescence maps had bright spots against a black back-
O ground) and to transform halftone images into very informa-
tive and/or easily read landscape plots (see the example
N-Ethylmaleimide presented in Figure 7). This type of graphical rendition
N
makes it easier to appreciate (both visually and on a quantita-
O tive basis) compositional and/or structural differences among
samples as reflected in their thiol accessibility maps.
O CH3
Hg It has to be noted that the very same 2-D gel can subse-
Phenylmercury acetate
O quently be stained for proteins by any of the procedures used
in standard proteomics studies (see Figure 8). Image analysis
Figure 3 Examples of common thiol reagents suitable for covalent of the protein-stained gel can provide additional information
labeling of proteins. on not only the occurrence and abundance of specific protein
158 PROTEINS | Thiolomics of the Gluten Protein Network of Wheat Dough
O OH
O
O
I
N O
H
OH
Iodoacetamide
fluorescein (IAF)
Cysteine
thiols
Interprotein
disulfide bonds
Hydrophobic
contact regions
Figure 4 A graphical rendition of the events related to covalent binding of the fluorescent probe IAF to wheat protein thiols in the absence/presence
of denaturants.
Mamone G, De Caro S, Di Luccia A, Addeo F, and Ferranti P (2009) Proteomic-based wheat processing properties. Advances in Food and Nutrition Research
analytical approach for the characterization of glutenin subunits in durum wheat. 45: 219302.
Journal of Mass Spectrometry 44: 17091723. Singh H and MacRitchie F (2001) Application of polymer science to properties of
Morel MH, Redl A, and Guilbert S (2002) Mechanism of heat and shear mediated gluten. Journal of Cereal Science 33: 231243.
aggregation of wheat gluten upon mixing. Biomacromolecules 3: 488497. Skylas DJ, Van Dyk D, and Wrigley CW (2005) Proteomics of wheat grain. Journal of
Shewry PR, Tatham AS, Forde J, Kreis M, and Niflin BJ (1986) The classification and Cereal Science 41: 165179.
nomenclature of wheat gluten protein: A reassessment. Journal of Cereal Science Veraverbeke WS and Delcour JA (2002) Wheat protein composition and properties of
4: 97106. wheat glutenin in relation to bread-making functionality. Critical Reviews in Food
Shewry PR, Halford NG, Tatham AS, Popineau Y, Lafiandra D, and Belton PS (2003) The Science and Nutrition 42: 179208.
high molecular weight subunits of wheat glutenin and their role in determining Wieser H (2007) Chemistry of gluten proteins. Food Microbiology 24: 115119.
Carbohydrates
Contents
Carbohydrate Metabolism
Starch: Chemistry
Starch: Synthesis
Starch: Starch Architecture and Structure
Starch: Analysis of Quality
Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides
Grains other than Cereals: Non-starch Polysaccharides
Resistant Starch and Health
Health Effects of b-Glucans Found in Cereals
Glycemic Index, Glycemic Load, and Their Health Benefits
Carbohydrate Metabolism
RN Chibbar, S Jaiswal, M Gangola, and M Baga, University of Saskatchewan, Saskatoon, SK, Canada
2016 Elsevier Ltd. All rights reserved.
Topic Highlights two-thirds to three-quarters of dry matter and are the major
components of storage organs. Cellulose, the most abundant
Carbohydrates are one of the major components of food plant carbohydrate, provides structural integrity to plants and is
grains. used for fiber and fuel by humans. In seeds, carbohydrates
Carbohydrates are mainly categorized on the basis of their provide energy and substrates required for the germination
degree of polymerization, structure, and function. and initial growth of seedlings. Carbohydrate in the form of
Biosynthesis of different sugars occurs in different plant cell starch is the major reserve polysaccharide present in grains.
organelles. Carbohydrates like starch and cellulose form the basis of several
Carbohydrates support seed germination and early seedling important industries in the food, feed, and fuel sectors. They
development. also provide renewable and environmentally friendly raw mate-
Carbohydrates, besides being an energy source, also pro- rials for industrial applications such as biodegradable plastics,
vide structural integrity to plant cells. adhesives, and ethanol-based fuels. With the development
Carbohydrate biosynthesis can be targeted to improve food of genetic modification technologies, the biosynthesis of car-
grain quality for human consumption. bohydrates can be altered in planta to generate novel products
Carbohydrates are also involved in abiotic stress tolerance. for various food and industrial applications. This article will
describe the structure, occurrence, and distribution of some of
the common storage and structural carbohydrates present in
Learning Objectives the grains of major crops. The biochemical reactions occurring
during the biosynthesis of carbohydrates and their utilization
To achieve understanding of the structural diversity of grain during seed germination will be discussed.
carbohydrates, their biosynthesis, storage in grains, and
mobilization during seed germination.
To appreciate the influence of structural diversity of grains Carbohydrates
on their end use for human food, animal feed, and indus-
trial applications. Definition and Classification
The term carbohydrate, or hydrates of carbon, is derived from
its basic elemental formula [Cx(H2O)y], in which carbon is
Introduction joined to hydrogen and oxygen present in the same ratio as in
water. Chemically, carbohydrates are polyhydroxy aldehydes or
Carbohydrates are one of the most important nutrients for ketones, their simple derivatives, or their polymers. Carbohy-
both plants and animals. In plants, carbohydrates constitute drates in grains are classified based on their chemical structures
or their digestibility when consumed by humans as food or by a disaccharide. Upon hydrolysis of disaccharides, the glycosidic
livestock as feed. Based on the polymeric nature of carbohydrates, bond is split into yield the component monosaccharides. Sucrose
they are classified as monosaccharides, disaccharides, oligosac- and maltose are the most commonly occurring disaccharides
charides, and polysaccharides (Figure 1). Monosaccharides are in the majority of grains. Oligosaccharides include the sugars
the simplest of all sugars, contain at least three carbon atoms, and that contain 320 sugar molecules joined by glycosidic bonds.
are the building blocks of all carbohydrates. The most common The most common oligosaccharides are soluble a-galactosides,
monosaccharides in grains are the hexoses (fructose, glucose, and characterized by galactose moieties joined by a(1,6) linkages.
galactose) and the pentoses (arabinose and xylose). Two sugar In legumes, raffinose, stachyose, and verbascose are the most
moieties joined by a glycosidic bond (an oxygen bridge) form common oligosaccharides and constitute 618% of the legume
Monosaccharides
H H H
O H O O O
H H H
O O
O O O O O O O O
H H H H
O
H O
H H O O
H H
O O O O O O
O O H H
O O
H H H
H
-D-Glucose -D-Fructose -D-Xylose -D-Galactose -L-Arabinose
Disaccharides
H H
O H H
O O O
H H H
H H O O O O
O H
O O O
H H
O
O H O H
O O O O O
O O O O
O O O O O
H H H H H H
O H O O
O O O O O O O
H H H H H H H
O O O O O
H H H H H
Oligosaccharides
HO
HO H
H O O HO
OH OH O
H
H
H H O OH H H H H
O O
O H O O O O O
O O H HO O H
O O O OH
H O H O OH HO
O O O O O H O H O H
H O O H O
O O
HO O
O HO O O
OH
H O O H OH
O O O O HO O O
HO O
OH H O O
O O O O
H O O H O O
O
H O H H
H H H O O HO
O
H
Raffinose Manninotriose Stachyose H
HO
H Verbascose
HO H
O
H H H H
H O O O O O O
H H H H
O O O
O
O O O H O
O O O O O H
O H O
H O HO H O O
H
O O O H
O H H H
O O O HO O O
H OH O
H O H O
O O
H O O H
O O
O OH H
H O
O O
H O O
O O O H O
H O
O H OH H H
O
O O
O O
O H H
Lychnose H
O H Stellariose Mediose
H
Figure 1
CARBOHYDRATES | Carbohydrate Metabolism 163
Polysaccharides
H
H
O
Main chain O OH
CH2OH CH2
CH2OH CH2
O O H O OH H O
O
OH O OH O O O O O
O O
OH OH H O OH
O OH CH2 CH2
-D-Glc(14) linkage n
H O O
H O-H H
HO O H
Amylose O O
O
O O
O
HO O H
CH2 CH2
O O
CH2OH CH2OH H H
O O
Branch
O OH O OH Cellulose units linked with inter and intra chain H bonds
OH OH O -D-Glc(16) branch point
CH2 CH2OH
CH2OH CH2OH
O O O O O OH
O O O
O
OH OH O O
O OH OH O O
O
OH OH OH
OH OH OH OH
OH OH OH
OH
Main chain
-D-Glc(14) linkage OH OH -1, 3-Xylan
n
Amylopectin
Fructans
CH2OH
O
OH
CH2OH
O
CH2OH CH2OH O OH
CH2OH CH2OH O CH2
O O
O OH
OH OH OH OH
CH2O CH2OH
O CH2O OH CH2OH
OH OH O OH (n)
OH CH2OH
OH (n) CH2 O
O OH
OH
CH2OH
O
OH OH
OH
Fructan-levan type
Fructan-inulin type
Sugar alcohols
O H
H
O O
H
H H H H
H O O O O
O O O
O O
O O O O H H H
H H H H O O
O O O O O
O O O H H H H
H H
O
H O
H
Figure 1 (Continued) Classification and chemical structures of representative carbohydrates found in grains of crops. Structure database source:
http://www.ncbi.nlm.nih.gov/pccompound. CID number reference: a-D-glucose, 79025; b-D-fructose, 439709; a-D-xylose, 6027; a-D-galactose, 439357;
a-L-arabinose, 641630; sucrose, 5988; a-D-maltose, 439341; galactinol, 11727586; lactose, 6134; raffinose, 439242; manninotriose, 440660;
stachyose, 439531; verbascose, 441434; lychnose, 25079986; stellariose, 46926270; mediose, 46926113; ciceritol, 44237401; sorbitol, 5780; and
mannitol, 6251.
seed dry weight. The presence of these flatulence-causing Some of the most common polysaccharides present in grains
a-galactosides is one of the major reasons restricting consumption are starch, cellulose, and xylans such as pentosans, b-glucans,
of legume-based food products. The absence of a-galactosidase and arabinoxylans. In relation to human and animal nutrition,
in the intestinal tract of humans causes fermentation of carbohydrates are often classified as available and unavailable.
a-galactosides by intestinal bacteria that contribute to the flatu- The available carbohydrates represented by sugars and starches
lence effect. Polysaccharides are polymers with more than 20 are broken down and absorbed in the human and animal diges-
monosaccharides joined together by glycosidic linkages. These tive tracts to provide energy for the body. Structural cell wall
are complex molecules because of the diversity in monosaccha- carbohydrates such as cellulose, hemicellulose, and nonstarch
ride units and type of linkages present in the polysaccharides. complex carbohydrates, for example, xylans, are categorized as
164 CARBOHYDRATES | Carbohydrate Metabolism
Table 1 Chemical composition of seeds of some important cereal grains (Table 3). Chickpeas especially have a high concen-
grain crops tration of soluble sugars, of which sucrose is the major compo-
nent. Lentils and peas have higher concentrations of stachyose
Composition (percentage dry
and verbascose as compared with other legumes. Starch is the
weight)
major storage carbohydrate in cereal grains, where it forms two-
Crop Carbohydrate Protein Lipid thirds to three-quarters of the dry weight. However, in oats, the
starch concentration is slightly less, ranging from 50% to about
Cereal crops 60%. Legume seeds also contain 50% starch, except lupins
Wheat (Triticum aestivum) 82 14 2 and soybeans in which only 0.51.4% of the dry weight is starch.
Barley (Hordeum vulgare) 80 9 1 The structural carbohydrates or functionally unavailable carbo-
Corn (Zea mays) 84 10 5
hydrates (dietary fibers) range in distribution from slightly above
Rice (Oryza sativa) 88 8 2
Oat (Avena sativa) 67 28 1
5% in rice to almost one-third of the grain dry weight in barley.
Sorghum (Sorghum vulgare) 82 12 4 In grain legumes, chickpeas have a higher concentration of total
Rye (Secale cereale) 82 14 2 nonstarch polysaccharides, whereas the concentration of total
Teff (Eragrostis tef) 83 11 2.5 dietary fiber corresponds to that of other legumes (Table 3). The
Pearl millet (Pennisetum glaucum) 81 12.5 5 major nonstarch complex carbohydrate in grain legumes is cel-
Foxtail millet (Setaria italica) 82 9.9 2.5 lulose, which represents about one-eighth of the grain dry weight
Finger millet (Eleusine coracana) 88 7 1.7 in most of the grain legumes (Table 3).
Kodo millet (Paspalum 84 11.5 1.3
scrobiculatum)
Proso millet (Panicum miliaceum) 82 10.6 4 Metabolism
Canary (Phalaris minor) 61 20.3 11
Triticale (xTriticosecale) 72 13 2.1
Autotrophic plants have the ability to convert simple nutrients
Cereal like non-Poaceae family grains such as carbon dioxide, water, and inorganic ions into interme-
Buck wheat (Fagopyrum 83 11 2 diates leading to the biosynthesis of complex molecules, such as
esculentum) nucleic acids, proteins, lipids, polysaccharides, and secondary
Quinoa (Chenopodium quinoa) 78 14.5 5.2 metabolites needed for growth and development (Figure 2).
Pulse crops Through the process of photosynthesis, plant cells capture light
Lentil (Lens culinaris) 65 27 1 energy to fix reduced carbon dioxide and water into a simple
Pea (Pisum sativum) 68 27 2 carbohydrate backbone, CH2O. The reaction takes place in chlo-
Chickpea (Cicer arietinum) 65 23 5 roplasts, wherein CO2 entering the Calvin cycle is incorporated
Kidney bean (Phaseolus vulgaris) 70 24 2
into 3-phosphoglycerate (3-PGA) molecules, which are subse-
Mung bean (Vigna radiata) 69 26 1
Pigeon pea (Cajanus cajan) 65 21 1
quently converted to triose phosphates. In addition, the
Soybean (Glycine max) 32 38 20 absorbed light energy is used for extracting reducing equivalents
Cowpea (Vigna unguiculata) 60 24 1 from water and ATP production required for processes involved
Lupin (Lupinus albus) 37 38 20 in plant growth and development. The reactions associated with
Oilseed crops light absorption are diurnal, which results in large variations in
Rape (Brassica napus) 25 23 48 nutrients available to plants during light and dark periods. How-
Sunflower (Helianthus annuus) 48 20 29 ever, plants have developed considerable flexibility in their
Flax (Linum usitatissimum) 32 26 38 metabolism to cope with fluctuations in nutrient concentration.
Peanut (Arachis hypogaea) 25 27 45 Plant cells can be envisioned to have a series of metabolic inter-
Sesame (Sesamum indicum) 19 20 54
mediate pools, which are interconnected to each other through
Hemp (Cannabis sativa) 27 29 41
enzyme reactions or transport mechanisms that are often revers-
ible (Figure 2). According to the physiological need(s) of a plant,
unavailable carbohydrates because they are not assimilated by the metabolites can be added to or withdrawn from these pools. Two
digestive tract. The unavailable carbohydrates are also often metabolite pools of the reductive pentose phosphate pathway
referred to as dietary fibers. and its intermediates are (1) the triose phosphate (3-PGA and
dihydroxyacetone phosphate (DHAP)) and (2) the hexose phos-
phate (glucose 1-P, glucose 6-P, fructose 1-P, and ADP-glucose)
Distribution in Important Grains
pools (Figure 2).
In general, carbohydrates dominate as the major constituent of The triose phosphates are either transported by triose phos-
the dry weight of seeds (Table 1). However, in oilseed grains, phate transporters to the cytosol or converted to other phos-
such as Brassica spp., lipids and proteins exceed carbohydrates as phorylated compounds, such as fructose 6-phosphate, in the
the major storage components by comprising three-quarters of plastid. The plastidial fructose 6-phosphate is used for both
grain dry weight. In cereal grains, the total concentration of regenerating ribulose-1,5-bisphosphate and producing glucose
mono-, di-, and oligosaccharides varies from < 1% to 3% 1-phosphate. Glucose 1-phosphate is converted to ADP-
(Table 2). Corn kernels contain the highest percentage of glucose glucose by ADP-glucose pyrophosphorylase (AGPase), the
and fructose, whereas in barley, sucrose is the major free sugar. immediate precursor for starch biosynthesis. Starch biosynthe-
Wheat grains have the highest concentration of raffinose among sis occurs inside plastids, but its functionality depends on the
the cereal grains (Table 2). Legume crops in general have a higher plastid type (chloroplast/amyloplast) and source plant tissue.
concentration of di- and oligosaccharides as compared with Starch synthesized in the chloroplast during the day is known as
CARBOHYDRATES | Carbohydrate Metabolism 165
Total
dietary
Crop Total Glucose Fructose Sucrose Raffinose Total Amylose fiber Pentosan b-Glucan Cellulose
Wheat 1.41.3 0.020.03 0.020.04 0.570.80 0.540.70 6372 2328 14.6 6.6 1.4 2.0
Barley 2.03.0 0.10.2 0.1 1.92.2 0.59 5759 2226 1923 5.9 37 4.3
Rice 11.1 0.120.13 0.110.13 0.60.66 0.10.2 66 1633 3.9 1.2 0.11 1.72.0
(brown)
Teff 1.8 0.73 0.47 0.62 NA 73 2026 8 NA NA NA
Sorghum 1.32.5 0.21.1 0.61.6 0.150.39 63 24 6 NA NA NA NA
Pearl 2.22.8 0.080.16 1.31.8 0.650.84 5370 2032 13.3 NA NA NA NA
millet
Finger 0.590.69 0.140.19 0.200.24 0.060.08 55.763.4 14.438.6 18 NA NA NA NA
millet
Foxtail 0.46 0.10 0.15 0.04 5269 11.232 NA NA NA NA NA
millet
Proso NA NA 0.66 0.08 6168 17.232.6 NA NA NA NA NA
millet
Corn 1.03.0 0.20.5 0.10.4 0.91.9 0.10.3 6478 24 13.4 5.86.6 0.1 2.1
Oats 1.4 0.05 0.09 0.64 0.19 4361 1627 9.6 7.7 3.96.8 6.012.9
Rye 3.2 0.08 0.10 1.9 0.4 69 2431 14.6 8.5 1.92.9 1.5
Canary 1.75 0.06 0.07 0.83 NA 61 17.2 6.0 NA 1.12.3 NA
Triticale NA 0.23 3.1 0.21.9 NA 5970 23 1316 NA 0.51.0 1.72.5
Quinoa 2.9 0.02 0.02 NA NA 63 1112 9.6 NA NA NA
transitory starch, as it is degraded to maltose, glucose, hexose each other, and their exchange is often restricted. Therefore, the
phosphates, and triose phosphates at night. This periodical regulation of carbohydrate metabolism in plants is more com-
starch degradation involves two major steps: initiation, which plex than in other organisms, where such subcellular compart-
involves the activation of starch, and starch digestion. Transient mentalization of hexose phosphates is not present.
starch degradation is mainly attributed to b-amylases, and the
product of its action is transported back to the cytosol, where it
Sucrose
undergoes further modifications.
Some of the starch degradation products are converted to Sucrose is the most abundant disaccharide and the major
sucrose, the major transport sugar. Sucrose through symplastic product of photosynthesis. It is a nonreducing sugar with
or apoplastic transport enters the phloem, through which it is limited chemical reactivity and thus is utilized as a transport
mobilized to the storage tissues, where symplastic or apoplastic and storage molecule in most plants. Sucrose synthesis is pre-
uptake into sink cells takes place. The transported sucrose is dominant in leaves, but the ability to synthesize sucrose is
further converted to substrates that give rise to the diverse carbo- fairly widespread among plant cells. Sucrose is synthesized in
hydrates present in grains of different crops as described earlier. the cytosol by the consecutive activities of two enzymes:
The hexose phosphate pool also plays an important role in starch sucrose-phosphate synthase (SPS) and sucrose phosphatase
and sucrose utilization by plants. In the glycolytic pathway, (SPP). SPS catalyzes the reversible transfer of glucosyl units
which acts in reverse of the triose phosphate synthetic pathway from UDP-glucose to fructose 6-phosphate, synthesizing
during photosynthesis, starch and sucrose are oxidized to pro- sucrose 6-phosphate that is dephosphorylated by SPP to yield
vide energy and hexose phosphates that on phosphorylation sucrose in an irreversible reaction. SPS is the key regulating step
contribute to the hexose phosphate pool. In addition, the glyco- of sucrose biosynthesis. In plants, SPS can be activated alloste-
lytic reactions provide several intermediates that contribute to rically by glucose 6-phosphate and inhibited by inorganic
the synthesis of proteins, lipids, and organic acids (Figure 2). phosphate (Pi). In addition, SPS also contains several regula-
These compounds are important grain components and are tory phosphorylation sites. Sucrose synthase (SucS), present in
described in other articles. both soluble and membrane-bound forms, is responsible for
The hexose phosphate pool is an important converging catalyzing both the synthesis and degradation of sucrose. How-
point for both carbohydrate synthesis and degradation and ever, sucrose degradation is mainly attributed to SucS in sink
thus is considered to play an important role in carbohydrate tissues participating in sucrose degradation.
metabolism in plants. The main components, glucose 1-phos- The translocation of synthesized sucrose from source
phate, glucose 6-phosphate, and fructose 6-phosphate, are to sink tissues is essential for proper plant growth and devel-
readily interconvertible when present in the same system. How- opment. Sucrose can be translocated either symplastically
ever, in plants, the hexose phosphates are compartmentalized (through plasmodesmata) or apoplastically (through apoplast-
to the plastids and cytosol. In photosynthetic tissues, the plas- s with the help of sucrose transporters). In sink tissues, sucrose
tidial and cytosolic hexose phosphate pools are independent of can be stored inside cell vacuoles through transporters located
166
CARBOHYDRATES | Carbohydrate Metabolism
Table 3 Carbohydrate composition of some grain legumes
Common 2.09.6 1.63.9 0.22.5 0.23.9 0.11.8 0.48.0 5151.9 22.136.0 6.420.4 11.227.5 0.55.6 3.213.1
bean
Peas 3.513.8 0.95.4 0.42.3 0.34.2 04.3 2.39.6 24.757.4 23.533.1 NA 16.121.6 0.912.4 0.913.3
Lentils 3.39.5 1.13.0 0.10.8 1.14.0 06.4 1.87.5 40.157.4 20.745.5 6.914.7 11.021.4 1.215.7 3.514.8
Chickpeas 4.614.2 2.86.9 00.3 0.42.0 trace0.4 2.07.6 43.059.0 31.845.8 5.535.4 8.224.0 0.616.0 1.113.7
Faba beans 2.28.5 0.13.8 0.11.5 0.21.6 1.12.4 1.04.5 39.247.2 22.035.0 17.5 17.123.8 1.68.9 8.314.3
Mung bean 3.97.2 0.32.0 0.32.6 1.22.8 1.73.8 NA 37.053.6 13.835.0 NA 15.7 0.39.1 2.54.6
Black gram 3.07.1 0.71.5 0.01.3 0.93.0 3.43.5 NA 32.247.9 43.9 NA 22.4 10.7 5.0
Red gram 3.510.2 2.7 1.01.1 2.73.0 4.04.1 NA 40.448.2 38.6 NA 18.7 10.1 7.3
Black gram: Vigna mungo; red gram: Cajanus cajan; faba bean: Vicia faba.
CARBOHYDRATES | Carbohydrate Metabolism 167
Ribulose-1,5-
Purines
bisphosphate
Ribulose5-bisphosphate Triose-phosphate pool Tryglycerides
Nucleic acids 3-PGA, GAP, DHAP
Amino acids
Pentose phosphate pathway
Tryglycerides
Delta-aminolevulnic acid Oxidative
phosphorylation Amino acids
Chlorophyll, NAD+
Cytochrome, FAD
phytochrome ATP
Figure 2 A schematic overview of the carbohydrate metabolism and its interrelationship with other biochemical pathways to show the
production of major plant cell constituents.
at tonoplasts or can be hydrolyzed by invertase providing Caryophyllaceae and Lamiaceae plants (Figure 1). The second
glucose and fructose to the cell. The translocation of sucrose group of a-galactosides includes galactosyl cyclitols. Ciceritol is
from source to sink tissues is very active during grain develop- the most common of the galactosyl cyclitols and was first
ment when storage carbohydrates are synthesized in the grains. reported from chickpea (Cicer arietinum), followed by lentil
A boost in sucrose concentration in seeds is observed during (Lens culinaris).
seed germination when stored carbohydrates are broken down RFO biosynthesis occurs in the cytosol and begins with the
to provide energy to the germinating embryo. formation of galactinol from myo-inositol and UDP-galactose in
a reaction catalyzed by galactinol synthase. In the next step,
raffinose synthase supports galactinol to donate its galactosyl
Alpha-Galactosides
unit to sucrose yielding raffinose. Raffinose and stachyose,
Alpha-galactosides are low-molecular-weight, nonreducing oli- together with galactinol, synthesize stachyose and verbascose
gosaccharides and rank next to sucrose among abundant soluble in two separate reactions catalyzed by stachyose synthase and
sugars. These sugars constitute about 618% of the dry weight of verbascose synthase, respectively. In a galactinol-independent
mature legume seeds. Alpha-galactosides are sucrosylgalacto- pathway, already present RFOs react with each other to
sides and are characterized by the presence of an a(1! 6) link- synthesize higher members of RFOs. Galactan/galactan
age between the galactosyl residue and the C-6 of the glucose galactosyltransferase is the sole enzyme catalyzing the reaction.
moiety of sucrose. Alpha-galactosides can further be classified RFOs participate in plant physiological processes like seed
into two groups. Raffinose family oligosaccharides (RFOs) con- desiccation tolerance, seed germination, photosynthate trans-
stitute the first group, and the first member of this group, raffi- location (mostly in Cucurbitaceae, Verbenaceae, Lamiaceae,
nose (degree of polymerization (DP) 3), is the main RFO in Oleaceae, and Scrophulariaceae families), and stress tolerance.
most monocotyledon seeds, while its higher homologues, sta- However, in the human diet, food rich in RFOs causes stomach
chyose (DP 4), verbascose (DP 5), and ajugose (DP 6), discomfort, flatulence, and diarrhea in humans and mono-
accumulate predominantly in seeds of dicotyledons. Higher gastric animals, as they lack a-galactosidase, a hydrolyzing
members like ajugose are generally found in trace quantities in enzyme needed for RFO breakdown. RFOs when consumed
seeds. Some alternative RFOs such as lychnose, stellariose, in lower concentrations are considered as prebiotic and sup-
mediose, and manninotriose have been recently reported from port the growth of beneficial intestinal microflora.
168 CARBOHYDRATES | Carbohydrate Metabolism
formation of glucose 1-phosphate. In storage tissues, sucrose is or other B-chains; and the central C-chain. The backbone of
converted into glucose 6-phosphate in the cytosolic region starch, that is, the C-chain from which B-chains are connected
from where it is imported into the amyloplasts through a hex- within a granule, is oriented toward the center or hilum of the
ose phosphate/Pi antiporter. The glucose 6-phosphate is con- granule. Amylopectin glucan chain length distribution influ-
verted to glucose 1-phosphate, which acts as a substrate for ences functional properties like gelatinization, retrogradation,
AGPase, to synthesize ADP-glucose as in chloroplasts and act and pasting properties of starch.
as the primary building block required for chain elongation to
initiate starch synthesis. AGPase is an allosterically regulated
enzyme, whose activity is generally stimulated by 3-PGA and
Cellulose
inhibited by Pi. This allosteric regulation is further influenced
by plant species, plastid type, and subcellular location. Cellulose is the major component of cell walls and accounts for
Studies of starch mutants have so far produced a general 2830% of dry matter in forage grasses and 4245% of wood.
consensus that AGPase, soluble starch synthases (SSs), starch- This extracellular polysaccharide-based outer matrix of the
branching enzymes (SBEs), starch-debranching enzymes plant cell is complex and dynamic and has enormous strength.
(DBEs; pullulanase and isoamylase), and possibly dispropor- It is the most abundant renewable biomass synthesized on
tionating enzyme (D-enzyme) catalyze the final steps leading to Earth and has since the early days of human civilization been
amylopectin synthesis. The five major classes of SS include used for fuel, timber, fiber, and forage. Today, cellulose has
GBSSI, SSI, SSII, SSIII, and SSIV, of which all but SSIV have additional uses as a raw material for the production of paper,
been shown to have distinct roles in determining starch com- pulp, and many chemicals. Cellulose has a paracrystalline struc-
position and structure. Granule-bound starch synthase 1 ture formed by b-(1 ! 4)-D-glucan chains, formed individually
(GBSSI) is the only enzyme committed to amylose synthesis and crystallized to form cellulose microfibrils (an average thick-
with limited contribution toward amylopectin as indicated in ness of 36 chains through inter- and intramolecular hydrogen
some recent reports. Since several of the enzymes exist in bonds and van der Waals forces). The chains contain a few
different isoforms, some of which vary in their subcellular thousand glucose units giving an individual length of 23 mm,
distribution, enzyme specificity, temporal activity, and interac- but the chains start and end at different points of the microfi-
tion with other enzymes; the starch biosynthesis pathway is bril, thus extending the length of the microfibril to hundreds of
complex. Hilum, also known as starch granule core, indicates micrometers. The length, angle, and crystallinity of these micro-
the presence of a priming mechanism for starch granule for- fibrils influence plant cell wall strength.
mation. The priming mechanism for starch glucan polymeri- In higher plants, cellulose synthesis occurs in a plasma
zation in plants is unclear, but it likely involves short-chain membrane-bound complex known as rosette TC. The structure
maltodextrins that are extended and branched by the action of contains six subunits, each composed of six glucan (cellulose)
GBSSI, SBEI, and SSI to form a molecule with an amylopectin- synthase molecules that are symmetrically arranged into a
like backbone. Glycogenin-like self-glycosylating protein has rosette. The subunits of the complex simultaneously synthesize
also been proposed. The primary glucan polymer is further 36 1,4-b-D-glucan chains that are bundled into a microfibril
polymerized through cycles of chain extensions and branching emerging from the rosette TC. The basic biochemical under-
catalyzed mainly by SSI and SBEII. DBEs have been proposed standing of cellulose synthase (CesA) has mainly been obtained
to play a role in starch biosynthesis by trimming the ends of the from studies of the bacterium, Acetobacter xylinum. In plants, the
water-soluble preamylopectin molecules to produce amylopec- number of CesA genes varies depending upon the species.
tin that can be crystallized and packaged into granules. Two CesAs categorized under the glycosyltransferase-2 (GT-2) super-
distinct DBE families (isoamylase and R-enzyme/pullulanase/ family with eight-transmembrane topology consist of at least
dextrinase) with substrate specificity have been identified in two subunits, one with a catalytic role and the other with a
plants. DBEs are also suggested to degrade water-soluble glu- regulatory role. The catalytic subunit of CesA is a progressive
cans synthesized by starch synthases and starch-branching glycosyltransferase that binds the substrate, UDP-glucose,
enzymes to prevent accumulation of phytoglycogen, which derived from sucrose by the action of a membrane-bound
cannot form granules. Observations made in Chlamydomonas SucS. Recent studies proposed that these subunits are placed
suggest that the D-enzyme is also involved in starch synthesis in the loop between transmembrane domains 2 and 3 with a D,
and granule formation in addition to starch degradation. D,D,QXXRW motif. The present model of cellulose synthesis
The polymerization of amylose is initiated from soluble postulates that the first two conserved aspartic acid residues
maltooligosaccharides that serve as primers for GBSSI. Starch interact with UDP, the third aspartic acid residue forms the
biosynthesis can be visualized as a wavelike formation, where catalytic base for the glucan extension, and the last QXXRW
high and low nodes result in the formation of regions of residues form a binding site for the terminal glucan parts of the
different crystallinities visible as growth rings (e.g., potato chain. At the initiation of polymerization, two UDP-glucose
starch). Many reports have indicated that the presence of starch molecules are present in the substrate-binding site, and as the
biosynthetic enzymes as protein complexes and positioning of chain elongates, glucose is added to the nonreducing end of the
SS, SBE, and DBE in these complexes influence amylopectin chain. In addition to Ces genes, plants have a number of cellu-
architecture. Amylopectin chains distributed in polymodal pat- lose synthase-like (CSL) genes that share homology with Ces
terns form clusters with architectural specificity. These chains and belong to the GT-2 superfamily. CSLs are grouped into ten
are designated as A-chains (DP 612), the outermost chains; families designated from A to J. Many of these are associated
B-chains, named B1, B2, and B3 depending on their degree of with b-glycan polymer synthesis, for example, CSLF for
polymerization (DP 1324 up to 50), which support A-chains b-(1 ! 3;1 ! 4) glucans.
170 CARBOHYDRATES | Carbohydrate Metabolism
In addition to Ces, membrane-anchored endo-1,4- Arabinoxylans and b-glucans affect the quality and utiliza-
b-glucanases may also play a role in cellulose biosynthesis, tion of cereal grains due to their presence in the endosperm cell
based on studies with the Arabidopsis KORRIGAN mutant and walls. The predominant component of starchy endosperm cell
the cellulose-producing bacterium Agrobacterium tumefaciens. walls in barley, oats, and maize is b-glucan, whereas arabinox-
ylans are the major component found in rye and wheat. These
compounds have a significant impact on the utilization of the
grains for both food and animal feed. In wheat flour, pentosans
Other Cell Wall Carbohydrates
absorb water and significantly affect the quality of baked prod-
Other important polysaccharide components of the cell wall ucts. The cell wall carbohydrates are not completely digested in
include cross-linking glycans, which are hydrogen-bonded to the stomach or small intestine of humans or other monogastric
cellulose microfibrils. These glycans coat the cellulose micro- animals but are broken down by bacteria in the large intestine.
fibrils and link two or more cellulose microfibrils to form a They tend to reduce the absorption of mono- and disaccharides
network in the primary cell walls of all plants. The two major in the gut by raising the viscosity of the gut contents, thus
classes of the cross-linking glycans are xyloglucans (XyGs) decreasing the levels of blood glucose and insulin. b-Glucans
and glucuronoarabinoxylans (GAXs), of which XyGs are pre- have been shown to significantly reduce blood cholesterol
sent in the cell walls of most dicots and almost half of all levels, when present in the diet at high concentrations. In addi-
monocot species. The XyGs are made up of linear chains of tion, they have also been shown to have antibacterial and anti-
1,4-b-D-glucan with numerous a-D-Xyl units linked to the O6 viral effects and accelerate wound healing. In animal feed, both
position of the glucan units. In some cases, the xylosyl units glucuronoarabinoxylans and b-glucans have antinutritional
are further substituted for by arabinose or galactosyl units. In effects, as these may decrease the absorption of nutrients due
addition to the two xyloglucans, some members of the Poa- to their high viscosity and indigestibility.
ceae family, which includes the majority of cereal crops, such Cell wall polysaccharides are synthesized from sucrose
as wheat and barley, contain a third cross-linking glycan, b- through the action of SucS, UDP-glucose dehydrogenase, UDP-
glucan. This is a linear polymer that consists of 3-D- glucuronate decarboxylase, xylan synthase, and arabinosyltrans-
glucopyranosyl monomers joined by both (1 ! 3) and ferase that sequentially give rise to UDP-glucose, UDP-glucuro-
(1 ! 4) linkages, normally in a 2:1 ratio. In comparison nate, UDP-xylose, 1,4-b-xylan, and arabinoxylan, respectively.
with other cereals, barley and oat grains are enriched with The action of three epimerases further converts UDP-glucose to
b-glucans. The b-1,4-linked molecules cellotriose (DP3) and UDP-galactose, UDP-glucuronate to UDP-galacturonate, and
cellotetraose (DP4) are randomly joined by b-1,3 bonds, UDP-xylose to UDP-arabinose.
which results in the formation of kinks in the molecule. The
DP3/DP4 ratio affects polymer solubility. The highest content
of soluble (1,3;1,4)-b-glucan is associated with DP3/DP4 Carbohydrate Breakdown
ratios in the 1.52.5 range. Both the ratio and concentration
of b-1,4-linked molecules in the grain cell wall, especially in Germination is initiated when the dry seed imbibes water,
oat and barley, influence their end use in food, feed, or and the process is completed with the elongation of the
industrial applications. embryonic axis (Figure 3). One of the first signs of metabolic
Husk
Pericarp
H2O Testa
Aleurone
Endosperm
-amylase GA receptor
GA
Starch
Maltose GAMyb
Aleurone cell
Glucose
-amylase
H2O GA
Scutellum Scutellum
Coleoptile
Embryo Shoot
Radicle
Coleorrhiza
Root
Figure 3 A generalized diagram of a germinating cereal grain to show the various parts of a grain and the major biochemical activities during
germination.
CARBOHYDRATES | Carbohydrate Metabolism 171
activity in a germinating seed is the activation of enzymes The a(1,6) glucan linkages in the starch molecule are
that hydrolyze storage reserves. This process has been well hydrolyzed by starch DBEs, which have also been implicated
characterized in cereal grains (Figure 3). A group of plant in starch synthesis as described earlier. This group of enzymes
growth regulators known as the gibberellins (GAs) are the hydrolyze a(1,6) linkages, thus producing additional end
first to be synthesized in the scutellum of the embryo. The groups for action by amylases and phosphorylases. Concerted
GAs diffuse into the aleurone layer and induce the produc- action of these enzymes results in the production of maltose,
tion of hydrolytic enzymes such as a-amylases active on the glucose, or glucose 1-phosphate. Maltose rarely accumulates
starch granules in the endosperm. The GA-inducible R2R3- during starch digestion as it is broken down to glucose by
MYB transcription factor known as GAMYB was first identi- maltase. Starch breakdown is influenced by its structure and
fied in the cereal aleurone. It has been proposed that a GA composition. Amylopectin fine structure and amylose both
receptor present on the aleurone membrane binds to GA, play important roles in effecting its breakdown by metabolic
which induces GAMYB activation, which further binds to enzymes.
the GA response element (GARE), 50 AACA/TA-30 , to activate
hydrolase genes including a-amylase involved in starch break-
down and mobilization for embryo growth. The hydrolysis
Raffinose Family Oligosaccharide
products are taken up by the embryo to initiate its growth.
Carbohydrates stored in the endosperm or cotyledons are the RFOs, accumulated in seeds by phloem loading and transport,
primary source of energy in the developing embryo until it are rapidly utilized during the early stages of seed germination.
forms leaves, and the photosynthetic process takes over to Alpha-galactosidase cleaves the terminal nonreducing
supply the necessary energy and chemical constituents for the a(1 ! 6)-linked galactose residues of a-galactosides, thus pro-
developing plantlet. viding readily available energy and carbon to the germinating
seed. Two types of a-galactosidases have been reported (1)
acidic (found in protein storage vacuoles (PSVs)) and (2)
alkaline (active in cytosol). Acidic a-galactosidase (activated
Starch Breakdown
by the pH shift during seed imbibition) was proposed to be
In cereal grains, starch is the major carbohydrate that is broken responsible for the breakdown of RFOs in PSVs during the
down during germination. Starch degradation occurs in three early stages of seed development, providing the main energy
distinct phases: (1) reduction of starch granules to soluble sources for seed metabolism. The resulting increase in sucrose
maltodextrins, (2) degradation and debranching of larger mal- concentration during the transition phase, from germination
todextrins to glucose and glucose 1-phosphate, and finally (3) to plant growth, was attributed to the induction of expression
transport of the glucose or glucose 1-phosphate from the stor- of alkaline a-galactosidase, resulting in the mobilization of the
age organs to the hexose phosphate pools of developing plant remaining RFOs. The galactose unit is thereafter phosphory-
cells for further metabolism into chemical constituents and lated by a galactokinase to produce a-D-galactose 1-P, which is
energy. In germinating seeds, the initial degradation of starch further metabolized through UDP-galactose to produce either
granules is associated with an increase in the a-amylase activ- UDP-glucose or glucose 1-P, which enters the cellular hexose
ity. Intact starch granules are attacked by a-amylase resulting phosphate pool. Depending on the metabolic activities of the
in the formation of random pits on the granules. During tissues, the carbon may flow into the respiratory pathways as in
the a-amylase attack, a(1,4) bonds in both amylose and the case of germinating tissues or to the synthesis of starch,
amylopectin are randomly broken to release large glucan sucrose, oligosaccharides, or other cellular components such as
polymers. Subsequent degradation occurs with assistance glycolipids or glycoproteins (Figure 2) in young meristematic
from b-amylases, phosphorylases, maltases, and starch DBEs. tissues. Most of the RFO-degrading enzymes have been puri-
a-Amylase action on amylopectin, which contains both a(1,4) fied from seeds of pulse crops, and increased activities have
and a(1,6) glucan linkages, results in maltose and branched been shown to be associated with seed germination. However,
dextrins of short chain length as a-amylase cannot break the strategies to modulate the activities of RFO-degrading enzymes
a(1,6) linkages. b-Amylases first act only on the nonreducing are complicated by the presence of two distinct pathways
ends of the glucan polymer and hydrolyze starch into for the metabolism of galactose 1-P to UDP-glucose and
b-maltose, which is rapidly converted into natural mixtures of glucose 1-P.
a- and b-isomers by mutarotation. Starch phosphorylases are a
group of phosphorolytic enzymes that degrade starch mole-
cules from the nonreducing end of the glucan chain. The
Fructans
reaction results in the complete breakdown of amylose to
glucose 1-P units, but in amylopectin, the degradation stops Fructans are hydrolyzed by fructan 1-exohydrolases (1-FEH) or
within a few glucose residues of the branch point to give fructan 6-exohydrolases (6-FEH) that cleave the terminal
dextrins. Like plant amylases, starch phosphorylases are also b(2,1)- or b(2,6)-linked fructosyl moiety, respectively. These
widely distributed in plants and have an important role in enzymes are also proposed to participate in fructan biosynthe-
starch degradation in leaves and seeds of many plants. sis by modulating the chain length and branching pattern of
However, in germinating cereal grains, starch phosphorylase the polymer. A b-fructofuranosidase from Jerusalem artichoke
activity is not detected, giving the a- and b-amylases a domi- tubers successively cleaves fructose units from inulin until a
nant role in starch degradation. In the seeds of other species mixture of fructose and the terminal sucrose unit remains.
and in leaves, starch phosphorylases also contribute toward Fructose can directly enter the respiratory pathway, whereas
starch degradation. sucrose must be broken down to glucose and fructose first.
172 CARBOHYDRATES | Carbohydrate Metabolism
Sucrose and abiotic stresses. It is often found that a single reaction in the
carbohydrate metabolic pathways is catalyzed by multiple iso-
Invertase and SS are the two enzymes capable of cleaving
forms of an enzyme. Differences in the temporal and spatial
sucrose present in plants. Invertases catalyze the irreversible
expression of genes encoding these isoforms present several
hydrolysis of sucrose to free glucose and fructose. Based on
challenges to modifying the carbohydrate metabolic pathways
optimum pH of their activity, invertases are grouped into acidic
by genetic engineering. To date, it has been possible to alter the
(optimum pH 4.05.5) and alkaline/neutral types (optimum
expression of genes encoding starch biosynthetic enzymes to
pH 7.08.0). Acidic invertases are mainly found in cytosol and
produce crops with modified grain carbohydrates that have
occasionally in cell walls, whereas alkaline/neutral types are
health benefits for human beings and/or grain processing
abundant in vacuoles and cell walls. Alkaline/neutral invertases
advantages. Some advances have also been made in changing
have recently been reported in chloroplasts and mitochondria.
the cell compartment and/or tissue in which a specific polysac-
Alkaline invertases are sucrose-specific, while neutral invertase
charide is stored. As knowledge of the carbohydrate metabolic
from carrot can also mediate raffinose and stachyose hydrolys-
pathway expands, novel strategies to alter the carbohydrate
es. The cell wall invertases hydrolyze the incoming translocated
biosynthetic or degradation pathways will emerge, which can
sucrose into glucose and fructose molecules.
be utilized to develop crop cultivars with novel products in the
SucS catalyzes the reversible reaction converting sucrose
grain. These new crops will diversify the uses of the grain to
and UDP to fructose and UDP-glucose. Fructose is available
benefit both the producers and consumers.
for respiration and the UDP-glucose can enter the hexose
phosphate pool for further metabolic processes. SS, compart-
mentalized in the cytosol, is the main enzyme that degrades
sucrose in starch storage organs such as developing seeds or in Learning Outcomes
rapidly growing tissues that are converting sucrose to cell wall
structural polysaccharides. In slow-growing and mature cells,
Hexose phosphate pool serves as a central component of
invertase is the major enzyme hydrolyzing sucrose, which pro- carbohydrate metabolism.
vides substrates for respiration.
Sucrose and raffinose family oligosaccharides maintain
sourcesink relationship by photosynthate translocation.
Starch, the most abundant storage polysaccharides, in
Cell Wall Polysaccharides plants exists as a temporary/transient form in leaves and a
stable storage form in grains and tubers.
The cell wall polysaccharides have a physiologically different
Cellulose and glycans are important for structural integrity
role in seed germination. The process of germination and of cell wall.
seedling development requires the participation of several
endo- and exohydrolases that degrade or modify the cell wall
Complex and highly regulated enzymatic machinery is
involved in structural and storage carbohydrate biosynthe-
structure. During germination, (1,3)- and (1,4)-b-D-glucan sis and metabolism. Its key modulator steps with possibil-
endohydrolases hydrolyze the (1,4) linkages adjacent to the ities of genetic alterations provide ample opportunities of
(1,3)-b-D-glucosyl residues to release large oligosaccharides. crop improvement for food and industrial usage.
The resulting oligosaccharides are further hydrolyzed by
broad-specificity exohydrolases to produce glucose, which can
subsequently be used in numerous ways (Figure 2). Some Exercises for Revision
other cell wall polysaccharides, such as arabinoxylans, can be
degraded to oligosaccharides that subsequently follow the oli-
gosaccharide degradation pathway as described earlier.
Question 1: Differentiate oligosaccharide from polysaccha-
ride with example. Which group of food grains has the
highest concentration of oligosaccharides?
Question 2: What are dietary fibers and what role do they
Concluding Remarks play in human nutrition?
Question 3: Describe two major metabolite pools of the
In most grains, carbohydrates are the major components that reductive pentose phosphate pathway with their major
provide the energy needed during seed germination and early functions inside the plant system.
seedling development. Carbohydrates also play a major role in Question 4: Describe the major steps of starch biosynthesis,
determining the quality of the grain and its subsequent use by with emphasis on the enzymes involved in each step.
human beings. Carbohydrate metabolism in plants differs from Question 5: In a germinating seed, what is the major starch-
most other organisms as it occurs in two distinct cellular com- metabolizing enzyme and how is it activated?
partments, the plastid and the cytosol. Most of the biosynthetic Question 6: How do tropical grasses differ from temperate
activities take place in the plastids, chloroplasts in leaves, and grasses in carbohydrate storage?
amyloplasts in the storage organs. The carbohydrate metabo-
lism in the cytosol communicates with that of the plastids
through metabolic carriers in the plastid envelope. The station- Exercises for Readers to Explore the Topic Further
ary nature of the plant forces it to withstand changes in the
external environment, and therefore, plants have developed How do plantenvironment interactions affect carbohy-
inherent flexibility in their metabolism to cope with both biotic drate metabolism in plants?
CARBOHYDRATES | Carbohydrate Metabolism 173
What are the mechanisms attributed for stress tolerance by Buchanan BB, Gruissem W, and Jones RL (eds.) (2002) Biochemistry and Molecular
sugars? Biology of Plants. Rockville, MD: American Society of Plant Physiologists.
Burton RA, Collins HM, Kibble NAJ, et al. (2010) Over-expression of specific HvCslF
How does genetic modification of one biosynthetic path-
cellulose synthase-like genes in transgenic barley increases the levels of cell wall
way affect the hexose phosphate pool and the biosynthesis (1,3,1,4)-b-D-glucans and alters their fine structure. Plant Biotechnology Journal
of other carbohydrates? 9: 119.
What are the critical limits (minimum and maximum) of Chibbar RN, Ambigaipalan P, and Hoover R (2010) Molecular diversity in pulse seed
carbohydrates concentrations required for normal plant starch and complex carbohydrates and its role in human nutrition and health. Cereal
Chemistry 87: 342352.
growth and development? Chibbar RN and Baga M (2003) Genetic modification of primary metabolism
How are sugars involved in the genetic regulation of meta- Carbohydrates. In: Thomas B, Murphy DJ, and Murray BJ (eds.) Encyclopedia of
bolic pathways? Applied Plant Sciences, pp. 449459. Oxford: Elsevier.
Cui SW and Wang Q (2009) Cell wall polysaccharides in cereals: Chemical structures
and functional properties. Structural Chemistry 20: 291297.
Cummings JH and Stephen AM (2007) Carbohydrate terminology and classification.
Acknowledgments European Journal of Clinical Nutrition 61: S5S18.
Edwin JR, Emilie AR, Veronique A, Lailiang C, and Robert T (2009) Phloem loading
We gratefully acknowledge the following for supporting our strategies in three plant species that transport sugar alcohols. Plant Physiology
149: 16011608.
research: Canada Research Chairs, Natural Sciences and Engi-
Hedley CL (2001) Carbohydrates in Grain Legume Seeds: Improving Nutritional Quality
neering Research Council, Western Grains Research Founda- and Agronomic Characteristics. Wallingford: CABI Publishing.
tion, Saskatchewan Winter Cereals Development Commission, Hrmova M and Fincher GB (2001) Structure-function relationships of b-D-glucan endo-
Winter Cereals Manitoba Inc., Agriculture and Agri-Food and exohydrolases from higher plants. Plant Molecular Biology 47: 7391.
Canada, Saskatchewan Pulse Growers, and Saskatchewan Jaiswal S, Ganeshan S, Baga M, and Chibbar RN (2010) In Planta modification of starch
quantity and quality. In: Singh BP (ed.) Industrial Crops and Uses, pp. 236258.
Agriculture Development Fund. Wallingford, UK: CAB International.
Lunn JE (2008) Sucrose metabolism. In: Encyclopedia of Life Sciences. Chichester:
Wileyhttp://dx.doi.org/10.1002/9780470015902.a0021259.
See also: Carbohydrates: Cereals: Chemistry and Physicochemistry Martinez-Villaluenga C, Frias J, and Vidal-Valverde C (2008) Alpha-galactosides:
of Non-starchy Polysaccharides; Grains other than Cereals: Non-starch Antinutritional factors or functional ingredients? Critical Reviews Food in Science
and Nutrition 48: 301316.
Polysaccharides; Starch: Analysis of Quality; Starch: Chemistry; Starch: McFarlane HE, Doring A, and Persson S (2014) The cell biology of cellulose synthesis.
Synthesis; Fats: Lipid Chemistry; Processing of Grains: Starch: Annual Review of Plant Biology 65: 6994.
Modification; Starch: Uses of Native Starch; Proteins: Protein Yan D, Duermeyer L, Leoveanu C, and Nambara E (2014) The functions of the
Synthesis and Deposition; The Basics: The Grains that Feed the endosperm during seed germination. Plant and Cell Physiology 55: 15211533.
Zeeman SC, Kossmann J, and Smith AM (2010) Starch: Its metabolism, evolution and
World.
biotechnological modification in plants. Annual Review of Plant Biology
61: 209234.
Further Reading
Relevant Websites
Aoki N, Hirose T, and Furbank RT (2012) Sucrose transport in higher plants: From
source to sink. In: Eaton-Rye JJ, Tripathy BC, and Sharkey TD (eds.) http://www.chem.qmul.ac.uk International Union of Pure and Applied Chemistry and
Photosynthesis: Plastid Biology, Energy Conversion and Carbon Assimilation. International Union of Biochemistry and Molecular Biology; IUPAC-IUBMB Joint
Advances in Photosynthesis and Respiration, vol. 34, pp. 703729. Berlin: Springer Commission on Biochemical Nomenclature (JCBN) Nomenclature of
Science Business Media B.V. Carbohydrates.
Blochl A, Peterbauer T, Hofmann J, and Richter A (2008) Enzymatic breakdown of http://www.fao.org Includes classification and chemistry of carbohydrates,
raffinose oligosaccharides in pea seeds. Planta 228: 99110. carbohydrate metabolism in fish, factors affecting it and energy transformation.
Starch: Chemistry
FL Stoddard, University of Helsinki, Helsinki, Finland
2016 Elsevier Ltd. All rights reserved.
Topic Highlights It is thus not as efficient an energy store where water avail-
ability is limited, such as in a dry, dormant seed.
Starch is a widespread polymer in the plant kingdom, Starch is deposited in granules between 1 and 100 mm in
composed of chains of a-1,4 linked glucose in helices, diameter within membrane-bound organelles. In the actively
bridged by a-1,6 bonds. photosynthetic parts of the plant, these organelles are the
It develops in granules within chloroplasts and amyloplasts. chloroplasts, and in storage organs, they are the modified
Two classes of starch molecule are recognized, highly chloroplasts called amyloplasts. Granule structure has a num-
branched amylopectin and poorly branched amylose. ber of features that are consistent across species, while granule
Packing of amylopectin double helices confers diagnostic size, morphology, and composition vary widely among spe-
x-ray diffraction patterns. cies. These aspects and starch composition affect the processing
Starch granule morphology differs between families of properties of the starch.
plants.
In the presence of heat and moisture, starch helices open
and the granules are disrupted. The Monomer and Linkages
Cooked, cooled starch forms a paste or gel with properties
that depend on amylose and amylopectin concentration The building block of starch is D-glucose, in its hexagonal
and structure. pyranose ring conformation (Figure 1). Carbon 1 of this ring
Cooked starch gradually recrystallizes (retrogrades), losing is a highly reactive, aldehydic, reducing end. Most commonly,
water (syneresis). during polymer formation in a plant, it attaches to carbon 4 of
Industry requires starches with different compositions another glucopyranose residue (Figure 2). When the mono-
(granule size, amylose content, and amylopectin chain mers are a-D-glucopyranose, the resulting polymer is a chain of
length) for different uses. starch, and when they are b-D-glucopyranose, it is a chain of
cellulose.
An a-1,4 linked glucan is at its most stable in a left-handed
Learning Objectives helix, with a complete turn taking six residues (Figure 3) and
occupying 2.1 nm. Hydrogen bonding between adjacent glu-
To understand the role of chain length, branching pattern, cose residues helps to stabilize the helix. The helix can allow a
and association in starch, as these affect economically second parallel helix to fit into it, creating a double helix. The
important industrial and food uses helical structure of starch is an important factor in its proces-
sing properties and in its enzymatic degradation during diges-
tion. Cellulose, in contrast, is highly crystalline and ribbonlike
Introduction rather than helical, and relatively few organisms have the
appropriate enzymes to hydrolyze it.
Starch is the major energy reserve of higher plants and hence a The presence of a-1,6 bonds (Figure 2) allows the develop-
major energy source in the diet of humans and many other ment of much larger and less soluble polymers through the
animals, including livestock animals grown for their meat and introduction of branching points connecting glucose chains of
milk and pest animals that consume grains in storage. Starch various lengths. Starch is an excellent example of the evolu-
comprises 70% of the dry matter of the endosperm of a tionary conservation of a successful structure throughout the
mature cereal grain and 16% of the fresh matter of a mature plant kingdom.
potato tuber.
Starch may be loosely defined as a polymer of glucose that
is distinguished from other glucans by a number of key Starch Polymer Types
features.
First, it is a-linked, unlike cellulose and other b-linked Two classes of starch, amylose and amylopectin, have been
glucans. The majority of these links are a-1,4 (also written as widely recognized, differing in chain length, branching pat-
a-(1 ! 4)), leading to chains of varying lengths, and the chains tern, and overall mass. Both have a single reducing group per
are joined together by a-1,6 linkages. These branching points molecule and a variable number of nonreducing ends. Most
occur in a highly ordered fashion, leading to double-helix plant starches comprise about one-fifth to one-third amylose
formation and a type of crystallinity, minimizing water- and the remainder amylopectin (Table 1). Plant breeders and
holding capacity. This feature is in significant contrast to geneticists have considerably enhanced this range since the
glycogen, an a-(1 ! 4), (1 ! 6) mixed-link glucan found in beginning of the twentieth century.
animals and bacteria. Ordering in glycogen is very limited, so Amylose is usually described as being essentially linear. A
it holds much more water than an equivalent mass of starch. typical amylose molecule comprises 211 chains of 200700
Source Granule average diameter (mm) Amylose concentration (%) Lipid concentration (%) Protein concentration (%)
Flexible spacers in
the amorphous
lamella (~2.7 nm)
B2 chain
Amylopectin
double helices in
the crystalline
lamella (~6.3 nm)
C chain
Figure 4 Representation of the amylopectin molecule as a liquid crystal. The shaded blocks represent double helices. In the dry state (left), they are
disordered, and in the hydrated state (right), they are aligned. Adapted from Donald AM, Perry PA, and Waigh TA (2001) The impact of internal
granule structure on processing and properties. In: Barsby TL, Donald AM, and Frazier PJ (eds.) Starch: Advances in Structure and Function, pp. 4552.
Cambridge: Royal Society of Chemistry, with permission.
chromatography is a modern method for amylose determina- polysaccharide that can accumulate in a grain, so when it dries
tion with a reasonable balance of throughput and accuracy. down at maturity, it is shrunken and wrinkled.
Intermediate material was a term used in maize starch
chemistry for many years, for polymers having some of the
features of amylose and others of amylopectin due to the Levels of Organization
combined effects of different mutations in starch-synthesizing
enzymes, for example, amylopectin-type branching but with The next order of structure is an alignment of the short, stiff
much longer chains. double helices of the amylopectin molecules. This takes place
Phytoglycogen is found in certain plant mutants such as in two ways. First, they align parallel to each other, along the
sweet corn. It is due to a mutation in the gene for starch- radius of the starch granule. This is what gives the polarization
debranching enzyme or isoamylase. Its structure is very com- of light, leading to a characteristic Maltese cross appearance
parable to that of animal glycogen, and its lack of higher-order when starch granules are viewed between crossed polarizers in
structure, such as double helix formation, allows it to remain a microscope. Second, the double helices pack together in an
largely water-soluble. This aspect limits the amount of storage energy-minimizing fashion (Figure 5). In some starches,
CARBOHYDRATES | Starch: Chemistry 177
Figure 5 Packing of amylopectin double helices: left, monoclinic crystal giving A-type x-ray diffraction pattern; right, open crystal giving B-type x-ray
diffraction pattern.
generally those from tubers and stems, the packing is hexago- Starch Granule Morphology
nal with a lumen in the center, which leads to a consistent
pattern, termed B-type, in x-ray diffraction. In other starches, Starches of wheat, rye, barley, and many of their wild relatives
mostly those from grains, the double helices pack much more in the cereal tribe Triticeae have biphasic granule initiation,
closely, without a lumen, leading to a different, A-type, x-ray rather than biphasic growth. One major starch granule, termed
diffraction pattern. Starches from grain legumes and from most A-type, originates in each amyloplast. About 15 days into grain
roots have an intermediate diffraction pattern, termed C-type, filling, membrane tubules appear around the equator of the
which implies that some of the hexagons have a lumen and A-granule and several subsidiary, B-type, granules form. At
others do not. A final x-ray diffraction pattern, V-type, is given grain maturity, the A-granules are approximately lenticular or
by amylose that encloses a lipid chain. High-amylose maize lentil-shaped, with a long diameter about twice the short dia-
gives a B-type diffraction pattern, suggesting that amylose does meter and with a detectable equatorial groove. Mean A-granule
not pack as tightly as amylopectin. Some species of yam (Dios- diameter can be as low as 15 mm in diploid wheats such as
corea) show A-type diffraction patterns and some B-type. Triticum monococcum or as large as 30 mm in some tetraploid
Starches with the B-type pattern generally retain or take up and hexaploid Aegilops species but is 2025 mm in cultivated
more water than those with an A-type pattern. rye, barley, durum wheat, and common wheat (Table 1). The
The crystalline double helices are thus aligned in a crystalline B-granules are roughly spherical and average 56 mm in diam-
lamella. This alignment is possible because the branches in eter in most of these species. There are other differences
amylopectin are clustered, leading to an amorphous (non- between A- and B-granules. B-granules lack the central cavity
crystalline) region (Figure 4). The thickness of this amorphous/ or hilum found in A-granules and also in the granules of other
crystalline ring is commonly 9 nm. This is commensurate with species. They also lack at least one granule-bound protein, and
a helix of 18 residues (6.3 nm) plus a few more residues to they can have amylose contents 24% lower than A-granules.
join with the B-chain in the amorphous lamella. This, in turn, With their greater surface area, they adsorb more water per unit
accounts for the 1530 unit length of the majority of amylo- mass than A-granules. The factors determining A-granule size
pectin chains. and B-granule concentration or initiation are the subjects
Alteration of packing can occur in the presence of gentle of continuing research. Many scientists view a third population
hydration, leading to the model that in the hydrated starch of small B-granules as a distinct C-granule type. This bimodal
granule, the rodlike double helices on their flexible attach- (or trimodal) granule size distribution is not found in any
ments function as a liquid crystal (Figure 4). other group of plant species.
Stacks of 16 amorphous/crystalline lamellae form semi- In maize, starch granules are spherical when isolated from
crystalline layers of 140 nm in thickness. These layers are mealy regions of the mature grain where there is space
separated by amorphous layers of comparable thickness. This between the granules but are polyhedral when isolated from
level of order was visible in early electron micrographs and vitreous regions where the granules have been in close phys-
considered to be growth rings. Early research found that they ical contact since partway through grain filling. Faceted granule
disappeared in wheat grown in continuous light, but this is not surfaces are thus attributable to restricted growing space rather
always the case and they are generally attributed to physical than active differences in granule development. A-type gran-
mechanisms. Most starch granules show the alternating semi- ules of wheat and barley often show indentations where other
crystalline and amorphous layers across their entire cross sec- granules have impinged on them during development. Oat
tion. Certain double mutants in maize, however, show clearly and rice have compound starch granules. Grain legume starch
biphasic growth. In ae du and ae wx lines, the core 3060% (by granules are generally ovoid or elliptical, and root or tuber
volume) of the starch granule shows the typical concentric starches are usually round to ovoid.
rings, but the remaining outer mass of the granule is shapeless Zero-amylose (waxy) starches have a fairly normal gross
and disorganized. It was proposed that there was a change in morphology. Granules of high-amylose starches, in contrast,
the relative expression of the starch synthetic enzymes and in such as those of amylomaize, pure-amylose barley, or
substrate concentration. wrinkled pea, are shapeless, fissured, and nonbirefringent.
178 CARBOHYDRATES | Starch: Chemistry
These features demonstrate the critical role of amylopectin The total protein concentration of extracted starch is higher
in determining granule crystallinity and the much less critical in cereal grains than in roots or tubers (Table 1), although the
role of amylose. exact figure depends heavily on the extraction method.
As mentioned in the preceding text, long amylose chains and Gelatinization has been defined as the collapse (disruption)
lipids can form complexes. The concentration of lipid in starch of molecular order within the starch granule, manifested in
is very strongly correlated with the amylose concentration of irreversible changes in properties such as granule swelling,
the starch. The amyloselipid complexes resist both their own native crystalline melting, loss of birefringence, and starch
leaching from the starch granule and also the entry of water solubilization. Pasting is what happens after gelatinization
into the granule. This latter aspect may be the biological impor- granule swelling, exudation of molecular components from
tance of starch-bound lipids in the quiescent or germinating the granule, and, eventually, total disruption of the granules.
seed. Starch processing properties are also affected by the pres- Starch granules in cold water can reversibly absorb 30% of
ence of bound lipid that increases gel temperature and the their weight in water. When starch is heated in an excess of
temperature of maximum viscosity, decreases gel strength, water, a sequence of events occurs as the kinetic energy of the
and delays staling by interfering with retrogradation. Removal polymer increases and the hydrogen bonds rupture. At the
of lipids lowers gel temperature and increases peak viscosity. pasting or gelatinization temperature, between 60 and 70 C,
Starch-bound lipids are generally either free fatty acids energy is absorbed in decrystallizing the crystalline regions of
(commonly linoleic) or lysophospholipids. The ratio of these the molecule. It also leads to an endotherm, as seen in the
two classes differs among species in the cereals (Table 2), but differential scanning calorimeter (DSC), due to the absorption
there is much less information on the pulses and pseudo- of energy. Typical values of the heat of gelatinization or gelati-
cereals. The lipid concentration of extracted starch varies up nization enthalpy are 812 J g1 but are as high as 32 in
to tenfold depending on the extraction method. amylomaize. The loss of crystallinity leads to a rapid swelling
of the granule combined with a considerable loss in structure,
as seen in a rapid increase in viscosity in a pasting curve
Starch-Bound Proteins (Figure 6). Amylose leaches from the granule but amylopectin
remains associated.
Numerous types of protein are found in close association with Further heating without stirring in an enclosed vessel, such
starch granules, some on the surface and others bound within. as a DSC, causes dissociation of amylose from lipid. This
The biological functions of some of these proteins remain to be process also absorbs energy and shows as an endotherm
determined. The surface proteins tend to be smaller than the between 102 and 110 C.
internal proteins. Altered sequences in the tryptophan-rich If, however, the heating is in the presence of stirring in a
surface proteins, puroindoline-a and puroindoline-b, have nonpressurized container, such as a Brabender Visco-Amylo-
been strongly linked with difference in grain hardness, but Graph or a Rapid Visco Analyzer (RVA), the viscosity of the
their causative role in this difference is controversial. suspension continues to increase as the amylose molecules get
Granule-bound starch synthase (GBSS), which has a rela- more dispersed and the granules become more porous, gelati-
tive molecular mass of 60 kDa, is responsible for synthesiz- nized, and swollen. Further stirring at a high temperature
ing amylose and is thus absent from the so-called waxy results in shear thinning, where the molecular entropy is not
mutants identified in a wide range of crops including maize, enhanced but granule structure is gradually further destroyed
barley, wheat, rice, pea, and cassava. Confocal laser scanning until a new plateau is reached. This phase is known as break-
microscopy of fluorescent-dyed proteins has shown that GBSS down. As the paste is cooled and stirred, viscosity reaches a
is localized in concentric spheres coinciding with the amor- final value. The RVA takes 15 min for the process, instead of
phous layers of the growth rings in the starch granules of about an hour for the Visco-Amylo-Graph. The RVA has the
maize, wheat, and potato. Other unidentified proteins were significant advantage that it gives readings in SI units (cP), in
shown by this method to be concentrated in the hilum of the contrast to the arbitrary units of older models and the Visco-
granule. A large, 140152 kDa form of starch branching Amylo-Graph.
enzyme I (SBE) was found to be associated only with In a set of rice starches of equivalent amylose content, most
A-granules, not B-granules, of wheat. RVA parameters were not significantly affected, but breakdown
was correlated negatively with the proportion of long amylo-
Table 2 Composition of starch-bound lipids of several cereals pectin chains and positively with the proportion of short. In a
separate study, increased firmness of cooked rice was associ-
Species Free fatty acids Lysophospholipids ated with increased content of long amylopectin chains. The
short chains are types A and B1, the long B2B4. Perhaps, the
Wheat, barley, and rye 10 90
presence of more interconnections between crystalline lamel-
Rice and oat 30 70
Millet and sorghum 45 55 lae, given by a higher content of B3 and B4 chains especially,
Maize 60 40 helps to maintain structure of the gelatinized starch granule.
Furthermore, the radius of rotation of the chain increases with
Data from a variety of sources. its degree of polymerization, and the viscosity increases as the
CARBOHYDRATES | Starch: Chemistry 179
Figure 6 Starch pasting profile, as determined with an RVA (Newport Scientific Pty Ltd., Warriewood, NSW, Australia). Key measurements are named
on the graph. The thin black line shows temperature profile and the thick green line shows viscosity profile.
Table 3 Classification of resistant starch material is used as an additive to white bread by an Australian
bread manufacturer in order to increase insoluble dietary fiber
Type Description Example
content without adversely affecting the color or texture of the
1 Physically inaccessible Grain fragments product.
2 Native granules Uncooked banana
3 Retrograded after heat/moisture Cooled, cooked
treatment potato
4 Chemically or thermally modified Acetylated starch Mutant Starches
and other metabolic problems related to insulin, glycemia, or functional foods and various articles on breeding and
obesity. genetics of cereals, pulses, and pseudocereals.
Many mutations affect maize starch synthesis, including ae What are the starch quality requirements (e.g., granule size
(amylose extender), du (dull), h (horny), sh1 and sh2 and amylose content) for bread, or biscuits, or noodles, or
(shrunken), and su1 and su2 (sugary). The double mutant ae pasta? See articles on the relevant food item.
wx maize has no true amylose, owing to the wx mutation, but The initiation of starch granules, particularly of the second-
has longer amylopectin chains (average 52 instead of 30 glu- ary starch granules in wheat, barley, and rye, remains an
cose residues) than the wild type due to the ae mutation. enigma, as does the regulation of granule size and shape.
Starches of other double or triple mutants of maize have desir-
able properties that emulate or exceed those of chemically
modified starches. These include good freezethaw stability See also: Barley, Rice and Maize Processing: Barley: Malting;
(wx sh1) and low viscosity when hot during processing but Breeding of Grains: Barley: Genetics and Breeding; Maize: Breeding;
high viscosity when cool, which is desirable for canning (du Wheat Breeding: Exploiting and Fixing Genetic Variation by Selection
h, du ae, du su2, and ae su2). Starch of the ae du wx triple mutant and Evaluation; Carbohydrates: Carbohydrate Metabolism; Resistant
has high paste viscosity, shear resistance, and acid resistance, Starch and Health; Starch: Synthesis; Food Grains and Well-being:
and the paste has a creamy texture making it a suitable fat Functional Foods: Overview; Genetics of Grains: Maize: Genetics;
substitute. Chemical analysis of chain length and branching Rice: Genetics; Processing of Grains: Starch: Modification; Starch:
pattern has not yet been at sufficiently high resolution to Uses of Native Starch; The Cereal Grains: Durum Wheat: Overview;
provide mechanistic explanations of these differences in pro- Maize: Overview; Oats: Overview; Rice: Overview; Rye Grain: Its
cessing properties. Genetics, Production, and Utilization; Wheat: An Overview of the Grain
That Provides Our Daily Bread; The Legumes and
Pseudocereals: Amaranth: Overview; Beans: Overview; Chickpea:
Conclusions Overview; Grain Legumes and Their Dietary Impact: Overview; Pea:
Overview; Pseudocereals: Overview; Quinoa: Overview; Wheat
Numerous techniques have been used to elucidate aspects of Processing: Chemistry of Cake Manufacturing; Wet Milling of Wheat;
the physical and functional chemistry of starch. Although it is a Wheat: Dry Milling; Wheat-Based Foods: Noodles: Starch; Pasta.
homopolymer consisting entirely of glucose, its structure is
extremely variable, showing differences in chain length,
branching pattern, and the incorporation of other molecules Further Reading
such as proteins and lipids. These differences lead to a wide
range of processing properties. Aspects of the nomenclature Atwell WA, Hood LF, Lineback DR, Varriano-Marston E, and Zobel HF (1988)
The terminology and methodology associated with basic starch phenomena.
may be confusing to the novice, in particular the use of the
Cereal Foods World 33: 306311.
letters A, B, and C to indicate such diverse traits as different BeMiller JN and Whistler RL (2009) Starch Chemistry and Technology, 3rd edn.,
types of amylopectin chain, granule size, or x-ray diffraction 904 pp. New York: Academic Press.
pattern. Many questions remain to be answered about the Bertolini AC (2009) Starches: Characterization, Properties and Applications, 288 pp.
various levels of organization within the starch granule, includ- Boca Raton, FL: CRC Press.
Donald AM, Perry PA, and Waigh TA (2001) The impact of internal granule structure on
ing the nature of the initiation of the starch granule itself. processing and properties. In: Barsby TL, Donald AM, and Frazier PJ (eds.) Starch:
Advances in Structure and Function, pp. 4552. Cambridge: Royal Society of
Chemistry.
Exercises for Revision Morrison WR (1995) Starch lipids, starch granule structure and properties.
In: Schofield JD (ed.) Wheat Structure, Biochemistry and Functionality,
pp. 261270. Cambridge: Royal Society of Chemistry.
Distinguish between amylose and amylopectin. Parker R and Ring SG (2001) Aspects of the physical chemistry of starch. Journal of
What happens to starch during gelatinization and Cereal Science 34: 117.
retrogradation? Sonnewald U and Kossmann J (2013) Starches From current models to genetic
engineering. Plant Biotechnology Journal 11: 223232.
Describe the ABCs of starch granule size, chain length, and
x-ray diffraction.
Describe the various levels of crystallinity in a starch
granule. Relevant Websites
http://www.corn.org Corn Refiners Association, Inc, Washington DC, USA.
http://www.jic.bbsrc.ac.uk The John Innes Centre, Norwich, UK.
Exercises for Readers to Explore the Topic Further http://www.mri.bund.de/en/de/institute/sicherheit-und-qualitaet-bei-getreide.html
Department of Safety and Quality of Cereals, Max-Rubner Institut, Detmold,
Germany.
How are we manipulating the genes of starch synthesis to http://www.starch.dk International Starch Institute, Aarhus, Denmark.
make new types of starch for food and industrial purposes? http://eu.wiley.com/WileyCDA/WileyTitle/productCd-2041.html Starch/die Starke
See articles on starch synthesis, uses of native starch, and (Journal home page), Wiley Interscience Publishing.
Starch: Synthesis
A Regina, CSIRO Food Futures National Research Flagship, Canberra, ACT, Australia; CSIRO Plant Industry, Canberra, ACT, Australia
S Rahman, Monash University Malaysia, Bandar Sunway, Malaysia
Z Li, CSIRO Food Futures National Research Flagship, Canberra, ACT, Australia; CSIRO Plant Industry, Canberra, ACT, Australia
MK Morell, International Rice Research Institute, Los Banos, Philippines
2016 Elsevier Ltd. All rights reserved.
Table 1 Typical properties of starch from different sources Waxy starches are also often preferred as thickeners because of
the greater solubility of starches without amylose. In cereals,
Starch Starch high-amylose starches are best known in maize among cereals
granule granule Apparent Gelatinization
where amylose content to 90% has been achieved. More
size range size amylose onset
recently, high-amylose wheat, barley, and rice have also been
Source (mm) distribution content temperaturea( C)
developed. The starch granules with elevated amylose content
Wheat 335 Bimodal 25 57 appear to be distorted. High-amylose starches are in demand
Barley 235 Trimodal 22 56 because such starches are useful starting points for the produc-
Rice 25 Normal 18 70 tion of resistant starches in foods and can be used in packaging
Maize 520 Normal 28 64 materials and adhesives.
a An important property of starch granules is the gelatiniza-
Gelatinization temperature measured by differential scanning calorimetry using starch
mixed with water in a ratio of 2 parts of water to 1 part of starch. The heating rate was
tion onset temperature, the temperature at which the granules
10 C min1 over a temperature range of 25150 C. begin to lose internal order and crystallinity. The gelatinization
Source: Data compiled from Jane J, Chen YY, Lee LF, et al. (1999) Effects of temperature of starches can vary considerably, from 57 C
amylopectin branch chain length and amylose content on the gelatinization and pasting for barley starches to over 75 C for rice. Mutations in some of
properties of starch. Cereal Chemistry 76: 629637; Rahman S, Li Z, Batey I, Cochrane the starch synthases (SSs) (GBSS and SSIIa) have been shown
MP, Appels R, and Morell M (2000) Genetic alteration of starch functionality in wheat. to lower the gelatinization onset temperature to 46 C in
Journal of Cereal Science 31: 91110. mutants of SSIIa in wheat and barley. On the other hand, the
suppression of starch branching enzyme (SBE) IIa and IIb
by weight, and the more numerous smaller granules, which are tends to increase gelatinization onset temperature to 60 C.
started later, contain only 2030% of the total starch in grain. The differences in gelatinization correlate with the differences
There have been some reports of slight variation in amylose in the branch lengths of starches from different sources and in
content between the A and B granules and between different the packaging of branches in the granule.
regions of the granule. Another important property is the viscosity of the gelati-
The amylose/amylopectin ratio does not normally vary nized starch. This is related to phosphate content and is a
greatly between sources, with a ratio of 1:3 being common. principal difference between starches from cereals and tubers.
However, mutations where amylose is completely missing are Potato starches have 0.5% phosphorylated glucose residues
fairly common and such starches are known as waxy starches. compared with 0.05% for cereal starches. Usually, the
In some food preparations, such starches are preferred, for phosphate is linked to the carbon at position six in glucose
example, in the preparation of many Asian sticky rice desserts. molecules. The difference in the phosphorylation imparts
Wheat Barley
Maize Rice
Figure 1 Starch granules from common cereals. Scanning electron micrographs by CSIRO Plant Industry by scanning electron microscopy.
CARBOHYDRATES | Starch: Synthesis 183
tuber starches with properties significantly different to cereal (1) ADPG pyrophosphorylase, which produces ADP-glucose
starches in terms of viscosity and flow properties. It has also from glucose-1-phosphate and ATP.
been shown that starches with more short and intermediate (2) SSs, which elongate glucan chains by using ADP-glucose to
glucan chains have a high starch viscosity. add a glucose residue at the nonreducing end.
The lipid content of cereal starches is low, 1%. However, (3) SBEs, which introduce a-1,6 branches to the glucan chain.
even this can be divided into three operational classes: non- These branches can then be extended by SSs.
starch, surface, and internal lipids. The internal lipids of wheat (4) Debranching enzymes remove branches from starch mol-
consist entirely of lysophospholipids, which are complexed ecules. The role of debranching enzymes in starch synthe-
inside linear glucan chains. The fatty acid composition of lipids sis is still being debated, although the genetic evidence for
differs slightly between wheat and rice, with wheat being richer a role for these enzymes is unambiguous.
in 18:2 (linoleic acid) and rice having more 16:0 (palmitic
Other enzymes are also capable of influencing the properties of
acid) fatty acids. How lipids accumulate in cereal starches is
the starch produced and these will be discussed later.
not yet clear, but the hydrophobic environment in the interior
Maize has been the most extensively characterized model
of the glucan chain leads to complexes of lipids with amylose.
system for the isolation of mutations affecting starch biosyn-
thesis. Analysis of these mutants (along with those from other
model systems such as Arabidopsis and Chlamydomonas) has
greatly contributed to the exploration of the key enzymes and
Pathway from Sucrose to Starch Granule
their isoforms involved in starch biosynthesis. In maize, muta-
Pathway Preceding Starch Biosynthesis tions in genes affecting starch synthesis such as the waxy and
amylose extender mutations can be brought about by the
Starch synthesis requires a hexose phosphate supply. Plants
insertion of transposable elements, a DNA sequence capable
produce sugars in photosynthetic tissues (e.g., leaves and peri-
of moving from one location to another within a genome.
carp of the grain) where they can support starch synthesis. Some
Such mutations often lead to alterations in the structure and
of these sugars are exported as sucrose. In the endosperm,
properties of starch. Specialty maize varieties such as waxy
incoming sucrose is either converted into uridine diphosphate
(granule-bound starch synthase (GBSS) I mutant), amylose
(UDP)glucose and fructose by a UDP-dependent sucrose
extender (SBEIIb mutant), sugary-2 (SSIIa mutant), sweet
synthase or converted to hexoses by invertase, as observed in
corn (sugary-1, isoamylase mutant), and dull (SSIII mutant)
maize. The UDPglucose or hexose phosphate derived from
find applications for specific food and industrial uses.
fructose is interconverted to glucose-1-phosphate by other
enzymes such as UDPglucose pyrophosphorylase, hexoki-
nase, and phosphoglucomutase. This glucose-1-phosphate
can be used in various metabolic pathways; however, once the ADPG Pyrophosphorylase
glucose-1-phosphate is utilized to form adenosine diphosphate
(ADP) glucose (ADPG), a commitment is made to the synthesis ADPG pyrophosphorylase is believed to be the key enzyme
of starch. that determines the rate of starch synthesis in leaves. Genetic
engineering has been used to introduce the ADPG pyropho-
sphorylase from Escherichia coli (which is not subjected to
feedback inhibition) into potatoes, and tubers with a higher
Biochemistry of Starch Biosynthesis
content of starch were obtained. It is not clear, however,
A consensus view of starch biosynthesis in cereal endosperm is whether ADPG pyrophosphorylase is also the rate-limiting
shown in Figure 2. Starch biosynthesis requires four types of enzyme for starch biosynthesis in the grain, as the purified
enzymes, which are as follows: enzyme from leaves and grains shows differences in properties.
ADPG pyrophosphorylase from both leaves and endosperm
is composed of four subunits, in which two of each type are
large and small. The large subunit has a mass of 55 kDa,
Isoamylase whereas the small subunit is slightly smaller, 50 kDa. The
SBE IIb
SBE IIa sequence of amino acids of these subunits is 50% identical.
ADPG SBE I Mutations in the sequence of either of these subunits can
75% Amylopectin
PPase destroy or reduce the activity of the enzyme, and then grains
SS III
or seeds with a shrunken appearance (due to decrease in starch
ATP PPi SS II
SS I synthesis) are produced. Such mutations in ADPG pyropho-
G1P ADPG 25% Amylose
sphorylase were given the names brittle-2 (mutation in small
ADP subunit) and shrunken-2 (mutation in large subunit) when
ADPG GBSS
they were discovered in maize (Table 1).
Recent evidence shows that ADPG pyrophosphorylase in
the cereal grain exists in both the cytoplasm and amyloplast of
Amyloplast endosperm cells; however, most of it is in the cytoplasm. This
enzyme in the cereal grain differs from that in the seeds of
Figure 2 A consensus view of the starch biosynthetic pathway in dicots and leaves of all the plants investigated so far, where
cereals. ADPG pyrophosphorylase appears to exist exclusively in the
184 CARBOHYDRATES | Starch: Synthesis
plastids. Evidence from both monocots (such as barley and source. The enzymes from the leaves of all plants examined,
maize) and dicots (such as peas) suggests that there is more maize endosperm, and potato tubers are allosterically activated
than one gene for the large and small subunits of ADPG by 3-phosphoglyceric acid and inhibited by inorganic phos-
pyrophosphorylase, and different versions of these may code for phate. Pea and bean embryo and wheat and barley endosperm
the forms that are expressed in the cytoplasm and in the plastid. ADPG pyrophosphorylases, on the other hand, are much less
The annotation of the rice genome sequence also indicates sensitive, or insensitive, to modulators compared with their
multiple gene sequences coding for ADPG pyrophosphorylase. leaf counterparts. The allosteric properties of ADPG pyropho-
The transfer of ADP-glucose from the cytoplasm of endo- sphorylases are of potential importance in determining its role
sperm cells to the amyloplast is important as the other enzymes in controlling the rate of starch synthesis. Altered starch bio-
in the process are located there. The Brittle-1-encoded ADPG/ synthetic rates resulting from altered allosteric properties due
ADP antiporter facilitates this transfer in maize. Mutations are to mutations in ADPG pyrophosphorylase subunits have been
known where this transfer is affected and this too produces demonstrated in maize, barley, and the green algae Chlamydo-
grain where starch synthesis is affected (Table 2). monas. Alterations in the properties of ADPG pyrophosphory-
ADPG pyrophosphorylases exhibit complex regulatory lase have been utilized in the plants grown in the greenhouse
properties that vary depending on the species and tissue to increase starch production and thus seed weight.
Source: Data compiled from the references in Further Reading section. ? not clear if the gene responsible has been identified.
CARBOHYDRATES | Starch: Synthesis 185
Starch Synthases clearly different from normal. The barley mutants have high
amylose (70%); the wheat mutants, less so (35%). It is not
SSs elongate preexisting glucan chains in the amyloplast by clear if there are differences in the structure of the amylose. The
adding the glucosyl moiety from ADP-glucose to the nonredu- starch granules are of distorted appearance and have a lower
cing end of an existing a-1,4-glucan. gelatinization temperature. The proportion of branches of
Ten different SS genes have been identified in plants that 725 glucose units in length is decreased, while the proportion
encode two classes of SSs. One class of SS is only found bound of branches of 46 glucose units in length is increased. There
to starch granules and these are known as GBSS. The other class are also effects on the abundance of other starch biosynthetic
of SS is present either in the amyloplast stroma alone or dis- enzymes, so it is difficult to know how many of the effects on
tributed between the stromal and granular fractions, and these starch properties are the direct result of the lack of this enzyme
are known as soluble SSs. Four types of soluble SS have been and how many are due to the other enzymes affected. The
reported in crop plants: SSI, SSII, SSIII, and SSIV. In monocots, differences in the starch between indica and japonica rice
the SSII has been further subdivided into the IIa and IIb forms, cultivars appear to be due to a combination of alterations in
SSIII into SSIIIa and SSIIIb forms, and SSIV into SSIVa and SSIIa and GBSS.
SSIVb forms. The primary structures of soluble SSs from different SSIII has been subgrouped into SSIIIa and SSIIIb. SSIIIb is
sources share a distinguishing feature, an N-terminal extension mainly expressed in the pericarp and does not appear to be
with little homology between sequences from different sources. easily detected in the endosperm. SSIIIa is a polypeptide of
Comparison of the deduced amino acid sequences of SSs 180 kDa. The absence of this enzyme in maize leads to a
revealed that maize SSI, SSIIa, SSIIb, and SS IIIa contained slightly higher amylose content, and the starch looks dull
N-terminal extensions of 93, 176, 144, and 656 amino acids, compared with normal. Mutants in rice and barley have a
respectively, compared with GBSS. Pea SSII also contains a flex- reduced amylopectin content, a reduced starch content, but
ible 162-amino acid N-terminal arm. It has been shown through an increased amylose content. The mutation of SSIIIa increases
expression analysis using N-terminally truncated SSII that the the expression of SSI and GBSSI in the endosperm. An inter-
N-terminal extension is not essential for the catalytic activity of mediate glucan fraction ( 15%), which is distinguished from
the enzyme but is probably related to substrate binding. amylose and amylopectin, occurs in starch from maize lines
There are two types of GBSSs: I and II. GBSSII is similar in missing SSIIIa.
amino acid sequence to GBSSI but is not expressed in the grain. SSIV has been subdivided into SSIVa and SSIVb isoforms in
GBSSI is a single polypeptide, 60 kDa in mass. If the grain cereals. Both isoforms are expressed in the endosperm,
lacks an active GBSSI, then starch is composed almost entirely although SSIVa is expressed at a higher concentration than
of amylopectin. Such starch is called waxy starch (Table 2). SSIVb in rice. The importance of either of the SSIVs for starch
Clearly then, GBSSI is essential for the synthesis of amylose synthesis in the endosperm is unknown at present. Mutation of
and the other SSs cannot substitute for it. SSIV in Arabidopsis dramatically reduces the number of starch
The hexaploid nature of wheat allows a range of starches of granules per plastid, leading to a large increase in starch gran-
intermediate amylose content to be developed, an opportunity ule size in the leaf, and it appears that SSIV has a critical role in
not possible in diploids such as barley. In wheat, the GBSSI regulating the number of starch granules.
gene has been mapped as a triplicate set of single-copy home-
oloci on chromosome arms 7AS, 4AS, and 7DS. A reduction in
the proportion of amylose is observed when the GBSSI activity
is reduced, and these starches are known as low-amylose or Starch Branching Enzymes
partial waxy starches. The loss of only one of the three iso-
forms, the 4A form of GBSS, has clearly been shown to yield a SBEs, as their name suggests, are required for the addition of
starch with increased ability to swell in water on heating. This branches to linear glucans. Their impact is clearly more on the
property is desirable for udon noodle production, and such synthesis of amylopectin than amylose, although branching
low-amylose starches from wheat are preferred for this end use. enzymes may also have a subtle role in amylose synthesis.
The soluble SSs are also all single polypeptide enzymes. SSI There are two broad types of SBEs: SBEI and SBEII. Expres-
is 75 kDa in mass and is found both free in the amyloplast sion of these enzymes in bacteria has shown that SBEI adds
and bound to the starch granule. Starches of SSI mutants of longer branches than SBEII. They are both single polypeptide
Arabidopsis and rice and SSI RNAi silenced wheat have reduced chains of 85 kDa. SBEI and SBEII share 60% sequence
amounts of amylopectin, low peak viscosities, and high peak identity over the middle third of the molecules.
gelatinization temperatures. Studies in these modified lines The importance of SBEI for starch biosynthesis in the grain
have shown that SSI is involved in the synthesis of short chains is not clear. Mutations in SBEI in maize and wheat did not
of the amylopectin (DP 812). produce a clear phenotype. A number of isoforms of SBEI may
SSII has been subdivided into IIa, IIb, and IIc forms on the exist in wheat. However, in rice, an SBEI mutant contains
basis of gene sequence comparisons; however, the IIb and IIc amylopectin with significantly decreased proportions of long
forms appear not to be expressed or expressed at very low chains of DP 37 and above and short chains of DP 1221.
concentrations in the cereal endosperm. The IIa form is However, the proportion of short chains of DP 10 and below
85 kDa in mass, although the wheat enzyme appears to be and intermediate chains of DP 2434 increased. Starches from
over 100 kDa by electrophoresis. Barley and wheat lines lack- the rice SBEI mutant had a low gelatinization onset tempera-
ing SSIIa have been produced, and the starch from these lines is ture. SBEI in rice may also play a crucial role in the synthesis of
186 CARBOHYDRATES | Starch: Synthesis
amylopectin molecules by interacting with phosphorylase 1 to were observed for the sugary mutants ranging from no starch
form branched maltodextrin primers. and only phytoglycogen to starch in the outer layer of the
SBEII is divided into two isoforms in cereals: IIa and IIb. endosperm and phytoglycogen internally. In contrast, in barley,
The amino acid sequences of IIa and IIb isoforms are very both phytoglycogen and starch were reported in the same
similar (over 80% sequence identity within a species). In endosperm cells. The maize sugary mutants also contain both
maize, the IIb form makes up 90% of the starch branching phytoglycogen and starch. In barley, it has been reported that
enzyme II activity in the endosperm, but the proportion of the the lack of isoamylase leads to an alteration in the initiation
two isoforms is nearly equal in wheat. Maize lines lacking pattern of starch granules.
starch branching enzyme IIb produce a very high (70% and While it is established that isoamylase plays a significant
higher) proportion of amylose. Rice SBEIIb mutants also con- role in starch biosynthesis, the mechanism by which it acts is
tain high-amylose starch, but the proportion is much lower not resolved. Two mechanisms have been proposed. One pro-
(40%) than in maize. In wheat and barley, suppression of posal known as the glucan trimming mechanism suggests that
SBEIIb expression using RNAi technology has no effects on isoamylase removes moderately and loosely spaced branches
starch amylose content. Suppression of both SBEIIa and SBEIIb from a soluble preamylopectin structure that interferes with
is required for the production of amylose (>70%). In rice the crystallization of amylopectin and promotes the formation
suppression of SBEI, IIa and IIb are required to achieve amy- of a stable amylopectin structure with regularly packed glucose
lose contents of more than 50%. High-amylose starches are units. In the absence of isoamylase activity, the preamylopectin
linked to the formation of resistant starches in products. Resis- structure is further branched by branching enzymes and results
tant starch has been shown to have very beneficial effects if in the production of phytoglycogen. The second model is
present in the diet. based on the assumption that a competition exists between
polysaccharide aggregation into starch and the nonproductive
formation of water-soluble polysaccharides (WSPs), in turn
Debranching Enzymes resulting in a competition for both carbon source and soluble
enzymes involved in amylopectin synthesis. The role of isoa-
Debranching enzymes remove branches from branched glu- mylase here would be to clear the stroma of WSPs and prevent
cans. Two types of debranching activities, pullulanase and phytoglycogen formation. A role in starch granule initiation
isoamylase, have been described from developing endosperms and growth, apart from phytoglycogen suppression, for isoa-
of rice and maize. They differ in the type of glucan they can mylase is also suggested based on the analysis of barley
debranch with pullulanase debranching more efficiently isoamylase mutants.
open structures (e.g., pullulan) and isoamylase debranching
the more branched structures (e.g., glycogen). The role and
importance of these enzymes for starch biosynthesis are still
being debated. Other Enzymes with Potential Roles in Synthesis
Pullulanase is a single polypeptide enzyme of 100 kDa in
mass. Grains from a rice pullulanase mutant contain starch Many of the new insights into starch synthesis have come from
with a reduced proportion of short chains (DP 13 and the study of simpler systems, such as Chlamydomonas and
below) and a slightly lower gelatinization temperature (lower Arabidopsis. Such studies have indicated other loci that are
by 13 C). Such mutations in rice do not change the amylose important in starch synthesis in these organisms. It is possible
content. A pullulanase mutant in maize contains significant (although by no means certain) that genes encoded by such
quantities of low-molecular-mass a-1,6-polyglucans in the loci are also important for starch synthesis in the grains of
endosperm. Similar to rice, mutants in maize pullulanase con- cereals.
tain starch with unaltered amylopectin structures and amylose/ D-Enzyme or disproportionating enzyme has the property
amylopectin ratios. of breaking a glucan chain and adding one portion to the
Isoamylase activity in plants is controlled by three gene nonreducing end of a preexisting chain. In this way, one
products, designated as ISA1, ISA2, and ISA3. Of these, ISA1 chain is made shorter but the other one is made longer. In
and ISA2 are involved in starch biosynthesis and ISA3 is Chlamydomonas mutants where the gene for D-enzyme is
involved in starch breakdown. ISA1 is more catalytically active affected, the starch produced appears to have somewhat more
than ISA2 and can form complexes with it. ISA1 is a single amylose. These mutants are required to be grown in constant
polypeptide enzyme of 90 kDa in mass. It can debranch light. Genes for D-enzyme have been described from Arabidop-
highly branched structures like amylopectin and glycogen. sis, rice, maize, barley, and wheat.
The lack of this enzyme in rice and maize is associated with Starch phosphorylase is another enzyme that could have an
the production of a very highly branched and unusual poly- impact on starch biosynthesis in the grain. This enzyme breaks
saccharide called phytoglycogen. Such lines also contain high down starch and was originally considered to be important for
levels of free sugars and are known as sugary mutants. How- degradation. There are at least two isoforms in the plant a
ever, there is also a reduction in the level of pullulanase in cytosolic form and a plastidic form; however, Chlamydomonas
these sugary mutants, so it is difficult to know which of the appears to contain three isoforms. The lack of one of these
effects are directly due to the lack of ISA1. The effect of the loss isoforms leads to a starch excess phenotype and the production
of ISA2 is slight in rice. of larger starch granules. In rice, the same phenotype is
There are differences between the isoamylase mutants observed, but this is more obvious at low growth temperatures
described in rice, maize, and barley. In rice, different phenotypes (20 C). In wheat, plastidic phosphorylase appears to be
CARBOHYDRATES | Starch: Synthesis 187
associated with leaf and seed endosperm accumulation, not The processing of the initial transcript to the final RNA is
with the degradation of reserve starch. clearly a complex process. Many of the mutants in GBSSI are
The initiator protein for glycogen is called glycogenin. Glyco- due to errors or inefficiencies in the processing of the RNA. In
genin is an autocatalytic glycoprotein, and the glucose residues barley, the waxy mutations have been reported that produce
provide the starting point for glycogen synthesis. By analogy, low amylose, rather than zero amylose. In these mutant starch
with this, a number of glycogenin-like proteins have been granules, the outer cell layers of the endosperm contain more
described from plants, but again, the importance of these pro- amylose than the inner layers. Investigation of these mutants
teins to starch biosynthesis in the grain is yet to be elucidated. revealed that the change in expression of GBSSI in these lines is
The R1 protein is important for starch phosphorylation in due to a 413-bp deletion of part of the promoter and 50 -
potatoes. If this protein is downregulated in potatoes, then the untranslated region of the gene. Two other barley waxy families
starch produced is sharply reduced in phosphate content and analyzed have no detectable level of amylose due to a 1-bp
increased in amylose content. The peak viscosity was also alteration in the GBSS gene that completely eliminates GBSS
reduced. As cereal starch contains naturally far less phosphate, activity. In rice, a single nucleotide polymorphism at the leader
the importance of this protein for cereal starch biosynthesis intron 50 splice site of the GBSS gene has been shown to be
needs to be ascertained. However, sequences similar to the associated with changes in amylose content due to altered
gene for the R1 protein are found in the cereal genome. In efficiency of GBSS mRNA processing. A sequence of AGGTATA
Arabidopsis, potato, and wheat, the absence of the R1 protein at this site is found to be associated with cultivars having
leads to a decrease in starch breakdown and the consequent an amylose content of above 18% as against a sequence of
production of the starch excess phenotype. AGTTATA that is present in cultivars having lower amylose
content.
The locations of most of the starch biosynthetic enzyme
genes are known in wheat and shown in Figure 4. Many of
Protein Interactions in Starch Biosynthesis
these genes are on chromosome 7. In rice, the locations of the
gene are known from the genome sequence, and the locations
Recent studies in maize and wheat have shown that the key
of the wheat genes are consistent with the established relation-
enzymes involved in starch biosynthesis congregate into protein
ships of the chromosomes in these two grasses.
complexes during starch biosynthesis. These complexes may act
as functional units to perform the task of granule formation.
Several such interactions are now known. For example, SSIIa
forms a complex with SSI and SBEIIb in maize. The composition
Future Work
of such assemblies may change when there are changes in the
Genomics has essentially provided the whole genome
expression of key enzymes. For example, in the amylose
sequence of Arabidopsis and rice. The sequencing of the wheat
extender mutant of maize, in which the SBEIIb activity is lack-
and barley genomes is also nearing completion. It should soon
ing, SBEI, SBEIIa, and starch phosphorylase are recruited into
be possible to identify all putative starch biosynthetic enzyme
the complex involving SSIIa and SSI.
genes in the genomes of these species.
The function of such putative candidate genes can be ana-
lyzed by performing conditional knockouts of selected genes,
Genes and Genetics in Wheat and powerful RNAi technology can be used. TILLING has also
been used to obtain desired mutants in starch biosynthesis.
The structure of the genes for the principal starch biosynthetic Additionally, more sophisticated gene-editing techniques
enzymes mentioned earlier has been described from wheat using DNA nucleases based on technologies such as TALENs
(Figure 3). The genes are all complex in intron/exon structure. and CRISPRs may be applicable for such experiments. Con-
The genes differ considerably in length, varying from 6 to 11 kb. versely, gene inactivation through tagging will also yield infor-
mation about the role of candidate genes. This can arise either
GBSS through the identification of the tagged gene and analysis of
the phenotype or by observing a phenotype and then identify-
SSI
ing the gene inactivated. Both approaches will be highly
SSII informative.
SSIII Analysis of expression of all of the genes in a tissue by
means of microarrays and transcriptomics can also be per-
SBEI
formed. The ability to study the pattern of transcription of
SBEIIa the whole genome by means of chips and the transcriptome
SBEIIb will facilitate the study of linked genes and pathways. If tagged
mutants are compared with the wild type, then the total
Isoamylase changes in transcription can be identified, and pathways can
begin to be deciphered. For untagged mutants or natural phe-
0 2 4 6 8 10 11 12 notypes, all the genes affected in a particular phenotype can be
Figure 3 Structure of wheat starch biosynthetic genes (data from identified from microarray or sequence abundance analysis of
authors laboratory). Exon sequences are illustrated by the thick lines; the expressed genes in bulked segregants differing in the phe-
intron sequences by the narrow line. notype studied.
188 CARBOHYDRATES | Starch: Synthesis
WxB1
WxA1 WxD1
SSI
SSII
Isoamylase
SSIII SBEIIb
SBEIIa AGP-S
SBEI
1 2 3 4 5 6 7
Figure 4 Chromosomal location of wheat starch biosynthetic genes. Data from authors laboratory observations and from the suggested readings.
New Starches in Cereals able to precisely manipulate the starch structure and function-
ality to suit specific end uses.
It is clear that with increasing knowledge of the enzymes
involved in starch biosynthesis, the ability to alter the starch
produced is also gained. New products and processes will need Exercises for Revision
to be developed to capture any advantages occasioned by these
new starches. Although such new phenotypes can be produced Why is starch important for our daily life?
most easily by using transgenic technology, current public How many classes of enzymes are involved in starch
opinion would make it prudent to also produce starches by synthesis?
conventional breeding. The development of molecular Which starch synthetic enzymes are important in the deter-
markers and rapid breeding technologies (such as the produc- mination of starch structures?
tion of doubled haploids and embryo culture and the ability to Which starch synthetic enzymes are important in the deter-
quickly extract DNA from small samples) will undoubtedly mination of starch properties?
also speed up the production of novel types of starch by How can novel starches be produced in cereals?
conventional breeding strategies. What approaches can be used for the study of the roles of
the enzymes in starch synthesis?
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Improving human health through understanding the complex structure of glucose Roles of resistant starch and nonstarch polysaccharides. Physiological Reviews
polymers. Analytical and Bioanalytical Chemistry 405: 89698980. 3: 10311064.
James MG, Denyer K, and Myers AM (2003) Starch synthesis in the cereal endosperm. Tuncel A and Okita TW (2013) Improving starch yield in cereals by over-expression of
Current Opinion in Plant Biology 6: 215222. ADPglucose pyrophosphorylase: Expectations and unanticipated outcomes. Plant
Jane J, Chen YY, Lee LF, et al. (1999) Effects of amylopectin branch chain length and Science 211: 5260.
amylose content on the gelatinization and pasting properties of starch. Cereal Utsumi Y, Utsumi C, Sawada T, Furjita N, and Nakamura Y (2011) Functional diversity
Chemistry 76: 629637. of isoamylase oligomers: The ISA1 homo-oligomer is essential for amylopectin
Morrison WR (1988) Lipids in cereal starches: A review. Journal of Cereal Science 8: 115. biosynthesis in rice endosperm. Plant Physiology 156: 6177.
Morrison WR (1993) Seed storage compounds: Biosynthesis, interactions and Yu TS, Kofler H, Hausler RE, et al. (2001) The arabidopsis sex 1 mutant is defective in
manipulation. 12. Cereal starch granule development and composition. the R1 protein, a general regulator of starch degradation in plants, and not in the
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pp. 175190. Oxford: Oxford University Press. biotechnological modification in plants. Annual Review of Plant Biology
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Advances in Plant Pathology Biotechnology of Cereals, vol. 34, pp. 165236.
San Diego: Academic Press. http://www.ncbi.nml.nih.gov A resource for molecular biology information.
Tetlow IJ (2011) Starch biosynthesis in developing seeds. Seed Science Research http://www.tigr.org/tdb Provides databases containing DNA and protein sequences
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Starch: Starch Architecture and Structure
V Vamadevan, University of Minnesota, St. Paul, MN, USA; Agriculture and Agri-Food Canada, Guelph, ON, Canada
Q Liu, Agriculture and Agri-Food Canada, Guelph, ON, Canada
2016 Elsevier Ltd. All rights reserved.
Topic Highlights common. Cereal starches are generally small and have round
or polyhydric shapes with a concentric hilum and a unimodal-
Granule morphology, structure, and composition of size distribution. Wheat, triticale, barley, and rye starches,
starches from different genetic backgrounds. however, exhibit bimodal size distributions, with large
Organization of glucan chains at different structural levels (A-type) and small (B-type) granules. Starch granules from
of starch granules. tubers are usually large and oval-shaped with an eccentric
Cluster model of amylopectin versus building block back- hilum. Pulse starches are normally medium-sized, with a bean-
bone model. like or ovoid shape and an elongated or starred hilum. High-
Unit chain categories of amylopectin. amylose starches generally appear as irregular, filamentous, or
Branching point distribution pattern in amylopectin. rod-shaped granules (Figure 1).
Structural parameters of amylopectin. Corn, sorghum, millet, wheat, rye, and barley starch gran-
ules are characterized by nonuniformly distributed surface
Learning Objectives pores. Some granules have numerous pores, others feature
only a few, and the starch granules of rice, oat, pulse, potato,
To understand the complex hierarchical structure of starch tapioca, canna, yam, and arrowroot appear to have none at all.
granules. In some starch granules, such as maize, sorghum, and millet,
To understand the concepts underlying proposed amylo- radially oriented channels that connect to a central cavity have
pectin models. been observed. Which biomechanism determines the size of
To understand the internal structure of amylopectin. the granules in different plants and how the pores and chan-
To identify unresolved structural aspects of starch granules. nels are formed are still unclear.
Introduction
(a) (b)
(c) (d)
Figure 1 Light micrographs (1000 ) of maize (a), canna (b), mung bean (c), and Hylon VII maize starches (d).
Reducing
f
end
Cluster
(short
chains)
A
Long chain
(a)
C Short
chains
A f
Amorphous Reducing
backbone end
(b) (long chains)
Figure 2 (a) Schematic representations of the cluster model (adapted from Robin JP, Mercier C, Charbonniere R, and Guilbot A (1974)
Lintnerized starches. Gel-filtration and enzymatic studies of insoluble residues from prolonged acid treatment of potato starch. Cereal Chemistry 51:
389406) and (b) the backbone model of amylopectin (reproduced from Chauhan F and Seetharaman K (2013) On the organization of chains in
amylopectin. Starch/Starke 65: 191199.).
associated with B-chains indicate the number of clusters of the crystalline lamellae, whereas internal chains are present
they span (Figure 3). According to the divisions obtained by primarily in the amorphous lamellae. The internal unit chain
Hizukuri using HPSEC, the periodicity of the CL is approxi- profile can be obtained from f,b-limit dextrins by removing
mately dp 27 to dp 28, and the average CL is dp 17 to dp 26. the external amylopectin chains using the exo-acting enzymes
Hanashiro et al. observed a shorter periodicity (dp 12) with phosphorylase and b-amylase. Since exo-acting enzymes can-
HPAEC and named the chain fractions fa- (dp 6 to dp 12), fb1- not bypass branching points, the resulting limit dextrin repre-
(dp 13 to dp 24), fb2- (dp 25 to dp 36), and fb3- (dp > 36) sents the internal amylopectin structure. In f,b-limit dextrins,
chains, which presumably represent A-, B1-, B2-, and B3- all A-chains and the external segments of B-chains appear as
chains, respectively. Using this division, B2-chains are consid- maltosyl and glucosyl stubs, respectively. The characterization
ered a subdivision of the B1-chains categorized by Hizukuri. of A- and B-chains from f,b-limit dextrin provides a relatively
The very shortest amylopectin chains consist of six glucose accurate estimate of the ratio of A-chains to B-chains, which,
residues. The limit dextrin profile of amylopectin reveals that on a molar basis, is typically in the range of 0.81.5.
A-chains (fa-chains) overlap with B1-chains (fb1-chains), so The internal amylopectin structure consists of two major
the fa fraction therefore represents only part of the A-chains. groups of B-chains: short B-chains (dp < 24) and long B-chains
A-chains consist of a subgroup of chains called fingerprint (dp > 24). Short internal B-chains (BS chains) are subdivided
A-chains (dp 6 to dp 8), which are too short to form double into fingerprint B-chains (Bfp) with dp values 7 and a major
helices and can cause structural defects within the crystalline group of short B-chains (BSmajor) with dp values from 8 to 25.
lamellae. On a molar basis, the ratio of short to long unit Long internal B-chains are classified as B2-chains (dp 26 to dp
chains varies from 4 to 19, depending on the starch source. 50) or B3-chains (dp > 50). Short B-chains are part of clusters,
Although the unit chain profile is useful for describing many of and long internal B-chains function as cluster-connecting
the structural features of the amylopectin molecule, it fails to chains. Based on the quantity and distribution pattern of the
explain how glucan chains are organized within the amylopec- short and long internal B-chains, amylopectin from different
tin molecule. botanical sources is one of four different types: type 1, type 2,
type 3, or type 4 (Figure 4). Type 1 amylopectin is character-
ized by large numbers of short internal chains, mainly BSmajor-
Organization of Amylopectin Internal Chains
chains, with a broad distribution pattern, whereas type 4
The organization of the unit chains in amylopectin determines amylopectin has a large number of long internal chains.
the unique architecture of the starch granule. Unit chains can Types 2 and 3 represent intermediate distributions. Type 2
be categorized as either external or internal chains. External amylopectin exhibits a significant number of Bfp chains in
chains are composed of A-chains and a portion of the B-chains. clusters. Based on this grouping, cereal starches are considered
The AB-chain segment that has branching points is designated to be types 1 and 2, and root and tuber starches to be primarily
an internal chain. External chains participate in the formation types 3 and 4.
CARBOHYDRATES | Starch: Starch Architecture and Structure 193
Reducing
A-chain B2-chain B3-chain B1-chain C-chain
end
Dp 1216
Figure 3 Chain categories of amylopectin. Adapted from Hizukuri S (1986) Polymodal distribution of the chain lengths of amylopectins, and its
significance. Carbohydrate Research 147: 342347.
Bfp BSmajor B2 B3
4
Carbohydrates (wt %)
0
0 20 40 60 80 100 120
Degree of polymerization (DP)
Figure 4 Internal unit chain profile of debranched f,b-limit dextrins from four different types of amylopectin, as obtained by high-performance
anion-exchange chromatography (HPAEC). Adapted from Bertoft E, Piyachomkwan K, Chatakanonda P, and Sriroth K (2008) Internal unit chain
composition in amylopectins. Carbohydrate Polymers 74: 527543, with permission.
The cluster structure of amylopectin is examined primarily based on the number of chains (2, 3, 4, or more) in the unit.
with the use of enzymatic methods. Clusters, or macro a-limit Adjacent building blocks within the cluster are connected by
dextrins, are isolated from amylopectin using limited hydroly- interblock chain segments, which are in the range of 58
sis with a-amylase. These clusters are defined as groups of short glucose residues. Computer simulation and crystalline
chains separated by internal chain segments with dp values stability studies have revealed that the CL between adjacent
< 9. This definition is somewhat arbitrary because the hydro- building blocks is one of the most critical parameters govern-
lysis pattern is dependent on the source of the a-amylase and ing the alignment of the external chains and the crystalline
its kinetics as well as on experimental conditions. Clusters packing. The number of building blocks within the cluster,
from different types of amylopectin differ with respect to size the interblock chain length (IB-CL), and the cluster size and
and CL and are made up of tightly branched units, known as the number of chains in the cluster and the length of the
building blocks. Building blocks are micro a-limit dextrins, external chains are all used to characterize different types of
which are released from clusters through extensive hydrolysis amylopectin. A schematic model illustrating the internal struc-
with a-amylase. The size of the building block is determined ture of amylopectin is provided in Figure 5. Cereal starches
194 CARBOHYDRATES | Starch: Starch Architecture and Structure
External chains
Interblock
Building blocks Inter-cluster chains length
chains (DP > 9) (IB-CL)
A-type
B-type
C-type
0 5 10 15 20 25 30 35 40
2q ()
Figure 7 Wide-angle x-ray scattering patterns of A-, B-, and C-type starches.
Amorphous
background
Crystalline
lamella (C)
Amorphous
lamella (A)
9 nm
Cluster
Starch granule Growth rings Blocklets
(1 100 mm) (120 400 nm) (20 100 nm)
Double helix
C 2.1 nm
5 6 nm
9 nm
A
3 4 nm
the daytime and amorphous rings being formed during the amylose and disordered amylopectin. A semicrystalline growth
night. However, whereas constant light treatment has been ring consists of a number of alternating crystalline and amor-
shown to affect ring formation in barley and wheat, it has no phous lamellae with a repeat distance of 8.511 nm. Amor-
effect on ring formation in potatoes. While this finding sug- phous lamellae are thought to contain amylose plus long
gests that a circadian rhythm related to biological mechanisms chains and branching points of amylopectin. Crystalline lamel-
influences ring formation, no definite explanation has yet been lae are formed primarily from the double helices of short
established for the occurrence of alternating zones of growth amylopectin chains. Each double helix consists of six anhydro-
rings in starch granules. glucopyranose units per turn with a pitch height of 2.1 nm2.
While the detailed structure of amorphous growth rings is The birefringence pattern of starch granules revealed under
still to be determined, such rings are thought to consist of polarized light microscopy is attributable to their
196 CARBOHYDRATES | Starch: Starch Architecture and Structure
(a) (b)
Figure 9 Bright-field microscopy (a) and polarized microscopy (b) of iodine-treated waxy potato starch granules.
semicrystalline nature, and the distinctive Maltese cross indi- biosynthetic enzymes, and the structurefunction relationship
cates that the ordered amylopectin chains in the crystalline of starch in a variety of applications.
lamellae are radially oriented (Figure 9).
Images obtained using high-resolution scanning electron
and atomic force microscopy indicate another level of organi-
zation called a blocklet, which appears to be present in both Exercises for Revision
semicrystalline and amorphous growth rings. Blocklets are
more or less spherical, with sizes varying from 20 to 100 nm, How does amylopectin build up the semicrystalline
depending on the location of the granule and the botanical lamellae?
source. Potato starch blocklets are generally larger than those Define the structural parameters of amylopectin.
in cereal starches. The presence of the larger blocklets, coupled Explain how a,b-limit dextrin has improved the unit chain
with its semicrystalline nature, is commonly linked to the analysis of amylopectin.
relative resistance to enzymatic hydrolysis of the potato starch How does the internal structure of amylopectin determine
granule. According to the blocklet concept, growth rings are the glucan chain alignment in crystalline lamellae?
organized in the form of blocklets that occur in a semi-
crystalline growth ring consisting of a stack of lamellae. How-
ever, the exact nature of both the blocklet and its relation with
the rest of the structural levels of a starch granule remains open Exercises for Readers to Explore the Topic Further
to speculation.
How can the validity of the proposed amylopectin models
be tested?
Does the nature of the branching pattern influence the
Conclusion polymorphic pattern?
Does amylose influence the assembly of crystalline
In starch granules, amylopectin chains are organized into a lamellae?
hierarchical structure. Although existing sophisticated imaging How does the molecular structure of amylopectin in the
and analytic techniques have enabled the observation of semicrystalline region of starch granules differ from the
different levels of order in starch granules, a substantial amylopectin in the amorphous region?
knowledge gap remains with respect to understanding how Do blocklets build up the super helical structure of starch
glucan chains are arranged in amylopectin and how the granules?
structural levels are interconnected. The crystalline lamellae
and unit CL distribution of amylopectin have been more
extensively analyzed than other structural features. However, See also: Carbohydrates: Starch: Analysis of Quality; Starch:
the organization of glucan chains cannot be explained based Chemistry; Starch: Synthesis; Genetics of Grains: Maize: Other
on the average composition of the unit chains. Detailed anal- Maize Mutants; Processing of Grains: Starch: Modification; Starch:
ysis of the internal amylopectin structure is therefore of Uses of Native Starch.
paramount importance for providing insight into the crystal-
line stability, the polymorphic pattern, and the architecture
of the building blocks, which are considered to be the funda-
mental structural unit in amylopectin. Enhanced knowledge Acknowledgment
of the granular architecture will provide additional opportuni-
ties for uncovering the processes that govern the initiation The authors thank Dr Eric Bertoft for his critical review of this
of starch synthesis, the control mechanism involved in article.
CARBOHYDRATES | Starch: Starch Architecture and Structure 197
Further Reading Nikuni Z (1978) Studies on starch granules. Starke 30: 105111.
OSullivan AC and Perez S (1999) The relationship between internal chain length
Bertoft E (2013) On the building block and backbone concepts of amylopectin structure. of amylopectin and crystallinity in starch. Biopolymers 50: 381390.
Cereal Chemistry 90: 294311. Perez S and Bertoft E (2010) The molecular structures of starch components and
Bertoft E, Piyachomkwan K, Chatakanonda P, and Sriroth K (2008) Internal unit chain their contribution to the architecture of starch granules: A comprehensive review.
composition in amylopectins. Carbohydrate Polymers 74: 527543. Starch 62: 389420.
Buleon A, Colonna P, Planchot V, and Ball S (1998) Starch granules: Structure and Robin JP, Mercier C, Charbonniere R, and Guilbot A (1974) Lintnerized starches.
biosynthesis. International Journal of Biological Macromolecules 23: 85112. Gel-filtration and enzymatic studies of insoluble residues from prolonged acid
Chauhan F and Seetharaman K (2013) On the organization of chains in amylopectin. treatment of potato starch. Cereal Chemistry 51: 389406.
Starch/Starke 65: 191199. Shannon JC and Garwood DI (2009) Genetics and physiology of starch development.
French D (1984) Organization of starch granules. In: Whistler RL, BeMiller JN, and In: Whistler RL and BeMiller JN (eds.) Starch: Chemistry and Technology, 3rd edn.,
Paschall EF (eds.) Starch: Chemistry and Technology, 2nd edn., pp. 183207. pp. 2586. San Diego, CA: Academic Press.
San Diego, CA: Academic Press. Thompson DB (2000) On the non-random nature of amylopectin branching.
Hanashiro I, Abe J, and Hizukuri S (1996) A periodic distribution of the chain length Carbohydrate Polymers 43: 223239.
of amylopectin as revealed by high-performance anion-exchange chromatography. Vamadevan V, Bertoft E, and Seetharaman K (2013) On the importance of organization
Carbohydrate Research 283: 151159. of glucan chains on thermal properties of starch. Carbohydrate Polymers
Hizukuri S (1986) Polymodal distribution of the chain lengths of amylopectins, and its 92: 16531659.
significance. Carbohydrate Research 147: 342347. Zobel HF (1988) Starch crystal transformations and their industrial importance. Starch
Imberty A, Chanzy H, Perez S, Buleon A, and Tran V (1988) The double-helical 40: 17.
nature of the crystalline part of A-starch. Journal of Molecular Biology
201: 365378.
Starch: Analysis of Quality
A Gunaratne and H Corke, The University of Hong Kong, Hong Kong, Peoples Republic of China
2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition, volume 3, pp. 202212, 2004, Elsevier Ltd with revisions made by the editor.
Topic Highlights will not survive in the competitive market. Methods that are
implemented for the analysis of starch functional properties
Starch as the main constituent of food grains and the reason should be simple and fast, technically feasible, nondestructive,
for many of the functional properties of grains. and able to provide reproducible results with high accuracy.
Starch occurs as granules in the endosperm of cereal grains; Although several methods have been developed recently to
granule shape is distinctive for the species. monitor some of the functional properties, individual methods
The functional properties of starch are affected by botanical are sometimes not capable of providing adequate data on all
source, structure and physicochemical properties, and pre- the dimensions of a functional property, especially in the
treatments during processing. determination of retrogradation and gelatinization. Therefore,
The two major compounds of starch are amylose (mainly it is often worth applying at least two methods to get comple-
linear polymers of a(14)-linked glucose) and amylopectin mentary data. This article aims to discuss some functional
(usually the major component, being highly branched properties of starch from the perspective of the applicability
polymers). of methods that are currently employed to determine those
Amylose forms a distinctive blue color complex with properties.
iodine. Better analysis is provided by high-performance
size-exclusion chromatography (HPSEC) and various
other methods.
Near-infrared (IR)-reflectance analysis can be used to deter- Properties, Functions, and Analysis of Starch
mine amylose content rapidly for mutant lines that differ Macromolecules
significantly in amylose:amylopectin ratio.
Amylose
Direct testing of starch function involves the pasting test
whereby viscosity changes are monitored for a starch sus- The two major compounds of starch are amylose and amylo-
pension heated in excess water, using equipment such as pectin. The two polymers are structurally different, the minor
the Viscoamylogram or the Rapid ViscoAnalyzer. component, which is linear, consisting mainly of a(14)-
Starch retrogradation is the key physical change associated linked glucose. However, in some starches, amylose chains
with bread staling. have shown a slight degree of branching. Incomplete conver-
sion to maltose by b-amylase suggests the presence of branched
points in the amylose polymer. The branched chains can also
Learning Objectives act similar to unbranched chains. Amylose isolated from tuber
and root starches such as potato and cassava has a larger
Understand the chemical properties of starch that affect its molecular weight than that isolated from cereal starches such
functional utilization. as maize and wheat. Although amylose is a linear molecule, its
Understand how and when to apply the range of testing conformation tends to change due to the large number of
procedures for starch chemistry and function. hydroxyl groups that can produce higher hydrogen bonding
capability with strong internal forces. The conformation of
amylose has been the subject of controversy and has been
Introduction shown to vary from helical to an interrupted helix, to a random
coil. In alkaline solutions (KOH) and dimethyl sulfoxide
Starch is the main polysaccharide produced in photosynthetic (DMSO), amylose probably has an expanded coil conforma-
tissues. Recently, there has been a growing demand for starch tion, while in water and neutral aqueous potassium chloride
in both food and nonfood industries because of its wide array solutions, it is a random coil with short, loose helical segments.
of applicability in diverse systems. Functionality is the key for Although helical conformation is common in amylose, double
the industrial utilization of starch. Functional properties can be helices form when different helices are packed together. The
defined as the characteristics that govern the behavior of a food helical arrangement of amylose forms a hydrophobic core
component during processing, storage, and preparation. Fac- inside the amylose molecule, which permits complexing with
tors such as botanical source, structure and physicochemical guest molecules such as lipids and iodine. The amount and
properties, and pretreatments of starch directly affect func- molecular characteristics of amylose have a significant influ-
tional properties of starch. Analysis of starch-related function- ence on starch functionality. Higher amylose content decreases
ality should provide a clear picture of the relevant property that the granular swelling power, whereas increasing amylose con-
will allow better use of starch in food and nonfood systems, for centration decreases the gel stickiness but increases the gel
example, to maintain consistent quality and functionality of firmness. Reassociation of amylose chains on cooling of a
starch-based products in every batch, otherwise the product starch paste shrink the starch gel resulting in water
Starch Granular size (mm) Granular shape Amylose (%) Swelling power at 95 C Solubility at 95 C Taste
Adapted from Collado LS and Corke H (2003) Starch properties and functionalities. In: Kaletunc G and Breslauer KJ (eds.) Characterization of Cereals and Flours, pp. 473506.
New York: Marcel Dekker.
accumulation on an aging gel, which decreases the storage Table 2 Amylose content in maize mutant starches as estimated with
stability and usually the quality of an affected starch-based size exclusion chromatography (SEC), Concanavalin (Con. A), differential
food product. scanning calorimetry (DSC), and iodine binding capacity (IBC)
Amylose (%)
Methods used in amylose estimation
Starches SEC Con. A DSC IBC
Conventional quantitative determination of amylose is based
on its ability to form a deep blue color complex with iodine. ae wx nd 7 nd 14
Amylose content of starch therefore can be quantified either by Du 27 45 55 45
spectrophotometric means or by potentiometric titration. su2 24 50 58 45
Table 1 shows amylose content of some common starches. du su2 34 58 66 60
However, interference from amylopectin and other intermedi- ae du 30 56 64 56
ate materials in the blue color formation reaction biases the Ae 54 63 60 63
estimation of actual content of amylose. Amylopectiniodine nd: not detected.
complex is not stable due to shorter unit chain length of Source: Planchot V, Gerard C, Bertoft E, and Colonna P (2001) An approach to
amylopectin, but long chains of amylopectin, e.g., long B structural analysis of granules using genetically modified starches. In: Barsby TL,
chain of waxy maize amylopectin can bind with iodine in a Donald AM, and Frazier PJ (eds.) Advances in Structure and Function, pp. 104128.
similar way to amylose. This would lead to an overestimation. UK: The Royal Society of Chemistry.
Despite shortcomings such as this, and the time-consuming
nature of the assay, the amyloseiodine reaction is still the
basis of the most widely used method to determine amylose component and the resulting supernatant after centrifugation
content of starch because of its accuracy, reproducibility, and contains only the amylose component. However, a compara-
easy operation. However, the excessive time needed limits its tive study of size exclusion chromatography (SEC), differential-
use in quality control applications. Instead of conventional scanning calorimetry, iodine-binding capacity (IB), and
thermal dissociation of amylose in DMSO, low-temperature concanavalin-A, where SEC was used as the reference method,
gelatinization in CaClB2B could be employed to reduce the has revealed that IB, differential-scanning calorimetry (DSC),
time requirement. High performance size exclusion chroma- and concanavalin-A resulted in an overestimation in the
tography (HPSEC), that has shown a good correlation with determination of amylose content of mutant maize starches
blue value measurements of amylose content, can be employed (Table 2). As other alternative methods to the conventional
to separate starch and de-branched starch. Ability to directly iodineamylose complex formation, spectroscopic methods
monitor the effect of de-branching on the molecular size have been developed recently. These techniques provide advan-
distribution of starch and high-molecular weight linear amy- tages such as ease of sample preparation, speed, and applica-
lose content from this technique provides the estimation of bility to online monitoring in the process control situation
long-chain amylopectin chains that contribute to the apparent compared with other techniques. Near-infrared (IR)-reflectance
amylose content. Recently another technique that is able to analysis has been successfully introduced in determining the
eliminate the effect of long-chain amylopectin in the estimation amylose content of waxy, normal, and high amylose maize,
of amylose content has been introduced. In this method, starch unground brown or milled rice, and whole grain maize sam-
completely dissolved in DMSO is subjected to de-fatting with ples. Raman spectroscopy on the other hand is becoming more
ethanol. The precipitate after de-fatting is dissolved in acetate/ popular as a quantitative analytical technique in the food
salt solution. A special reaction mixture (concanavalin-A) is industry and has shown potential applicability in the estima-
then added to the solution to precipitate the amylopectin tion of amylose content in maize starches.
200 CARBOHYDRATES | Starch: Analysis of Quality
Amylopectin and root starches show no sign of any pores on the granular
surface. However, pores are present on the surface of maize,
Amylopectin, the highly branched molecule, is usually the
sorghum, and millet starch granules. Pores on granule surfaces
major component in the starch granule with a(14)-linked
increase accessibility of a-amylase to the granule interior result-
glucose linear chains and a(16)-linked branch points. Crys-
ing in increased enzymatic hydrolysis. Microscopic analysis
talline domains of the starch granules are due to the clustered
provides information on surface features and size of starch
branches of amylopectin chains that are packed together,
granules.
whereas the free amylose, amylose complexed with lipids,
and branch points of the amylopectin are found in the amor-
phous region. Alternative arrangement of crystalline and amor-
Characterization and Analysis of Starch Thermal
phous region was proposed for the semicrystalline starch
Properties
granule. However, there is no clear demarcation between
amorphous and crystalline regions. Crystalline region is less Gelatinization
susceptible to enzymatic hydrolysis, water penetration, and
Gelatinization is one of the most important processes affecting
other chemical reactions than amorphous region. Amylopectin
starch. It occurs when starch is heated in excess water, when
has a lesser tendency to gelation, retrogradation, and syneresis
irreversible granular swelling, native crystallites melting, loss of
because of the branched structure. The amount of amylopectin
granular order, loss of birefringence, and starch solubilization
varies among different starches. Waxy varieties contain almost
take place. The extent of these changes depends on type of
100% amylopectin. The extent of functional characteristics of
starch, starch concentration, temperature, presence of other
starch (viscosity, gelatinization, solubility, texture, gel stability,
solutes, and shear applied during the gelatinization process.
retrogradation, shear resistance) are directly affected by the
As a result of the above changes during gelatinization, the
amylose/amylopectin ratio. Although amylopectin is the
starch paste develops viscosity, the basis of most technological
major component of the starch granule, there is no convenient
usefulness of starch as an ingredient in food and nonfood
method developed for direct estimation of amylopectin and
applications. In the majority of food applications, starch func-
studies on amylopectin are dependent on development of
tions (e.g., imparts texture on consumption) in a gelatinized
enzymatic and instrumental methods. Average structural prop-
form. Several concepts have been proposed to explain the
erties of whole molecule and impact of internal structure
gelatinization process. Granular swelling acts primarily as a
of clusters on crystallization have been studied by means of
driver to destabilize the starch crystallites. Recently, a model
enzymatic and SEC. It has been shown that the combined
was proposed to explain the phenomena involved in gelatini-
application of preparative and analytical size exclusion chro-
zation and hydration of starch based on the side-chain liquid
matography with multiple detection, precipitation techniques,
crystalline. In this model, lamellae in starch are considered in
and enzymatic de-branching provides the most detailed
terms of three components (1) backbone, (2) side-chain, and
analytical insight on the microstructural properties of
(3) double helices. The degree of mobility of these three com-
amylopectin.
ponents, coupled with the helixcoil transition, give starch its
distinctive properties in gelatinization. Gelatinization is influ-
enced by many factors such as, botanical source of starch, water
Function Related to Morphological Characteristics content, and added solutes. Several methods have been pro-
of Starch Granules posed to study the effect of water content on thermal stability.
Depending on the water content available during gelatiniza-
Granules are the basic physical structural unit of starch. The tion, changes in endothermic transition have been detected in
size, shape, and other morphological characters of starch gran- terms of number and the position of the differential scanning
ules are extremely diverse (Table 1). The shape of the starch calorimetric endotherms. Only one endotherm is detectable at
granule can be spherical, disk, polygonal, or elongated. In higher levels of moisture content while two endothermic tran-
general, cereal starches have smaller granules compared to sitions are reported at the low-moisture-content level for some
tuber and root starches. Most tuber and root starch granules starches where low temperature and higher temperature endo-
have a simple size distribution. In some starches, such as barley therms are designated as G1 and M1 respectively (Figure 1). In
and wheat, two different granular size populations exist, i.e., a addition to two endothermic transactions at lower moisture
bimodal distribution. Granular size and morphology have content, peaks are shifted to higher temperature, indicating
recently received more attention, e.g., the size of granules is need for higher thermal energy to melt the starch crystallites
important in determining taste and mouth feel of some starch- at low moisture content. If there is sufficient water available in
based fat mimetics. Some specific industrial applications of gelatinization, then crystallites melt cooperatively, resulting
starch are related to size and the size distribution of starch in a single endotherm. Two endothermic transition peaks in
granules. For example, the small size of rice starch granules insufficient water could be attributed to the differences in the
makes it highly suitable for laundry use. Most of the physico- stability of starch crystallites, which melt over different tem-
chemical properties of native starch are highly correlated with peratures, less stable crystallites melt first and the others are
the size of the starch granules. Larger size granules tend to swell next causing the second endotherm. Redistribution of water
more than smaller granules, whereas smaller size granules are between crystalline domain and gelatinized starch could also
more susceptible to enzymatic hydrolysis. The outer surface of result in two endothermic transitions, in which the first endo-
starch granules also plays a key role in many applications of therm could be attributed to melting of starch crystals in excess
starch. Electron microscopic images have revealed that tuber water. Much research has been conducted to investigate the
CARBOHYDRATES | Starch: Analysis of Quality 201
Temperature (C)
To Tc
Hcx
M2
Hgel
G M1
Tm
Figure 1 Schematic representation of DSC thermograms obtained for wheat starch heating. (Eliasson A-C (2003) Utilization of thermal properties for
understanding baking and salting processes. In: Kaletunc G and Breslauer KJ (eds.) Characterization of Cereals and Flours, pp. 65115. New York:
Marcel Dekker.)
To onset temperature ( C); Tp peak temperature ( C); Tc conclusion temperature ( C). DH enthalpy (J g 1).
Sources: Gunaratne A and Hoover R (2002) Effect of heat-moisture treatment on the structure and physicochemical properties of tuber and root starches. Carbohydrate Polymers 49:
425437; and Hoover R, Vasanthan T, Senanayake NJ, and Martin AM (1994) The effects of defatting and heat-moisture treatment on the retrogradation of starch gels from wheat, oat,
potato, and lentil. Carbohydrate Research 261: 1324.
influence of added solutes such as sugars, emulsifiers, and resonance, small angle X-ray scattering, and small angle neu-
electrolytes (such as sodium chloride), which are widely tron scattering, Brabender viscoamylography (BV), and Rapid
applied in food industry on starch behavior. Sugars have Visco Analysis. DSC, the most widely used method, measures
shown to increase the gelatinization temperature but not to the dissociation parameters, TBoB (onset), TBpB (peak), TBcB
affect endothermic heat absorption. Quality of various food (conclusion), and DH (endothermic heat absorption), of
products, bread and cakes, thickening and gelling of sauce, pie starch crystallites in gelatinization (Figure 1). DH is the area
filling, and extrusion of cereals are highly dependent on below the transition endotherm. Gelatinization parameters of
starchwater interaction. Increased gelatinization, when sugars starches from different botanical sources measured by DSC are
are added to starch paste could be due to decrease of water presented in Table 3. In addition, DSC can be used to study the
availability for the gelatinization, binding water to sugar mol- glass transition temperatures of various starchy food products.
ecules. However, it was reported that sugar acts as an antiplas- Gelatinization temperatures are influenced more by the gran-
ticizer rather than gelatinizer through the effect on water- ular architecture than the amyloseamylopectin ratio and
binding capability. Increased sugar concentration decreases depend on the degree of starch crystallite perfection, whereas
the plasticization effect requiring more heat energy to achieve endothermic heat absorption reflects the quality and amount
the gelatinization. The extent of the influence of sugars on of starch crystallites. Analysis of gelatinization parameters of
gelatinization, swelling, and viscosity differs depending on starch using DSC was first reported in 1971. Gelatinization
the type of sugars and salts used when compared on a molar parameters measured by DSC can be affected by sample prep-
basis. Monosaccharides are less effective than disaccharides aration, baseline application, pan selection, method of adding
except maltose; among disaccharides, sucrose is more effective. water, sample mass, heating rate, and moisture equilibration
time. Thermal behavior of amyloselipid complex as well as
Methods in the analysis of starch gelatinization formation of amyloselipid inclusion with various molecules
There are several methods to analyze the gelatinization process, such as fatty acids and emulsifiers can also be detected by DSC.
such as DSC, Kofler hot-stage microscopy, light microscopy, Characterization of amyloselipid complex in cereal starches
electron microscopy, X-ray crystallography, enzymatic analysis, and influence of amyloselipid complex on starch properties
nuclear magnetic resonance (NMR), pulsed nuclear magnetic are well documented. Some of the internal lipids in cereal
202 CARBOHYDRATES | Starch: Analysis of Quality
starch is thought to be complexed with amylose to form the method of observation, granular type, and heterogeneity. Past-
amyloselipid complex, which is dissociated at higher temper- ing is defined as the phenomena following gelatinization in
ature (M2 endotherm in Figure 1) than that of normal starch the dissolution of starch, involving granular swelling, exuda-
crystallites (Figure 1 and Table 4). Although DSC is widely tion of molecular components from the granule, and eventu-
used to study gelatinization, the small sample size used can ally total disruption of the granules. The schematic
limit its real applications. In a comparative study of maize representation of granular changes and viscosity development
starch gelatinization with DSC and NMR, it was reported that that occurs during pasting is shown in Figure 2.
NMR could provide much better performance in the analysis of
gelatinization for larger samples than DSC. Gelatinization tem- Measurement of pasting properties
perature of Peruvian carrot, potato, and maize starch measured The BV is the most widely established method for determining
by Rapid Visco Analyzer (RVA) and BV, and DSC have shown pasting properties, although it has some methodological and
different values for different methods. This indicates the need geometrical shortcomings. This apparatus measures the devel-
for multiple analytical techniques for a meaningful under- opment of viscosity when a starchwater suspension is sub-
standing of gelatinization along with careful specification jected to a programmed heating and cooling cycle under a
(and preferably standardization) of experimental conditions shear force. The temperature at which viscosity begins to
used. Table 5 shows the differences in gelatinization parame- increase is termed pasting temperature. With further heating,
ters measured by different techniques for the same starch. granular swelling increases the viscosity to reach a peak. Fur-
ther heating at elevated temperature under shear force tends to
disintegrate the swollen granules resulting in decrease of vis-
Pasting properties cosity. Upon cooling the starch paste in the next stage, there is a
Starch heated in excess water undergoes various changes as a tendency, mainly due to amylose chain reassociation, to
result of heat and moisture transfer. Gelatinization and pasting increase viscosity. Therefore, a typical pasting profile exhibits
occur in the same system and have often been used to describe
all the changes that occur. Gelatinization may be used to refer
to early changes whereas pasting includes later changes. Starch
gelatinization is defined as the collapse of the starch granule
manifested in irreversible melting, loss of birefringence, and
starch solubilization. The point of gelatinization, and the range
over which it occurs is governed by starch concentration,
Viscosity
Table 5 Gelatinization temperature range of some starches as measured with different techniques: DSC, NMR, BV, and RVA
three distinct viscosity developments, peak viscosity, hot paste (starches >55% amylose). The above four different categories
viscosity, and cold paste viscosity (Figure 3). Pasting curves of starch exhibit the following pasting properties respectively:
vary according to botanical source of the starch, starch concen- high peak followed by rapid and major thinning during cook-
tration, and the programmed heatingcooling cycle chosen. ing; lower peak and much less thinning; no peak but maintain
Among the native starches, potato shows the highest peak high viscosity during cooking; and no swelling to gave a vis-
viscosity, and generally, low peak viscosity and higher setbacks cous paste at normal concentration.
can be detected in normal cereal starches compared to tuber Because of some technical shortcomings with the BV, such
and root starches. Waxy cereal starches, on the other hand, as large sample size requirement and inability to program the
behave more like the tuber and root starches (Figure 4). temperature profiles, a more recent equipment, the RVA, has
According to the characteristics of pasting curves, starch has become popular for analyzing pasting properties. The RVA
been categorized into four groups as, high swelling starch differs from the Brabender instrument due to rapid heating
(potato, cassava, waxy cereals, ionic starch derivatives), mod- rate and stronger mixing action. However, controlled heating
erate swelling starch (normal cereal starch), restricted swelling rate to 1.5 C min 1 in RVA provides similar results to those
starch (cross-linked starch), and highly restricted starch observed in the BV. Comparative study of pasting properties of
cassava, sago (sagu), and arrowroot using Brabender and RVA
has shown that both techniques provide similar pasting pat-
terns (Table 6). Pasting curves of some of the starches deter-
250 100
mined by BV and RVA is presented in Figures 4 and 5.
Temperature Cool paste viscosity
profile
200
Peak viscosity
80
Swelling and Solubility
Peak time
Native starch granules are insoluble in water. Although small
Setback amounts of water can be absorbed at room temperature, gran-
Temperature (C)
Breakdown
Viscosity (RVU)
1000
starch to be obtained. Swelling volume is a modified version in
Waxy maize
which programmed shaking of starch suspension is used
500
instead of stirring, and measurement is of the volume
Wheat expanded due to granular swelling. Swelling factor is the ratio
0
of volume of swollen starch granule to the volume of the dry
50 95 95 95 50 50 50 C
starch. Swelling factor measures only the intragranular water
Heat Hold Cool Hold
content. Calculation of the swelling factor is based on starch
Temperature weight adjusted to 10% moisture assuming the density of
Figure 4 Brabender Viscoamylogram of starches from different 1.4 mg ml 1. The basic principle of this method is the mea-
botanical sources. surement of the volume of water absorbed by starch granules
204 CARBOHYDRATES | Starch: Analysis of Quality
Table 6 Pasting properties of cassava, sago, and arrowroot as determined by BV and RVA
GT gelatinization temperature range; PV peak viscosity; FV final viscosity; V viscosity; BD breakdown; SB setback.
a
Starch concentration 4.3%.
b
Starch concentration 8%.
Source: Perez EE, Breene WM, and Bahnassey YA (1998) Gelatinization profiles of cassava, sagu, and arrowroot native starches as measured with different thermal and mechanical
methods. Starch/Starke 50: 1416.
Sweet potato has been found that retrogradation is the key physical change
associated with bread staling. Retrogradation sometimes aids
300 Maize 60 processing in some food products such as hardening of par-
boiled rice and to improve textural characteristics of certain
types of noodles.
150 40
Determination of starch retrogradation
Rice Several methods have been developed to determine starch
Wheat
0 retrogradation, because of its great influence on industry-
0 5 10 15 20 25 based food and nonfood products, such as DSC, X-ray analysis,
Time (min) rheological methods, and spectroscopic methods.
Heating steps
Bread crumb
1
2
Wheat starch
Endothermic heat flow
Amylose
0.05 mcal s1
To Tc transition temperature range; DH enthalpy; AMP amylopectin. Sources: Zobel (1988b) and Gunaratne A and Hoover R (2002) Effect of heat-moisture
a
Each number indicates the temperature for one day of storage; 6/40 means one day at treatment on the structure and physicochemical properties of tuber and root starches.
60 C, followed by one day at 40 C. Carbohydrate Polymers 49: 425437.
Source: Silverio J, Fredriksson H, Andersson R, Eliasson A-C, and Aman P (2000) The
effect of temperature cycling on the amylopectin retrogradation of starches with different
amylopectin unit-chain length distribution. Carbohydrate Polymers 42: 175184. crystallinity is found mainly in cereal starches. Most tuber and
root starches exhibit typical B type X-ray pattern and C type is
sample size and inability to determine retrogradation in dilute intermediate between A and B types, shown in legumes, cas-
starch paste. More often DSC coupled with X-ray analysis pro- sava, and some varieties of sweet potato (Table 8). X-ray anal-
vides better performance in characterizing retrogradation. ysis of retrograded starch has shown that aging gels form B-type
crystals irrespective of the native starch. Some research using
wide range and small angle X-ray diffraction, shows that crys-
X-ray analysis talline formation in starch paste occurred primarily due to
X-ray analysis can be applied to detect the presence and nature amylopectin aggregation, whereas amylose provides a template
of crystallinity in native starch granules as well as crystals effect and highly ordered amylose aggregation does not neces-
formed in aging starch gels. In native starch, the crystallinity sarily possess a crystalline nature but accelerates the amylopec-
is due to the amylopectin components and the crystal domains tin reassociation. Recently, more information on bread staling
are constructed mainly of A chains and outer B chains of has been revealed by X-ray analysis. Aggregation of starch poly-
amylopectin. Native starches can be categorized into three mers that formed crystals in bread staling was shown from
groups according to their X-ray diffraction pattern. The A type X-ray analysis; however, NMR and FTIR techniques are better
206 CARBOHYDRATES | Starch: Analysis of Quality
at detecting minor differences of starch polymer aggregation Describe the process of starch pasting and the methods for
than X-ray analysis. its analysis. How does this property affect the uses of starch
in the food industry?
Rheological methods What is meant by starch retrogradation? Explain the result-
Measurement of pasting parameters, analysis of textural proper- ing beneficial and undesirable consequences for food
ties of starch paste using texture analyzer, and small deforma- products.
tion dynamic techniques oscillatory rheometry can be applied What are the events of the starch-aging process? How do
to the behavior of starch. Pasting behavior of starch indicates the these phenomena affect the functional use of starch in food
trend of starch retrogradation. In a typical pasting profile, the and non-food uses?
magnitude of setback reflects the tendency of mainly amylose
polymer reassociation in a starch paste. Several investigations Exercises for Readers to Explore the Topic Further
have been performed to study the properties of aging breads
using viscometry and have observed a tendency of decreased Explore the range of mutant types of grain species for which
peak viscosity when bread staling progresses. Recently there is a wide differences in amylose:amylopectin ratio have been
growing interest, especially in industry, in use of texture ana- developed. What are the resulting differences in functional
lyzers that can directly and rapidly measure the textural proper- properties and thus in applications in utilization?
ties such as hardness, stickiness, cohesiveness, adhesiveness, Starch is insoluble in water. Do you agree with this state-
fracturability of starch paste (Table 9). Long-term reliability ment? Under what circumstances and for which forms of
and accuracy, direct measurement, time saving, and technical starch is the statement true and untrue? Explain how the
feasibility of the texture analyzer increases its attractiveness solubility (or not) of starch is critical to its utilization.
among food technologists. The main issue which requires con- Explore how the presence of starch-degrading enzymes
tinuing work is that of instrumental to sensory correspondence, (such as amylase enzymes) increases starch solubility and
i.e., whether the texture analysis profile is adequately predictive thus its functional properties.
of the sensory traits important to the consumer. What lessons might be learned from research into the chain
Many additional methods can be employed to detect retro- lengths and degree of branching of the various starch
gradation. No individual method is able to fully characterize molecules.
all the events of the starch-aging process. DSC measures only
the transition of heat energy in the melting of crystal aggre-
gates, X-ray measures the presence and characteristics of retro- See also: Carbohydrates: Carbohydrate Metabolism; Glycemic
graded starch crystals, and texture analysis provides textural Index, Glycemic Load, and Their Health Benefits; Resistant Starch and
parameters for the aged starch gel. Therefore, it is clear that a Health; Starch: Chemistry; Starch: Starch Architecture and Structure;
multiple analytical technique approach for the advance detec- Starch: Synthesis; Food-quality Testing: Noodles: Testing for
tion of retrogradation behavior is necessary. For example, a Quality; Pasta: Quality Testing Methods; Rice: Eating Quality;
study of bread staling with multiple techniques, DSC, FTIR, Genetics of Grains: Maize: Other Maize Mutants; Maize: Quality
and NIR, led to greater understanding of the process than Protein Maize; Grain Composition and Analysis: Standardized Test
would be possible with each individual technique used alone. Methods for Grains and Grain-Based Products; The Composition of
Food Grains and Grain-Based Products; Processing of Grains:
Starch: Modification; Starch: Uses of Native Starch; Wheat
Exercises for Revision Processing: Baked Product Staling: Mechanisms, Determinations,
and Anti-staling Strategies; The Basics: Grain and Plant Morphology
of Cereals and how characters can be used to identify varieties.
What are the chemical and functional differences between
amylose and amylopectin?
Explain the terms waxy and non-waxy in terms of starch
composition and function. Further Reading
Describe the range of methods available to determine amy- Collado LS and Corke H (2003) Starch properties and functionalities. In: Kaletunc G and
lose:amylopectin ratio. What situations might suit the use Breslauer KJ (eds.) Characterization of Cereals and Flours, pp. 473506. New York:
of one of these methods versus another? Marcel Dekker.
CARBOHYDRATES | Starch: Analysis of Quality 207
Daniel JR and Whistler RL (1985) Principal changes in starch during food processing. Phillips DL, Xing J, Liu H, Pan DH, and Corke H (1999) Potential use of Raman
In: Richardson TR and Finley JW (eds.) Chemical Changes in Food During spectroscopy for the determination of amylose content in maize starch. Cereal
Processing, pp. 305326. Westport, CT: AVI Publishing Co. Chemistry 76: 821823.
Eliasson A-C (2003) Utilization of thermal properties for understanding baking and Planchot V, Gerard C, Bertoft E, and Colonna P (2001) An approach to structural
salting processes. In: Kaletunc G and Breslauer KJ (eds.) Characterization of Cereals analysis of granules using genetically modified starches. In: Barsby TL, Donald AM,
and Flours, pp. 65115. New York: Marcel Dekker. and Frazier PJ (eds.) Advances in Structure and Function, pp. 104128. UK: The
Fredriksson H, Silverio J, Andersson R, Eliasson A-C, and Aman P (1998) The influence Royal Society of Chemistry.
of amylose and amylopectin characteristics on gelatinization and retrogradation Praznik W, Huber A, Watzinger S, and Beck RHF (1994) Molecular characteristics of
properties of different starches. Carbohydrate Polymers 35: 119134. high amylose starches. Starch/Starke 46: 8894.
Gonera A and Cornillon P (2002) Gelatinization of starch/gum/sugar systems studied by Silverio J, Fredriksson H, Andersson R, Eliasson A-C, and Aman P (2000) The effect of
using DSC, NMR, and CSLM. Starch/Starke 54: 508516. temperature cycling on the amylopectin retrogradation of starches with different
Gunaratne A and Hoover R (2002) Effect of heat-moisture treatment on the structure and amylopectin unit-chain length distribution. Carbohydrate Polymers 42: 175184.
physicochemical properties of tuber and root starches. Carbohydrate Polymers Tester RF and Morrison WR (1990) Swelling and gelatinization of cereal starches 1.
49: 425437. Effects of amylopectin, amylose and lipids. Cereal Chemistry 67: 551557.
Hoover R, Vasanthan T, Senanayake NJ, and Martin AM (1994) The effects of defatting Yu L and Christie G (2001) Measurement of starch thermal transitions using differential
and heat-moisture treatment on the retrogradation of starch gels from wheat, oat, scanning calorimetry. Carbohydrate Polymers 46: 179184.
potato, and lentil. Carbohydrate Research 261: 1324. Zobel HF (1988) Molecules to granules: a comprehensive starch review. Starch/Starke
Jane J, Kasemsuwan T, Leas S, Zobel HF, and Robyt JF (1994) Anthology of starch 40: 4450.
granules morphology by scanning electron microscope. Starch/Starke 46: 121.
Karim AA, Norziah MH, and Seow CC (2000) Methods for the study of starch
retrogradation. Food Chemistry 71: 936.
Liu H, Ramsden L, and Corke H (1999) Physical properties and enzymatic digestibility
of hydroxypropylated ae, wx, and normal maize starch. Carbohydrate Polymers Relevant Website
40: 175182.
Perez EE, Breene WM, and Bahnassey YA (1998) Gelatinization profiles of cassava, starch.dk http://www.starch.dk Website of the International Starch Institute, Denmark,
sagu, and arrowroot native starches as measured with different thermal and with useful links to other starch-related sites.
mechanical methods. Starch/Starke 50: 1416.
Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides
GB Fincher, Australian Research Council Centre of Excellence in Plant Cell Walls, University of Adelaide, Glen Osmond, SA, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights that are eventually found in shoots and roots. At the interface
of the embryo and the starchy endosperm lies the scutellum,
Structure and function of cell wall polysaccharides. which delivers nutrients to the developing endosperm and
Fine structure of wall polysaccharides and how these vary in plays a dual role during germination; in the first instance, the
different tissues and at different stages of development. scutellar epithelial layer secretes degradative enzymes into the
How fine structure affects physicochemical properties, proximal region of the starchy endosperm, but later, it trans-
including viscosity, solution behavior, and gelation. locates digestion products of the endosperm reserves into the
Genes involved in the biosynthesis, remodeling, and deg- developing embryo.
radation of cell wall polysaccharides. The cell wall compositions and organization of the cell
The importance of solubility and viscosity in cereal proces- types found in the tissues of the grain reflect these different
sing and human health. functions and, in turn, determine the nutritional impact and
The benefits of dietary fiber from cereal grains in reducing behavior of various grain parts in technical operations during
the risk of serious human diseases, including type II diabe- grain utilization and processing. Together, the nonstarchy
tes, colorectal cancer, and cardiovascular disease. polysaccharides usually constitute <10% by weight of the
grain (Table 1) but can nevertheless be key determinants of
grain quality. Although the precise physical relationships
Learning Objectives between individual nonstarchy polysaccharides and other
wall components have not been described, it is generally con-
To know how chemical structure affects solution properties. sidered that in the wall, microfibrils of cellulose are embedded
To understand how cell wall polysaccharides contribute to in a matrix phase that consists mainly of noncellulosic poly-
dietary fiber. saccharides and smaller amounts of protein. Lignification of
To know the structurefunction relationships in cereal pro- walls in the outer layers of the grain is observed, but little or no
cessing and nutrition. lignification is found in most cells of the endosperm and
embryo. Wall integrity is maintained predominantly through
extensive noncovalent interactions, especially hydrogen bond-
Introduction ing, between the matrix phase and the microfibrillar constitu-
ents. In the walls of some grain tissues, covalent associations
The nonstarchy polysaccharides of cereal grains include cellu- between heteroxylans, lignin, and proteins are present. Indeed,
lose, (1 ! 3, 1 ! 4)-b-D-glucans, heteroxylans (arabinoxylans), there have been several recent reports of previously unrecog-
glucomannans, xyloglucans, pectic polysaccharides, callose, nized covalent linkages between noncellulosic polysaccharides
fructans, and arabinogalactan proteins (AGPs). With the excep- of plant cell walls more generally. The extent of covalent asso-
tion of the fructans and the AGPs, all are key components of ciations between components also varies with the wall type
the walls of cells in the various tissues that comprise the grain. and genotype. The relative ease of extraction of noncellulosic
In this article, the chemistry and physicochemical properties of polysaccharides from the different wall types in cereal grains is
nonstarchy polysaccharides from cereal grains, which are strongly influenced by both noncovalent and covalent interac-
important in human and animal nutrition and which have tions. Cell walls have been isolated and analyzed from some
an impact in grain utilization and in cereal technology, will cereal grains, in particular aleurone and starchy endosperm
be described. walls from wheat and barley (Table 1).
The cells of grain tissues have diverse functions during Noncellulosic polysaccharides, especially heteroxylans and
development, during dormancy, and after germination. The (1 ! 3, 1 ! 4)-b-D-glucans, constitute a relatively high propor-
pericarp and seed coat tissues are concerned with the protec- tion of the walls of the aleurone and starchy endosperm and
tion of the seed during development and dormancy. However, probably also of the scutellum. Cellulose contents are corre-
at grain maturity, these tissues are nonliving and consist almost spondingly lower (Table 2). The generally low cellulose con-
entirely of cell wall residues. The nucellar tissue between the tent of these walls, together with the fact that the walls of the
seed coat and the aleurone surface is involved in the transfer of endosperm contain no lignin, might be related functionally to
nutrients to the developing grain but collapses at maturity to a limited requirement for structural rigidity of walls in central
leave compressed cell wall remnants in the mature grain. Cells regions of the grain and to a requirement for rapid depolymer-
of the starchy endosperm are nonliving in the mature grain, ization of wall components following germination of the
but are packed with starch and storage protein and, in most grain. There is some indirect information on the composition
cereal species, have relatively thin cell walls. In contrast, the of starchy endosperm walls from rice, which are extremely thin
thick-walled, nucleated, aleurone cells are living at grain and appear to consist mostly of cellulose and heteroxylans. The
maturity and are packed with membrane fragments, protein (1 ! 3, 1 ! 4)-b-D-glucan levels appear to be low, and varying
bodies, and lipid droplets. The living embryonic tissues of the amounts of pectin, xyloglucan, and glucomannan have been
grain are at least partly differentiated into functional cell types reported in rice endosperm walls.
Component
(1 ! 3, 1 ! 4)-
Cereal Origin of cell wall Cellulose Glucomannan b-D-Glucan Heteroxylan Pectin Lignin Proteina
Table 2 Content of (1 ! 3, 1 ! 4)-b-D-glucan and heteroxylan in whole cereal grains (gram per 100 g DW)
(1 ! 3, 1 ! 4)-b-D-Glucan Arabinoxylan
?, data unavailable.
In walls of cells in the multiple tissue layers of the mater- polymerization of up to 6000 in primary walls and 14 000 in
nally derived, pericarpseed coat, which provides protection secondary walls. The molecular chains have an extended,
for the embryo and endosperm and which is not mobilized ribbonlike conformation that allows parallel packing of chains
during germination, cellulose and lignin contents are much into three-dimensional, fibrillar aggregates stabilized by exten-
higher and the concentrations of noncellulosic polysaccharides sive intermolecular hydrogen bonding and van der Waals
are correspondingly lower. interactions (Figure 1(b)). These aggregates, termed microfi-
brils, are easily visible under the electron microscope and may
reach diameters of 3 nm in primary walls and 510 nm in
secondary walls. Within microfibrils, individual cellulose mol-
Cellulose ecules are mostly packed into parallel, highly ordered, crystal-
line arrays, although there are usually regions where molecular
Cellulose is a component of all cell walls in cereal grains. alignment is less ordered. In particular, the surface molecules
However, its concentration varies widely. Cellulose is usually may have a different conformational and hydrogen-bonding
a minor constituent of the primary walls of starchy endosperm arrangement compared with molecules within the more crys-
and aleurone cells of cereal grains, but its content is much talline interior. Long-held views that the cellulosic microfibrils
higher in secondary walls of the pericarp and seed coat cells of higher plant walls contain about 36 individual cellulose
(Table 1). It is a linear homopolymer composed of molecules have recently been challenged, and it appears likely
b-D-glucopyranosyl residues, all of which are (1 ! 4)-linked that the number of individual cellulosic chains per microfibril
(Figure 1(a)). Individual molecules have a degree of is considerably lower than originally believed.
210 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides
Figure 1 Cellulose structure. (a) Portion of a cellulose molecule. (b) Crystal structure of cellulose I: (i) projection down the fiber axis c showing the
layers (sheets) of cellulose chains hydrogen-bonded in the ac plane but lacking intersheet bonding. The intersheet bonding involves van der Waals
interactions between the hydrophobic faces of the glucose units. (ii) View of a layer approximately perpendicular to the ac plane showing the two
intramolecular hydrogen bonds in the direction of the fiber axis c and the interchain hydrogen bonds. Reproduced from Kroon-Batenburg LMJ and Kroon
J (1995) The crystal and molecular structures of cellulose. Carbohydrates in Europe 12: 1519, with permission.
Cellulose microfibrils may also associate with water and Iceland moss (Cetraria islandica). There have been recent reports
matrix polysaccharides such as the (1 ! 3, 1 ! 4)-b-D-glucans, of the presence of (1! 3, 1 ! 4)-b-D-glucans in the walls of the
heteroxylans (arabinoxylans), and glucomannans. Cellulose is horsetail fern (Equisetum spp.), in some bryophytes, various
insoluble in water but swells in concentrated sodium hydroxide algal species, and the walls of certain fungal pathogens.
solutions and may be brought into solution by powerful
hydrogen bond-breaking reagents such as N-methylmorpholine
N-oxide. Thermal and shear treatments, followed by alkaline
Structure
peroxidation and shearing to remove lignin, have been used to
convert cellulose-rich maize bran (Table 1) to a cellulosic gel for (1 ! 3, 1 ! 4)-b-D-Glucans are linear, unbranched polysaccha-
use as a dietary fiber supplement. rides in which b-D-glucopyranosyl monomers are polymerized
through both (1 ! 4)- and (1 ! 3)-linkages (Figure 2(a)). The
ratio of (1 ! 4) to (1 ! 3)-linkages is generally fairly constant,
and in the range 2.22.6:1, although in the (1 ! 3, 1 ! 4)-b-D-
(1 ! 3, 1 ! 4)-b-D-Glucans glucan from sorghum endosperm, the ratio is 1.15:1. The two
types of linkages are not arranged in regular, repeating
(1! 3, 1 ! 4)-b-D-Glucans, also referred to as mixed-linkage or sequences. Single (1 ! 3)-linkages are usually separated by
cereal b-glucans, occur almost exclusively in members of the two or more (1 ! 4)-linkages. Regions of two or three adjacent
monocotyledon family Poaceae, to which the cereals and grasses (1 ! 4)-linkages predominate, but there is no regularity in the
belong, and in related families of the order Poales (sensu stricto). arrangement of these units. Thus, cereal (1 ! 3, 1 ! 4)-b-D-
They are important constituents of the walls of the starchy glucans may be considered as (1 ! 3)-b-linked copolymers of
endosperm and aleurone cells of many cereal grains, where cellotriosyl (G4G4GRed), cellotetraosyl (G4G4G4GRed) units,
they can account for up to 70% by weight of the walls (Table 1). and longer (1 ! 4)-b-D-oligoglucosyl units (Figure 2(b)). The
The concentration of (1! 3, 1 ! 4)-b-D-glucans in cereal grains ratio of tri- to tetrasaccharide units varies between cereal spe-
varies across the grain and depends on the genotype, the posi- cies, as follows: wheat (3.04.5:1), barley (2.93.4:1), rye
tion of the grain on the spike and environmental factors such as (2.7:1), and oats (1.82.3:1). The ratios vary according to
planting location, climatic conditions during grain develop- the temperature and conditions of extraction. However,
ment, and soil nitrogen. Barley, oat, and rye grains are rich up to 10% of the chain may consist of longer stretches of
sources of (1! 3, 1 ! 4)-b-D-glucans, whereas wheat, rice, and 520 adjacent (1 ! 4)-linkages, and at low frequency, single
maize have lower concentrations of the polysaccharide (1 ! 4)-linkages are detected. An important point related to
(Table 2). The (1! 3, 1 ! 4)-b-D-glucans are relatively minor (1 ! 3, 1 ! 4)-b-D-glucan structure is that the single (1 ! 3)-
components of walls in vegetative tissues of cereals and grasses. b-D-glucosyl residues will introduce molecular kinks into the
A structurally related (1! 3, 1 ! 4)-b-D-glucan, lichenin, is main chain of the polysaccharide and thereby interrupt what
found in the walls of the fungal component of the lichen, would otherwise be an extended, straight cellulose molecule
CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides 211
Figure 2 (a) Portion of a (1 ! 3, 1 ! 4)-b-D-glucan molecule. (b) Distribution of linkages in a (1 ! 3, 1 ! 4)-b-D-glucan. (G, b-glucosyl unit);
3 (1 ! 3)-linkage, 4 (1 ! 4)-linkage; red, reducing end, arrows, site of hydrolysis by (1 ! 3, 1 ! 4)-b-D-glucan endohydrolase (EC 3.2.1.73).
(c) Perspective drawings of computer-generated, instantaneous conformations of b-glucans: (i) (1 ! 4)-b-D-glucan (cellulose), (ii) (1 ! 3)-
b-D-glucan (callose), (iii) (1 ! 3, 1 ! 4)-b-D-glucan, () (1 ! 4)-linked residues, and () (1 ! 3)-linked residue. Reproduced from Fincher GB
and Stone BA (1986) Cell walls and their components in cereal grain technology. Advances in Cereal Science and Technology 8: 207295, with
permission.
(Figure 2(c) (i) cf. Figure 2(c) (iii)). First-order Markov chain
analyses have been used to show that the DP3 and DP4 units ..G 3 G 4 G 4 G 3 G 4 G 4 G 4 G 3 G 4 G 4 G 3 G ..
of the water-soluble barley (1 ! 3, 1 ! 4)-b-D-glucan are
arranged at random (Figure 3), which means that the molec- 1 p
ular kinks imposed by (1 ! 3)-b-D-glucosyl residues in the
main chain of the polysaccharide will also be irregularly
p Cellotriosyl Cellotetraosyl 1 q
spaced.
q
Solubility r=p+q
The extractability of (1 ! 3, 1 ! 4)-b-D-glucans from walls of
cereal grains is partly a function of their degree of self- r = autocorrelation (interdependence between
association and their association with other wall polysaccha- DP3 and DP4 units)
rides and proteins. In addition, extractability depends on the Figure 3 Determination of the arrangement of cellotriosyl (DP3) and
molecular mass and linkage distribution in the (1 ! 3, 1 ! 4)- cellotetraosyl (DP4) residues in a water-soluble (1 ! 3, 1 ! 4)-
b-D-glucan chains. Extensive association with other polymers b-D-glucan from barley grain. The polysaccharide was hydrolyzed
and very high molecular masses will render the (1 ! 3, 1 ! 4)- (vertical arrows) with a specific (1 ! 3, 1 ! 4)-b-D-glucan
b-D-glucans more difficult to extract from the grain. endohydrolase until the penultimate oligosaccharides of DP6, DP7, and
The average molecular masses reported for cereal (1 ! 3, DP8 predominated. The autocorrelation coefficient (r) is defined in
terms of the probabilities that a cellotriosyl residue is followed by
1 ! 4)-b-D-glucans range from 48 000 (degree of polymeriza-
another cellotriosyl residue (p) or a cellotetraosyl residue (q), which
tion, DP, 300) to 3 000 000 (DP 1850), depending on the
can be estimated from the abundance of the oligosaccharide products.
cereal species, cell wall type, extraction procedure, and the If the polysaccharide is treated as a first-order Markov chain, a r value
method used for molecular mass determination. It is likely of 1 indicates an adjacent arrangement of DP3 and DP4 units, a r
that in many cases, the DP of the polysaccharide has been over- value of 1 indicates an alternating arrangement of DP3 and DP4 units,
estimated, particularly where gel permeation chromatography is and a r value of 0 indicates a random arrangement of DP3 and DP4
used for size determination, because standards used to calibrate units. Values very close to 0 were obtained for two barley (1 ! 3,
the columns often have a more globular conformation than the 1 ! 4)-b-D-glucans.
212 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides
highly asymmetrical wall polysaccharides. The (1 ! 3, 1 ! 4)-b- Table 3 Adoption of (1 ! 3, 1 ! 4)-b-D-glucan in grain is correlated
D-glucans are invariably polydisperse with respect to molecular with fine structure
mass, and this is illustrated by a weight average to number
Typical properties of
average molecular mass ratio (Mw/Mn) of 1.18 for barley
Species (1,3;1,4)-b-glucans in grain
(1! 3, 1 ! 4)-b-D-glucan. Certain barley (1 ! 3, 1 ! 4)-b-D-
glucans are covalently associated with small amounts of protein Oats (Avena sativa) Ratio in grain 2:1
and have estimated molecular masses of up to 40 000 000. 80% soluble in water (40 C)
Apart from their molecular mass, the fine structure of the 68% (w/w) grain
(1 ! 3, 1 ! 4)-b-D-glucans is a key determinant of their solu- Barley (Hordeum vulgare) Ratio in grain 2.8:1
bility. There are two components of fine structure that need to 20% soluble in water (40 C)
be considered in this context. Firstly, the longer stretches of 410% (w/w) grain
Wheat (Triticum aestivum) Ratio in grain 3.2:1
520 adjacent (1 ! 4)-linkages in the polysaccharide will be
0% solubility in water (40 C)
cellulosic in nature and would allow intermolecular aggrega-
0.52.3% (w/w) grain
tion and hence reduced solubility. However, these longer
blocks of adjacent (1 ! 4)-linkages are not abundant and As the DP3:DP4 ratio increases, the solubility of the (1 ! 3, 1 ! 4)-b-D-glucan
their effects on solubility have not been quantitated. The sec- decreases and the amount in the grain also appears to decrease.
ond and probably more important element of polysaccharide
fine structure that will affect solubility is the relative propor- that is water-soluble at 40 C varies within and between species.
tions of the major subunits of the (1 ! 3, 1 ! 4)-b-D-glucans, For example, waxy (high-amylose) barleys have a higher pro-
namely, the cellotriosyl (DP3) and cellotetraosyl (DP4) units portion of water-soluble (1 ! 3, 1 ! 4)-b-D-glucan than normal
(Figure 2(b)). The ratio of these tri- to tetrasaccharide units barleys. (1! 3, 1 ! 4)-b-D-Glucans extracted from barley at
(DP3:DP4 ratio) as illustrated in Figure 2(b) is proving to be a 40 C have a slightly lower tri- or tetrasaccharide ratio (1.7:1)
useful predictor of (1 ! 3, 1 ! 4)-b-D-glucan solubility, insofar than those extracted at 65 C (2.0:1). Complete extraction of
as the polysaccharide appears to become less soluble in aque- cereal (1! 3, 1 ! 4)-b-D-glucans from grain requires the use of
ous media as the ratios move away from 1:1. As the number of alkaline extractants such as 4 M NaOH or aqueous Ba(OH)2,
DP3 units in the (1 ! 3, 1 ! 4)-b-D-glucan increases, the containing NaBH4 to prevent alkali-induced degradation from
molecular kinks imposed by the single (1 ! 3)-b-D-glucosyl the reducing terminus. Alkali-extracted barley (1 ! 3, 1 ! 4)-
residues will become more evenly spaced, and the polysaccha- b-D-glucan fractions have higher molecular masses, higher ratios
ride chains will be able to align over longer regions. Thus, of (1! 4):(1 ! 3)-linkages, more contiguously linked (1 ! 4)-
solubility will decrease as the DP3:DP4 ratio increases to larger linked segments, and higher trisaccharidetetrasaccharide ratios
values. Similarly, as the number of DP4 units in the (1 ! 3, than their water-extractable counterparts. Other extractants,
1 ! 4)-b-D-glucan increases, the molecular kinks imposed by such as dimethyl sulfoxide, hot perchloric acid, trichloroacetic
the single (1 ! 3)-b-D-glucosyl residues will again become acid, N-methylmorpholine N-oxide, and dimethylacetamide
more evenly spaced, and solubility will decrease as the DP3: LiCl, have also been used to solubilize (1 ! 3, 1 ! 4)-
DP4 ratio decreases to low values. This key argument for b-D-glucans, but all are liable to cause some depolymerization
explaining (1 ! 3, 1 ! 4)-b-D-glucan solubility in terms of or degradation of the polymer. Thus, care must be exercised in
DP3:DP4 ratios only holds if the major DP3 and DP4 units comparing structural differences in cereal (1 ! 3, 1 ! 4)-
of the polysaccharide are arranged in an essentially random b-D-glucans extracted using different solvents. Once extracted
fashion along the (1 ! 3, 1 ! 4)-b-D-glucan chain. This has with hot water or alkali, the (1 ! 3, 1 ! 4)-b-D-glucans are
been demonstrated for very few (1 ! 3, 1 ! 4)-b-D-glucans, soluble at neutral pH and room temperature. However, upon
but a random arrangement of the DP3 and DP4 units does cooling, the (1! 3, 1 ! 4)-b-D-glucans slowly aggregate and
occur in a water-soluble (1 ! 3, 1 ! 4)-b-D-glucan from barley precipitate.
grain (Figure 3).
The variations in the DP3:DP4 ratios between (1 ! 3,
1 ! 4)-b-D-glucans from different tissues and at different devel-
Molecular Conformations
opment stages indicate that the biosynthetic enzymes respon-
sible for (1 ! 3, 1 ! 4)-b-D-glucan synthesis are able to adjust In aqueous media barley (1 ! 3, 1 ! 4)-b-D-glucans adopt a
the fine structure and hence the physicochemical properties of partially extended, somewhat rigid, reptate (snakelike) chain
the polysaccharide. There is circumstantial evidence that the conformation (Figure 2(c)) with a cross-sectional diameter of
amount and solubility of (1 ! 3, 1 ! 4)-b-D-glucan in grain are 0.45 nm and a lengthwidth (axial) ratio of 100. It is likely
correlated (Table 3). However, it needs to be emphasized that that other (1 ! 3, 1 ! 4)-b-D-glucans adopt a similar confor-
the solubility of the (1 ! 3, 1 ! 4)-b-D-glucan will also depend mation in aqueous media, but there are insufficient conforma-
on intermolecular interactions with other (1 ! 3, 1 ! 4)-b-D- tional data available for (1 ! 3, 1 ! 4)-b-D-glucans from a
glucan molecules and with other polysaccharides and proteins range of sources to allow meaningful comparisons. It would
in the grain during cereal processing, as noted earlier. be particularly interesting to compare the molecular confor-
A portion of the (1! 3, 1 ! 4)-b-D-glucan from barley, oat, mations of (1 ! 3, 1 ! 4)-b-D-glucans with different DP3:DP4
and rye flours is extracted by water at pH 7.0 and 40 C. These ratios.
flours consist predominantly of aggregates from starchy In the solid state, x-ray fiber diagrams of the water-soluble
endosperm cells. Further fractions can be solubilized at higher barley, (1 ! 3, 1 ! 4)-b-D-glucan can be interpreted as a
temperatures. The proportion of total (1 ! 3, 1 ! 4)-b-D-glucan (1 ! 3)-linked cellotriosyl polymer with a threefold screw
CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides 213
axis and a fiber repeat of 4.2 nm. The shape of the chain is (1 ! 4)-b-D-glucosyl units, not present in lichenin, may also
reminiscent of cellulose in the cellotriosyl and longer cello- participate in junction zone formation. The induction time for
oligosaccharide units, but the (1 ! 3)-linkages provide some gelation depends on mobility and diffusivity. It increases with
helical character. the molecular mass of (1 ! 3, 1 ! 4)-b-D-glucan and decreases
as the concentration increases. Gelation is repressed at high
and low temperatures, where molecular motion is high and
Viscosity of Solutions of (1 ! 3, 1 ! 4)-b-D-Glucans
low, respectively. Gelation may be induced by homogeniza-
The intrinsic viscosities of (1 ! 3, 1 ! 4)-b-D-glucan solutions tion, centrifugation, and freezing and thawing.
are dependent on the cereal species from which the polysac-
charide is extracted and on the solvent used. Intrinsic viscosity
StructureFunction Relationships in Cereal Processing
values for barley (1 ! 3, 1 ! 4)-b-D-glucans in water range
and Nutrition
from 4.6 to 6.9 dl g1 and from 2.0 to 9.6 dl g1 for oat
(1 ! 3, 1 ! 4)-b-D-glucans. The extractability, solubility, and The viscosity-generating properties of soluble cereal (1 ! 3,
viscosity characteristics of cereal (1 ! 3, 1 ! 4)-b-D-glucans 1 ! 4)-b-D-glucans are critical determinants in many aspects
can be understood by reference to their fine structures, as of cereal processing. For example, incompletely degraded
noted earlier. Long runs of adjacent (1 ! 4)-b-D-glucosyl (1 ! 3, 1 ! 4)-b-D-glucans from malted barley and cereal
units in the (1 ! 3, 1 ! 4)-b-D-glucan chain will form extended adjuncts can contribute to wort and beer viscosity and are
ribbonlike cellulosic stretches, but the insertion of a (1 ! 3)- associated with problems in wort separation and beer filtra-
linked glucosyl unit between them causes a kink in the mol- tion. In other applications, (1 ! 3, 1 ! 4)-b-D-glucans can be
ecule (Figure 2(c)). Chain flexibility arises principally from the cast as biodegradable films, have foam- and emulsion-
isolated (1 ! 3)-b-linkages. Because the (1 ! 3)-linked stabilizing ability, and, when used as a flour supplement,
b-glucosyl units do not occur regularly along the chain, the increase water absorption in doughs. Soluble cereal (1 ! 3,
polysaccharides are not able to align over extended regions and 1 ! 4)-b-D-glucans have antinutritive effects in monogastric
hence remain dispersed and soluble in aqueous media. The animals such as pigs and poultry. The antinutritive effects
overall extended polysaccharide conformation is due to the have been attributed to the increased viscosity of gut contents,
predominance of (1 ! 4)-linkages and results in the occupancy which is believed to slow both the diffusion of digestive
of a high volume of solvent (high hydrodynamic volume). This enzymes and the absorption of degradative products of
results in aqueous solutions of high viscosity. enzyme action. This, in turn, leads to slower growth rates.
The viscosity of solutions of cereal (1 ! 3, 1 ! 4)-b-D- Moreover, in dietary formulations for poultry, high (1 ! 3,
glucans depends strongly on concentration. At concentrations 1 ! 4)-b-D-glucan concentrations are associated with sticky
of oat (1 ! 3, 1 ! 4)-b-D-glucan where the total volume occu- feces, which are indicative of the poor digestibility of the
pied by the chains is, in aggregate, less than the total volume of (1 ! 3, 1 ! 4)-b-D-glucans and which may present major han-
the solution, there is contact between individual chains, but dling and hygiene problems for producers.
the viscosity of the solution is independent of shear rate. At Cereal (1 ! 3, 1 ! 4)-b-D-glucans are important compo-
concentrations above the critical overlapping concentration, nents of dietary fiber in human and animal diets. Humans
there is interpenetration of the chains in response to increasing and monogastric animals produce no enzymes that degrade
space occupancy. The viscosities of such aqueous solutions are (1 ! 3, 1 ! 4)-b-D-glucans, although there are indications that
reduced with increasing shear rate, a phenomenon known as some depolymerization occurs in the stomach and small
shear thinning. These different concentration-dependent rhe- intestine, presumably due to the activity of commensal
ological properties of (1 ! 3, 1 ! 4)-b-D-glucan solutions may microorganisms. By comparison, the soluble (1 ! 3, 1 ! 4)-
be of significance in their physiological responses as food b-D-glucans and other nonstarchy polysaccharides are readily
ingredients and in their use in food formulations. fermented by colonic microorganisms and make a small con-
tribution to digestible energy. In contrast to their antinutritive
effects in monogastric animals, oat and barley (1 ! 3, 1 ! 4)-
Gelation
b-D-glucans at high concentrations in human foods have
(1 ! 3, 1 ! 4)-b-D-Glucan solutions at concentrations of 5% beneficial effects, especially for noninsulin-dependent dia-
(w/v) or more form elastic gel networks. The gels are thermo- betics, by flattening glucose and insulin responses that follow
reversible, exhibit broad melting transitions, and show syner- a meal. High concentrations of (1 ! 3, 1 ! 4)-b-D-glucans
esis. Their melting temperatures depend on the source of (20% w/v) in food have also been implicated in the reduction
(1 ! 3, 1 ! 4)-b-D-glucan as follows: lichenin, 73 C; barley, of serum cholesterol concentrations, by lowering the uptake of
65 C; and oats, 62 C. These values are again related to dietary cholesterol or resorption of bile acids from the intes-
trisaccharidetetrasaccharide ratios, which in these examples tine. There is now a good deal of information on the micro-
are 22:1, 3:1, and 2:1, respectively. In the gel state, the (1 ! 3, biota of the human large intestine, on how the relative
1 ! 4)-b-D-glucan chains interact to form three-dimensional abundance of constituent microorganisms is controlled by
networks. Association between chains at junction zones, the carbon source embodied in dietary fiber, and on how the
through pairs of consecutive cellotriosyl units, is proposed. fermentation products of specific bacteria affect bowel health
This accounts for the higher melting temperature of lichenin and reduce the risk of serious human diseases, including type II
gels, where cellotriosyl units make up most of the molecule. In diabetes, cardiovascular disease, and colorectal cancer. More
the cereal (1 ! 3, 1 ! 4)-b-D-glucans, consecutive cellotriosyl recently, connections between dietary fiber and protection
units are less frequent, although the longer runs of adjacent against inflammatory diseases are emerging.
214 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides
(1 ! 3, 1 ! 4)-b-D-Glucans, in common with a number of locus (QTL) mapping has enabled regions of the barley
other polysaccharides, in particular (1 ! 3)-b-D-glucans, mod- genome that control (1 ! 3, 1 ! 4)-b-D-glucan concentrations
ify immunologic responses in humans by a process that is to be identified and placed on high-density genetic maps.
mediated through binding to receptors on cells of the reticulo- Furthermore, the conservation of gene order (synteny) across
endothelial system (leucocytes and macrophages). In addition, cereal genomes means that regions controlling (1 ! 3, 1 ! 4)-
they may have the capacity to activate the proteins of the b-D-glucan concentrations in cereals other than barley can
human complement pathway, a system that is invoked as a often be deduced from the genetic positions of barley QTLs.
first line of defense before circulating antibodies are produced. Using a comparative genomics approach based on barley QTLs
and the rice genome sequence, it has been shown that (1 ! 3,
1 ! 4)-b-D-glucan synthesis in the cereals is mediated by the
Analysis and Detection
cellulose synthase-like (Csl) genes of the CslF, CslH, and CslJ
Specific reagents for the analysis and detection of cereal (1 ! 3, clades. The CslF gene subfamily is the largest of these clades.
1 ! 4)-b-D-glucans include enzymes, fluorochromes, and There are ten CslF genes in barley and similar numbers in
monoclonal antibodies. The (1 ! 3, 1 ! 4)-b-D-glucan endo- wheat, sorghum, and rice. The CslH and CslJ gene subfamilies
hydrolase (EC 3.2.1.73) from Bacillus subtilis has been adopted are generally much smaller. The multiple members of the CslF
for the specific quantitation of (1 ! 3, 1 ! 4)-b-D-glucan in gene subfamily are distributed across cereal genomes, although
grain or grain extracts. This enzyme cleaves only (1 ! 4)- there is a cluster of CslF genes on chromosome 2H of barley
linkages that adjoin (1 ! 3)-substituted glucosyl units in the and in syntenic regions of other cereal genomes. The function
glucan chain and therefore liberates a series of oligoglucosides of the gene cluster is not known.
with (1 ! 3)-linked reducing units (Figure 2(b)). These may Expression analyses indicate that the CslF6 gene is the most
be quantified by b-glucosidase cleavage and measurement of highly transcribed member of the gene subfamily in develop-
the released glucose with the glucose oxidaseperoxidase ing barley grain and in most other tissues of barley. The CslF6
reagent, by separation and quantitation by high-performance gene also appears to be important in most other cereals. Over-
liquid chromatography (HPLC), or by matrix-assisted laser expression of the CslF6 gene in transgenic barley led to signif-
desorption ionization mass spectrometry (MALDI-TOF-MS). icantly elevated levels of (1 ! 3, 1 ! 4)-b-D-glucan in the grain
The latter procedures allow an assessment of the profile of and to increased dietary fiber values. This work also suggested
oligosaccharides making up the primary structure of the (1 ! 3, that different CslF isoenzymes synthesize (1 ! 3, 1 ! 4)-b-D-
1 ! 4)-b-D-glucan and, in particular, the trisaccharide glucans with different fine structures, as measured by DP3:DP4
tetrasaccharide ratio. ratios. Genome-wide association studies are showing that addi-
b-D-Glucans, including (1 ! 4)-b-D-glucan, cellulose, tional genes are likely to participate in the regulation of (1 ! 3,
(1! 3)-b-D-glucan, callose, and cereal (1 ! 3, 1 ! 4)-b-D-glu- 1 ! 4)-b-D-glucan levels in grain.
cans, form complexes with the diphenyldiazo dye, Congo red,
and members of the calcofluor family of diaminostilbene sul-
fonates. These reagents complex poorly with heteroxylans and Heteroxylans
AGPs. Irradiation of the complexes induces a brilliant red fluo-
rescence in the case of Congo red (emission lmax, 590 nm; green The cereal heteroxylans, or pentosans, are of two main types:
excitation, 470 nm) and a bright blue fluorescence with the arabinoxylans and the glucuronoarabinoxylans. They are
calcofluor (emission lmax 420, 442 nm; blue excitation, characteristically abundant in walls of vegetative tissues of
350 nm). The fluorochromes complex with and precipitate sol- cereals and grasses, where they represent the core non-
uble b-glucans. Although complexing with the fluorochromes is cellulosic polysaccharide in walls of the Poaceae. The arabinox-
not specific for (1! 3, 1 ! 4)-b-D-glucans, it is nevertheless ylans are the major noncellulosic polysaccharides in walls of
useful in quantifying soluble (1! 3, 1 ! 4)-b-D-glucans with the starchy endosperm cells and of the aleurone layer of cereal
molecular weights of > 10 kDa in grain extracts and in products grains, whereas the glucuronoarabinoxylans are characteristi-
such as wort, where cellulose will not be present. cally found in walls of the pericarpseed coat tissues (Table 1).
The fluorochromes and a (1 ! 3, 1 ! 4)-b-D-glucan-specific Within a cereal species, heteroxylan concentration is influ-
monoclonal antibody can be used to locate (1 ! 3, 1 ! 4)- enced by both genotypic and environmental factors.
b-D-glucans in tissue sections in histological studies. Although
near-infrared reflectance (NIR) spectra of polysaccharides are
Structure
complex, making their direct interpretation difficult, principal
component analysis of the spectra of different polysaccharides Cereal heteroxylans have a (1 ! 4)-b-D-xylan backbone that
enables characteristic NIR wavelengths to be determined. NIR has the same shape as the (1 ! 4)-b-D-glucan, cellulose
spectroscopy has been applied to the determination of (1 ! 3, (Figure 1). The b-xylopyranosyl (Xylp) units of the xylan back-
1 ! 4)-b-D-glucans in ground barley grain and this method bone are modified by side-chain branching (substitution)
forms the basis of rapid, routine (1 ! 3, 1 ! 4)-b-D-glucan (Figure 4). The major side chains comprise single a-L-
estimation. arabinofuranosyl (Araf) units, situated predominantly at
C(O)3, but in certain species also at C(O)2 of the Xylp units.
In some cases, Araf substitutions occur at both C(O)3 and
Genetics
C(O)2. The frequency of Araf substitution depends on the
(1 ! 3, 1 ! 4)-b-D-Glucan concentrations in grain are influ- cereal species and wall type and is reflected in a very wide
enced by both genotype and environment. Quantitative trait range of Xylp:Araf ratios. The Xylp units of cereal arabinoxylans
CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides 215
may be acetylated. In sorghum heteroxylans, acetyl groups are Other monosaccharides found on the xylan backbone
present on both the C(O)2 and the C(O)3 of xylopyranosyl include D-glucuronic acid (GlcAp) and its 4-O-methyl ether,
residues. Heteroxylans from cell walls in vegetative tissues, linked to the C(O)2 of Xylp units. GlcAp units are found on
especially lignified walls such as those from parenchyma, scle- endosperm and bran heteroxylans of rice and sorghum and on
renchyma fibers, and the pericarpseed coat tissues that are bran heteroxylans of wheat, rye, and maize. The barley husk
found in cereal brans, have low degrees of Araf substitution arabinoxylan has 4% by weight GlcAp. Various oligomeric
and therefore high Xylp:Araf ratios. On the other hand, hetero- chains are also found as substituents on arabinoxylans
xylans from the aleurone layer and starchy endosperm have (Table 4).
relatively low Xylp:Araf ratios. Characteristically, a proportion of the Araf units in the arabi-
The distribution of Araf units along the xylan chain is not noxylans of cereals, grasses, and several other families of
regular. In one wheat arabinoxylan fraction, three major types monocotyledons are esterified with the hydroxycinnamic acids,
of substitution patterns have been identified in different ferulic acid (FA), and, to a lesser extent, its nonmethoxylated
regions of the xylan backbone (Figure 5). In region I, isolated analog p-coumaric acid (pCA) (Figure 4). These hydroxycinna-
unsubstituted Xylp units are separated by one or two mono- or mates are found at C(O)5 of Araf units that are linked to C(O)3
disubstituted units. Region II contains high amounts of Araf of the Xylp units. Hydroxycinnamates constitute 1.8% (w/w) in
units linked at C(O)3 and region III contains high amounts of walls of wheat aleurone cells and the FA:pCA ratio is 9:1. In
contiguous (up to six and possibly more) unsubstituted Xylp contrast, walls of the starchy endosperm cells contain only
units. 0.04% (w/w) FA, with a trace of pCA. During wall formation,
enzyme-catalyzed, radical (oxidative) coupling of feruloyl units
on neighboring arabinoxylan chains occurs and leads to their
HO cross-linking. At least five different cross-linking FA dehydrodi-
HO O OH
OH O mers are encountered in fiber from wheat, barley, rye, spelt, oats,
O
O
O rice, and millet. Their structures are shown in Figure 6.
O OH O
In walls of pericarp and seed coat tissues, dehydrodiferulate
O
O bridges also cross-link arabinoxylans, and in lignified walls, the
OH arabinoxylans are cross-linked to lignin through FA and dehy-
drodiferulate esterether bridges (Figure 7). Thus, the lignified
O O CH2
OH
a-L-Araf-(l-
H3CO 4-Me-D-GlcpA-(l-
OH D-GlcpA-(l-
b-D-Galp-(l ! 5)-L-Araf-(l-
Figure 4 Structure of portion of a (1! 4)-b-D-xylan chain substituted at
b-D-Xylp-(l ! 2)-L-Araf-(l-
O3-L-arabino of one residue with a 5-O-trans-feruloyl-L-arabinofuranosyl
b-D-Xylp-(l ! 3)-L-Araf-(l-
substituent. Reproduced from Fincher GB and Stone BA (1986) Cell walls
b-D-Galp(l ! 4)-b-D-Xyl-(l ! 2)-L-Araf-(l-(or L)
and their components in cereal grain technology. Advances in Cereal
4Me-D-GlcA-(l ! 4)-D-Xylp-(l ! 4)-D-Galp-(l-
Science and Technology 8: 207295, with permission.
Region I ~ ~
>
Region II ~ ~
>
Region III ~ ~
>
Figure 5 Model of distribution of substituents on an arabinoxylan fraction from wheat endosperm showing three regions with different substitution
patterns. Reproduced from Izydorczyk MS and Biliaderis CG (1995) Cereal arabinoxylans Advances in structure and physico-chemical properties.
Carbohydrate Polymers 28: 3348, with permission.
216 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides
O O
Ara
O O O
Ara Ara
O
O O O
Ara Ara O Ara 8
O H 8 O H 5 OMe
5 O 4 H O
5 OMe OMe
OH
OH
OMe OMe
OH OMe
1 (5-5) 2 (8-O-4) OH 3(8-5)
OH
OMe O O
O Ara
O O Ara
Ara O
H 8
8 H
Ara 4 5
O O O OMe
MeO OMe OH
OH
4 (8-8) 5 (4-O-5)
Figure 6 Ferulated arabinoxylan chains in cell walls of grasses become cross-linked by radical coupling of ferulate monomers into ferulate
dehydrodimers (structures 16). Dotted arrows indicate potential sites for further radical coupling with hydroxycinnamyl alcohols or lignin
oligomers, resulting in cross-linking of arabinoxylans to lignin. Ara is an arabinofuranosyl residue on an arabinoxylan. Reproduced from Grabber
JH, Hatfield RD, Ralph J, Zon J, and Amrhein N (1995) Ferulate cross-linking in cell walls isolated from maize cell suspension cultures. Phytochemistry
40: 10771082, with permission.
Direct ether OH
O O
(a) (c) C=O
acch
(d)
C=O
Polys
O OH Hydroxycinnamic
O O acid ether
Lignin
=
=
O O C
O C (e)
O
OH O H
C
O Ferulic acid
=
O (f)
(h) bridge
de
Po O=C
hari
lys OH O
ac O
sacc
ch O
=
Dehydrodiferulic ari C C
de O
Poly
acid diester-ether
=
O O
bridge H
(g)
Dehydrodiferulic
acid diester bridge
Figure 7 Schematic diagram showing possible covalent cross-links between polysaccharides and lignin in walls. () p-Coumaric acid; ( ) ferulic
acid; ( ) dehydrodiferulic acid. (a) Direct ester linkage; (b) direct ether linkage; (c) hydroxycinnamic acid esterified to polysaccharide; (d)
dehydrocinnamic acid esterified to lignin; (e) hydroxycinnamic acid etherified to lignin; (f) ferulic acid esterether bridge; (g) dehydrodiferulic acid
diester bridge; and (h) dehydrodiferulic acid diesterether bridge. Reproduced from liyama K, Lam TB-T, and Stone BA (1994) Covalent cross-links in
cell wall. Plant Physiology 104: 315320, with permission.
CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides 217
walls in the pericarpseed coat layers and husk of cereal grains (Figure 8(a)). Araf substitution of the backbone Xylp units
contain significant concentrations of ligninheteroxylan com- does not substantially change the basic threefold helical con-
plexes. Wheat bran, which consists largely of pericarpseed formation (Figure 8(b) and 8(c)). However, substituted arabi-
coat and aleurone, contains 0.090.18% (w/w) diferulate res- noxylans cannot assume the twofold helical conformation in
idues. This arises almost exclusively from the pericarpseed solution because of significant steric interactions between
coat, because the contribution of FA dimers from the aleurone the Araf units and the xylan backbone. Nevertheless, (1 ! 4)-
is very low (0.03% w/w). b-D-xylans with low degrees of substitution interact reversibly
The heteroxylans in maize bran contain 1.3% (w/w) dehy- with the surfaces of cellulose microfibrils and may do so in
drodiferulates, which represent 15 dehydroferulate cross brid- the twofold helical conformation. With increased Araf
ges per heteroxylan molecule. Ligninheteroxylan associations substitution, the affinity for cellulose decreases, presumably
through esterether FA and di-FA cross-links also occur in ligni- due to the shift from twofold to threefold helical conforma-
fied walls in stem tissues of cereals and grasses. In walls of wheat tion, which would be unfavorable for association with cellu-
aleurone, highly branched arabinoxylan appears to be cova- lose, and because of the steric hindrance to interaction
lently associated with protein, possibly through ester-linked FA imposed by the substituents themselves.
that is linked to tyrosine in wall proteins by radical coupling. The length of the backbone chains of arabinoxylans is
Heteroxylans in maize and rye brans are also covalently associ- 35 nm. In solution, the arabinoxylans behave as partially
ated with protein. The covalent bridges between heteroxylans, stiff, wormlike, cylindrical molecules whose flexibility is largely
wall proteins, and lignins are important determinants of the unaffected by Araf:Xylp ratios, which are generally in the range
cohesiveness of the wall fabric and of the resistance of lignified 0.390.82:1.
walls to digestion by polysaccharide and ester hydrolases.
Solution Properties
Molecular Parameters
The intrinsic viscosities of wheat arabinoxylans range from 0.8
Average molecular masses of cereal heteroxylans range from to 5.5 dl g1, depending on the degree of substitution. These
65 000 to 5 000 000 (DP 50038 000) and, as is the case with values may be compared with 0.21 dl g1 for dextran, 0.19 for
the (1 ! 3, 1 ! 4)-b-D-glucans, the value depends on the cereal beet arabinan, 0.120.25 for gum arabic, and 4.66.9 and
species, cell wall type, extraction procedure, and the method 2.09.6 for barley and oat (1 ! 3, 1 ! 4)-b-D-glucans, respec-
used for molecular mass determination. The cereal heteroxy- tively. The behavior of the arabinoxylans in solution is influ-
lans are highly polydisperse with respect to molecular mass as enced not only by the overall asymmetry of the molecules and
indicated by Mw/Mn values that range from 1.3 to 4.2 for alkali- their DP but also by the specific arrangement of Araf units
soluble wheat arabinoxylan, 4.1 for water-extractable wheat along the xylan backbone. Thus, stretches of unsubstituted
arabinoxylan, and 8.5 for rye arabinoxylan. Xylp units may permit intermolecular alignment over these
As with the (1 ! 3, 1 ! 4)-b-D-glucans, a portion of the sequences and lead to the formation of chain associations
arabinoxylans from walls of cereal grains is soluble in water that are stabilized by hydrogen bonds. As with (1 ! 3, 1 ! 4)-
at 40 C and higher temperatures, but other extractants are b-D-glucans, the properties of arabinoxylan solutions are
required to bring all the arabinoxylan into solution. Whereas strongly concentration-dependent. At low concentrations,
0.05 MNa2CO3, 8 M urea, and dimethyl sulfoxide extract only their viscosities are not shear-dependent but become so at
a small part of the water-insoluble arabinoxylan, most can be higher concentrations. This behavior is typical of polysaccha-
extracted with 1 M NaOH, 1 M hydroxylamine hydrochloride, rides with extended, asymmetrical conformations. Removal of
saturated Ba(OH)2, or 4-methylmorpholine N-oxide. Of these, Araf units, either by dilute acid or by a-L-arabinofuranosidase
saturated Ba(OH)2 containing NaBH4 is the most successful action, leads to aggregation and precipitation of the residual
extractant. Oxidative degradation with chlorite or alkaline (1 ! 4)-b-D-xylan or of partially substituted arabinoxylan
H2O2 is effective in extracting arabinoxylan cross-linked to molecules.
lignin from cereal brans. Arabinoxylans extracted under alka-
line conditions may lose all or part of their acetyl and hydro-
Gelation
xycinnamate substituents.
At high concentrations, arabinoxylan solutions form thermo-
reversible gels that are stabilized by noncovalent interactions.
Solid-State and Solution Conformations
Gelation of arabinoxylan solutions can also be induced by
Unsubstituted (1 ! 4)-b-D-xylans can adopt a twofold, helical dimerization (cross-linking) of feruloyl units on neighboring
conformation with only one hydrogen bond (O50 H-O30 ) arabinoxylan chains by radical coupling (Figure 6). The dimer-
between adjacent Xylp units. This conformation is more flexi- ization can be catalyzed by peroxidase/hydrogen peroxide,
ble than the twofold helix of the (1 ! 4)-b-D-glucan, cellulose, manganese-dependent peroxidase, laccase, ammonium persul-
which has two hydrogen bonds (O20 H-O60 and O50 H-O30 ) fate, ferric chloride, and other metal chlorides, halogens, and
between adjacent glucosyl units (Figure 1(b)). The absence of linoleic acid/lipoxygenase, but not by iodate or bromate. Well-
cooperative interresidue associations in xylans allows the mol- developed, three-dimensional gel networks are obtained with
ecules to be conformationally more versatile. Indeed, in the arabinoxylans containing high FA contents, high molecular
solid state, the unsubstituted (1 ! 4)-b-D-xylan chain preferen- masses, and relatively unsubstituted xylan backbones. The lat-
tially exists as a fully extended, threefold (i.e., three Xylp units ter facilitate the initial contact between feruloyl groups on
per helical turn), left-handed helix or a slowly twisted ribbon neighboring arabinoxylan chains. The rate of gelation depends
218 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides
Figure 8 Arabinoxylan conformations. Projections perpendicular (top) and parallel (bottom) of (a) (1 ! 4)-b-D-xylan backbone, (b) backbone with
single arabinofuranosyl side groups, and (c) backbone with two arabinofuranosyl side groups. Hydrogen bonds are shown by dotted lines. In each case,
the backbone is a left-handed, threefold helix. Reproduced from Atkins EDT (1992) Three dimensional structure, interactions and properties of xylans.
In: Visser J, Beldman G, Kusters-van Someren MA, and Voragen AGJ (eds.) Xylan and Xylanases, 576 pp. Amsterdam: Elsevier, with permission.
on temperature, pH, and concentration of oxidizing agent. The are mainly arabinoxylan, whereas those from barley and oats are
gel-forming abilities of water-extractable arabinoxylans from mainly (1! 3, 1 ! 4)-b-D-glucans. Ingestion of arabinoxylan-
rye and barley are higher than for those from wheat and enriched, wheat flour by-products reduces the glucose response
triticale, presumably because of fine chemical differences of following a meal in normal subjects. Water-extractable cereal
the type listed earlier. Covalently cross-linked arabinoxylans fractions have a very low content of dehydrodiferulate and are
may hold up to 100 g water per 1 g polysaccharide. Gels partially fermented by anaerobic bacteria in the colon to pro-
formed by treatment of maize bran with peroxide/peroxidase duce short-chain fatty acids. Water-unextractable cereal fractions
are brittle, probably due to the high content of cellulose. contain variable proportions of dehydrodiferulate; maize and
millet have the highest content and oat wheat and spelt the
lowest. Water-unextractable cereal fractions are poorly fermen-
StructureFunction Relationships in Cereal Processing
ted in the colon, which may be related to the relatively high
and in Nutrition
diferulate cross-linking of arabinoxylan chains that prevents
Arabinoxylans, together with (1! 3, 1 ! 4)-b-D-glucans, con- swelling and attack by bacterial polysaccharide hydrolases or,
tribute to wort and beer viscosity, which may impede wort more generally, because of lower rates of pentose fermentation.
separation and beer filtration. Arabinoxylans are also compo- High concentrations of water-unextractable fractions in the diet
nents of some beer hazes. The water-soluble arabinoxylans in are effective in increasing stool weight, through its bulking
barley, rye, and wheat contribute an antinutritive effect in capacity. In recent pig feeding trial experiments, diets supple-
dietary formulations for poultry. In human diets, cereal arabi- mented with wheat arabinoxylans were associated with
noxylans are components of the dietary fiber fraction. Water- increased viscosity of small intestine digesta, changes in large
extractable fractions from rye, wheat, and spelt (Triticum spelta) intestine microbiota composition that led to large increases in
CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides 219
short-chain fatty acids, reduced damage to DNA from Table 5 Genes that have been implicated in heteroxylan biosynthesis
colonocytes, and reduced serum triglyceride levels. These in higher plants, including proposed roles in xylan backbone synthesis
changes can be linked with human health benefits that are and the addition of the numerous substituents of the backbone chain
attributed to dietary fiber, including reduced glycemic index,
Gene family Proposed role
reduced risk of colorectal cancer and cardiovascular disease,
and reduced risk of type II diabetes. GT8 (IRX8 and GAUT) GluA transferases
Arabinoxylans have well-demonstrated impacts in bread GT43 (IRX9 and IRX14) (1,4)-b-Xylan synthase
making. Added water-extractable arabinoxylans have positive GT47 (IRX10 and IRX7) (1,4)-b-Xylan synthase
effects on the water absorption of doughs, especially after GT61 Ara transferases or Xyl transferases
oxidative gelation, and also enhance loaf volume. In both BAHD/PMT acyltransferases Feruloyl transferases
cases, the amounts and molecular masses of the added arabi- DUF579 (IRX15) O-Methyl transferase (GluA)
AXY O-Acetylation
noxylan are important. Added water-extractable arabinoxylans
are reported to slow starch retrogradation and to produce less
firm breadcrumbs. The effect on breadcrumb texture is attrib-
uted to the increased moisture content of the samples. Water- have been identified in barley, and although ADF consists pre-
extractable arabinoxylans from wheat bran are good emulsion dominantly of cellulose and lignin from the pericarpseed coat,
stabilizers. They increase the strength and elasticity of gluten some of the effects might be related to arabinoxylan and/or
starch films surrounding gas bubbles in doughs. This leads to a (1! 3, 1 ! 4)-b-D-glucan content. Three QTLs, with relatively
higher retention rate of CO2 in the bubbles and produces a large effects, map close to one another on chromosome 2H,
positive effect on the fineness and homogeneity of crumb whereas the other two QTLs are on chromosomes 4H and 1H.
texture. On the other hand, water-unextractable arabinoxylans Overall, however, interpretation of the QTL data in terms
have detrimental effects on dough rheology and loaf volume. of identifying genes involved in the biosynthesis of arabinox-
Water-extractable rye arabinoxylans are inhibitors of ice crystal ylans has usually been complicated by the phenotyping
growth and are thus candidate cryostabilizers. Arabinoxylan assays. For example, QTLs for water-extractable arabinoxylans
lipid mixtures may be cast as films. might be attributable either to xylan synthase activity or to
arabinofuranosidase/glucuronidase activity, because solubil-
ity will be affected by the degree of substitution of the xylan
Analysis and Detection
backbone. Phenotyping on the basis of Araf or Xylp levels or
The determination of arabinoxylans in cereal and cereal prod- Araf:Xylp ratios can also be misleading, because both mono-
ucts relies on the phloroglucinolHCl or orcinolHCl colori- saccharides will be found in polysaccharides other than ara-
metric reactions that show a high, but not absolute, specificity binoxylans, such as xyloglucans, pectin, and AGPs.
for pentoses. In addition, NIR can be used to estimate arabi- Based largely on the analyses of Arabidopsis mutants, a
noxylan content in grain, and NIR peak intensity ratios at 1164 number of genes have been implicated in heteroxylan biosyn-
and 990 cm1 allow the degree of Araf substitution in arabi- thesis (Table 5). However, the roles of these genes in arabinox-
noxylans to be defined. Arabinoxylans in grain tissues can be ylan synthesis in cereal grains have not always been defined
sensitively detected by UV-induced, bright blue fluorescence and the identification of genes that mediate the biosynthesis of
(lmax, 415445 nm; excitation, 310 nm) due to esterified FA the (1 ! 4)-b-D-xylan backbone of cereal arabinoxylans still
residues. Lignins also fluoresce under UV irradiation but may requires confirmation.
be distinguished from ferulate esters by examining walls at a
high pH, where the lignin continues to fluoresce blue, but
ferulate esters fluoresce green. Arabinoxylans may also be Glucomannans
located in tissue sections through immunogold labeling with
polyclonal antibodies raised against 4-O-methyl glucuronoxy- Glucomannans are linear copolymers of b-D-glucopyranose
lan or arabinoxylan or with gold-labeled xylanases. Polyclonal ( 30%) and its 2-epimer, b-D-mannopyranose ( 70%),
antibodies raised specifically against highly or sparsely joined by (1 ! 4)-linkages (Figure 9(a)) to form linear chains
substituted glucuronoarabinoxylans can also differentiate with DPs ranging from < 100 to several thousand. In some
these types of arabinoxylan in cell walls. examples, the backbone has (1 ! 6)-linked b-D-
galactopyranosyl substituents and is generally esterified with
acetyl groups. The shape of the glucomannan polymer is sim-
Genetics
ilar to cellulose (Figure 1(a)), and it has therefore been sug-
As high-density genetic maps for barley and wheat have gested that the chains may associate strongly with surfaces of
become available, QTLs for arabinoxylan (or pentosan) con- cellulose microfibrils. The glucomannans are extractable from
tent in grain have been mapped. A QTL accounting for 35% walls with alkaline borate solutions that act by complexing
of variation in both Araf: Xylp ratio and extract viscosity has with the mannopyranosyl units.
been found on the long arm of chromosome 1B of bread Glucomannans are minor constituents of walls of most
wheat. The observation that there is no apparent change in aleurone and starchy endosperm cells of cereals (Table 1),
arabinoxylan concentrations in the wheat 1BL/1RS transloca- but in certain Oryza indica rice cultivars, values of up to 17%
tion lines suggests that the genes involved are located outside glucomannan have been reported in the endosperm walls. The
the segment of that chromosome that was introgressed from CslA genes have been shown to mediate glucomannan biosyn-
rye. A total of five QTLs controlling acid detergent fiber (ADF) thesis in some plants.
220 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides
Gal
6
--Man 4 Man 4 Glc 4 Man 4 Man 4 Man 4 Man 4 Man 4 Man 4 Glc 4 Glc 4 Man--
6
Gal
Man = -D-mannopyranosyl unit
Glc = -D-glucopyranosyl unit
Gal = a-D-galactopyranosyl unit
4 = (14)-linkage
(a) 6 = (16)-linkage
Fuc Fuc R R R
2 2 4 4 4
Gal Gal --4 Gal A 2 Rha 4 Gal A 2 Rha 4 Gal A 2 Rha 4--
2 2
Gal A = -D-galaclopyranosyl unit
Xyl Xyl Xyl Xyl
Rha = a-L-rhamnopyranosyl unit
6 6 6 6
4 = (14)-linkage
--Glc 4 Glc 4 Glc 4 Glc 4 Glc 4 Glc 4 Glc 4 Glc 4 Glc 4 Glc 4 Glc--
2 = (12)-linkage
6 6
R = (14)-linked -D-galactopyranosyl chain
Xyl Xyl
or
Glc = -D-glucopyranosyl unit (13,16)-linked -D-galactopyranosyl chain
Xyl = a-D-xylopyranosyl unit or
Gal = -D-galactopyranosyl unit (15)-linked a-L-arabinofuranosyl chain with
Fuc = a-L-fucopyranosyl unit (c) (12) and (13)-side-branching
4 = (14)-linkage
6 = (16)-linkage
(b) 2 = (12)-linkage
Figure 9 Structural features of three noncellulosic polysaccharides found as minor constituents in cell walls of cereal grains: (a) glucomannans,
(b) xyloglucans, and (c) pectic polysaccharides.
tissue. In the nucellar projection and the vascular tissue, callose specific enzymatic procedure, and their size distribution deter-
deposition might be related to the control of assimilate trans- mined by HPLC or MALDI-TOF-MS.
port into the developing endosperm.
Arabinogalactan Proteins
Fructans
AGPs are widely distributed in plants. They consist of a protein
Fructans from cereals consist of short chains of up to 45 b-D- backbone that is rich in hydroxyproline and to which arabino-
fructofuranosyl units in (2 ! 6)-b-linkage to the fructosyl res- galactan polysaccharides are covalently attached, although
idue of the disaccharide sucrose. Some branching through there are many structural subclasses, some of which are
(2 ! 1)-b-linkages may also occur. Cereal fructans are soluble membrane-anchored. In wheat grains, the major AGP is an
in water and in boiling 80% (v/v) ethanol. During the early arabinogalactan peptide, with a peptide backbone that is
development of wheat grain, fructans can account for as much much smaller than most AGPs. The arabinogalactan peptide
as 20% dry weight. In mature grains, the contents are low and from wheat flour constitutes 0.270.38% on a dry-weight basis
variable: wheat, 1.32.5%; rye, 4.66.6%; barley, 0.8%; and and has an average molecular mass of 22 000, although higher-
oats, 0.1%. In the grain, fructans are found in both the endo- molecular-mass AGPs have been detected in other cereal
sperm and the pericarp/embryo, with higher concentrations in grains, including barley. The polysaccharide portion of the
the latter fraction. wheat arabinogalactan peptide constitutes 92% of the
Fructans are important carbohydrate reserves in the vegeta- molecule and consists of branch-on-branch b-D-galactosyl
tive tissues of temperate cereals, particularly in young seedlings units (Galp) linked by (1 ! 3)- and (1 ! 6)-linkages
where they can account for 70% of the dry weight. Cereal (Figure 10). The branched b-D-galactan backbone is
fructans are not digested in the monogastric stomach or small substituted by single a-L-arabinofuranosyl units. The arabino-
intestine but are readily fermented by microorganisms in the galactan has an Araf:Galp ratio of 0.630.72:1. In wheat endo-
lower gut. Thus, they may be considered to be components of sperm, the arabinogalactan component of the molecule is
cereal dietary fiber. Fructans may be quantified by NIR or by a covalently linked to a peptide of molecular mass 7800 at
+
H3N Tyr Ala Glu Val Hyp Ser Hyp Ala Ala Gly Ala Hyp Thr Ala Asp COO
Figure 10 Structure of wheat arabinogalactan peptide. () (1 ! 3)-Linked-b-D-galactopyranosyl unit, ( ) (1 ! 6)-linked b-D-galactopyranosyl unit,
and ( ) terminal a-l-arabinofuranosyl unit. Structure based on data of Fincher GB, Sawyer WH, and Stone BA (1974) Chemical and physical
properties of an arabinogalactan-peptide from wheat endosperm. Biochemical Journal 139: 535545; Van den Bulck K, Looseveld A-MA, Courtin CM,
Proost P, Van Damme J, Robben J, Mort A, and Delcour JA (2002) Amino acid sequence of wheat flour arabinogalactan-peptide, identical to part
of grain softness protein GSP-1 leads to improved structural model. Cereal Chemistry 79: 329331.
222 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides
each of the three hydroxyproline residues in the peptide used to prevent microfibril formation in arabinoxylans
sequence. The same peptide sequence is found at the NH2- and (1 ! 3, 1 ! 4)-b-D-glucans?
terminus of a protein associated with the surface of wheat Summarize current knowledge on the genes that mediate
starch granules. Among eight Canadian wheat cultivars, the arabinoxylan and (1 ! 3, 1 ! 4)-b-D-glucan biosynthesis in
peptide constituted between 6.5% and 14.3% of the AG- cereal grains.
peptide molecule. Summarize the functions of arabinogalactan peptides in
The wheat AG peptide is very soluble in water and is not wheat grains.
precipitated at saturating ammonium sulfate concentrations. Show the detailed metabolic pathways leading to the struc-
Its solutions are of low intrinsic viscosity (0.0450.062 dl g1). tures shown in Figure 7.
Neither its subcellular location in the endosperm nor its phys-
iological function is known. The wheat AG peptide, unlike its
larger AGP counterparts, does not bind to the phenylazo dye, Exercises for Readers to Explore the Topic Further
b-glycosyl Yariv reagents. The effects of wheat AG peptides on
bread-making properties are of equivocal significance. Although there have been significant advances in our under-
standing of the metabolism of cereal grain nonstarchy poly-
saccharides, there remain a number of unanswered questions
Concluding Remarks that will undoubtedly attract research attention in the imme-
diate future. Examples of these key questions include the
There is a great deal of information available on the chemistry following:
and physicochemistry of nonstarchy polysaccharides in cereal
grains. This is directly attributable to the central importance of What is the precise molecular mechanism through which
these polysaccharides in large-scale food processing activities (1 ! 3, 1 ! 4)-b-D-glucans are synthesized and how are the
that include brewing, baking, and stock feed manufacture. fine structures and hence physicochemical properties
Moreover, the nonstarchy polysaccharides of cereals have altered in response to changing functional requirements?
attracted renewed interest in recent years because of their What genes are involved in the biosynthesis of arabinoxy-
potentially beneficial effects in human nutrition. Despite this lans in cereal grains?
interest, there remain major gaps in our knowledge of the Can we identify regulatory genes that control and amounts
genes and enzymes that control nonstarchy polysaccharide and fine structures of (1 ! 3, 1 ! 4)-b-D-glucans and arabi-
biosynthesis in the cereal grain, particularly the genes that noxylans in cereal grains?
control arabinoxylan biosynthesis. In the case of cereal Can we define the cellular aspects of (1 ! 3, 1 ! 4)-b-D-
(1 ! 3, 1 ! 4)-b-D-glucans, where genes involved in the bio- glucans and arabinoxylan synthesis, including sequential
synthesis of the polysaccharide have been identified, attention assembly events and the organelles involved?
is now turning to genes that regulate levels of grain (1 ! 3, Is there evidence for the two phase assembly of cell wall
1 ! 4)-b-D-glucan and eventually to an understanding of the polysaccharides, whereby oligosaccharides are synthesized
factors that control fine chemical structure of the polysaccha- first and subsequently assembled into polysaccharides?
rides and hence their solubility and general solution behavior
during cereal processing. This information can be confidently
See also: Carbohydrates: Health Effects of b-Glucans Found in
expected to provide additional molecular markers through
Cereals; Food Grains and the Consumer: Grains and Health; The
which breeders will track key genes in their breeding programs,
Basics: Grain: Morphology of Internal Structure; The Cereal Grains:
to allow breeders to improve quality characteristics related to
Barley: An Overview of a Versatile Cereal Grain with Many Food and
nonstarchy polysaccharides in commercially important pro-
Feed Uses; Wheat: An Overview of the Grain That Provides Our Daily
cesses, and to further enhance the role of cell wall polysaccha-
Bread.
rides as components of dietary fiber that carry human health
benefits.
Further Reading
Exercises for Revision
Atkins EDT (1992) Three-dimensional structure, interactions, and properties of xylans.
In: Visser J, Beldman G, Kusters-van Someren MA, and Voragen AGJ (eds.) Xylan
What differences would you expect in the solubility of and Xylanases, p. 576. Amsterdam: Elsevier.
wheat flour arabinoxylans with xylosearabinose ratios of Bamforth CW (1994) b-Glucans and b-glucanases in malting and brewing: Practical
1.2 and 1.5, respectively, and how would you rationalize aspects. Brewers Digest 69: 1216.
those differences in terms of chemical fine structure? Belobrajdic DP, Bird AR, Conlon MA, et al. (2011) An arabinoxylan-rich fraction from
wheat enhances caecal fermentation and protects colonocyte DNA against diet-
What differences would you expect in the solubility of
induced damage in pigs. British Journal of Nutrition 107: 19.
barley (1 ! 3, 1 ! 4)-b-D-glucans with DP3:DP4 ratios of Burton RA, Collins HM, Kibble NAJ, et al. (2011) Over-expression of specific HvCslF
2.5 and 3.0, respectively, and how would you rationalize cellulose synthase-like genes in transgenic barley increases the levels of cell wall
those differences in terms of chemical fine structure? (1,3;1,4)-b-D-glucans and alters their fine structure. Plant Biotechnology Journal
9: 117135.
Cereal grains produce noncellulosic cell wall polysaccha-
Burton RA, Gidley MJ, and Fincher GB (2010) Heterogeneity in the chemistry, structure
rides, such as arabinoxylans and (1 ! 3, 1 ! 4)-b-D-glucans, and function of plant cell walls. Nature Chemical Biology 6: 724732.
that will not form microfibrils of the type produced by Burton RA, Wilson SM, Hrmova M, et al. (2006) Cellulose synthase-like CslF genes
cellulose. What are the two different chemical strategies mediate the synthesis of cell wall (1,3;1,4)-b-D-glucans. Science 311: 19401942.
CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides 223
Carpita N and Gibeaut D (1993) Structural models of primary cell walls in flowering Harris PJ and Fincher GB (2009) Distribution, fine structure and function of (1,3;1,4)-b-
plants: Consistency of molecular structure with the physical properties of the walls glucans in the grasses and other taxa. In: Stone BA, Bacic T, and Fincher GB (eds.)
during growth. Plant Journal 3: 130. Chemistry, Biochemistry, and Biology of (1,3)-b-Glucans and Related
Collins HM, Burton RA, Topping DL, Liao M-L, Bacic A, and Fincher GB (2010) Polysaccharides, pp. 621654. Philadelphia, PA: Elsevier.
Variability in the fine structures of non-cellulosic cell wall polysaccharides from Iiyama K, Lam TB-T, and Stone BA (1994) Covalent cross-links in the cell wall. Plant
cereal grains: Potential importance in human health and nutrition. Cereal Chemistry Physiology 104: 315320.
87: 272282. Izydorczyk MS and Biliaderis CG (1995) Cereal arabinoxylans Advances in structure
Cui SW (2001a) Cereal non-starch polysaccharides I: (1 ! 3) (1 ! 4)-b-D-glucans. and physico-chemical properties. Carbohydrate Polymers 28: 3348.
In: Cui SW (ed.) Polysaccharide Gums from Agricultural Products, pp. 104165. Kroon-Batenburg LMJ and Kroon JJ (1995) The crystal and molecular structures of
Lancaster: Technomic Publishing. cellulose. Carbohydrates in Europe 12: 1519.
Cui SW (2001b) Cereal non-starch polysaccharides II: Pentosans/Arabinoxylans. Skendi A, Biliaderis CG, Lazaridou A, and Izydorczyk MS (2003) Structure and
In: Cui SW (ed.) Polysaccharide Gums from Agricultural Products, pp. 167227. rheological properties of water soluble b-glucans from oat cultivars Avena sativa
Lancaster: Technomic Publishing. and Avena bysantina. Journal of Cereal Science 38: 117.
Evers T and Millar S (2002) Cereal grain composition and development: Some Stone BA and Clarke AE (1992) The Chemistry and Biology of (1 ! 3)-b-Glucans.
implications for quality. Journal of Cereal Science 36: 261284. Victoria: Latrobe University Press.
Fincher GB, Sawyer WH, and Stone BA (1974) Chemical and physical properties of an Van den Bulck K, Looseveld A-MA, Courtin CM, et al. (2002) Amino acid sequence of
arabinogalactan-peptide from wheat endosperm. Biochemical Journal 139: 535545. wheat flour arabinogalactan-peptide, identical to part of grain softness protein GSP-
Fincher GB and Stone BA (1986) Cell walls and their components in cereal grain 1 leads to improved structural model. Cereal Chemistry 79: 329331.
technology. Advances in Cereal Science and Technology 8: 207295. Vinkx CJA and Delcour JA (1996) Rye (Secale cereale L.) arabinoxylans A critical
Grabber JH, Hatfield RD, Ralph J, Zon J, and Amrhein N (1995) Ferulate cross-linking review. Journal of Cereal Science 24: 114.
in cell walls isolated from maize cell suspension cultures. Phytochemistry Visser J, Beldman G, Kusters-van Someren MA, and Voragen AGJ (eds.) (1992) Xylan
40: 10771082. and Xylanases, p. 576. Amsterdam: Elsevier.
Grains other than Cereals: Non-starch Polysaccharides
L Ramsden, University of Hong Kong, Hong Kong, China
2016 Elsevier Ltd. All rights reserved.
Non-starch polysaccharides (NSP) have a diverse composi- In common with all plant tissues, the tissues forming a grain
tion and are widely distributed in plants, are all are composed of cells, each of which is defined by a cell wall.
indigestible, but can strongly influence texture. The cell is the basic structural unit of plants and cellulose is the
Different types of NSP are produced in monocots and basic structural component of the plant cell wall. All plant cell
dicots. walls contain not only cellulose but a number of other poly-
NSP may be accumulated as a reserve polysaccharide inde- saccharides that are important in maintaining the structural
pendently or as part of the cell wall. integrity of the wall and controlling wall permeability. Other
NSP may influence the absorption of carbohydrates, lipids, components may be involved in interactions with neighboring
and proteins. cells and the environment.
The legume family is an important source of non-cereal The cell walls of plants display a composition that is charac-
grains. teristic of the taxonomic group in which the plant is found. In
the angiosperms, there are consistent differences in the pattern
of wall polysaccharides present between dicots and monocots.
Thus, cereal plant cells possess walls typical of monocot grasses,
Learning Objective and this also applies to the endosperm cells in the cereal grain.
The cell wall is formed from two types of wall, the primary and
the secondary. Primary walls are those first laid down after a cell
To appreciate the diversity, abundance, and potential value
of NSP resources in non-cereal grains. is formed and are thinner and more flexible than secondary
walls that are found in mature cell types. Both primary and
secondary walls contain cellulose, but the cellulose fibrils of
the primary wall are thinner and more loosely arranged; in the
secondary wall, the cellulose fibrils are closely packed into
Introduction organized layers that are linked to each other by cross-links.
Endosperm cells are typically larger and have thinner walls
All plant tissues contain a large variety of polysaccharides, than other tissues in the plant so the proportion of cellulose
many of which serve structural or reserve functions in the present can be lower than that in other tissues.
plant. However, the most important polysaccharide obtained Seed-bearing plants deposit energy-containing reserves to
from grain crops is starch and the focus in starch producing support growth of the embryo within the seed. While the major
grain crops is naturally on the yield of starch. Any other poly- reserve polysaccharide is indisputably starch, there are a num-
saccharides present are considered to be of secondary impor- ber of other polysaccharides that are used particularly within
tance as non-starch polysaccharides (NSPs), a term that covers the legume plant family (Fabaceae). Starch reserves are stored
a great variety of biological functions and chemical structures. in the cytoplasm inside amyloplasts, but the non-starch reserve
There is a growing awareness that these polysaccharides can polysaccharides in seeds are deposited in highly thickened
have considerable value both in human nutrition and for the walls of endosperm cells without a structural function.
food industry. In particular, non-cereal grains show an inter-
esting diversity of polysaccharides that are less familiar than
the better studied NSPs of the cereal grain. Non-cereal grains
include a great variety of potential species, many of which have Distribution of Grain NSPs
not been analyzed for NSP content. Consequently here, it is
only possible to present a limited selection together with some There is a clear distinction between monocot- and dicot-grain
details of the structure and their value in products derived from NSPs, which reflects the differences between the monocot and
grains. dicot cell walls. The actual levels present in any grain sample
Polysaccharides in a grain can serve two major biological can also vary dependent on the growth conditions and partic-
functions. They can act as the energy reserve in the endosperm ular genotype. The distribution of NSPs in dicot grains is more
or cotyledons or they may be involved in forming the structure difficult to assess due to the variability in designation of what
of the grain tissues. Other functions in which polysaccharides constitutes a dicot grain. In dicots, the major monocot NSPs,
can play essential roles are in the regulation of water balance the mixed linkage b-glucans, are absent (Figure 1), and arabi-
during dormancy and imbibition and in the protection of the noxylans, though they may be present in dicot cell walls,
seed against predation and attack by pathogens. In cereals, should generally be expected to be largely replaced by xyloglu-
only starch has a major reserve function, but in dicot grains, can that is the main hemicellulose in dicots. Xyloglucans can
other polysaccharides are found to act as reserves, often be found as a reserve polysaccharide in a number of seeds.
together with starch. Galactomannans are found as seed reserve polysaccharides in
14 glucose 13 glucose
several dicot families; though their commercial importance as Table 1 Dietary fiber in cereal and legume grains
food additives is derived from their presence in large tree seeds,
they are also present in many smaller grains. Total fiber
Wheat 14.6
Uses of Grain NSPs Barley 22.6
Rice 2.4
Sorghum 10.1
The presence of high levels of NSPs in grain products will
Millet 8.5
reduce the nutritional value of the grain, a factor which is of
Maize 13.4
significance in the animal feed industry. An important feature Oats 9.6
of NSPs is their ability to bind water that allows them to act as Rye 14.6
hydrocolloids and disproportionately influence the rheology Triticale 18.1
of aqueous systems. In baking, the NSPs of wheat can enhance Chickpea 13.5
water retention and texture of bread and cakes. Purified NSPs Black gram 7.1
can be used as raw materials for the food additive industry that Mung bean 12.8
is a major user of polysaccharide texture modifiers, stabilizers, Pigeon pea 8.1
and gelling agents. Currently, traditional grains are not major Soybean 4.96
Lupin 4.0
sources of industrial polysaccharide hydrocolloids, but high
Lima bean 6.3
demand on traditional supplies makes this a competitive area
Kidney bean 6.2
for development of new sources. However, one of the major Faba bean 8.0
benefits of NSPs arises after they have been consumed as part Sunflower 4.2
of the diet, when they contribute to the dietary fiber.
Figures given are the highest levels reported. Actual values may vary considerably
depending on the variety grown and the cultivation conditions.
Dietary Fiber Data from Shelton DR and Lee WJ (2000). In: Kulp K and Ponte JG (eds.) Handbook of
Cereal Science and Technology, 2nd edn., pp. 385415. New York: Marcel Dekker;
The NSPs together form the major part of the dietary fiber of Sathe SK (1996). In: Nwokolo E and Smartt J (eds.) Food and Feed from Legumes and
grain crops. Dietary fiber is the fraction of a food consumed that Oilseeds, pp.1232. London: Chapman & Hall.
is not degraded in the small intestine, and as human digestive
enzymes can only cleave a14 glucan bonds, all polysaccharides less well suited to the modern high-fat, energy-rich, low-volume
other than starch are part of the dietary fiber. Dietary fiber can be diets of industrialized countries. This deficiency in the modern
both soluble and insoluble dependent on the molecular struc- diet can be overcome through the consumption of grain prod-
ture of the polysaccharides. There are also a smaller number of ucts high in dietary fiber. NSP can lower the availability and
other polymers present in plant tissues, such as lignin, which glycemic index of starch by complex formation and restricting
also form part of the dietary fiber. However, the simplest avail- access for digestive enzymes. Polysaccharides can also bind to
able measure of the quantity of NSP in grains is the level of dietary lipids and reduce their uptake into the body, a factor that
dietary fiber present (Table 1), and often, this is the only figure has been observed in the ability of high-fiber diets to lower
available for many grains that have not been analyzed in ade- blood lipid levels with potentially beneficial consequences for
quate detail for polysaccharide composition. the incidence of cardiovascular disease.
Although unable to contribute to human nutrition in terms
of provision of energy, the dietary fiber is well recognized to
form an important component of our diet for the correct func-
tioning of the digestive system. Polysaccharides through their Insoluble Cell Wall Polysaccharides
high water-binding capacity play an important part in providing
bulk to the gut contents to allow easy passage through the The main structural component of any cell wall is cellulose, a
intestine. The human digestive system developed to cope with b14 linked polymer of glucose (Figure 2). It is the worlds most
a diet rich in high fiber plant material with a large volume and is abundant polymer, followed by its sister glucan starch, with the
226 CARBOHYDRATES | Grains other than Cereals: Non-starch Polysaccharides
-Arabinose
Figure 3 The arrangement of sugar units in arabinoxylan. a-Arabinose and a-glucuronic acid linked 12 or 13 to a backbone chain of b14 xylose units.
14 glucose Galactose
Xylose
Figure 4 The arrangement of sugar units in xyloglucan. a-Xylose linked 16 to a backbone chain of b14 glucose units, with some b-galactose
linked 12 to the xylose units.
Reserve Polysaccharides galactose residues (Figure 5). The ratio of galactose to man-
nose is roughly constant for a given species. The distribution of
Seed-bearing plants deposit energy-containing reserves to sup- galactose along the mannan chain is not uniform but tends to
port growth of the embryo within the seed. While the major be clustered in blocks of high substitution (rough regions) that
seed reserve polysaccharide is without doubt starch, there are a are separated by intervening stretches with few galactose resi-
number of other polysaccharides that are used as reserves, dues (smooth regions).
particularly within the legume family. Unlike starch, which is Purified galactomannans for industrial usage are obtained
stored in amyloplasts in the cytoplasm, the major non-starch from four main plant sources in the legume subfamily Ca-
reserve polysaccharides in seeds are deposited in the cell wall esalpinioideae that are well characterized. There are guar gum
but do not serve a structural function. from the seeds of Cyamopsis tetragonoloba, locust bean gum
Apart from starch, there is only one other major polysaccha- from seeds of Ceratonia siliqua (carob tree), tara gum from
ride used by plants as a reserve outside the cell wall. These are seeds of Caesalpinia spinosa, and cassia gum from seeds of
polymers of fructose, the fructans, which can have chain lengths Cassia obtusifolia. Extraction of galactomannans involves de-
of up to 250 sugar units but are generally shorter and highly hulling of seeds and crushing to remove the embryo, followed
water-soluble. Many grasses are known to use fructans for stor- by milling of the endosperm to produce a crude flour. The
age and for frost resistance. Potentially, fructans may occur in flour can be purified by dissolving in hot water, followed by
the grains but there is little evidence for significant quantities filtration and precipitation with isopropanol to remove impu-
and most sources of fructans are from tubers or stem tissue. rities. The properties of galactomannans from different sources
vary, depending on the structure of the galactomannan that is
characteristic of the source species.
Galactomannans The use of locust bean galactomannan (LBG) in the Medi-
terranean and the Middle East has been part of traditional food
Many seeds of the legume family contain high levels of reserve preparation for hundreds of years. LBG has a galactose to
galactomannans. The seed galactomannans all possess the mannose ratio of 1:4 and a molecular weight of around
same basic structure of an a14 linked mannan backbone 300 000. The cold water solubility of LBG is low and disper-
chain with varying degrees of substitution with a16 linked sions need to be heated to 85 C to achieve good dissolution,
228 CARBOHYDRATES | Grains other than Cereals: Non-starch Polysaccharides
will be encountered. Many of these new grains have not been Using literature sources, try and find a new non-cereal grain
fully investigated for the composition of minor polysaccharides crop that is not currently exploited for NSP and predict
present in the endosperm that may have a significant impact on from DF content if it would be a useful source.
the properties of grain-derived products. In particular, grain
legumes as well-developed crops with high levels of NSP
would have potential for further investigation to establish the See also: Carbohydrates: Carbohydrate Metabolism; Cereals:
composition of dietary fiber fractions and permit exploitation as Chemistry and Physicochemistry of Non-starchy Polysaccharides;
defined NSP sources. Similarly, there are opportunities for the Health Effects of b-Glucans Found in Cereals; Food Grains and
development of grain crops to obtain NSP resources that are Well-being: Functional Foods: Dietary Fibers, Prebiotics, Probiotics,
presently obtained from related species with much lower yields. and Synbiotics; Functional Foods: Overview; Processing of Grains:
Eventually, we may see NSPs no longer being regarded as a b-Glucans: Measurement and Processing; The Basics: Grain:
nuisance reducing the value of the grain but instead representing Morphology of Internal Structure; The Legumes and
the major economic product. Pseudocereals: Grain Legumes and Their Dietary Impact: Overview;
The Oilseeds: Oilseeds: Overview.
Topic Highlights steps. First, the enzymes diffuse into the starch matrix of the
food. In the second step, the enzymes bind to the substrate,
What is resistant starch? and finally, the enzymes cleave the a-1,4-glycosidic linkages of
Resistant starch intake; measurement and consumption. the substrate (starch). The structure of the food matrix and the
Effect of resistant starch intake on gut health. structure of the starch itself influence the kinetics of the amy-
Metabolic effects of resistant starch intake. lase hydrolysis.
Resistant starch (RS) has been defined as the starch and
products of starch digestion that are not absorbed in the small
Learning Objectives intestine of healthy individuals. Based on the source of the
enzyme resistance, RS has been classified into five different
To get a basic understanding of what resistant starch is and types (Table 1).
where it is found. In type 2, type 3, and type 5 RS, the enzyme resistance is due
To learn about how its consumption has been linked to a to the physical structure of the starch molecules. Type 2 RS is
variety of potential beneficial health outcomes in the area found in raw starch granules and the enzyme resistance is due
of gut health and various metabolic effects such as blood to the natural organization of the starch within the starch
glucose, insulin, and lipid levels. granules. Significant amounts of type 2 RS can be found in
green banana, potato, and high-amylose maize.
In type 3 RS, the enzyme resistance is due to the physical
What Is Resistant Starch? structure of starch chains that have undergone some type of
restructuring due to retrogradation or heat treatment. Different
Starch is the major source of carbohydrates in the human diet. factors, such as amylose/amylopectin ratio, chain length, lipid
Starch is present in many different fruits, vegetables, roots, and content, and processing conditions, have been shown to influ-
grains. Starch and starch derivatives are a nutritive, abundant, ence the amount and quality of type 3 RS. Annealing and heat
and economical food source. Starch can be consumed unpro- moisture treatment are two types of heat treatments that are
cessed in the form of raw fruits and vegetables or in the form of often used to create RS. Both of these treatments, as well as
more shelf-stable processed foods. Food starches contribute to gelatinization, either fully or partially melt crystalline structure
the characteristic viscosity, texture, mouthfeel, and consistency present in the native starch granules. After heat treatment, linear
of many food products. amylose molecules and linear regions of amylopectin molecules
In the human body, starch is digested by a-amylases. First, can organize into a mix of amorphous and crystalline areas with
salivary a-amylase in the mouth catalyzes the hydrolysis of varying degrees of enzyme resistance. Formation of the crystal-
amylose to maltose, maltotriose, and maltotetraose and the line structures usually takes place above the glass transition
hydrolysis of amylopectin to the same products plus two temperature and below the melting temperature, and any com-
a-limit dextrins. The partially digested starch then passes into ponents present that influence the glass transition temperature
the stomach. After some resident time in the low pH environ- can therefore be expected to influence the formation (yield and
ment of the stomach, the partially hydrolyzed starch then quality) of the formed type 3 RS. The amylose content of starch
passes into the small intestine where it is neutralized. The has been positively correlated with RS yield and the formation
majority of hydrolysis of the starch is then accomplished by of type 3 RS is strongly related to the crystallization of amylose.
pancreatic a-amylase that is secreted from the pancreatic duct The amount of RS formed is also dependent on the water
via a multiple attack mechanism. a-Amylases hydrolyze the content and temperature used during the heat treatment.
starch into small mono-, di-, and oligosaccharides that then Water does act as a plasticizer in the system and a minimum
need to be further broken down to be absorbed as glucose. Two water content is necessary to achieve the chain mobility needed
other enzymes are necessary to convert the hydrolysis products to form crystalline structure resistant to enzyme digestion. At
of the a-amylases into glucose, which can be actively trans- high-starch concentrations, the starch chains interact more eas-
ported across the small intestine membrane. These two ily, leading to increased crystal and RS formation. The presence
enzymes are the brush-border glucogenic enzymes maltase of lipids has been shown to decrease the formation of type 3 RS
glucoamylase and sucraseisomaltase. The resulting D-glucose due to formation of amyloselipid complexes (type 5 RS).
is then actively transported across the luminal membrane of The enzyme resistance in type 5 RS is due to the molecular
the small intestine and passes into the blood via the sodium structure of amyloselipid complexes that can be either present in
glucose cotransporter, which is located at the luminal surface the native starch or formed by controlled reactions using non-
of enterocytes. granular starch and lipids to form the resistant amyloselipid
Several factors influence the rate and extent of starch diges- complex.
tion. The main hydrolysis of starch is performed by the The enzyme resistance in type 4 RS is due to chemical mod-
a-amylases. Starch digestion by a-amylases requires a series of ification of the starch. Chemical modification of starch creates
Table 1 Classification of resistant starch (RS) data, but these measurements are difficult and intrusive and
can therefore not be done on large scales.
RS It is generally accepted that any in vitro method used to
type Source of enzyme resistance Examples
measure RS should give values in line with those obtained
1 Whole grains Seeds, grains with ileostomy patients. In 1986, Berry developed a new
2 Raw starch granule structure Green banana, high in vitro method by modification of a method previously devel-
amylose, potato oped by Englyst, Wiggins, and Cummings in 1982. This mod-
3 Structures formed on Cooked pasta, cooked ified method was subsequently confirmed to give results in line
reassociation or retrogradation potato with healthy ileostomy subjects. In 1992, Englyst developed a
of starch after heat treatment method that could measure rapidly digestible starch (RDS),
4 Chemical modifications of the Modified starch slowly digestible starch (SDS), and RS. This method employed
starch
incubation of the samples at 37 C with pancreatic amylase
5 Starch lipid complex Amylose-containing starch
and amyloglucosidase and measures the amount of RS by
subtracting the sum of RDS and SDS from the total starch
chain irregularities or branches in the starch chains. Cross- present in the sample. Several new methods for the measure-
linking of starch covalently links two starch chains together, in ment of RS were developed during the European Research
effect creating a branch point on both chains. Chemical substi- Program EURESTA. Even though many different methods
tution introduces a bulky side group to the starch chains. The were developed during the 1990s for the measurement of RS,
introduction of these chemical groups may create a steric hin- none of these methods were successfully subjected to interla-
drance to one or more human digestive enzymes. boratory evaluation to be adopted as an official AOAC method
of analysis. In 2002, McCleary and Managhan developed a new
RS method by modifying existing methods to obtain a method
that would yield robust, reproducible results that correlated
RS Intake with results from ileostomy patients. This method underwent
interlaboratory evaluation and was later adopted as AOAC
Quantification of RS Official Method 2002.02. An Integrated method for the mea-
To determine the effect of RS intake on health, it is important surement of total dietary fiber was later developed and was
to determine the amount of RS consumed as accurately as published in 2007. This method allows the accurate measure-
possible. Ideally, RS consumption would be determined ment of all dietary fibers including RS. The enzymatic incuba-
in vivo by quantification of undigested starch in healthy indi- tion step uses pancreatic a-amylase and more closely simulates
viduals. This is, however, very difficult to do. Several in vivo digestion in the human digestive tract and yields RS values in
approaches have been used for measurement of RS. These line with those obtained with AOAC Official Method 2002.02
in vivo methods include the breath hydrogen test, direct collec- and with results from ileostomy patients. This method was
tion of ileal effluent from ileostomy patients, and direct col- accepted as AOAC Method 2009.01.
lection of the ileal effluent from healthy subjects using a long
triple lumen tube.
Sources of RS and Intake Levels
The measurement of breath hydrogen to determine the
amount of RS in a diet is based on the concept of fermentation Of all unprocessed foods, the green banana has the highest RS
of dietary fibers including RS. When RS enters the colon, it may content of between 47% and 57%. While flour can be prepared
be fermented by the colon microbiota that leads to the pro- from green bananas and incorporated into different foods as a
duction of short-chain fatty acids (SCFAs) (acetate, butyrate, source of RS, it is important to keep in mind that food proces-
and propionate) and gases like hydrogen (methane and carbon sing methods such as cooking may gelatinize the starch gran-
dioxide). Most of the gases produced by colonic fermentation ules and destroy the structures responsible for the enzyme
are absorbed into the circulation, but about 10% is present in resistance originally present and responsible for RS formation.
the breath on exhalation. Since breath hydrogen is not specific Raw potato starch is also high in RS, but since raw potato starch
for only RS fermentation, but does indeed measure fermenta- is seldom consumed, it is not a good source of RS. However,
tion from any dietary fiber source, care has to be taken when cooling of cooked potatoes or potato products can lead to
interpreting these results for RS content in a food/diet. retrogradation of the starch and may lead to formation of
Direct collection of ileal effluents would be expected to give new RS (type 3) if the right conditions (heat, moisture, time,
more accurate results than breath hydrogen measurement and starch concentration) are met. The level of RS in different
especially when different forms of dietary fibers and RS may legumes can vary widely and RS content in the legumes as
be present in a diet. Analysis of the ileal effluent allows for consumed depends on processing and legume variety.
quantification of undigested carbohydrate and potential iden- Murphy et al. examined the RS intake in the United States
tification of the different sources of dietary fiber within the by comparing reported RS contents of different foods with
effluent. Most ileostomy models are, however, based on the dietary recall records from participants in the 19992002
use of patients that had their colon removed due to health National Health and Nutrition Examination Survey. They
issues and their digestive patterns may therefore not be repre- showed that the RS concentration varies greatly, even within
sentative of a healthy population. Analysis of ileal effluent the same food category. This could be due to natural differ-
from healthy individuals should provide the most accurate ences (cultivar), differences in food preparation methods, and
232 CARBOHYDRATES | Resistant Starch and Health
also differences in analytic methods used to determine the Butyrate has been shown to induce apoptosis, decreasing
amount of RS present. The RS content of different breads varied the risk of cancer development. Animal studies have shown
between 0.1% in wheat rolls and 4.5% in pumpernickel bread. that an increased supply of butyrate to the large bowel
Ready-to-eat breakfast levels contained between 0% (rice decreases the induction of large bowel tumors in rats. How-
cereal) and 6.2% (puffed wheat) of RS and cookies/crackers ever, despite the fact that the data strongly suggest the link
had between 0.2% (oatmeal) and 2.8% (crackers and crisp between butyrate and reduced risk of colonic cancer, studies
bread) of RS. The category of cooked cereal and pastas had examining the link between RS intake and colonic cancer have
the widest range of RS contents between 0.4% (noodles and not been able to prove such a relationship. One reason for this
chow mein) and 11.3% (oats and rolled uncooked). Within may be the importance of RS intake levels. Studies examining
the vegetable category, the lowest RS content was shown for the butyrate production from RS intake have shown significant
sweet corn (cooked/canned) and potatoes (slow-cooked) with increase in fecal butyrate when 22 g of RS was consumed per
0.3% while the highest content was observed for fried potatoes day, but no changes in butyrate or pH levels were observed
with 2.8%. In the legume category, RS content ranged from when 12.5 g of RS was consumed per day. The level of butyrate
0.6% for chickpeas (cooked/canned) to 4.2% for white beans produced may also depend on the type of RS consumed.
(cooked/canned). Another aspect of gut health often linked to dietary fibers is
Comparison of the RS content of different foods and the laxation and fecal bulk. An increase of fecal bulk is thought to
food intake records showed that Americans on average con- be beneficial and is often reported as a potential health benefit
sumed an estimated 38 g of RS per day. In Europe, the intake of different dietary fibers. Increases in fecal mass are important
of RS has been estimated to be between 3 and 6 g per day while for relief of constipation and prevention of diverticulosis and
the intake in developing countries has been estimated to be other disorders such as hemorrhoids. The effect of RS on fecal
between 30 and 40 g per day. bulk has been studied and an increase of fecal output was
reported from different studies. Phillips et al. reported an
average increase in fecal output of 42% and also showed that
Health Benefits of RS the fecal output increased with increased RS intake. This
increase fecal output could be due to increased water in the
Gut Health
fecal material, undigested and unfermented starch and dietary
By definition, RS escapes digestion in the human small intes- fibers, or possible bacteria from the digestive tract. Increased
tine. RS therefore passes into the colon where like other dietary fecal nitrogen was observed in diets high in RS indicating either
fibers, it may be fermented. Fermentation of carbohydrates in an increase in bacterial mass or protein excretion.
the colon leads to the production of SCFAs. SCFAs are the
metabolic products of anaerobic bacterial fermentation and
Metabolic Effects
consist of butyrate, propionate, and acetate.
The SCFAs produced by bacterial fermentation in the Several studies have shown that replacement of digestible car-
human gut are the preferred fuel for the cells lining the colon bohydrate with RS leads to a reduced blood glucose response.
(colonocytes) and have been shown to lower luminal pH, This has formed the basis for approval of a European Food
increase the excretion of bile acids, and help prevent the devel- Safety Authority (EFSA) claim for RS (EFSA, ID 681, April
opment of abnormal colonic cells. A lower pH has been asso- 2011). While this claim was granted, there was insufficient
ciated with protection against colorectal cancer and has been evidence for the effects of RS when the glycemic load (amount
shown to inhibit the conversion of primary to secondary bile of digestible carbohydrate) remains constant. In 2012,
acids. Secondary bile acids are thought to promote tumors as Robertson reviewed several studies on the effect of RS addition
they are cytotoxic to colonic cells. It is therefore reasonable to to the diet when the available carbohydrate load was held
assume that RS may confer health benefits to gut health by constant and noted a significant inconsistency between the
production of SCFA, which in turn have a positive effect on pH studies reviewed. While some found no effect of RS on glyc-
and reduced production of secondary bile acids. A low pH has emia and significant reduction in insulin secretion, others
also been found to suppress the growth of potentially patho- found the opposite, that is, a reduction in glycemia and an
genic organisms and reduce the absorption of toxic com- elevation in postprandial insulinemia. The difference in these
pounds (such as ammonia). results could be due to differences in the RS tested or simply be
Individual SCFAs have also been shown to have more spe- due to the difficult relationship between glucose absorption,
cific actions in the colon. Most RS is readily fermented into clearance, and hepatic release, which makes accurate measure-
SCFA and many are a good source of butyrate. Butyrate is a ments and conclusions difficult. Another important factor to
preferred energy source for colonocytes and helps drive the consider is the sources of RS used in different studies. RS can
uptake of electrolytes and water. Intake of butyrate producing come from foods such as beans, high-amylose corn starch, or
RS as part of oral rehydration solutions has been shown to potatoes, and these foods all have different physiochemical
significantly reduce the recovery time for children with watery properties that can affect the response to RS ingestion. The fat
diarrhea. Butyrate has also been shown to help maintain bar- content of the diets is also important as fat is known to have a
rier function in the gut by enhancing mucin production. The significant impact on the glycemic response to a meal. A careful
barrier function is important to reduce bacterial translocation, review of studies with appropriate fat and protein levels
which may be a key element in the development of diseases showed that RS consumption does indeed lead to a small
such as inflammatory bowel disease (IBD). Intake of RS has decrease in postprandial glycemia and a larger attenuation of
been suggested to be beneficial for the reduction of IBD. postprandial insulinemia. These acute changes in plasma
CARBOHYDRATES | Resistant Starch and Health 233
glucose and insulin concentrations caused by the addition of expect some effect on satiety and/or total energy intake when
RS to the diet could have important metabolic implications. RS is used to replace digestible starch in the diet. Animal
Both hyperglycemia and hyperinsulinemia are affected in the studies have shown the production of satiety hormones
development of insulin resistance. Increases in plasma glucose GLP-1 and PYY when RS was added to the diet, which suggests
concentrations have been linked to increased concentrations of an effect of RS on satiety. However, only a few human studies
free fatty acids. Glucose uptake in the muscles and increased have been conducted to examine the effect of RS on satiety and
glucose uptake in the adipose tissue have been linked to the results were mixed. Some studies have shown an increase in
hyperinsulinemia. satiety after RS consumption, but others have shown little or
The potential chronic effects of RS intake on metabolism no effect. Satiety seems to be linked to changes in plasma
and associated diseases are even more interesting than the glucose concentrations and studies that showed a decrease in
acute effects discussed earlier. There are animal data showing plasma glucose/insulin response to a high RS meal also
a significant relationship of RS inclusion in the diet and showed an increase in satiety while those studies that showed
improved insulin sensitivity. Decreased insulin sensitivity, or no change in plasma glucose also showed no difference in
insulin resistance, has severe metabolic consequences as it is satiety. Overall, there is indication that there is at least a weak
strongly associated with several diseases known as metabolic association between RS consumption and satiety. Some animal
syndrome. Metabolic syndrome describes diseases such as type studies have shown reduced energy intake when RS was added
2 diabetes, obesity, coronary heart disease, hypertension, and to the diet. Aziz et al. showed that body weight was reduced by
dyslipidemia. 40% in obese rats that were fed with a diet high in RS com-
Even if RS only leads to reduction in blood glucose pared with rats fed with a low-RS diet. While this study seems
response when digestible carbohydrates are replaced, such an to indicate a real positive effect of RS on energy intake and
effect may have significant benefits to human health. Diabetes potentially obesity management, the levels of RS used for this
affects about 8% of US population and is increasing globally. study (23.4%) may not be realistic in a human diet. Long-term
Diabetes is characterized by hyperglycemia that can subse- studies in humans with realistic RS inclusion levels are
quently lead to systemic tissue toxicity. Risk factors for diabetes required to get a better understanding of the potential for RS
include increased glucose response (both fasting and postpran- to help in the management and reduction of the current obe-
dial), decreased insulin sensitivity, and obesity. These risk sity epidemic.
factors may be reversible with lifestyle modifications that
have been shown to be effective in delaying the onset of type
2 diabetes. One lifestyle change could be to reduce the glyce- Summary/Conclusions
mic load of the diet that could be achieved by replacing ordi-
nary starch in foods with RS. RS is a dietary fiber and can be used to replace digestible starch
RS has been shown to have significant effects of serum in foods. Many animal studies have been conduced to examine
triglyceride and cholesterol concentrations. Several animal the effect of RS consumption. RS may be fermented in the
studies have shown significantly lower plasma cholesterol and colon and the resulting SCFA production has been linked to a
triglyceride concentrations when RS is included in the diet in number of health benefits such as reduced pH, which has been
replacement of digestible starch. A decrease in fasting choles- associated with the suppression of growth of potentially path-
terol and triglycerides was observed when RS was consumed ogenic organisms and reduction of the absorption of toxic
chronically (514 weeks). The mechanism responsible for compounds. Fermentation of RS has been shown to produce
these effects seems to be increased bile acid excretion caused relatively high levels of the SCFA butyrate that has been linked
by consumption of RS, but some study showing reduced to reduced risk of colon cancer. RS fermentation has also been
plasma cholesterol and triglyceride concentrations even com- linked to benefits in the risk reduction of IBD by strengthening
pared to the bile acid sequestrant cholestyramine suggests that of the gut barrier via increased production of mucin. RS has
the effects of RS extend beyond just bile acid secretion. The been shown to increase fecal bulk, which is important for relief
effects of RS on cholesterol and blood triglyceride levels seem of constipation and prevention of diverticulosis and other
to be mediated through a combination of enhancement of bile disorders such as hemorrhoids.
acid secretion, a significant decrease in cholesterol absorption, An EFSA claim was granted in 2011 for reduced blood
and an increase in hepatic LDL receptor expression. Both high glucose response when RS is used to replace digestible carbo-
blood lipid and cholesterol levels are associated with coronary hydrate in the diet (EFSA, ID 681, April 2011). The effect of RS
heart disease and the effects of RS consumption may therefore on glycemia and insulinemia is less clear, and studies have
have important health ramifications. shown that different effects are possible due to study designs
The effects of RS consumption on satiety and weight man- (e.g., amount of fat and protein in diets) and amount of RS
agement are not as clear. One major reason for the current tested. Overall, it does seem that RS consumption does indeed
global obesity epidemic is the overconsumption of energy, and lead to a small decrease in postprandial glycemia and a more
therefore, strategies that can manage the energy intake may be significant attenuation of postprandial insulinemia that could
successful in reducing obesity. Dietary fiber is one way to have important metabolic implications. Both hyperglycemia
decrease the energy value of foods if it is used to replace and hyperinsulinemia are affected in the development of
digestible carbohydrates that have a higher caloric value. insulin resistance, which in turn has been linked to several
Some dietary fibers have also been linked to increased satiety diseases known as metabolic syndrome, for example, type 2
and lower BMI, but not all fibers have the same effects. RS is a diabetes, obesity, coronary heart disease, hypertension, and
type of dietary fiber and it would therefore be reasonable to dyslipidemia.
234 CARBOHYDRATES | Resistant Starch and Health
RS has been shown to have significant effects on serum of the consumption of resistant starch in man. European Journal of Clinical
triglyceride and cholesterol concentrations. The effects of RS Nutrition 46: S1S148.
Aziz AA, Keeney LS, Goulet B, and Abdel-Aal ES (2009) Dietary starch type affects body
on cholesterol and blood triglyceride levels seem to be medi-
weight and glycemic control in freely fed but not energy-restricted obese rats.
ated through a combination of enhancement of bile acid secre- Journal of Nutrition 139: 18811889.
tion, a significant decrease in cholesterol absorption, and an Berry CS (1986) Resistant starch: Formation and measurement of starch that survives
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Giczewska A and Borowska J (2003) Nutritional value of broad bean seeds. Part 1:
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and insulin responses. Cereal Chemistry 87: 257262.
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Higgins JA (2004) Resistant starch: Metabolic effects and potential health benefits.
More human studies with careful study designs are needed in Journal of AOAC International 87: 761768.
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and chronic inflammatory diseases. International Journal of Colorectal Disease
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Jenkins DJA, Vuksan V, and Kendall CWC (1998) Physiological effects of resistant
See also: Carbohydrates: Carbohydrate Metabolism; Health Effects
starches in fecal bulk, short chain fatty acids, blood lipids and glycemic index.
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Consumer Trends in Grain Consumption; Grains and Health: Keenan MJ, Zhou J, and McCutcheon KL (2009) Effects of resistant starch, a
Misinformation and Misconceptions; Grains and Health; Food Grains non-digestible fermentable fiber, on reducing body fat. Obesity (Silver Spring)
and Well-being: Functional Foods: Overview; Grains Around the 14: 15231534.
Knowler WC, Barrett-Connor E, Fowler SE, et al. (2002) Reduction in the incidence of
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EURESTA: European FLAIR concerned action no. 11 on physiological implications Chemistry and Technology, pp. 575590. New York: Academic Press.
CARBOHYDRATES | Resistant Starch and Health 235
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Nichols BL, Avery SE, Sen P, Swallow DM, Hahn D, and Sterchi EE (2003) The maltase- Cereal Chemistry 68: 8691.
glucoamylase gene: Common ancestry to sucrase-isomaltase with Silvester KR, Englyst HN, and Cummings JH (1995) Ileal recovery of starch from whole
complementary starch digestion activities. Proceeding of the National Academy diets containing resistant starch measured in vitro and fermentation of the ileal
of Sciences in the United States of America 100: 14321437. effluent. American Journal of Clinical Nutrition 62: 4034113.
Nugent A (2005) Health properties of resistant starch. Nutrition Bulletin 30: 2754. Smirnov A, Perez R, Amit-Romach E, Sklan D, and Uni Z (2005) Mucin dynamics and
Pencek RR, Koyama Y, and Lacey DB (2002) Transporter-mediated absorption microbial populations in chicken small intestine are changed by dietary probiotic
is the primary route of entry and is required for passive absorption of intestinal and antibiotic growth promoter supplementation. Journal of Nutrition
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Phillips J, Muir JG, Birkett A, Lu ZX, Jones GP, and ODea K (1995) Effect of resistant Swallow DM, Poulter M, and Hollox EJ (2001) Intolerance to lactose and other dietary
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carbohydrates. European Journal of Clinical Nutrition 46(supplement 2): S77S90. supplementation: Effects on luminal, inflammatory, epigenetic, and epithelial
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Health Effects of b-Glucans Found in Cereals
SM Tosh, Agriculture and Agri-Food Canada, Guelph, ON, Canada
S Shea Miller, Agriculture and Agri-Food Canada, Ottawa, ON, Canada
2016 Elsevier Ltd. All rights reserved.
Figure 1 Fluorescence micrographs of glycol methacrylate sections of cereals. (a), (c), (e) and (g) stained with Congo red to show b-glucan (red)
in the endosperm and aleurone cell walls, as distinguished from the autofluorescence of phenolics (blue) in the aleurone cell walls (oriented at
the top of each micrograph). (b), (d), (f), and (h) stained with calcofluor white to demonstrate b-glucan in the cell walls of the central endosperm
(bluewhite fluorescence). Bars in (a)(c), (d), and (g) represent 20 mm. Bars in (e), (f), and (h) represent 10 mm. (a) and (b) oat; (c) and
(d) barley; (e) and (f) rye; (g) and (h) wheat.
Table 1 Content and characteristics of mixed-linkage b-glucan found in major cereal grains
Chemical Structure cellulose; thus, it is much more soluble in water than cellulose
(Figure 2).
Mixed-linkage b-glucan is a linear homopolymer of glucose, Of all the cereals, the b-glucan in oats tends to have the
similar in structure to cellulose ((1 ! 4)-b-D-glucan). However, longest molecules (highest molecular weight). In the native
there are single b-(1 ! 3) linkages interspersed with the grain (Table 1), the molecular weight is generally around
b-(1 ! 4) linkages that make up about 30% of the linkages. 22.5 million g mol1 (12 000 glucose units). Approximately
Because the b-(1 ! 3) linkages are not regularly spaced, 40% of the b-glucan in uncooked oatmeal, oat bran, or oat
b-glucan does not form crystalline structures and fibrils like flour is soluble in water at 37 C in the presence of digestive
238 CARBOHYDRATES | Health Effects of b-Glucans Found in Cereals
Cellotetraosyl
unit Cellotriosyl
unit
Figure 2 Schematic diagram depicting the chemical structure of a portion of a b-glucan molecule.
4
Health Benefits
Cholesterol Reduction
3
It has been well established that daily consumption of soluble 0 20 40 60 80 100 120
b-glucans can reduce total and LDL cholesterol. A recent meta- Time (min)
analysis determined that persons with mildly elevated serum Figure 3 Glycemic response to oat bran muffin containing 4 g of
cholesterol levels (3.05.0 mmol l1) who consumed at least b-glucan and whole-wheat muffin containing <0.1 g of b-glucan. After
3 g of oat b-glucan per day experienced an average reduction in consumption of the oat bran muffin, blood glucose levels increased
LDL cholesterol of 0.240.29 mmol l1. Similar benefits have more slowly, peaked at a lower level, and returned to baseline
been demonstrated with the consumption of barley b-glucan. more slowly than after consumption of the whole-wheat muffin.
Barley and b-glucan isolated from barley lowered LDL-
cholesterol concentrations by 0.210.39 mmol l1 compared enzymes with food, inhibiting the release of glucose from
with control. Health claims are permitted on food products starch. Intact cell walls in cooked whole kernels (including
containing physiologically significant levels of b-glucans from pearled and pot barley), steel-cut oat, and milled flakes also
oats or barley in the United States, Canada, the European protect starch from digestive enzymes.
Union, and Malaysia. Sugars, inherent in a meal or released from starch, take
The viscosity generated by the b-glucan in the gut increases longer to reach the intestinal wall in a high viscosity mass
excretion of cholesterol and bile acids. Bile acids are released than in low viscosity environment because of the slow mixing
into the small intestine to emulsify fat in ingested food. Nor- and diffusion rates. Therefore, glucose enters the bloodstream
mally bile acids are reabsorbed lower down in the digestive at a slower rate and over a longer section of the small intestine.
tract and recycled, but viscous fibers tend to physically entrap The protracted release of glucose into the bloodstream results
them and carry them out of the body. To replenish what was in a beneficial reduced insulin response; repeated spikes in
lost, in preparation for the next meal, the liver converts stored insulin concentrations increase the risk of developing glucose
cholesterol into bile acids, thus lowering the total cholesterol intolerance that in turn may lead to type 2 diabetes.
levels.
A A
* *
A
A
*
(a) (b) (c)
Figure 4 In vitro digestion and fermentation of oat bran. (a) Suspension of oat bran in buffer, stained with calcofluor to demonstrate b-glucan. Note
sheets of aleurone cells (A) and endosperm cell walls (*). (b) Suspension of oat bran after in vitro digestion with a-amylase and proteases to mimic
digestive enzymes, stained with calcofluor. Note that aleurone cell wall sheets (A) are unchanged by digestive enzymes but that the b-glucan in the
endosperm cell walls has been solubilized and appears as an amorphous blue mass (*), although some endosperm cell wall structure is visible (arrow).
(c) Suspension of oat bran/feces after in vitro fermentation, stained with calcofluor. The b-glucan has been completely fermented by the fecal flora.
Some autofluorescence of phenolics is visible in vascular elements from other plant-based foods (arrow), as well as in small fragments of blue cell walls
embedded in the larger fecal particle. Bars in (a)(c) represent 100 mm.
for studies measuring changes at a subsequent meal have been Oat and barley b-glucans are not absorbed into the blood-
mixed. Some studies observed a significant drop in calories stream. How are they able to influence concentrations of
consumed while others did not. glucose and cholesterol in the blood?
b-Glucan is a dietary fiber. What is the definition of dietary
fiber? What functions do soluble and insoluble dietary fib-
Fermentation and Laxation ers play in maintaining health?
Oat and barley fiber are not broken down by enzymes in the If wheat contains b-glucan, why does it not reduce
upper intestine and arrive in the colon intact. Intact oat bran cholesterol?
contains particles of aleurone and endosperm cells, which are
easily seen with the fluorescence microscope due to the auto-
fluorescence of phenolics in the aleurone layer and the blue Exercises for Readers to Explore the Topic Further
fluorescence of b-glucans stained with calcofluor (Figure 4(a)).
After an in vitro digestion with a-amylase and proteases meant How do different types of food processing affect the effec-
to mimic conditions in the upper GI tract, the b-glucans have tiveness of b-glucan? Does it matter whether oats and barley
been mostly solubilized, as demonstrated by the loss of dis- are consumed in extruded breakfast cereals, cooked cereals,
cernible cell wall structures and the appearance of amorphous breads, soups, snack bars, desserts, or beverages?
blue-staining material in Figure 4(b), but are still present in the How can consumers be encouraged to increase their intake of
digesta. Because of its solubility, b-glucan is susceptible to oat and barley foods to take advantage of their health benefits?
enzymes produced by gut microbiota and rapidly fermented, Oat and barley foods have been reported to increase and
as demonstrated by the loss of calcofluor-stained material in the extend satiety after a meal? Is the satiety effect related to b-
digesta (Figure 4(c)). glucan or glycemic response modulation?
During fermentation of b-glucan, gut bacteria produce
increased levels of butyrate, propionate, and acetate, the
short-chain fatty acids associated with improved gut health See also: Carbohydrates: Carbohydrate Metabolism; Cereals:
(and possibly satiety as well, in the case of acetate). Chemistry and Physicochemistry of Non-starchy Polysaccharides;
Oat bran improves laxation mainly because of the insoluble Glycemic Index, Glycemic Load, and Their Health Benefits; Food
fiber portion. Wheat bran is more effective at promoting laxa- Grains and the Consumer: Grains and Health; Food Grains and
tion because of its high insoluble fiber content. Well-being: Functional Foods: Overview; Processing of Grains:
b-Glucans: Measurement and Processing; The Cereal Grains:
Oats: Overview.
Exercises for Revision
Where are b-glucans mainly found? Name the plant family. Further Reading
Which food ingredients are the main sources?
Abdel-Aal EM and Wood PJ (eds.) (2005) Specialty Grains for Food and Feed. St. Paul,
Draw a schematic diagram of a b-glucan molecule includ-
MN: AACC International.
ing two tetrasaccharide and two trisaccharide segments. AbuMweis SS, Jew S, and Ames NP (2010) b-Glucan from barley and its lipid-lowering
Identify the 1 ! 3 and 1 ! 4 linkages. capacity: A meta-analysis of randomized, controlled trials. European Journal of
What are the linkage differences between cereal b-glucans, Clinical Nutrition 64(12): 14721480.
cellulose, and starch? What are the implications for struc- Aman P (2006) Cholesterol-lowering effects of barley dietary fibre in humans: Scientific
support for a generic health claim. Scandinavian Journal of Food and Nutrition
ture and physiological effect? 50(4): 173176.
What four health benefits have been attributed to oat and Anderson JW (1987) Dietary fiber, lipids and atherosclerosis. American Journal of
barley b-glucans? Cardiology 60: 17G22G.
240 CARBOHYDRATES | Health Effects of b-Glucans Found in Cereals
Braaten JT, Wood PJ, Scott FW, et al. (1994) Oat b-glucan reduces blood cholesterol Truswell AS (2002) Cereal grains and coronary heart disease. European Journal of
concentration in hypercholesterolemic subjects. European Journal of Clinical Clinical Nutrition 56: 114.
Nutrition 48: 465474. Webster FH and Wood PJ (eds.) (2011) Oats: Chemistry and Technology, 2nd edn. St.
Chu Y-F (ed.) (2013) Oat: Nutrition and Technology. Chichester: Wiley. Paul, MN: AACC International.
El Khoury D, Cuda C, Luhovyy BL, and Anderson GH (2012) Beta glucan: Health Whitehead A, Beck EJ, Tosh SM, and Wolever TMS (2014) Cholesterol-lowering effects
benefits in obesity and metabolic syndrome. Journal of Nutrition and Metabolism, of oat beta-glucan: A meta-analysis of randomized controlled trials. The American
Article number 851362. Journal of Clinical Nutrition 100: 14131421.
Kale M, Hamaker B, and Bordenave N (2013) Oat b-glucans: Physicochemistry and Wood PJ (2010) Oat and rye b-glucan: Properties and function. Cereal Chemistry
nutritional properties. In: Chu Y-F (ed.) Oat: Nutrition and Technology. Chichester: Wiley. 87(4): 315330.
Newman RK and Newman CW (2008) Barley for Food and Health: Science, Technology, Wood PJ (2011) Oat b-glucan: Properties and function. In: Webster FH and Wood PJ
and Products. St. Paul, MN: AACC International. (eds.) Oats: Chemistry and Technology, 2nd edn., pp. 219254. St. Paul, MN:
Othman RA, Moghadasian MH, and Jones PJH (2011) Cholesterol-lowering effect of oat AACC International.
b-glucan. Nutrition Reviews 69(6): 299309. Wood PJ, Arrigoni E, Shea Miller S, and Amado R (2002) Fermentability of oat and
Shewry PR and Ullrich SE (eds.) (2014) Barley: Chemistry and Technology, 2nd edn. wheat fractions enriched in b-glucan using human fecal inoculation. Cereal
St. Paul, MN: AACC International. Chemistry 79: 445454.
Tosh SM (2013) Review of human studies investigating the post-prandial blood- Wursch P and Pi-Sunyer FX (1997) The role of viscous soluble fiber in the metabolic
glucose lowering ability of oat and barley food products. European Journal of control of diabetes: A review with special emphasis on cereals rich in b-glucan.
Clinical Nutrition 67: 310317. Diabetes Care 20: 17741780.
Glycemic Index, Glycemic Load, and Their Health Benefits
D Dan Ramdath, Agriculture and Agri-Food Canada, Guelph, ON, Canada
2016 Elsevier Ltd. All rights reserved.
Topic Highlights their effects on blood glucose levels. In 1981, the glycemic (or
glycaemic) index (GI) was proposed as a numeric physiologi-
Carbohydrates are required as part of a healthy diet, but cal basis of classifying carbohydrate foods in order to improve
they vary in the rate at which they increase blood glucose blood glucose control in persons with diabetes. Studies that
following ingestion. preceded this concept demonstrated that different carbohy-
Glycemic index (GI) is a numeric physiological basis of drate foods raised blood glucose by varying amounts, which
classifying carbohydrate foods initially used to improve could not be predicted from chemical analysis. Indeed, differ-
blood glucose control in persons with diabetes. ences were measured among foods belonging to the same food
Glycemic load is the product of the carbohydrate content group. It is now widely accepted that the blood glucose raising
per serving of food and its GI. potential of carbohydrate-rich foods can be ranked in a stan-
Measurement of GI requires experienced users and a dardized matter using GI, and this feature can be used as an
method that is accurate and valid. adjunct in the dietary management of persons with diabetes.
Glycemic response is the blood glucose response following In 1997, a joint Food and Agriculture Organization/World
consumption of digestible carbohydrate and is measured as Health Organization Expert Consultation defined GI as the
the area under the blood glucose response curve. incremental area under the blood glucose response curve
Several food characteristics may affect derivation of GI (AUC) elicited by a 50 g available carbohydrate (AC) portion
values. of a test food expressed as a percentage of the response elicited
Regular consumption of diets that are low in GI and GL is by the same amount of AC from a standard food (either white
associated with lower risk for type 2 diabetes (T2DM) and bread or glucose) taken by the same person. GI is therefore a
cardiovascular disease (CVD). measure of how much a standard quantity of a food raises
blood glucose compared with a standard quantity of glucose
or white bread, with a value of 100 having been assigned to
Learning Objectives glucose. In practical terms, GI is derived from repeated testing
of a carbohydrate food in at least ten research participants, with
the reference food tested at on 2 occasions (see eqn [1]).
To describe the different types of carbohydrates in the
human diet and their effects on blood glucose. Although the significance of GI in managing persons with
diabetes is widely accepted, there have been numerous debates
To describe the concept of glycemic index and glycemic
load and their role in management of diabetes and health on the derivation of GI, including the accuracy and precision of
promotion. the method and its universal application to public health
promotion campaigns to guide healthy food choices:
To understand the derivation, required methodology, and
classification of GI.
Glycemic index food AUCfood =mean AUCreference 100 [1]
To calculate the GI and GL of a mixed meal.
To understand the food characteristics that may affect der- The derivation of GI is based on the amount of available (or
ivation of GI values. preferably glycemic) carbohydrates present in a carbohydrate-
To outline the difference between GI, GL, and glycemic rich food. GI compares equal quantities of carbohydrate and
response. provides a measure of carbohydrate quality but not quantity. It
To examine human studies on the role that GI and GL has been argued that GI is a property of carbohydrate-rich
might play in reducing risk for T2DM and CVD, and pro- foods, which is independent of age, gender, body mass index,
moting satiety and weight loss. ethnicity, and glucose tolerance status but may be influenced
by factors such as food variety, cultivars, processing,
preparation, and storage effects, among others. Using stan-
Introduction dardized methodology, the GI of carbohydrate-rich foods can
be assessed with a standard deviation of about 9, and the
Carbohydrates provide approximately 5060% of energy resulting values can be classified (with glucose as the standard)
intake in the normal human diet, and this requirement is met as high (GI 70), medium (GI 5669), or low (GI 55). See
through regular consumption of starchy foods such as cereals, Table 1 for a sample listing of GI values of different foods.
fruits, and vegetables. Following consumption, carbohydrates Diets containing high amounts of refined carbohydrates
are progressively broken down by digestive enzymes in the have been shown to increase plasma levels of triglyceride and
gastrointestinal tract where degraded starch structures give reduce high-density lipoprotein. Assessing this type of relation-
rise to glucose, which is then actively transported to various ship in human studies required an understanding of carbohy-
tissues to meet cellular energy requirements. About four drate quality (i.e., GI) and the quantity eaten in a meal or
decades ago, it was recognized that carbohydrates differ in during the course of the day. The concept of glycemic load
the rate at which they are digested and consequently vary in (GL) was therefore introduced in 1997 by researchers at
Apple, Golden Delicious 39 3 French fries, baked (OreIda) 64 6 Baked beans in tomato sauce 40 5
Banana (Canada) 62 9 Mashed potato, instant (Canada) 80 13 Chickpeas, canned 38 3
Cherries, dark, raw 63 6 Parsnips, peeled, boiled 52 5 Chickpea hummus 22 5
Kiwi 58 7 Potato, Irish, peeled, baked 83 6 Kidney beans, dried, soaked overnight, boiled 45 min 51 5
Mango 51 3 Potato, Irish, boiled 59 4 Lentils, brown, canned 42 5
Orange (South Africa) 33 6 Sweet potato, peeled, boiled 46 5 Lentils, red, dried, boiled 21 7
Papaya 56 6 Taro (China) 48 5 Mung bean, germinated 25 4
Pineapple 66 7 White yam, boiled (Jamaica) 79 4 Soya beans, canned (Canada) 14 2
Ice cream, vanilla (Sara Lee) 38 3 All-Bran (Canada) 51 5 Chips (Doritos original) 42 4
Milk, full-fat (Canada) 34 6 Cornflakes (Canada) 80 6 Chips, potato, plain, salted 51 7
Milk, skim (Canada) 32 5 Rice, Basmati, boiled 8 min 69 6 Chocolate, dark, DoveW 23 3
Soy milk, VitasoyW organic 43 3 Rice, brown 66 5 Chocolate, milk, DoveW 45 8
Yogurt, white peach (Finest) 34 7 Rice, sticky 87 2 Jelly beans (Allens) 80 8
Yogurt, Greek style, plain 12 4 Rolled oats, cooked 20 min 49 8 Mars BarW 62 8
Yogurt, frozen, vanilla Blue Menu 46 5 Spaghetti, white, durum wheat 58 7 Mixed nuts, roasted, salted 24 10
Yogurt, natural, fat-free 19 5 Wonder, white bread 72 4 Sports bar, Clif barW, Brownie 57 6
Harvard University to quantify the overall glycemic effect of a been included in the current International Organization for
portion of food (see eqn [2]). GL is defined as the product of Standardization methodology (ISO 26642:2010) for deter-
the carbohydrate content per serving of food and its GI. As mining the GI of foods. This well-defined set of guidelines
such, dietary intakes of carbohydrates can be weighted to pro- include mandatory procedures, recommendations for charac-
vide the GL, which is a summative value of the quantity of a teristics of study participants and pretesting preparation, and
carbohydrate-rich food ingested and the quality of the food internal validity checks, such as the use of a method to measure
and allows for a better understanding of how different-sized blood glucose with an analytic variation of <3% and exclusion
portions of carbohydrate foods compare in terms of their of outliers.
blood glucose raising effect. The higher the GL, the greater Typically, the ISO GI method calls for a minimum of ten
the expected elevation in blood glucose and in the insulino- healthy research participants who, following an overnight fast,
genic effect of the food. Foods can be classified as having GLs are randomly fed a 50 g AC portion of the test or reference food
that are low (GL 10), medium (10 < GL < 20), or high (white bread or glucose) using a randomized block design.
(GL 20). Before consumption of the test meal, two fasting blood sam-
ples are obtained via the finger prick method, and additional
GIfood available carbohydrate g=serving
Glycemic loadfood blood samples are collected at 15, 30, 45, 60, 90, and 120 min
100
after the first bite is taken. Persons with type 2 diabetes
[2]
(T2DM) or glucose intolerance can be included as research
participants, in which case additional blood samples are
taken at 150 and 180 min. Finger prick capillary blood is
preferred because (i) it is easier to obtain, (ii) the rise in
GI Derivation and Methodology blood glucose is greater than in venous plasma, and (iii) results
for capillary blood glucose are less variable than those for
The method for derivation of GI has evolved considerably over venous plasma. As such, differences in the blood glucose
the last two decades with many improvements in the tech- response between foods are larger and easier to detect statisti-
niques and prerequisites for testing. In an attempt to standard- cally using capillary blood glucose. Changes in the blood
ize and improve the precision of the method used to determine glucose concentrations above fasting are calculated as the
GI, two multilaboratory studies were undertaken with each lab incremental area under the glucose response curve using the
analyzing the same foods. These studies concluded that varia- trapezoid rule. The reference food is eaten by each research
tion in the GI values obtained by established laboratories participant on at least two separate occasions to derive a mean
could be accounted for by differences in data analysis and AUC for that individual. The AUC of each participant after each
within-subject variation of glycemic responses. Several test food is expressed as a percentage of the mean AUC elicited
improvements in the methodology were suggested, including by the reference food in the same participant. Next, the mean
duplicate measurements of fasting blood glucose, use of capil- of these values for all participants is calculated as the food GI. If
lary blood sample, and pretest restrictions on exercise and white bread is used as the reference food, the GI values are
alcohol consumption. Many of these improvements have multiplied by 0.71 to convert them to the glucose scale
CARBOHYDRATES | Glycemic Index, Glycemic Load, and Their Health Benefits 243
(i.e., the GI of glucose 100). The concept of GI testing and before the existence of a standardized method for GI and
derivation is illustrated in Figure 1. Given the care required to that in some instances, the GI values of the same food item
produce accurate and valid GI value, it is advisable that new have relatively large variation. Some of this variation could be
investigators adhere closely to the ISO method and perform a attributed to foods being tested in 10 persons; however,
minimum of three tests on the reference food. differences in GI values of the same food category may also
From the protocol outlined in the preceding text, it be explained by differences in processing, in determining
becomes clear that GI is not a simple ratio of two values, but the digestible carbohydrate content of the test foods, in GI
rather a summation of 3 or 4 values from each volunteer and testing, inherent botanical differences and growing conditions,
may be expressed mathematically as etc. Indeed, derivation of GI values is affected by many food
Xx1 factors (see Table 2), and this has been the focus of many
x
100 Fx =G disagreements on the relevance and application of GI to
GI n
[3]
n human health.
where Fx is the incremental blood glucose response (AUC) in
participant x elicited by 50 g AC from the test food and Gx is the
mean AUC in participant x elicited by 50 g glucose (or the Glycemic Carbohydrates
equivalent white bread) tested on two or three separate occa-
sions. In the hands of experienced investigators, this approach An imperative of GI testing is that both the reference foods and
with 12 healthy adult volunteers usually provides a power of test foods must contain the same amount of available or glyce-
80% to detect a 22% difference in AUC. Further, among 28 mic carbohydrate; typically 50 or 25 g portions are consumed by
centers participating in an interlaboratory study on GI meth- participants. The method used for determining available
odology, the between-laboratory standard deviation of the GI
values was 9.
Table 2 Food characteristics that may affect derivation of GI values
In order to reduce unnecessary repetition in the testing of
individual foods and to promote wider research and applica- Food factor Effect on GI
tion, an international table of GI and GL values was compiled
from the scientific literature in 1995 and updated in 2002 and Presence of monosaccharides Formulations with glucose will have
2008 to contain 2487 separate entries from 205 published Glucose; fructose; galactose higher GI than those containing
articles. Further, GI values derived from studies of 5 persons sucrose or high-fructose corn
syrup
are listed separately, and caution is given in using these values.
This resource, a sample of which is shown in Table 1, has been Type of starch Amylopectin has branched structure
instrumental in improving the quality of research examining Relative abundance of containing a1-6 linkages, which is
the relation between GI, GL, and health. It has been used by amylose and amylopectin digested faster and has higher GI
researchers in many human cohort and interventional studies Storage Many fruits and vegetables undergo
and in supporting dietary guidelines for high-fiber, low-GI Ripening ripening during which starch
foods for individuals with diabetes as a means of improving Moisture content decreases and sugars
postprandial glycemia and weight control. However, others Temperature increase
have been more reluctant to putting advice about GI values Retrogradation Moisture influences retrogradation
into practice over concerns about potentially adverse effects on and formation of resistant starch
Cold storage after cooking can
food choice and fat intake. Limitations in the use of the
promote recrystallization of the
international GI table include the fact that it was compiled starch chains
Processing Different cultivars often have
Starch particle size different starch structures
Cell wall integrity Large starch granules have larger
surface areas and higher GI
10
50 g glucose Disruption of cell wall during
Plasma glucose (mmol l1)
standard processing
Cooking Differences in the rate of
8 50 g AC Same product with different carbohydrate digestion and hence
Test food ingredients the GI value
Ingredients processed Degree of starch gelatinization and
6 differently consequently the GI values
Recipe variation; flour used,
moisture content, cooking
time
4
Other food components Soluble viscous fiber associated
Dietary fiber and resistant with lower GI
0 30 60 90 120 starch Presence of a-glucosidase inhibitors
Time (min) Antinutrients Delayed gastric emptying
carbohydrate is therefore critical. In fact, there have been sugges- measure routinely. There are suggestions that it may be more
tions that glucose should be used as the only reference food accurate to quantitate glycemic carbohydrates directly, rather
because of the ease of measuring 50 g of glucose. AC is defined than by difference; however, this measurement is only avail-
as the fraction of carbohydrate that can be digested by human able in specialized laboratories.
enzymes, is absorbed, and enters into intermediary metabolism.
It does not include dietary fiber, defined as intrinsic plant cell GI Versus Glycemic Response
wall polysaccharides, which can be a source of energy only after
colonic fermentation in the large intestine. It has been recom- One of the central issues driving the debate on the derivation
mended that the term AC be replaced by glycemic and application of GI to human health is possibly an appreci-
carbohydrate. Using this terminology, glycemic carbohydrates ation of the difference between glycemic response and GI. As
will include most monosaccharides and disaccharides, some shown in Figure 1 and outlined in eqn [3], a test or reference
oligosaccharides (maltodextrins), and rapidly digested starches. food will produce an individual glycemic response, defined as
Slowly digested starches, which are digested less rapidly to release the blood glucose response following consumption of digest-
glucose, are also considered to be glycemic carbohydrates. In ible carbohydrate and measured as the AUC. As outlined
contrast, nonglycemic carbohydrates include oligosaccharides, before, the food GI obtained in an individual is derived by
nonstarch polysaccharides (defined as macromolecules consist- dividing the test food AUC by the mean glycemic response
ing of a large number of monosaccharide residues joined to each obtained from 2 or 3 tests of the reference food in the same
other by glycosidic linkages), and resistant starch. Resistant individual. The food GI is then taken to be the average of the GI
starch (RS) is defined as the sum of starch and products of starch obtained in at least ten individuals. This approach reduces
digestion (such as maltose, maltotriose, and a-limit dextrins) variation and provides a more precise estimate of the true
that are not absorbed in the small bowel and can be classified mean. Studies have shown that when healthy or diabetic vol-
into four types: RS I, physically inaccessible starch mostly present unteers repeatedly consume carbohydrate foods (e.g.,
in whole grains; RS II, RS granules (B-type starches); RS III, spaghetti, rice, and bread) on different occasions, significant
retrograded starch (after food processing); and RS IV, modified variation in their glycemic response is observed, but mean GI
starches. values remain similar. Figure 2 probably best illustrates the
It is possible that earlier GI studies relied on generic food- difference between glycemic response and GI and the effect of
composition tables, data from food manufacturers, or even individual variation on the derivation of these indices. Follow-
food labels, which are insufficient to obtain an accurate mea- ing the consumption of 50 g AC portions of five different foods
surement of glycemic carbohydrate. Accurate laboratory mea- (sucrose, instant potato, bread, rice, and barley) by persons
surement of glycemic carbohydrate should be undertaken prior with varying levels of glucose tolerance, significant differences
to embarking of GI testing. Typically, this involves analysis of in their blood glucose response were observed, but the
moisture, fat, ash, and protein from which total carbohydrate between-individual variation in GI of the foods studied
P
is obtained as 100 ash protein moisture fat. In remained relatively constant. As such, it may be concluded
addition, measurement of total dietary fiber is required. that GI is a property of the food, whereas the glycemic response
Glycemic carbohydrate is then calculated as [total carbohy- is a variable individual characteristic.
drate by difference total dietary fiber]. This approach does The ecological validity of dietary GI and GL has been
not take into account resistant starch, which is difficult to assessed in a randomized trial of lifestyle modification in
Partitioning of variance
12 AUC
1200 Within-subjects
Variance (106)
Between-subjects GI
Glycaemic index (%)
Food
AUC (mmol min l1)
4
800
0
AUC GI
120
400 80
40
0 0
1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31
Subject ID number
Figure 2 Comparison of variance associated with AUC and GI in persons with varying glucose tolerance.
Incremental AUC and estimates of GI in 31 study participants (9 normal, 12 hyperinsulinemic, and 10 with type 2 diabetes) elicited by 50 g avCHO
portions of five different carbohydrate foods (instant potato, white bread, spaghetti, barley, and sucrose); values are mean s.d. for the five test meals in
each participant. AUC differs significantly among participants, but GI does not. Inset: the amounts of variance in AUC and GI due to the different foods,
the different study participants (i.e., between individuals), and day-to-day variation (within participants) from repeated-measures analysis of variance.
Expressing results as GI reduced variation between participants by 95%. Figure re-drawn using data derived from Lan-Pidhainy X and Wolever TMS
(2011) Are the glycemic and insulinemic index values of carbohydrate foods similar in healthy control, hyperinsulinemic and type 2 diabetic patients?
European Journal of Clinical Nutrition, 65, 727734.
CARBOHYDRATES | Glycemic Index, Glycemic Load, and Their Health Benefits 245
free-living overweight and obese individuals with T2DM, similar names (e.g., bread, cookies, and cakes) may have dif-
using continuous glucose monitoring to measure the glyce- ferent ingredients (e.g., type of flour used) or may have used
mic response to foods. In this study, participants consumed different processing methods (e.g., different cooking times
foods in self-selected amounts, at self-selected times, and in resulting in different moisture contents), which could influ-
self-selected combinations. Participants kept 3-day dietary ence the degree of starch gelatinization and rate of digestion
diaries from which intakes of energy, fat, protein, carbohy- and ultimately the GI value.
drates, sugar, and fiber were obtained and GI and GL calcu- Starch type and structure and relative abundance of amylose/
lated. After controlling for energy and carbohydrate intakes, amylopectin in different varieties of rice and potato can influ-
dietary GI accounted for 1018% of the variance in the ence their GI. Amylose is a form of starch with a long, straight
glycemic response indices assessed. Further, when dietary chain structure, whereas amylopectin is highly branched and
intake was adjusted for energy, GL and GI were significantly allows easier access by the digestive enzymes and is therefore
correlated with the glycemic response. The authors concluded associated with higher GIs. Various factors affect the amylose
that GI was the strongest and most consistent independent and amylopectin content of foods; many of these influence
predictor of glycemic response and variability. Less invasive retrogradation and resistant starch formation and include mois-
studies have also shown that the predominant determinants ture, cooking and storage temperature, reheating, and ripening.
of the glycemic response to a mixed meal are the amount of These factors are discussed in detail in other articles. Many fruits
carbohydrate and meal GI, which explain 90% of the varia-
tion in mean AUC. In fact, the glycemic response is depen-
dent on the relative amounts of carbohydrates and the
amounts and types of fat and protein in the mixed meal. Table 4 Glycemic index of a sample of relatively unprocessed foods
For example, it has been shown that addition of fat to a from different countries
potato test meal reduces the glycemic response by about
Food item Country GI (mean SEM if available)
50%. Other factors affecting glycemic response include nature
of monosaccharide components, starch structure and relative White flour The United Kingdom 59 11
abundance of amylose and amylopectin, degree of ripening of Canada 69 5
fruits and vegetables, and cooking and processing. It is there- South Africa 71 7
fore reasonable to conclude that eating a food as part of a Italy 89 12
mixed meal alters the glycemic response, but not its GI; this Rice India 43
China 83 1
differentiation therefore allows for the calculation of the GI
Canada 72 9
of a meal containing several carbohydrate foods as outlined
Banana Australia 47 5
in Table 3. Canada 62 9
South Africa 70 5
Lentils Canada 29 3
Australia 37 3
Effect of Food Processing on GI/GL The United Kingdom 21 7
Soya beans Canada 15 5
Examination of published data will show that some foods (e.g., Australia 20 3
rice and potatoes) have a wide variation in their GI values Canada 14 2
(Table 4). This is likely due to inherent botanical differences Sweet corn New Zealand 37 12
Australia 48
from country to country, compositional and processing differ-
China 55 2
ences, and possibly methodological differences in deriving GI
The United States 60
values (see Table 2). The majority of available GI values were South Africa 62 5
derived from commercially processed foods, and changes by
the food manufacturers in the ingredients or processing Source: Atkinson FS, Foster-Powell K, and Brand-Miller JC (2008) International tables
methods used will influence the GI. Manufactured foods with of glycemic index and glycemic load values: 2008. Diabetes Care, 31, 22812283.
GI values are based on the glucose scale, using GI of glucose 100. CHO, AC.
Source: Wolever TMS (2013) Is glycaemic index (GI) a valid measure of carbohydrate quality? European Journal of Clinical Nutrition, 67, 522531.
246 CARBOHYDRATES | Glycemic Index, Glycemic Load, and Their Health Benefits
and vegetables undergo ripening during which starch content had significantly higher relative risk of T2DM: 1.19 (CI
decreases and sugars increase; this will produce higher GI. Mois- 1.141.24) and 1.13 (CI 1.081.17), respectively. In contrast,
ture influences retrogradation and formation of resistant starch, a cohort study in eight European countries found no relation-
whereas cold storage after cooking can promote recrystallization ship between GI, GL, and T2DM. This difference may be attrib-
of the starch chains, which delays amylase activity. In this uted to the dietary assessment tools and methods used to assess
regard, potato that was precooked, refrigerated overnight, and dietary intakes but may also reflect the limitations in using the
reheated before consumption was shown to have a significantly international table of GI and GL values. As discussed earlier,
lower GI than potato that was cooked and eaten immediately. inherent botanical differences and growing conditions of crops
Processing of grains and cereals disrupts the outer germ and differences in formulation and composition of manufac-
layer, cell wall integrity, and starch granules, which in turn tured goods may limit the generalizability of GI values derived
influences starch particle size and exposes starch to enzymatic in one geographic region and used in another.
digestion, thereby affecting GI. In the case of finely milled
particles, starch is more available to amylase compared to larger
Randomized Controlled Trials
or intact grains. Similarly, a given food might be processed
differently to produce different products (e.g., whole fruit, There are many limitations in examining dietdisease associa-
whole fruit puree, and fruit juice). During the varied processing tion through epidemiological studies; this is overcome by con-
methods needed to produce these items, there will likely be ducting randomized clinical trials, which is widely accepted as
changes in starch structures, particle size and relative abun- an unbiased assessment of the effectiveness of an intervention.
dance of starch fractions; these will result in different GI values. Numerous RCTs have been conducted to examine the efficacy
The influence of food fiber content on GI is not well of low-GI and low-GL diets in reducing risk factors for T2DM
defined with many studies showing some or no effect. This and CVD and their effects on satiety and weight loss. Over the
relationship is complicated by observations that soluble and past 5 years, several systematic reviews have been conducted in
insoluble fiber appear to have varying effects on GI. In ran- order to assess the overall effectiveness of interventions with
domized controlled trial (RCT), treatment with a low-GI diet low-GI and low-GL diets in persons with diabetes. These meta-
for 6 months resulted in lower HbA1c levels (a valid marker of analyses showed that interventions of at least 4 weeks in dura-
glycemic control) compared with a high-cereal fiber diet. tion consistently resulted in improved glycemic control, with
Examination of dietary interventions in persons with T2DM additional benefit seen when intake of unavailable carbohy-
shows that high-fiber diets improve glycemic control. How- drates is high. Similar reviews of existing RCTs have shown that
ever, it was apparent that those incorporating purified viscous insulin sensitivity is improved by consumption of lower-GI,
fibers more frequently lowered fasting blood glucose and total lower-GL diets containing higher unavailable carbohydrates.
cholesterol than interventions with purified nonviscous fibers. In terms of CVD risk factors, after adjusting for confound-
These effects have been directly attributed to the viscosity of ing factors such as dietary fat intake, low GI was shown to
purified fibers and the resulting effects on digestion and reduce fasting triacylglycerol, which is one of the two major
absorption in the small intestine. lipids found in blood. Conversely, high dietary GI is positively
associated with triglycerides and negatively associated with
HDLcholesterol. When legumes are included in low-GI die-
GI, GL, and Health Outcomes tary interventions, clinically significant reductions in HbA1c
are achieved, along with reductions in blood lipids, blood
Epidemiological Studies
pressure, and body weight. There is inconsistency in the rela-
Examination of epidemiological cohort studies in human tionship between low-GI or low-GL dietary interventions and
populations consistently shows an inverse association between weight loss, although most studies show a nonsignificant trend
dietary whole grains and incident cardiovascular disease that favored these diets. When unavailable carbohydrate is
(CVD). Current guidelines recommend that diets rich in min- taken into account interventions with low GI or low GL, high
imally processed whole grains may lower risk for T2DM. How- legume diets are significantly associated with reductions in
ever, the relationship between GI, GL, and T2DM seems to body weight and are probably linked to an associated reduc-
differ by geographic location. In the United States, a recent tion in energy intake. In fact, preliminary evidence suggests
meta-analysis of three large studies confirmed previous find- that low-GI or low-GL diets may have superior satiety effects,
ings that dietary GI and GL are associated with increased risk of which may influence dietary energy intake.
T2DM. Further, the combination of low-GI or low-GL diets
with high cereal fiber intakes was associated with reduced
relative risk of T2DM. In fact, persons who consumed diets Summary
with high GI or high GL, along with a low cereal fiber intake,
had 40% higher risk of T2DM than persons who consumed GI can be used to classify the blood glucose raising potential of
low-GI or low-GL diets and had a high cereal fiber intake. This carbohydrate-rich foods, but its derivation can be influenced
analysis also showed that obese persons who consumed high- by many food factors. As such, several precautions must be
GI or high-GL diets had a >10-fold higher risk of T2DM. taken in order to obtain valid GI values. This could be achieved
Further, a meta-analysis of ten studies on GI and T2DM and within a regulatory framework where there is scrutiny over the
14 on GL and T2DM, conducted in the United States (n 6), generation of GI values. Widespread utilization of GI in public
Europe (n 4), and 1 each in the United Kingdom, China, and health promotion message has been limited due to concern
Japan, showed that persons with the highest dietary GI and GL that increasing consumption of low-GI or low-GL diets could
CARBOHYDRATES | Glycemic Index, Glycemic Load, and Their Health Benefits 247
lead to increased consumption of total and saturated fats, See also: Carbohydrates: Carbohydrate Metabolism; Cereals:
thereby increasing CVD risk. However, recent evidence indi- Chemistry and Physicochemistry of Non-starchy Polysaccharides;
cates that lower AC intake following reductions in GI was not Grains other than Cereals: Non-starch Polysaccharides; Resistant
accompanied with increased fat intake. Rather, they are associ- Starch and Health; Food Grains and the Consumer: Grains and
ated with improved satiety and lower dietary energy intakes. Health; Grains and Health: Misinformation and Misconceptions; Food
Grains and Well-being: Functional Foods: Dietary Fibers,
Prebiotics, Probiotics, and Synbiotics.
Exercises for Revision
Define glycemic index (GI) and explain the rationale for its
utilization. Further Reading
Define glycemic load (GL) and explain the rationale for its
Atkinson FS, Foster-Powell K, and Brand-Miller JC (2008) International tables of
utilization.
glycemic index and glycemic load values: 2008. Diabetes Care 31: 22812283.
Outline the method and precautions needed to derive the Bhupathiraju SN, Tobias DK, Malik VS, et al. (2014) Glycemic index, glycemic load, and
GI value for a food. risk of type 2 diabetes: Results from 3 large US cohorts and an updated meta-
Explain the derivation of the GI of a mixed meal. analysis. The American Journal of Clinical Nutrition 100: 218232. http://dx.doi.
org/10.39345/ajcn-079533.
Outline the food factors that may influence the GI value of
Fabricatore AN, Ebbeling CB, Wadden TA, and Ludwig DS (2011) Continuous glucose
a food.
monitoring to assess the ecologic validity of dietary glycemic index and glycemic
Discuss the precautions required to achieve accurate and load. The American Journal of Clinical Nutrition 94(6): 15191524.
valid GI values. Food and Agriculture Organization of the United Nations (1998) Carbohydrates in
Compare the findings of epidemiological and interven- Human Nutrition. Rome: Food and Agriculture Organization of the United Nations
tional studies that have focused on the health benefits of Food and Nutrition, Paper 66.
International Organization for Standardization (2010) Food products Determination
low-GI diets. of the glycaemic index (GI) and recommendation for food classification.
International Organization for Standardization, September 23, 2010 (ISO
26642:2010). Available from: http://www.iso.org/iso/catalogue_detail.htm?
csnumber43633 (cited 18 August 2014).
Exercises for Readers to Explore the Topic Further Jenkins DJA, Wolever TMS, Taylor RH, et al. (1981) Glycemic index of foods:
A physiological basis for carbohydrate exchange. American Journal of Clinical
Nutrition 34: 362366.
What additional steps are required to improve the method Lan-Pidhainy X and Wolever TMS (2011) Are the glycemic and insulinemic index
for GI so that it is universally accepted as being valid and values of carbohydrate foods similar in healthy control, hyperinsulinemic and type 2
reproducible? diabetic patients? European Journal of Clinical Nutrition 65: 727734.
Li SS, Kendall CW, Souza RJ, et al. (2014) Dietary pulses, satiety, and food intake: A
What is required to allow the widespread inclusion of GI
systematic review and meta-analysis of acute feeding trials. Obesity (Silver Spring)
values on food labels?
22(8): 17731780. http://dx.doi.org/10.1002/oby.20782.
How can GI values be better used for public health promo- Livesey G, Taylor R, Hulshof T, and Howlett J (2008) Glycemic response and health
tion of healthy eating? A systematic review and meta-analysis: Relations between dietary glycemic
Is there a need for further human trials of low-GI foods? If properties and health outcomes. The American Journal of Clinical Nutrition 87(1):
258S268S.
yes, what outcomes would be relevant?
Mirrahimi A, Chiavaroli L, Srichaikul K, et al. (2014) The role of glycemic index and
What is the evidence that low-GI diets can influence brain glycemic load in cardiovascular disease and its risk factors: A review of the recent
function, cognition, and satiety? literature. Current Atherosclerosis Reports 16(1): 110.
Salmeron J, Manson JE, Stampfer MJ, Colditz GA, Wing AL, and Willett WC (1997)
Dietary fiber, glycemic load, and risk of non-insulin-dependent diabetes mellitus in
women. JAMA 277: 472477.
Sluijs I, Beulens JW, van der Schouw YT, et al. (2013) Dietary glycemic index,
Acknowledgments glycemic load, and digestible carbohydrate intake are not associated with risk
of type 2 diabetes in eight European countries. Journal of Nutrition 143(1):
This article is supported by Agriculture and Agri-Food Canada 9399.
Venn BJ and Green TJ (2007) Glycemic index and glycemic load: Measurement issues
RBPI #1503, RBPI #2834/Agri-Innovation Program #068 with and their effect on dietdisease relationships. European Journal of Clinical Nutrition
Pulse Canada, and #T4.H29.V1 cluster program with Pulse 61(Suppl. 1): S122S131.
Canada. Wolever TMS (2013) Is glycaemic index (GI) a valid measure of carbohydrate quality?
Prof. Thomas Wolever kindly provided Table 3 and Figure 2. European Journal of Clinical Nutrition 67: 522531.
Wolever TMS, Brand-Miller JC, Abernethy J, et al. (2008) Measuring the glycemic index
Emily Padhi assisted with preparation of the other figures and
of foods: Interlaboratory study. The American Journal of Clinical Nutrition 87(1):
tables. S247S257.
Fats
Contents
Lipid Chemistry
Healthy Fats and Oils
Lipid Chemistry
L Day, AgResearch Ltd., Palmerston North, New Zealand
2016 Elsevier Ltd. All rights reserved.
Topic Highlights extrusion. On the other hand, oilseeds are primarily grown for
their oil content. Thus, the composition of the lipids (or oil) in
The major lipid class in cereal grains and oilseeds is various oilseeds is an essential consideration in determining
triacylglycerols. the use of the oil. Lipids are also essential for the nutrition and
The minor complex lipids are phospholipids, glycoglycer- consumer acceptance of finished food products as they are
olipids and sterols. carriers of fat-soluble flavor compounds and vitamins. The
The composition of the lipids (or oil) in oilseeds deter- terminology of lipid chemistry is largely dependent on the
mines its food or non-food applications. extraction conditions, including the solvents used and temper-
Extraction and quantification of different lipid classes are ature, and quantification methodologies.
achieved by solvent selections and various chromatograph-
ic methods.
Supercritical fluid carbon dioxide with or without cosolvent
Classification
is now applied for lipid analyses and for industrial extrac-
tion of oils.
Lipids can be classified based on their physical properties at
Mass spectrometry is increasingly being used for the deter-
room temperature (oils are liquid and fats are solid) or their
mination of lipids.
polarity (neutral lipids include fatty acids (FAs), glycerides, and
Lipids play an important technological role in cereal pro-
sterols and polar lipids include glycerophospholipids (phos-
duction, storage, processing and quality of cereal-based
pholipids) and glyceroglycolipids (glycolipids)). The most com-
foods.
mon classification of lipids is based on their structure simple
Plant lipids contain more essential polyunsaturated fatty
or complex. Simple lipids, composed of FAs and alcohol,
acids than animal fats, and their inclusion in food is posi-
include acylglycerols, ether acylglycerols, sterols and their esters
tive from a nutritional point of view.
that can be hydrolyzed to two different components usually an
alcohol and an acid. Complex lipids include phospholipids,
glycolipids and sphingolipids, which yield three or more differ-
Learning Objective ent compounds on hydrolysis.
The FAs (carboxylic acids) are the main building blocks in
To understand lipid classes in grains and oilseeds, their
lipid structures. Only a small portion of the total lipid fraction
analysis, their technological properties in processing, and
consists of free carboxylic acids, or free fatty acids (FFAs). Most
nutritional values of plant lipids.
of the carboxylic acids in the lipid fraction are found as esters of
glycerol, that is, as triacylglycerols (TAGs), commonly known
as triglycerides (Figure 1).
Introduction The most common TAGs are those with long-chain carbox-
ylic acids (C14C22). The positions of carboxylic acids in TAGs
After protein and carbohydrate, lipids (commonly known as are also important to their properties and utilization. The
fats or oils) are the third most important macronutrient in saturated forms of FAs are the most stable against oxidation.
human and animal diets. Although lipids are relatively minor They also have high melting temperatures in comparison with
constituents in food grains, they play an important technolog- unsaturated FAs with the same number of carbon atoms. In
ical role in the quality of grains upon storage: and in most plant TAGs, unsaturated FAs are predominantly located in the
aspects of processing, including milling, brewing, baking and 2 position. Monoacylglycerols (MAGs) contain only one long-
Glycerol
Ester Compositions and Distribution of Lipids in Cereal
Carbon
Linkage Grains and Oilseeds
Position
H O H O
Cereal Grains
1 H C OH + H O C R H C O C R
O O Lipids are minor components of the cereal grains. Wheat,
2 H C OH + H O C R H C O C R
barley, brown rice, rye and sorghum have lower lipid contents
O O than other cereal crops such as oats, pearl millet, and maize. All
cereal grain lipids are richer in nonpolar lipids than in polar
3 H C OH + H O C R H C O C R
lipids. Wheat contains the highest level of glycolipids among
H H all cereal grains, followed by triticale, rye, and barley; millet,
Glycerol Fatty acid Triglyceride maize, and sorghum lipids contain the lowest glycolipid con-
Figure 1 Triacylglycerol synthesis: R, R0 , R00 are usually long-chain
tent. In general, phospholipids are more abundant in wheat,
alkyl groups and R, R0 , R00 are typically all different and may also contain triticale and rye lipids and slightly lower in the lipids of barley,
one or more carboncarbon double bonds. maize, oat, sorghum and rice. Approximate lipid contents and
the proportions of nonpolar lipids and polar lipids (separated
into glycolipids and phospholipids) in each cereal are listed in
Table 1 General classification of lipids and their commonly used Table 2.
abbreviations TAGs are the major nonpolar lipids in cereal grains, repre-
senting from 50% of the total nonpolar lipids in wheat and
Lipid class Abbreviation
barley to as high as 90% in maize. They are deposited in
Simple lipids: compounds with two types of structural spherosomes (oil droplets) bound by a monolayer membrane
moiety and are usually stored by plants in this form. The remainder of
Glyceryl esters: esters of glycerol and fatty acids (see the nonpolar lipids are mainly di- and monoglycerides, FFAs,
Figure 1) and sterol esters. The major phospholipids in cereal grains are
Triacylglycerols: 3 fatty acids TAGs phosphatidylcholine (PC), phosphatidylethanolamine (PE),
Diacylglycerols: 2 fatty acids at any 2 positions DAGs and phosphatidylinositol (PI). Monoacyl PLs, or lysophospho-
Monoacylglycerols: 1 fatty acid at any position MAGs lipids (prefix L added to LP abbreviation, Table 1), are usually
Sterol esters: esters of sterol and fatty acids SEs regarded as degradation products of PL. The GLs are quantita-
Waxes: esters of long-chain alcohols and fatty acids
tively the major components of the glycolipids in whole cereal
Complex lipids: compounds with more than two types
of structural moiety
grains and starchy endosperm. The principal sugar in the GLs is
Phospholipids: glycerol esters of fatty acids, galactose, and glucose is either a minor component or absent.
phosphoric acid, and other groups containing The main galactosylglycerides are monogalactosyldiglyceride
nitrogen (MGDG) and digalactosyldiglyceride (DGDG). Small amounts
Phosphatidic acid: diacylglycerol esterified to of galactosylmonoglycerides (monogalactosylmonoglycerides
phosphoric acid (MGMG)) and digalactosylmonoglycerides (DGMG) are also
Phosphatidylcholine: phosphatidic acid linked to PC found in the endosperm of some mature cereal grains. Polar
choline lipids are found in all membranes including the amyloplast
Phosphatidylethanolamine PE membrane. Apart from these major acyl lipids, the other lipids
Phosphatidylserine PS
in cereals include sterols and lipid-associated compounds such
Phosphatidylinositol PI
Lysophosphatidylcholine, LPC, LPE,
as carotenoids and tocopherols. Although carotenoids are very
lysophosphatidylethanolamine, LPS, LPI minor constituents in cereal grains, color contributed by carot-
lysophosphatidylserine, lysophosphatidylinositol enoids is an important factor in the use of cereal grains in food
Glycoglycerolipids: 1,2-diacylglycerol joined by a production, particularly in the use of durum wheat for pasta
glycosidic linkage through position sn-3 to a making.
carbohydrate moiety The lipid content in cereals is influenced by the water
Monogalactosyldiglyceride MGDG content, the stage of maturity, and the variety of the grain
Digalactosyldiglyceride DGDG crop. Lipids are also unevenly distributed in various parts of
Monogalactosylmonoglyceride MGMG cereal grains. For example, in the wheat grain, the germ com-
Digalactosylmonoglyceride DGMG
prises only 4% of the total grain, by weight, but it has the
highest lipid content and the highest proportion of polar lipids
(Table 3), whereas the endosperm, the major fraction of wheat
grain, has significantly lower lipid content than the other
chain FA (saturated or unsaturated) at either the 1, or 2, or 3 fractions. Lipids in flour (except whole meal) can be subdi-
position. Similarly, diacylglycerols (DAGs) have two FA chains vided into nonstarch and starch lipids (Figure 2). The non-
at any of the two positions. However, the phosphoric acid starch lipids are further divided into free lipids and bound
residue of phosphoglycerides (PLs) and the first glycosidic lipids based on the extractability of lipids by solvents. The
residue of glycosylglycerides (GLs) are always in the 3 position. lipids that are easily extractable from flour or freeze-dried
The general classification of the principal lipids found in cereal dough, with a nonpolar solvent such as hexane, ether, or
grains and oilseeds, brief descriptions of their basic structures, petroleum ether, are called free lipids. Bound lipids refer to
and their common abbreviations are listed in Table 1. the nonstarch lipids that are only extractable with a very polar
250 FATS | Lipid Chemistry
Table 2 Approximate lipid contents and the distribution of lipid classes of the total lipids in cereal grains
TL, total lipids; NL, nonpolar lipid; GL, glycolipid; PL, phospholipid.
Grain fraction Grain composition (%) Lipid distribution (%) Total lipids in each fraction (% w/w) NL GL PL
TL, total lipids; NL, nonpolar lipid; GL, glycolipid; PL, phospholipid.
Wheat flour lipids PL, whereas starch lipids are predominantly PL and almost
exclusively lyso-PL in which LPC is the main lipid component.
The distribution of lipid classes in other cereal grains is
Nonstarch lipids Starch lipids qualitatively similar to those in wheat. The germ is the richest
(1.4%) (1.0%) source of lipids among all cereal grain fractions; more than
one-third of the total lipids is in the germ. In particular, maize
has a larger proportion of germ in the grain, 1125%, by
Free lipids Bound lipids
weight, with an exceptionally high lipid content, 3947% by
(0.8%) (0.6%)
weight, containing mostly TAGs, with a small amount of PL
and traces of GL. Maize germ is, therefore, more suited for
Nonpolar lipids Polar lipids
(<0.1%) (0.9%) edible oil production than any other cereal, after it has been
Nonpolar lipids Polar lipids
(0.6%) (0.2%) separated from the starch endosperm.
The FA composition of cereal lipids is generally similar for
wheat, barley, triticale, maize, and sorghum (Table 4). All
TAG, DAG, MAG, GL, PL LPC cereal grain lipids are rich in unsaturated FAs. Palmitic acid
FFA, SE (16:0) is a major saturated FA, and linoleic acid (18:2) is a
major unsaturated FA for all cereals expect for brown rice. Rye
Figure 2 The subdivision of wheat flour lipids. TAGs, triacylglycerols;
DAGs, diacylglycerols; MAGs, monoacylglycerols; SEs, sterol esters; lipids contain slightly higher levels of linolenic acid (18:3)
GLs, glycolipids; PLs, phospholipids; LPC, lysophosphatidylcholine. than those of other cereals. Oat lipids are similar to those of
brown rice; both are rich in oleic acid (18:1).
Lipid source Palmitic (16:0) Stearic (18:0) Oleic (18:1) Linoleic (18:2) Linolenic (18:3)
Table 5 Oil contents in oilseeds linoleic acid (C18:2). The FA composition of safflower oil is
similar to that of sunflower. The oil of commercial safflower
Oilseeds Oil in dry seed (% w/w) cultivars contains 5581% linoleic acid and 742% oleic acid
as major FAs, followed by stearic acid (110%) and palmitic
Canola 4050
Rapeseed/mustard 3050
acid (210%) as minor FAs. In peanut oil, oleic and linoleic
Cottonseed 1525 acids constitute 3371% and 1246% of the total FAs, respec-
Linseed 3545 tively. A high proportion of unsaturated FAs in an edible oil,
Peanut 3656 however, does decrease its storage stability and quality, due to
Safflower 1247 oxidation reactions associated with the double bonds in unsat-
Soybean 1522 urated FAs. In addition, more and more nutritional studies are
Sunflower 2550 showing that linoleic acid is not necessarily good for your
health. A new type of sunflower that has about 6571% oleic
acid has been produced to replace the traditional sunflower.
Unlike other oilseeds, FA composition in rapeseed oil is
leguminous oilseed, is used not only for edible oil production unusual in that it contains substantial amounts of long-chain
but also for direct consumption in various human foods, being FAs, in particular a significant amount of erucic acid (22:1) as
a good source of protein and oil. Another oilseed crop, linseed, 2050% of total lipids (Table 6). Major changes in the FA
produces oil that is not extensively used for food, but it is composition of rapeseed oils have arisen as a result of selective
nevertheless an important industrial oil. breeding for low levels of erucic acid. This is because of some
Lipid contents vary among the oilseed crops, shown as oil indications that erucic acid had antinutritional properties.
contents in Table 5, ranging from about 15% to 25% in soy- Consequently, varieties of rapeseed (called canola) have
bean and cottonseed, to as high as 50% in sunflower and been developed in which the level of erucic acid is virtually
canola/rapeseed, to 56% in peanut. It must be emphasized zero. Canola varieties are also characterized by markedly
that both oil content and component FA proportions in an increased levels of oleic acid with smaller increases also in
oilseed are subject to quite a wide variation. Such variations linoleic and linolenic acid content compared with those of
arise from genetic manipulation of seed varieties, climatic fac- the former rapeseed varieties due to the substantial reduction
tors and cultivation practices. of erucic acid content (Table 6). Canola is the oil with the
Lipids in oilseeds are predominantly TAG, 90% or more lowest content of saturated FAs. Linolenic acid content has also
of total lipids, and phospholipids being the main polar lipids. been reduced in some canola varieties in favor of linoleic acid,
Other minor lipid constituents are FFAs, sterols, hydrocarbons, through selective breeding. Low-linolenic acid content canola
etc. The FA composition of lipids is the main influence in terms varieties could have less than 2% linolenic acid and resemble
of the quality and the uses of the oil. Cottonseed oil has high olive oil in terms of their monounsaturated FA contents more
levels of saturated FAs among the oilseeds, 1825% palmitic than the high linolenic acid content oil variety (conventional
acid, and a small amount (13%) of myristic acid (Table 6). canola varieties).
The main unsaturated FAs are linoleic acid (3252%) followed Linseed oil is characterized by high levels of linolenic acid
by oleic acid (1032%). Cottonseed oil has a significant stor- (5260%), which undergoes autoxidation to give thin, cross-
age stability not only due to its lesser amount of unsaturated linked, and tough films, making it unsuitable for edible pur-
FAs content but also due to the presence of enough tocopherols poses. It is, however, an important drying oil used to produce
to reduce lipid peroxidation. oil-based paints.
Among the common oilseeds, safflower, sunflower, and
peanut oils contain high levels of unsaturated FAs. Sunflower
oil is characterized by its high concentration of linoleic acid
(6070%) followed by oleic acid (1535%). The high propor- Extraction and Quantification of Lipids
tion of polyunsaturated FAs makes sunflower oil a popular
source of essential FAs in the diet. Sunflower, safflower and Accurate and precise analyses of lipids are important, not only
soybean oils are the oils containing the highest levels of for the determination of their nutritive values for human
252 FATS | Lipid Chemistry
Lipid source Palmitic (16:0) Stearic (18:0) Oleic (18:1) Linoleic (18:2) Linolenic (18:3) Others
health benefits, but also for the understanding of the structural minimized by keeping lipids in solvents such as chloroform
characteristics and functional properties of lipids, which may and under a nitrogen atmosphere as well as by working at low
allow the development of tailor-made products that are temperatures if possible.
designed for a particular function or application. Analyses of
lipids involve several steps which usually include preparation
Supercritical Fluid Extraction
of the sample, solvent extraction of the lipids, removal of the
solvent and separation of the lipids into classes for qualitative Supercritical fluid extraction (SFE), which utilizes the solvation
or quantitative analyses. Sample preparation mainly involves power of the gases when compressed above critical point fluids
particle-size reduction of plant tissues or seed materials using at pressure and temperature, has been applied for lipid ana-
methods such as grinding. The purpose of this step is to lyses and for industrial extraction of oils. Carbon dioxide
increase surface area and facilitate subsequent lipid extraction. (CO2) is the most widely used supercritical fluid. The prepara-
In some cases, acid hydrolysis may be required to release lipids tion of the sample before extraction is extremely important. For
which are covalently bound to proteins and carbohydrates. solid materials, for example, grains, seeds, flours, or meals, the
material needs to be ground to produce smaller particles in
order to increase surface area and promote more efficient
Solvent Extraction
extractions. Supercritical CO2 is generally considered a more
The choice of organic solvents for lipid extraction largely selective solvent than hexane. The fact that supercritical CO2 is
depends on both the chemical nature of the sample and the a good solvent for nonpolar and moderately polar compounds
type of lipids to be extracted, that is, total lipids, starch lipids, makes it an excellent choice for extracting lipids without
and polar or nonpolar lipids. Neutral lipids are hydrophobi- extracting other more polar nonfat components. In this way,
cally bound and can be extracted readily by nonpolar solvents, it is possible to extract nonpolar TAG without the coextraction
whereas polar lipids, such as glycolipids and phospholipids, of PL. However, polar lipids such as PL can also be extracted by
which are predominantly bound by electrostatic forces and the supercritical CO2 method with the use of solvent modifiers
hydrogen bonding, require polar solvents capable of breaking such as ethanol, which increases the polarity of the supercrit-
such bonds. Hexane (petroleum ether) is the preferred solvent ical CO2. The SFE method uses no hazardous solvents, so it is
for extracting nonpolar, neutral lipids such as triglycerides and well suited for the process-control environment.
is the main solvent used for oilseed crushing/extraction for the The SFE-based test methods have been approved as an
industrial production of crude oil from oilseeds because of its official method by AOCS for the determination of process-
low cost. However, cereal grains and milled flour contain scale extractable oil or determination of total oil. The method
relatively large amounts of glycolipids and phospholipids, has two parts involving extraction with supercritical CO2 alone
and large amounts of cereal lipids (about half of the total or plus 15% ethanol. Although SFE has been used for wheat
lipids) are bound to polypeptides or polysaccharides. There- lipid extraction, there is no official SEF-based method for
fore, more polar solvents such as chloroform, alcohol, or extraction of lipids from wheat and other similar cereal grains.
chloroformmethanol (2:1) mixture are needed for extraction. Lipid extracts are complex mixtures of individual classes of
Hexane combined with an acid hydrolysis will give a complete compounds and require further separation. Many chromato-
extraction of the lipids from cereals. graphic and spectroscopic techniques are in use to separate and
Removal of the solvent from lipid extracts is normally con- quantify individual lipid components. The most commonly
ducted under vacuum in a rotary evaporator at or near room used techniques are thin-layer chromatography (TLC), high-
temperature. Care must be taken to minimize lipid oxidation performance liquid chromatography (HPLC) and gas chroma-
during sample preparation, solvent extraction and storage of tography (GC).
lipids. Cereal lipids characteristically contain large proportions
of polyunsaturated FAs together with carotenoids and tocols,
Thin-Layer Chromatography
all of which are highly susceptible to autoxidation. The lipids
are only slowly oxidized while the grain tissues are intact, but TLC provides the separation of a wide variety of compounds
once the tissues have been disrupted, oxidation must be with different polarities on a single plate and has long been
FATS | Lipid Chemistry 253
DGDG
900
1200
18:1 oleic
800
1000
700
Response (mV)
MGDG
800
Response (mV)
600
18:2 linoleic
500
600
400
18:3 linolenic
16:1 palmitic
400
18:0 stearic
300
PG
200 200
PC
0 100
10 15 20 25 30 35 40 0
Time (min) 20 25 30 35
Time (min)
Figure 4 HPLC analysis of wheat flour polar lipids extracted by
chloroform. Separation was carried out on a PVA-Sil threaded guard Figure 5 GC analysis of a canola oil. FAs methyl esters separated using
column (23 4.0 mm i.d.) with gradient elution at 0.5 ml min1 and a capillary GC column (SP2560, Supelco) and detected by an FID at
detected by a ELSD. Four steps of gradient were used for the separation 240 C.
010 min, 100% eluent A (2,2,4-trimethylpentane); 1020 min, 90%
eluent A 10% eluent B (tert-butylmethylether); 2035 min, 100%
eluent B; and 3540 min, 100% eluent C (methanol/n-ethylmorpholine/ flour lipids do affect the loaf volume and crumb texture of
glacial acetic acid (500:4.2:1.5 v/v)). For abbreviation of lipids, see bread. Fractions of polar and nonpolar lipids have opposite
Table 1.
effects on baking performance.
Incremental addition of the extracted lipid to a defatted
flour produces an unusual effect on the loaf volume and tex-
Since the principal FAMEs in cereal and oilseed lipids are 16:0, ture in a rapid baking test in which exogenous lipid is omitted.
16:1, 18:0, 18:1, 18:2, and 18:3, heptadecanoic acid (17:0) can Loaf volume decreases to a minimum at a lipid content inter-
be used as an internal standard. Identification and quantifica- mediate between those of the defatted flour and untreated
tion of individual FAs can be achieved by comparing and flour and thereafter increases. At high lipid contents (above
calibrating with relatively cheap FAME standards. Figure 5 the inherent lipid content of the flour), the loaf volumelipid
shows a typical GC profile of FA compositions in canola oil. curve tends to plateau. The changes in volume are paralleled by
FA compositions in lipid classes can also be obtained by GC changes in crumb grain, which, at first, also deteriorates and
analysis of lipids in each class separated or prepared by TLC or then improves. Fractions of polar and nonpolar lipids affect
other column chromatographic techniques. loaf volume and crumb grain in opposite ways. The polar lipid
Lastly, mass spectrometric detection is being used increas- fraction has generally beneficial effects, whereas the nonpolar
ingly for the detection and analysis of complex lipids. The fraction has deleterious effects. Thus, if the ratio of polar
technique can be used with direct inlet injection or linked to to nonpolar lipid is increased, the minimum of the loaf
HPLC or GC systems where the mass spectrometer serves both volumelipid content curve is shifted to lower lipid contents.
to detect lipids and to obtain information on their molecular If the percentage of polar to nonpolar lipid in a flour is varied
structure. from 0 to 100 at a constant lipid level, test-bake loaf volume
increases approximately in a linear manner. The addition of
different lipids to test their effects on dough has shown that the
Functionality of Lipids in Bakery Products unsaturated FAs such as linoleic acid are the components that
mainly contribute to the deleterious effects of the nonpolar
Apart from being used directly as a source of human food and fraction. Higher loaf volume and better crumb texture in bread
animal feed, a large proportion of cereal grains are milled to are favored by a high ratio of polar/nonpolar lipid and a high
flour, which is then used for producing various food products. content of flour lipid. However, the variation in loaf volume
Wheat has been the predominant cereal grain used for baked that can be attributed to lipid is relatively small, and by far, the
goods, largely due to its ability to form dough when flour is greatest variation is imposed by the effect of gluten protein
mixed with water. Rye ranks next in importance while other quality.
cereals assume a role in specialized products and in the prep- The role of the polar lipid appears to act as a complex
aration of composite flours. mixture of components with surface activities in stabilizing or
The rheological properties of doughs made from wheat destabilizing the gas bubble structure during expansion of the
flour are principally determined by the gluten protein, and loaf and thus the final texture. Upon heating (i.e., during
lipids (naturally occur in cereal grain, excluding added fat) baking), the dispersed lipids provide an interfacial film at the
appear to make little contribution. When protein is removed surface of gas cells, thus helping to modulate the growth,
from flour, dough and bread-making properties are lost. In expansion and coalescence of air cells. In addition to their
contrast, good dough properties and bread-making capacity effects in influencing volume and texture of baked products,
are retained after removal of the nonstarch lipids. However, lipids also play a role in staling mechanisms. Generally, the
FATS | Lipid Chemistry 255
presence of lipid reduces bread staling and enhances shelf life. Oils extracted from oilseeds, however, have a sufficient influence
This is largely due to the interaction between lipids and starch- in the total fat intake of the human diet since they are used
es. When monoacyl lipid monomers are present to form the directly for frying foods, as cooking ingredients, and as spreads.
lipidstarch complex, an insoluble amyloselipid film is likely Recent developments in plant molecular biology now allow the
to be formed on the surface of the starch granules. Such a film engineering of oilseeds with increased omega-3 long-chain
is capable of acting as a barrier against water transport involved PUFAs through complex multigene biosynthetic pathways.
in the staling process. Such development can have a considerable potential to increase
The effects of lipids have also been studied for Arabic bread, the nutritional values of plant-based oils.
steamed bread, cakes and biscuits. Despite differences in pro-
cessing, the general features with respect to lipids have much in
common. It is the crumb texture that is influenced most, and, Exercises for Revision
like bread, differences in lipids have not been found to account
for major variations in quality. However, the contribution of What are the different classes of lipids in cereal grains and
the lipid component to flour quality should not be ignored oilseeds?
when considering parameters to be used as a basis for selection What are the most common fatty acids in cereal grains and
in plant breeding programs even though protein composition oilseeds?
appears to be the major factor that determines flour quality. What are the different solvents used for the extraction of
The quality factors in flour lipid, as presently known, are a nonpolar or polar lipids?
high nonstarch lipid content, a high ratio of polar to nonpolar What is the most common gas used in the SFE-based test
lipid, and a low free fatty acid content. methods and how can the SFE extraction be used for the
determination of nonpolar TAG and total oil?
How to prepare lipid samples for gas chromatographic
Nutritional Values analysis?
What are the functional properties of lipids in bakery prod-
Lipids are the richest source of energy on a weight basis. They ucts? What are the different roles of nonpolar and polar
are essential components of our diet and play vital physiolog- lipids in cereal processing?
ical and biochemical roles in human health and well-being. Which polyunsaturated fatty acids are most common in
Lipids also increase palatability, including taste, aroma, and plant-based lipids?
texture of food, because they are carriers of fat-soluble flavors.
Fats and oils are also good carriers of fat-soluble vitamins such
as vitamins A, D, E, and K. Plant lipids contain tocopherols and
Exercises for Readers to Explore the Topic Further
tocotrienols, which are the most important sources of vitamin
E. Appropriate intake of lipids is essential for health mainte-
nance. However, a high consumption of fat, especially satu-
The multiple interactions of lipids with other constituents
(i.e., proteins during dough mixing and starch) are impor-
rated FAs, may be connected with several chronic diseases, such
tant for the processing and quality of cereal foods. How do
as heart disease and obesity.
different classes of lipids, for example, nonpolar and polar
Of the lipid components of a normal diet, the most impor-
lipids, influence the functions of gluten proteins and the
tant FAs are linoleic acid (n-6) and a-linolenic acid (n-3), the
starch-gelatinization mechanism?
two primary essential polyunsaturated fatty acids (PUFAs).
They play a role in stabilizing biological membranes by creat-
How can mass spectrometry be used to improve the detec-
tion and identification of lipids?
ing physical properties that are optimal for the transport of
substances across the membrane and for the biochemical reac-
How can generic breeding increase the nutritional values of
plant-based oils?
tions occurring in the membrane. Through metabolism, they
are converted into a whole range of oxygenated compounds
which exert a range of profound physiological activities involv- See also: Agronomy of Grain Growing: Soybean: Agronomy;
ing lowering plasma cholesterol, aggregating red blood cells, Barley, Rice and Maize Processing: Oil from Rice and Maize;
and smoothing muscle performance, all attributes that are Breeding of Grains: Soybean: Germplasm, Breeding, and Genetics;
required for good health. However, studies have shown that Wheat Breeding: Exploiting and Fixing Genetic Variation by Selection
too much linoleic acid (C18:2) can have adverse effects on and Evaluation; Carbohydrates: Starch: Starch Architecture and
health, especially causing inflammatory conditions. Since the Structure; Fats: Healthy Fats and Oils; Grain Marketing and
human body is unable to synthesize them, they must be Grading: Soybean: Grading and Marketing; Oilseed and Legume
obtained from dietary sources. Since plant lipids contain more Processing: Soybean: Processing; The Basics: Grain: Morphology
essential PUFAs than animal fats, their inclusion in food is of Internal Structure; The Cereal Grains: An Overview of the Family
positive from a nutritional point of view. The important source of Cereal Grains Prominent in World Agriculture; The Legumes and
of the essential FAs is oilseeds. Although cereal grains contain Pseudocereals: Peanuts: Overview; The Oilseeds: Cottonseed:
high amounts of linoleic acid, due to the generally low total Overview; Oilseeds: Overview; Wheat Processing: Analysis of Dough
lipid content of the grain, they are unlikely to have much contri- Rheology in Breadmaking; Baked Product Staling: Mechanisms,
bution to the essential FA intake in the human diet, except when Determinations, and Anti-staling Strategies; Bakeries: The Source of
the lipids are extracted from the lipid-rich germ and produced Our Unique Wheat-Based Food, Bread; Gluten and Modified Gluten;
as oil, such as maize, or rice germ oil or wheat bran oil. Ultrastructure of the Wheat Grain, Flour, and Dough.
256 FATS | Lipid Chemistry
Further Reading King JW (2004) Critical fluid technology for the processing of lipid-related natural
products. Comptes Rendus Chimie 7: 647659.
AOCS (2000) In: Official Methods and Recommended Practice of the American Oil Macritchie F (1981) Flour lipids Theoretical aspects and functional-properties. Cereal
Chemists Society. Champaign, IL: The Society. Chemistry 58: 156158.
Barnes PJ (1983) Lipids in Cereal Technology. London: Academic Press. Morrison WR (1978a) Cereal lipids. Advances in Cereal Science and Technology
Christie WW (2010) Lipid Analysis Isolation, Separation, Identification and Structural 2: 221348.
Analysis of Lipids, 4th edn. Bridgwater: The Oily Press. Morrison WR (1978b) Wheat lipid-composition. Cereal Chemistry 55: 548558.
Chung OK (1986) Lipidprotein interactions in wheat-flour, dough, gluten, Morrison WR (1978c) Cereal lipids. In: Pomeranz Y (ed.). Advances in Cereal Science
and protein-fractions. Cereal Foods World 31: 242244, 246247, 249252, and Technology, vol. II, pp. 221348. St. Paul, MN: American Association of Cereal
254256. Chemists.
Chung OK and Ohm JB (2000a) In: Lorenz KJ and Kulp K (eds.) Handbook of Cereal Morrison WR, Tan SL, and Hargin KD (1980) Methods for the quantitative-analysis of
Science and Technology, 2nd edn., pp. 417477. New York, NY: Marcel Dekker. lipids in cereal-grains and similar tissues. Journal of the Science of Food and
Chung OK and Ohm J-B (2000b) Cereal lipids. In: Karel K and Ponte Jr. JR (eds.) Agriculture 31: 329340.
Handbook of Cereal Science and Technology, 2nd edn. New York, NY: Marcel Papantoniou E, Hammond EW, Scriven F, Gordon MH, and Schofield JD (2001)
Dekker. Isolation of polar lipid classes from wheat flour extracts by preparative high-
Chung OK, Ohm JB, Ram MS, Park SH, and Howitt CA (2009) Wheat Lipids. St. Paul, performance liquid chromatography. Cereal Chemistry 78: 663665.
MN: American Association of Cereal Chemists Inc (AACC). Rocha JM, Kalo PJ, Ollilainen V, and Malcata FX (2010) Separation and identification of
Damude HG and Kinney AJ (2008) Engineering oilseeds to produce nutritional fatty neutral cereal lipids by normal phase high-performance liquid chromatography,
acids. Physiologia Plantarum 132: 110. using evaporative light-scattering and electrospray mass spectrometry for detection.
Eliasson A-C and Larsson K (1993) Cereals in Breadmaking. A Molecular/Colloidal Journal of Chromatography A 1217: 30133025.
Approach. New York, NY: Marcel Dekker. Salunkhe DK, Chavan JK, Adsule RN, and Kadam SS (1992) World Oilseeds
Gunstone FD (1996) Fatty Acid and Lipid Chemistry. Glasgow: Chapman and Hall. Chemistry, Technology, and Utilization. New York: Van Nostrand.
Hubbard JD, Downing JM, Ram MS, and Chung OK (2004) Lipid extraction from wheat Tweeten TN, Wetzel DL, and Chung OK (1981) Physicochemical characterization of
flour using supercritical fluid extraction. Cereal Chemistry 81: 693698. galactosyldiglycerides and their quantitation in wheat-flour lipids by high-
King JW (2002) Supercritical fluid technology for lipid extraction, fractionation, performance liquid-chromatography. Journal of the American Oil Chemists Society
and reactions. In: Lipid Biotechnology, pp. 663687. New York, NY: Marcel Dekker. 58: 664672.
Healthy Fats and Oils
SM Ghazani and AG Marangoni, University of Guelph, Guelph, ON, Canada
2016 Elsevier Ltd. All rights reserved.
Topic Highlights sources are important for their unique quality of healthy minor
components, fatty acid profile, or triacylglycerol composition
Healthy minor components in fats and oils. (e.g., high oleic oils for frying oil or omega-3-enriched oils). As
Fatty acids and triglyceride profiles of different oil sources. fats and oils are the highest source of energy in the human diet
Classification of fats and oils. and their main contribution is palatability of many food prod-
Major vegetable oils and beneficial components in them. ucts, selecting a good source of fats and oils based on the daily
Oil refining and its effects on healthy minor components. calorie need is important. Choosing an appropriate source of
oil and monitoring the energy intake may be helpful in con-
trolling obesity, which is considered a risk factor for many
Learning Objectives chronic diseases such as diabetes, cancer, and cardiovascular
diseases (CVDs).
To understand healthy minor components in oils.
To categorize oils and fats based on fatty acid composition.
To diagnose differences between desirable and undesirable Health and Nutrition Properties of Fats and Oils
minor components in oils.
To realize the main purpose of oil refining. Because fats and oils have different composition when it comes
to health, interest in specifying a fatty acid profile and healthy
minor components has been emerging. Oils and fats are pre-
Introduction dominantly a mixture of triacylglycerols as main organic com-
ponents that provide energy (9 kcal g1) accompanied by
Fats and oils are considered an important part of our daily diet other minor components that could be nutritionally important
intake because they provide a concentrated source of energy such as tocopherols, tocotrienols, sterols, polyphenols,
and essential fatty acids. They are carriers of fat-soluble vita- phospholipids (PLs), sphingolipids, and hydrocarbons. Also,
mins (A, D, and E) and make fundamental parts of cell walls other minor components are available and mostly removed
and other biological structures. In addition to biological func- during refining steps like waxes, free fatty acids, mono- and di-
tions and nutritional properties, they exhibit key functionality acylglycerols, flavoring, and coloring agents.
by playing an important role in the final quality of food More than the source of fats and oils, many other factors
products. Fats and oils can naturally be found in different such as varieties, growing conditions, maturity of seeds or
sources, but only a few of these sources are of economic fruits, harvesting time, storage conditions, handling, and pro-
importance. Oils and fats may mainly originate from plants cessing settings (extraction and refining oil) may affect the level
(seeds, beans, and fruits), animals (lard, tallow, fish oil, and of healthy minor components in oils. Based on current recom-
milk fat), and microbial sources (yeasts, algal, and bacteria). mendations, adults should get 2035% of daily calories from
Edible oils are mainly obtained from beans, seeds, and fruits by fats and oil sources. Currently we are emerging with two trends
mechanical and/or solvent extraction. Based on USDA statistic- in food intakefirst, increasing consumption of processed
al reports, total world vegetable oils production for 2012/2013 foods and snacks, and second, increased demand for virgin or
was 160.7 million tonnes. Meanwhile, the production levels of cold-pressed oils. As oils in the snack foods made up a rela-
four main vegetable oils in that period were palm oil (55.8 tively large portion intake of energy in industrialized countries,
million tonnes) followed with soybean oil (42.8 million choosing snacks could directly affect the health. In this article,
tonnes), canola oil (24.9 million tonnes), and sunflower oil first nutritional properties of healthy minor components will
(13.5 million tonnes). In that period, the total amount of be discussed followed by a comparison between fatty acid
produced animal fats from four main sources (lard, tallow, composition of fats and oils, and main sources of fats and
fish oil, and milk fat) was about 25 million tonnes (Table 1). oils will be reviewed.
It is important to consider two important points. First,
some part of produced oils is used for nonedible applications
(e.g., oleochemical industry, biodiesel, cosmetic), and second, Healthy Minor Components
daily dietary intake of fats and oils may go above the total
Phospholipids
amount of produced oils because other lipid-containing food
sources such as nuts, meat, and dairy products should be PLs are organic components with both hydrophilic and hydro-
considered in the diet as a source of fats and oils. Recent studies phobic moieties in molecule structures. They can be found in
in biotechnology and plant molecular biology led to the intro- all plant and animal cell membranes. The main food sources of
duction of numerous types of engineered oilseeds and micro- PLs are egg yolk (810%), butter (0.51.2%), and crude vege-
bial sources to produce novel fats and oils with new fatty acid table oils (0.53.7%). The six major types of PLs in vegetable
and triglyceride compositions and with a higher oil yield. New oils are phosphatidic acid (PA), phosphatidylcholine (PC),
phosphatidylethanolamine (PE), phosphatidyl glycerol, phos- esters, for instance g-oryzanol in rice bran oil, have unique
phatidylserine (PS), and phosphatidylinositol (PI). PLs nutritional properties such as antioxidant activity, hypolipi-
showed great positive functionality in several diseases such as demic effects, and stimulation of growth and hypothalamus
liver diseases, neurological disorders, CVDs, and hepatic and activity. Hypocholesteremic properties of phytosterols have
colon cancers. They also have immunological functionality for been studied, and based on the guidelines, 2 g of phytosterols
body system, and they can improve physiological and mental per day, along with 1025 g soluble fibers, was recommended.
performances. The amount of PLs in crude oils depends on the In animal fats, the main sterol is cholesterol. Although a
oil sources and extraction method. Solvent-extracted oils have trace level of cholesterol 2050 ppm could be found in vege-
higher phosphatides than mechanically extracted oils. table oils, it is much lower than the levels reported for animal
PLs have been shown to have effects on lowering blood fats (up to 1000 ppm), fish oils (up to 7000 ppm), dairy fats
cholesterol and TAG levels in serum. Many studies have shown (20003000 ppm), and egg yolk (12 500 ppm). It is important
that in athletes, PS can improve performance and decrease to note that cholesterol is not primarily associated with the
muscle damage during some sports like cycling, weight train- storage adipose (tallow or lard), so lowering the level of total
ing, golf, and endurance running. fat in meat (using lean or extra-lean meats) will not greatly
Among PLs, PI and PE are more effective than PC in increas- decrease the cholesterol content because it is a necessary mem-
ing oxidative stability of oils as a synergistic function with brane component.
a-tocopherol. In the digestive tract, PLs play an important In the human blood system, there are two main types of
functionality in the metabolism of lipids and other digestive cholesterol named high-density lipoprotein (HDL) cholesterol
fluids. Although PLs have health-promoting properties, from and low-density lipoprotein (LDL) cholesterol. HDL choles-
an industrial viewpoint, they are considered undesirable com- terol is good cholesterol because excess HDL cholesterol is
ponents. In the neutralization process, they work as an carried to the liver and then metabolized. LDL cholesterol is
emulsifier, hinder the separation of oil and water phases, and bad cholesterol because it remains in the blood plasma and
help to increase the oil loss in this step. Also, they may interfere can start to produce plaques in the arterial wall and lead to
with bleaching clays, poison nickel catalysts throughout hydro- obstruction of heart blood artery, causing a heart attack. So
genation, darkening oil during deodorization and decreased elevated total and HDL cholesterol levels are considered risk
shelf life of refined oils. PLs can be almost completely removed factors for the development of coronary heart disease that is
from oil through the degumming stage of oil refining; only one of the main health problem in developed countries. Phy-
they can be found in virgin or cold-pressed oils in different tosterols in vegetable oils can inhibit cholesterol absorption.
levels. Intake of phytosterols at the level of 1.53 g day1 can bring
total blood cholesterol and LDL cholesterol lower by 1015%.
Phytosterols usually have a lower absorption rate in the diges-
Sterols
tive tract than cholesterol; for example, only 5% of b-sitosterol
Sterols comprise of the main part of unsaponifiable matter of of the food supply can be absorbed, so phytosterols in foods
fats and oils. In fats and oils, they can be in two forms of free can compete with cholesterol absorption and leads to an
and esterified sterols. Free sterols can commonly esterify with increase in bile secretion and inhibition of cholesterol absorp-
fatty acids, glycosides, and/or ferulic acid to make sterol esters. tion from the gut.
The amount of phytosterols in most of common vegetable oils Much research has shown that phytosterols, especially
is usually between 1000 and 5000 ppm (15 g kg1). How- b-sitosterols, inhibited cancer growth and may offer protec-
ever, higher levels could be found in rapeseed oil tion from colon, prostate, and breast cancers.
(500011 000 ppm) and in corn oil (800022 000 ppm). Besides cholesterol, ergosterol (precursor of vitamin D2)
The major phytosterols in common vegetable oils are bras- can also be found in animal fat sources such as fish oil and
sicasterol, campesterol, stigmasterol, b-sitosterol, and avenas- butter fat.
terol. Canola oil, corn oil, sesame oil, and soybean oil are good Phytosterols are stable molecules, but after as long time
sources of phytosterols between vegetable oils. Some sterol being heated or exposed to oxygen during food processing
FATS | Healthy Fats and Oils 259
OH
O
O
-Carotene Sesamol
Squalene
HO
H3C O
CH3
-Tocopherol
HO HO
Cholesterol -Sitosterol
Polyunsaturated fatty acids (PUFAs) can be characterized and serve as the template for re-esterification by intestinal
based on a methylene group that can be central, conjugated, or cells to reform triacylglycerols. In vegetable oils, unsaturated
unconjugated. For instance, linolenic acid (C18:3) can be fatty acids are esterified in the sn-2 position, whereas in animal
found in two forms of a-linolenic -9, 12, 15-oic acid, (n 3) fats saturated fatty acids are in this position. In addition, selec-
or g-linolenic-6, 9, 12-oic acid (n 6). Eicosapentaenoic acid tive distribution of fatty acids on glycerol backbone is an
(EPA) C20:5 (n 3) and docosahexenoic acid (DHA) C22:6 important factor that leads to triacylglycerols diversity. For
(n 3) found in oily fishes are examples of PUFAs with nutri- example, palm oil contains six types of fatty acids, but only
tional characteristics. 14 diverse triacylglycerols can be detected.
Triacylglycerols comprise more than 95% of fats and oils Omega-3 and omega-6 PUFAs have different physiological
composition. Each triacylglycerol molecule consists of three properties in the human body system. Whereas n 3 fatty acids
fatty acids that attached to the glycerol molecule with ester have anti-inflammatory, antithrombotic, antihypertensive,
bonds. Normally triacyglycerols can categorize to simple tri- and anti-arrhythmic derivatives, n 6 fatty acids cause inflam-
glycerides with only one type of esterified fatty acid, for matory, thrombotic, hypertensive, and arrhythmic metabo-
example, triolein (OOO); di-acid triglycerides with two types lites. That is the reason for the importance of the n 6/n 3
of fatty acids esterified to glycerol molecule, for example, di- ratio balance in human diet.
palmito 1,3 olein 2 (POP); and tri-acid with three types of acyl The impact of saturated fatty acids on LDL cholesterol
groups, for example, palmito 1, oleo 2, and stearin 3 (POS). depends on the number of carbon atoms in the fatty acid
Glycerol molecules can also esterify with only one fatty acid to chain with lauric acid > myristic acid > palmitic acid > stearic
form monoacyl glycerols or with two same or different fatty acid. Clinical studies showed that saturated fatty acids lead to
acids to make diacylglycerols. The physical, chemical, an increase in total blood cholesterol and LDL cholesterol,
biological, and physiological properties of fats and oils related whereas mono-unsaturated and more effective
to triacylglycerol composition and presence of healthy minor polyunsaturated fatty acids lead to a decrease in the total
components. The reason for the importance of triacylglycerol serum cholesterol and LDL cholesterol and increasing HDL
composition is that two oils with the same fatty acid profile cholesterol in the bloodstream. The risk of coronary heart
may have different physical, chemical, and physiological prop- disease in the societies in which people consume olive or
erties. Also the position of fatty acids on glycerol molecules has canola oil (good source of oleic acid) and fish oil (good source
health and physiologically importance. For instance during fat of omega-3 fatty acids) is lower.
digestion, because human lipase is specific (hydrolyzing only Studies have shown that consumption of n 3 fatty acids
fatty acids in positions 1 and 3 of TAGs), the fatty acids (fish oil, walnut oil, and flaxseed oil) reduces mortalities
attached to carbon number 2 of glycerol structure (the sn-2 related to CVDs by lowering triacylglycerol accumulation in
position) are especially absorbed as the 2-monoacylglycerol the liver and circulation.
FATS | Healthy Fats and Oils 261
Classification of Oils and Fats Based on Fatty Acid Linoleic Acid Oleic Acid Subgroup
Profile Sunflower oil, corn oil, sesame oil, pumpkin seed oil, and
Medium-Chain Fatty Acids (C6:0 to C10:0) Subgroup borage oil are good examples of this subgroup. In this group,
borage oil has a high amount of g-linoleic acid (about 22% of
This group of oils (MCTs) is important in the case of providing
the total fatty acids).
an instant source of energy for the body. These fatty acids have
small size and higher solubility and are mainly transported
directly via portal circulation and metabolized in the liver, Conjugated Linoleic Acid
whereas longer-chain fatty acids are absorbed via the lymphatic
system after micellar transport at the intestinal wall. MCTs are Conjugated linoleic acid (CLA) is a term used for positional
neutral on LDL-, HDL-cholesterol or triacylglycerol concentra- and geometric isomers of linoleic acid that can be found in
tion in serum and may be beneficial in controlling obesity. animal fats such as beef, lamb, and dairy foods. CLA are
produced by microbial fermentation of PUFAs and isomeriza-
tion of linoleic acid in the rumens of ruminants. Whereas
Lauric Acid and Myristic Acid Subgroup double bonds in linoleic acid are between the ninth and
tenth carbons and the twelfth and thirteenth carbons, CLA
Among saturated fatty acids, lauric acid and myristic acid are has conjugated double bonds at carbon atoms 10 and 12 or 9
the most blood cholesterol and LDL-cholesterol increasing and 11, with possible cis and trans combinations. Many clin-
fatty acids. Although, the effect of myristic acid is higher than ical studies have shown that conjugated linoleic acid may have
lauric acid, the position of fatty acids on the glycerol backbone physiological effects, including anticarcinogenic, anti-
needs to be considered. Coconut oil and palm kernel oil are atherogenic, improvement of type II diabetes, and immuno-
categorized in this subgroup. modulating properties of CLA.
Some clinical studies on animals showed that conjugated
linoleic acid is quite effective to inhibit the growth and metas-
Palmitic Acid Subgroup tasis of breast and prostate cancers.
The effect of palmitic acid on increasing serum cholesterol and
LDL cholesterol is lower than lauric acid and myristic acid.
Trans Fatty Acids
Palm oil and lard are good examples of this group.
Two types of trans fatty acid (TFAs) isomers can be found in
fats as naturally and industrially produced fatty acids. The
Stearic Acid Subgroup presence of TFAs in animal fats was revealed some time ago.
TFA in tallow and milk fat is vaccinic acid C18:1 (n 7t). This
Among saturated fatty acids, stearic acid exhibited as quite
TFA is naturally produced during biohydrogenation and isom-
neutral with no deleterious effect on plasma lipids. Cocoa
erization of PUFAs in rumen guts.
butter and shea butter can be in this subgroup because they
Catalytic hydrogenation is an industrial modification pro-
contain up to 40% of stearic acid.
cess that is applied to reduce the amount of polyunsaturated
fatty acids and leads to improvement in oxidative stability of
oils and increasing hardness. This industrial process produces
Oleic Acid Subgroup (>60%)
TFA, which are almost unavailable in natural fats. TFA levels
Olive oil is the best representative oil of this subgroup, which and isomeric distribution depend on processing factors (nickel
contains about 80% of oleic acid and then canola oil with catalyst type and concentration, hydrogen pressure, stirring
about 65% oleic acids. In nuts oil, hazelnut oil and macadamia speed, and reaction temperature), fatty acid composition of
oil are rich in oleic acid. original oil, and degree of saturation (partially or fully
hydrogenation).
Partial hydrogenation produces mostly geometrical and
Linoleic Acid Subclass (>60%) positional trans-18:1 isomers (mainly elaidic acid) while
reducing the amount of PUFA. During deodorization of edible
The common vegetable oils gathered in this group are grape
oils, a small amount of trans-linoleic and trans-linolenic acid
seed oil, evening primrose oil, and safflower oil. As the human
are produced.
body cannot synthesize linoleic acid from food components,
A positive correlation between intake of a high amount of
presence of this fatty acid is essential for good health and
TFAs in a diet with increased risk of chronic diseases such as
termed an essential fatty acid. As mentioned earlier, the inflam-
cardiovascular, inflammatory, autoimmune diseases, and can-
matory effect of consuming n 6 fatty acids should be consid-
cer have been found. TFAs raise the amount of LDL cholesterol
ered and n 6/n 3 ratio should be balanced in the diet.
and lower the level of HDL cholesterol, thus resulting in dou-
bly detrimental effects. New regulations on TFA labeling,
which led to a ban on industrially produced trans fats in
Linoleic Acid Saturated Fatty Acids Subgroup
foods, caused a dramatic reduction in TFAs in food products,
Cottonseed oil and watermelon seed oil are clustered in this but they still can be found in some foods such as confectionery
group. Their fatty acid profiles mainly contain linoleic acid, coatings and fillings, biscuits (cookies), pie crusts, toffees, and
associated with saturated fatty acids (especially palmitic acid). analogs of dairy products such as butter, cream, and toppings.
262 FATS | Healthy Fats and Oils
Using naturally saturated fats such as palm oil and/or and was introduced to East Asia in 1884. Currently, Indonesia
applying fully hydrogenated oils or vegetable oils from new and Malaysia are the two main palm oil producers in the world.
varieties such as high oleic oils could be solutions for food Palm oil is rich in healthy minor components such as carbohy-
industries to substitute TFAs in their products. drates (carotenoids and squalene) and tocols. Crude palm oil
has about 4001000 ppm carotenoids (pro-vitamin A) but
after physical refining, the amount of carotenoids decreased
Omega-3 Fatty Acids
to nondetectable levels. The squalene content in crude palm
The omega-3 fatty acids consist of a-linolenic acid, EPA, and oil is between 200 and 500 ppm, which is dramatically higher
DHA. Several clinical studies showed a positive correlation than other common vegetable oil sources. Total tocols in crude
between consumption of omega-3 fatty acids in the diet and palm oil are about 800 ppm, 70% of which are tocotrienols.
an improved prognosis of CVD and nonalcoholic fatty liver The distribution of tocopherols in palm oil is a-tocopherol
disease. The main sources of a-linolenic are flaxseed oil and 22%, and b-, g-, and d- tocopherols represent 20%, 46%, and
walnut oil and oily fishes such as tuna, menhaden, anchovies, 12%, respectively. Crude palm oil contains 210620 ppm of
sardines, and mackerel are the main sources of EPA and DHA. phytosterols. The main phytosterols in palm oil are b-sitosterol
(120370 ppm), campesterol (46150 ppm), and stigmasterol
(2666 ppm).
Major Vegetable Oils Palm oil is a good source of carotenoids. Based on the
carotenoid content, crude palm oil color varies from yellowish
This section is mainly focused on the main sources of vegetable to dark red. The main carotenoids in palm oil are a-carotene
oils that would cover more than 95% of total vegetable oils (2442%) and b-carotene (5060%). Palmitic acid and oleic
consumed in the word (Table 1). acid are the main fatty acids in palm oil (more than 80% of
total fatty acids and almost in equal amount). Normally, palm
oil is fractionated into different derivatives (palm olein, palm
Palm Oil
stearin, palm super olein, and palm mid fraction) with specific
Palm oil is the main vegetable oil produced in the world. physical and chemical characteristics (Tables 2 and 3). Because
During 20132014 about 53.4 million tonnes of palm oil of fatty acid composition (about 50% saturated fatty acids and
were produced (Table 1). Palm oil originated from South Africa 50% oleic acid), crystallization in the form of b0 and the
Fatty acids (%) Palm oil Palm olein Palm super olein Palm top olein Palm stearin Palm mid fraction
Malaysian Palm Oil Board (MPOB) (2009) Pocketbook of Palm Oil Uses, pp. 341.
Fatty acids (%) Soybean oil Canola oil Sunflower oil Cottonseed oil Corn oil Peanut oil Olive oil
12:0
14:0 0.1 0.5-1.3 <0.1 <0.1
16:0 11.0 4.3 57 1731 813 813 7.520
16:1 0.1 0.2 <0.4 <1 <1 <0.3 0.33.5
18:0 4.0 1.8 46 13 14 34 0.55
18:1 23.4 62.3 1525 1321 2432 4866 5583
18:2 53.2 19.4 6270 3460 5562 1428 3.521
18:3 7.8 9.2 <0.2 <1 <2 <0.3 <1
20:0 0.3 0.6 <1 0.20.5 <1 12 <0.8
20:1 1.2 <0.5 <0.5 12
20:2 0.1
22:0 0.1 <1 <0.5 24 <0.3
24:0 12 <1
FATS | Healthy Fats and Oils 263
presence of natural antioxidants (tocols), this oil is widely used (100386 to 189753 ppm, respectively; data adapted from
in the food industry as a main part of base stock to formulate Codex Alimentarius, 2010). Green color of crude canola oil is
margarines, shortenings, frying oils, and confectionary fats. related to the presence of chlorophylls (550 ppm).
Plant breeders have developed canola oil with new fatty acid
composition such as high oleic canola oil (oleic acid raised to
Soybean Oil
about 85%), high lauric canola oil (up to 31% lauric acid), and
Soybean is the main oilseed produced in the world, and soy- high stearic acid canola oil (stearic acid content increased to
bean oil is the second-largest-produced vegetable oil after palm about 28%).
oil (Table 1). The worlds major producers of soybean oil are
the United States, China, Argentina, Brazil, EU-27, and India.
Crude soybean oil is a good source of phosphatides that after
Sunflower Oil
extraction and purification steps can be used as a natural
emulsifier (lecithin) in various food products. The three Russia, Ukraine, Turkey, European countries, and Argentina are
major phosphatides in crude soybean oil are PC, PE, and PI the main producers of sunflower oil. During 20132014,
present in the ratio of 55%, 26%, and 18%, respectively. Con- about 15.7 million tonnes of sunflower oil were produced
centration of phytosterols and tocopherols in soybean oil is (Table 1). Linoleic acid as an essential fatty acid (6270%)
23504050 ppm and 12571370 ppm, respectively. In soy- and oleic acid (1525%) are predominant fatty acids in this
bean oil, b-sitosterol (4759%) and g-tocopherol (4460%) oil. These amounts for new mutant of high oleic sunflower oil
are the major phytosterol and tocopherol in soybean oil. Soy- are 217% and 7591%, respectively. The main triglycerides in
bean oil contains about 68% a-linolenic acid. Research regular sunflower oil are LLL (36%) followed by OLL (29%).
showed that soybean oil consumption led to a reduction in Sunflower oil normally has about 32005100 ppm phytos-
total blood serum cholesterol and LDL cholesterol. terols (5864% b-sitosterol) and 4401200 ppm tocopherols
The major triacylglycerols in soybean oil are OLL (20%), (9197% a-tocopherol). It is also considered a good source of
LLL (19%), and PLL (13%). Crude solvent extracted soybean squalene (150200 ppm). Crude sunflower oil has 0.51.2%
oil typically has about 2%PLs. The main phytosterols in soy- PLs and around 180 ppm of total polyphenols. The main poly-
bean oil are b-sitosterol (52%), campesterol (25%), and stig- phenols in crude sunflower oil are caffeic acid, vanillic acid,
masterol (23%). Two major hydrocarbons in soybean oil are and chlorogenic acid.
squalene and hexahydrofarnesyl acetone.
Corn Oil showed that squalene in olive oil may have a preventive effect
on breast cancer, and it also has cardioprotective properties.
Corn oil is a by-product of corn meal- and starch-making
Total phytosterol content in virgin olive oil is between 980
companies. Fatty acid composition of corn oil comprises a
and 1840 ppm; meanwhile the main phytosterol is b-sitosterol
relatively high level of linoleic acid (5862%). The major
(7590%). Virgin olive oil is considered a rich source of poly-
triacylglycerol molecules in corn oil are LLL (25%), LLO
phenols. The amount of polyphenols in virgin olive oil
(22%), LLP (15%), OOL (11%), and PLO (10%).
changes from 100 to 300 ppm. Polyphenols in virgin olive
In vegetable oils, corn oil is categorized as one of the richest
oil (tyrosol, hydroxytyrosol, and oleuropein) have very strong
source of phytosterols (830025 500 ppm) and tocopherols
free radical scavenger properties and antioxidant activity.
(11301830 ppm). The main phytosterol and tocopherol in
Research showed that they may correlate with decreasing the
corn oil are b-sitosterol (6370%) and g-tocopherol
risk of CVDs and cancer. The level of phosphatides in olive oil
(6889%), respectively. Numerous clinical studies showed a
is lower than common seed oils (40135 ppm). Clinical stud-
correlation between corn oil consumption and total serum
ies have shown that olive oil consumption has lots of preven-
cholesterol reduction that may be related to a high concentra-
tive effects against chronic diseases such as obesity, CVDs,
tion of phytosterols in this oil. Research showed that a high
inflammatory diseases, and cancer.
ratio of g-tocopherol in corn oil has health benefits such as a
protection effect against DNA damage, blood pressure, platelet
aggregation, and diabetes. Coconut Oil
Average annual production of coconut oil is 34 million
tonnes, mainly in the Philippines, Indonesia, and India. Nor-
Peanut Oil mally, coconut oil is extracted after sun or fire drying of the
China and India are the main producers of peanut oil. Phos- coconut kernel. Dried kernel is named copra, and it has about
pholipid content of crude peanut oil is 0.62%, depending on 65% oil. Coconut oil contains high levels of saturated fatty
the maturity of peanuts. The main fatty acids in peanut oil are acids mainly lauric acid and myristic acid (more than 60% of
oleic acid (4553%), linoleic acid (2732%), and palmitic total fatty acids). Coconut oil and palm kernel oil are the
acid (1114%). This oil specifically has long-chain saturated richest natural source of medium-chain fatty acids. Because
fatty acids such as arachidic acid (12%), behenic acid the major fatty acids in coconut oil are saturated, the total
(1.54.5%), and lignoceric acid (0.52.5%). The main triacyl- tocopherol level in this oil is very low (55 ppm). The main
glycerols in peanut oil are OOL (19%), OLL (18%), POL fatty acid and triacylglycerol in coconut oil are lauric acid
(13%), OOO (12%), and POO (7%). Crude peanut oil con- (48%) and trilaurin (19%), respectively. The special flavor of
tains about 3400 ppm total phytosterols, whereas the main coconut oil is related to the presence of d- and g- lactones.
phytosterols are b-sitosterol (64%), campesterol (15%), stig- The range of total phytosterol in this oil is between 800 and
masterol (11%), and D5-avenesterol (8%). Tocopherols con- 2300 ppm. The major sterols are b-sitosterol (45%), D5-
centration in peanut oil ranges from 130 to1300 ppm with the avenasterol (30%), and stigmasterol (14%).
composition of a-tocopherol (4765%) and g-tocopherol The amount of hydrocarbons in coconut oil is high,
(3050%). This oil also has a high level of squalene whereas it only has a small amount of squalene (20 ppm),
(400700 ppm). but the level of polycyclic aromatic hydrocarbons (PAHs)
could reach to about 3000 ppm as result of contamination of
oil with petroleum oil or during fire drying. The main differ-
ence between fatty acid composition of coconut oil and palm
Olive Oil
kernel oil is that although the level of medium-chain fatty acids
Olive oil has been used for a long time as a food and a (caprylic and capric acids) in coconut oil is higher than palm
medicine. The world average annual production of olive oil is kernel oil, the amount of oleic acid in palm kernel oil is higher
about 3 million tonnes mainly in Mediterranean countries. than coconut oil. These oils have a sharp melting point that
Many epidemiological studies have shown that the lower makes them highly suitable to formulate different food prod-
rates of breast, ovarian, and colorectal cancers in Mediterra- ucts such as confectionary fats, ice creams, and icings (Table 4).
nean countries may be related to consumption of olive oil.
There are two kinds of olive oil in the world: one with a high
Sesame Oil
amount of oleic acid and low amount of linoleic acid
(European origin) and another with the opposite ratios For a long time, sesame was used as an oilseed crop because of
(North African origin). This diversity caused a wide range of high levels of oil (50%) and protein (2025%) in this seed.
fatty acid and triacylglycerol composition to be considered for Sesame oil is produced mainly in China, Japan, India,
this oil. In general, three major fatty acids in olive oil are oleic Singapore, and Mexico. In sesame oil, oleic acid and linoleic
acid (5583%), palmitic acid (7.520%), and linoleic acid acid are the main fatty acids with the same ratio (around 40%).
(3.521%). The greenish color of virgin olive oil is related to The major triacylglycerols in this oil are LLO (25%), LLL
the presence of chlorophyll in this oil, and their level ranges (20%), and LOO (15%). Total phytosterols and tocoph-
from 10 to 30 ppm. a-Tocopherol is the major tocopherol (up erols contents in sesame oil are 54006300 ppm and
to 95%) in olive oil, and it has a range from 100 to 300 ppm. 200500 ppm, respectively. b-Sitosterol and campesterol are
The level of squalene in olive oil is relatively high and in most the main phytosterols in this oil (more than 80% of total
of samples reported in the range of 200500 ppm. Research sterols), and g-tocopherol comprises more than 96% of
FATS | Healthy Fats and Oils 265
Table 4 Fatty acid composition of lauric oils oils, but g-tocopherols is the major tocopherol in chestnut,
pecan, walnut, heartnut, cashew, and Brazil nut oils. Macadamia
FA (%) Palm Kernel Coconut nut oil is rich in tocoterienols (about 60 ppm). Walnut oil has
6:0 <0.8 <0.1 the highest amount of total phytosterols (3000 ppm) followed
8:0 25 610 by peanut and pecan oils (2800 ppm) and almond oil
10:0 35 510 (2700 ppm). Macadamia oil has the lowest level of phytosterols
12:0 4451 3954 (1280 ppm).
14:0 1517 1523 Based on clinical studies, regular nut consumption or using
16:0 710 611 nut oils as salad oil is associated with reduced risk of coronary
16:1 <0.1 <2 heart disease, diabetes, and cancer.
18:0 23 14
18:1 1218 411
18:2 14 12
Fats and Oils Refining
18:3 <0.7 <0.1
20:0 <0.3 <0.2 The main purpose of crude oil refining is to remove undesir-
20:1 <0.5 <0.2 able minor components and contaminants from oils. Free fatty
acids, oxidation products, chlorophyll, waxes, and trace metals
Karkeskind A (1996) Sources of the main fats and relevant monographs. In: Oils and
from oils are generally undesirable because they may decrease
Fats Manual, pp. 117323. Paris: Lavoisier Publishing.
the quality of refined oil through darkening, foaming, smok-
ing, precipitation, development of off-flavor, and decreasing
total tocopherols. Sesame oil has a good oxidative stability.
oxidative stability or be potentially dangerous to health. The
More than tocopherols, the stability of this oil is related to
source of oil contamination could be from oilseeds during
the presence of the unique natural phenolic antioxidants of
harvesting and storage with trace metals, dioxins and poly-
sesamol and sesamolin (totally around 5000 ppm). In
chlorobiphenyls from industrial wastes, polycyclic aromatic
many countries, sesame oil is used as frying oil because
hydrocarbons, or mineral oil from vehicle exhausts. Another
of its high stability and also its nutty flavor.
source of oil contaminants could be the trace amount of insec-
ticides, fungicides, or herbicide extracted from oilseeds and
transferred to oils. Although most of these chemicals are
Rice Bran Oil
removed during refining, oil refining may also lead to the
Rice bran oil is mainly produced in Japan, Thailand, India, formation of another group of degraded products such as
China, and Vietnam. The major fatty acids in rice bran oil are polycyclic aromatic hydrocarbons (PAHs) or 3-monochloro-
oleic acid, linoleic acid, and palmitic acids (typically, 42%, propane-1,2-diol (3-MCPD) esters and glycidolesters.
32%, and 20%, respectively). Activity of lipase during rice Another important fact about oil refining is related to the
bran oil extraction leads to an increase in the amount of FFA, loss of micronutrients during refining. Fully refining (degum-
mono-, and diacylglycerols. Triglycerides totally comprise ming, neutralization, bleaching, and deodorization) of oils
8386% of total neutral lipids, whereas PLO, PLL, and OOO leads to change not only in the level of healthy components
are predominant triacylglycerols in this oil. The concentration but also in their chemical structure and composition by isom-
of phytosterols in rice bran oil is higher than common erization or polymerization. Research showed that traditional
vegetable oil (32 000 ppm) with the composition of 54% of refining leads to about a 20% loss in total amount of tocoph-
b-sitosterol, 20% of campesterol, and 11% of D5-avenesterol. erols and 24% in phytosterols level in canola oil.
Rice bran oil is a good source of tocopherols, and the average A typical specification for fully refined oil in general could
of tocopherol content in rice bran oil is 860 ppm. Research has be as taste (bland), moisture (max. 0.05%), phosphorus (max.
shown that rice bran oil has health benefits such as lowering 5 ppm), free fatty acids (max. 0.1%), peroxides (max.
serum cholesterol level, inhibiting platelet aggregation, 3 meq/kg), iron (max. 0.5 ppm), copper (0.05 ppm), lead
increasing fecal bile excretion, and decreasing early (max. 0.01 ppm), hexane (max. 5 ppm), benzopyrene (max.
atherosclerosis. 2 ppb), dioxins (0.75 pg), aflatoxins (6 ppb), and mineral oil
(LOD).
Nut Oils
The main worlds most popular nuts are peanut, almond, Conclusion
cashew, pistachio, walnut, hazelnut, and pecan. The oil content
of nuts varies from 0.8% for chestnuts to 67% for Brazil nuts. Although fats and oils contribute to palatability of many food
Fatty acid composition of nut oils is very diverse. The highest products, consumers are nowadays paying more and more
amount of saturated fats belongs to Brazil nut oil (around 23%), attention to their health and well-being. Based on WHO rec-
and the lowest level of saturated fat belongs to heartnut oil ommendation for dietary intake of fats and oils for adults, it is
(about 5%). Hazelnut oil and pistachio oil have the highest suggested that we get 2035% of our energy from fats and oils
amount of oleic acid (more than 75%), and walnut oil has the sources. In a selection of healthy oil sources, many factors
highest amount of n 3 and n 6 PUFAs. a- and g-Tocopherols should be considered such as fatty acid composition, healthy
are the main tocopherols in nut oils. a-Tocopherol is the minor components, process of oil extracted (solvent extracted,
predominant tocol in hazelnut, pistachio, almond, and peanut hot-pressed or cold-pressed), and minimally refined or fully
266 FATS | Healthy Fats and Oils
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Hammond EG, Johnson LA, Su C, Wang T, and White PJ (2006) Soybean oil.
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lipids. In: Processing and Nutrition of Fats and Oils, pp. 137154. West Sussex:
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Bioactives and Toxins
Contents
Bioactive Compounds in Wheat Bran
Bioactives: Antioxidants
The Antinutritional Components of Grains
Mycotoxins
The structure of the wheat kernel is described. Wheat, eaten as bread and other baked products, has been a staple
There is an important difference between wheat bran as it of the human diet for several thousand years. Wheat is a globally
exists in the seed, which can be hand-dissected, and wheat important agricultural commodity and ubiquitous food ingredi-
bran commonly obtained by milling. ent mainly because it is the only cereal grain capable of being
Food-grade bran is not a standardized product and can vary processed into leavened bread, an attribute that derives from its
in purity and composition of bioactive compounds due to unique gluten protein composition. While the use of whole
genetic factors (different wheat varieties or types), the wheat for food waned significantly in the early part of the twen-
wheat growing environment, and also how the milling is tieth century in favor of baked goods made with refined flour, the
carried out. past several decades have seen a remarkable resurgence in the
The unique protective role of bran from a seeds perspective consumption of whole wheat and whole grains in general. This
reflects its complement of bioactive compounds, particu- has been sparked by increasing consumer interest in healthful
larly those in fiber tissues. eating and awareness of the importance of the health-protecting
Bran is the most complex physical component of the wheat and health-promoting effects of whole grains. The functional
kernel containing many layers of tissues with more or less properties of whole wheat for health extend well beyond its
distinct composition of bioactive compounds, particularly nutritional properties and derive for the most part from bioactive
between the outer pericarp and inner aleurone. compounds in the bran fraction. Those compounds provide a
The aleurone layer of bran has a high concentration of the significant source of dietary fiber and micronutrients, along with
fiber compound arabinoxylan and many other bioactive a substantial complement of phytochemicals including those
phytochemical compounds such as minerals, B vitamins, thought to have a role in reducing the risk of chronic diseases
phytic acid, and ferulic acid. such as atherosclerosis, obesity, diabetes, and cancer. This article
Bran is a heterogeneous mixture of several different types of presents a compilation and discussion of the diverse array of
nonstarch polysaccharides, which together constitute the major bioactive compounds that reside in wheat bran. These
fiber fraction of bran that represents most of the fiber in compounds in wheat are generally qualitatively similar across
whole wheat. different cereal grains, but specific proportions and distribution
The typical contents of bioactive compounds that are sig- in the kernel can be very different. The health efficacies of these
nificantly enhanced in bran compared to levels found in compounds are not substantially discussed as several excellent
whole wheat are presented. reviews or books on the topic are available for further reading.
Hairs of
brush
Endosperm
(~82%)
Crease
Nucellar tissue
Seedcoat (testa)
Tube cells Bran
Cross cells Inner and outer (~15%)
Hypodermis pericarp
Epidermis
Germ
(3%) Scutellum
Figure 1 The structure of the wheat kernel sectioned longitudinally (left) and transversely (right). There are six bran layers plus the aleurone, which is
invariably part of conventionally milled bran. The crease, which runs the full length of the kernel, prevents fully efficient separation of bran from
endosperm. Typical weight proportions contributed by bran, germ, and endosperm are shown. Adapted from Dexter JE and Sarkar AK (2004) Wheat: dry
milling. In: Wrigley C, Corke H, and Walker C (eds.), Encyclopedia of Grain Science, pp. 363374. Oxford, UK: Elsevier Science.
Table 1 Proximate and select nutrient composition of the three different components of whole wheat grain, with flour, bran, and germ being
products of commercial millinga
Nutrients Whole wheat (%) Refined flour (endosperm %) Bran (%) Germ (%)
a
Moisture 10.5 11.9 9.9 11.1
Proteina 13.4 10.3 15.6 23.2
Total lipida 2.1 0.98 4.3 9.7
Ash (minerals)a 1.6 0.47 5.8 4.2
Total carbohydrates (by difference)a 72.3 76.3 64.5 51.8
Total dietary fibera 11.5 2.7 47.8 13.2
Sugarsa 0.41 0.27 0.41 37.6
Total bioactive phytochemicalsb 1.2 nd 6.2 nd
a
Data source: USDA National Nutrient Database for Standard Reference (US Department of Agriculture, 2015). Carbohydrates includes total dietary fiber (TDF), starch, and sugars; the
sugar value for germ carbohydrates TDF.
b
Values for whole wheat and bran (not determined, nd for endosperm and germ) correspond to the sum of major phytochemicals considered and reported in this article in descending
order of content in bran: phytic acid, betaine, ferulic acid ferulic acid dehydrodimers, alkylresorcinols, choline, phytosterols, sum of other phenolic acids, total B vitamins, and
lignan.
wall of a wheat floret. The pericarp, which represents about 50% scutellum) have been found to represent about 83%,
of the thickness of bran, should not be confused with the hull or 1315%, and about 3% by weight of the whole grain, respec-
husk of the grain. Modern wheat has evolved as a free-threshing tively. These three fractions of the wheat kernel are relatively
grain, that is, the caryopsis is naked and does not include the distinct in basic chemical composition (Table 1). The endo-
highly fibrous hull, which is easily removed as chaff in mature sperm is energy-rich as it is predominantly composed of starch
wheat when threshed or harvested. In this way, wheat is distinct but contains less than 10% of the nutritive value (minerals,
from some other grains like most barley types, as well as oats and vitamins, and dietary fiber) of whole grain. Germ is distinct in
rice, whose kernels naturally include the hull. its relatively high protein and lipid content and exceptionally
The three major components of the wheat kernel are the high concentrations of sugars. Bran contains nearly all the
germ, bran, and endosperm (Figure 1). Based on research bioactive compounds of wheat, that is, it is enriched in fiber
involving careful wheat kernel dissection, endosperm, bran components (nonstarch polysaccharides and lignin), minerals,
(pericarp, testa, and aleurone), and germ (embryo and B vitamins, and other phytochemical compounds (Table 2).
270 BIOACTIVES AND TOXINS | Bioactive Compounds in Wheat Bran
From the seeds perspective, the germ contains the embry- concentrated in commercial wheat bran, which includes the
onic plant and an organ called the scutellum, which separates bioactive-rich aleurone layer that represents about 50% by
the embryo from the endosperm. During germination of the weight of total bran (Figure 1).
seed, the scutellum absorbs nutrients from the endosperm to It is important to point out that commercial wheat bran is
provide food for the germinated seed and growing seedling. not the same as botanical bran, that is, as it exists in the original
The biological function of bran is very different, which reflects grain, which excludes the aleurone layer, which developmen-
its complex and multilayered structure as well as chemical tally forms the outer part of endosperm (Figure 1). Wheat bran
composition; it is to protect the seed from microorganisms can be derived from various stages of the milling process and,
and insects and from adverse weather conditions. depending on the setup of the mill in terms of flour extraction,
The wheat kernel, as with all cereals, contains a crease that can vary in purity, that is, endosperm contamination, and
runs parallel to its long axis, which covers about 25% of its hence can vary in its content of fiber, starch, protein, and
surface. This crease affects the utilization of wheat for food, other constituents, due to milling effects alone and also due
such that the endosperm can only be freed from bran by to genotype, wheat hardness, and environmental conditions
conventional roller milling using relatively low flour extraction during wheat kernel development. Wheat bran is therefore not
rates. The low extraction rate, in the context of milling, mini- a standardized product of milling and can be produced from
mizes significant contamination of the flour by bran and conventional dry roller milling or a combination of conven-
germ and results in the production of refined flour, which is tional milling and abrasion processing similar to that used in
used to make the so-called white bread. Low extraction rates rice polishing. An example of how the degree of flour extrac-
in flour milling also result in the exclusion of the full comple- tion in commercial milling affects the retention of the B vita-
ment of the great many bioactive compounds, which are min group of bioactive compounds is shown in Figure 2. A
milling extraction rate of 100% is the equivalent of whole
wheat flour.
Table 2 Average concentration of phytochemicals enhanced in
wheat bran compared to whole grain (mg/100g)a
Nature of Variation in the Content of Bioactive
Phytochemicals Whole grain Bran
Components in Bran
Lignan 0.4 4.75
Phytosterols 78 158 It is clear from the literature that there is considerable variation
Betaine 156 868 in the content of bioactive components in wheat or bran
Choline 111 172 depending on the type of wheat or variety (genotype effect)
B vitamins, total 9.02 33.5 and the environment in which the wheat is grown. Recent
B1 (thiamin) 0.56 0.65 science has nicely documented this variation for European
B2 (riboflavin) 0.18 0.51 wheats and has shown that whole grains can vary from about
B3 (niacin) 6.5 28
1.5- to 4-fold depending on the specific compound and more
B5 (pantothenic acid) 1.35 3.15
importantly that some compounds have ample heritability to
B6 (pyridoxine) 0.38 1.0
B9 (folate) 0.05 0.23 allow breeders to further increase levels if that is desired.
Regarding bran, however, there is another source of variation
a
Values are means of range data in Table 2 of Fardet A (2010) New hypotheses for the that creates issues in accurately documenting bioactive compo-
health-protective mechanisms of whole-grain cereals: What is beyond fibre? nent content, that is, milling. As noted in the previous section,
Nutrition Research Reviews 23: 65134. wheat bran is not a standardized product of milling. While it is
possible to obtain pure bran or even isolate specific bran
tissues by hand dissection, this is not the form of bran available
100 Thiamine
Proportion of vitamin retained (%)
outer
pericarp aleurone
cross
cells
endosperm
500 25
410 Whole wheat
400 20.0 Whole wheat
Bran 20
296 Bran
mg/100 g
mg/100 g
300 15
200 10 9.0
117 6.3
73 5 3.7 3.5 3.2
100 2.2
1.5 1.3 0.7
0 0
Ferulic + dimers Alkylresorcinols
(a)
(b)
Figure 6 Phenolic compounds significantly enhanced in wheat bran compared to whole grain. Ferulic acid plus ferulic acid dehydrodimers and
alkylresorcinols (a) are the predominant phenolic compounds, each having at least tenfold higher concentration compared to the total of all other
phenolic compounds including simple phenolic acids (b) and flavonoids (not shown). Phenolic acid data compiled from a variety of sources.
Alkyresocinol values are the mean of those reported by Fardet (2010). Reproduced from Mattila et al. (2005) Contents of phenolic acids, alkyl- and
alkenylresorcinols, and avenanthramides in commercial grain products. Journal of Agricultural and Food Chemistry 53: 82908295.
OH
-O O
OH P P O-
O
O O
O
P O-
O
R O O- O-
P
O O-
R = CnH2n+1 O O
n = 15, 17, 19, 21, 23, 25 O P O-
OH O P O-
O- OH
OH
OH HO
HO
A
A HO
HO
OH O
O O O O
HO O
O X O X X O X
HO O O O
O OH O O OH
HO A
O
OH
H3CO
O
F Arabinoxylan
(c) HO
R3
Cinnamic acid derivatives
ferulic acid, R1 = OCH3; R2 = OH
R2 CH = CH- COOH sinapic acid, R = OCH ; R = OH
1 3 2
p-coumaric acid, R2 = OH
R1
Phenolic acids
R3
Figure 8 Basic molecular structures or portions of the structure of four types of bioactive compounds in wheat bran: alkylresorcinol (a); phytic acid (b);
arabinoxylan (c) showing xylose (X) backbone with substituted arabinose (A) molecules, some of which are linked to the phenolic acid ferulic acid (F); phenolic
acids (d). (Figure 8c adapted from Saulnier et al. (2007) Plant cell wall polysaccharides in storage organs: Xylans (food applications). In: Kamerling J,
Boons G-J, Lee Y, Suzuki A, Taniguchi N, and Voragen AGJ (eds.) Comprehensive Glycoscience, vol. 2, pp. 653689. Oxford: Elsevier Science Ltd.)
and lignin (Figure 5), is to promote intestinal health. There are Wheat bran (fiber) is particularly relevant in this discussion
an amazing number of bacterial cells in the colon in particular, as it is highly indigestible, yet fermentable by bacteria in the
and estimates run from 1011 to 1012 cells per gram. This huge colon, which have enzymes necessary to hydrolyze the poly-
population of cells, often referred to as the microbiome, is meric compounds in bran such as polysaccharides. It has been
gaining considerable importance in science due to an increased known for a long time that the stimulation of fermentation
understanding of the profound health effects, both positive caused by consuming these fiber bioactives leads to an increase
and negative, that good or poor gut health, respectively, can in bacterial mass and consequently fecal mass, which has a
contribute. positive effect on laxation and regularity. What is only being
274 BIOACTIVES AND TOXINS | Bioactive Compounds in Wheat Bran
understood very recently is that fermentation causes alterations the subaleurone layer, while in barley and rye, b-glucans are
in the microbiome, which in turn alters bacterial metabolites located mainly in the endosperm. The b-glucan levels of wheat
(short-chain fatty acids), modulates inflammatory responses, bran are low (2%) and represent <1% of the total dietary
and likely leads to a decreased risk of inflammation-associated fiber content of whole wheat. Because of its low concentration
and other metabolic abnormalities such as atherosclerosis and in wheat, b-glucans likely contribute relatively little by way of
type II diabetes. Also, by producing short-chain fatty acids, health attributes, compared to oats and barley. In these latter
most notably butyric acid, and other metabolites, the intestinal grains, b-glucan is considered to be an important soluble fiber
microbiome is believed to contribute to a gut environment that that is associated with lowering serum cholesterol levels and
resists the proliferation of pathogenic bacteria. attenuating glycemic response. These effects originate in the
upper gastrointestinal tract and probably are related to the
viscosity of b-glucan.
Arabinoxylans and Ferulic Acid
nature and classified according to linkage and origin. Fructoo- Compared to simple phenolic acids, alkylresorcinols represent
ligosaccharide is a subgroup of inulin, consisting of linear a very distinct group of phenolic compounds in terms of molec-
chains of fructose sugar monomers linked by b-(12) bonds ular properties, that is, they are amphiphilic, meaning they have
with a low degree of polymerization (10). Whole wheat grain both polar and nonpolar properties. This bioactive compound
and bran fructan contents are similar to those reported for consists of a phenolic ring with two hydroxyl groups and a
lignin. Fructans are classified as a functional fiber that are hydrophobic and mostly saturated alkyl chain that can contain
believed to be extensively fermented, but with little effect on from 15 to 25 carbons (Figure 8). Wheat bran is a rich source of
stool weight. Fructans are known to have positive effects on alkylresorcinols and has been reported to contain levels of about
calcium uptake in the gut and hence calcium bioavailability. 0.3%, which is fourfold higher than its content in whole wheat
(Figure 6). Refined flour contains essentially no alkylresorcinol
content. Accordingly, alkylresorcinols can be used as biochem-
Phenolic Acids
ical markers of whole wheat and bran in foods.
Several studies have reported that alkylresorcinols have the
Phenolic acids are a very noteworthy constituent of wheat
ability to protect cellular lipid components from oxidative pro-
bran. Apart from flavonoids, which are found to a large extent
cesses. Accordingly, alkylresorcinols appear to behave as antiox-
in wheat germ and in relatively low concentration in bran,
idant compounds, but they are generally known as weak
virtually all other phenolic compounds (phenolic acids and
antioxidants, based on their hydrogen-donating and radical-
phenolic acid esters) reside in wheat bran tissues, most notably
scavenging abilities, compared to some other bioactive com-
aleurone. Phenolic acids can be subdivided into two major
pounds such as a-tocopherol (vitamin E). Their antioxidant
chemical groupings: hydroxybenzoic acids and hydroxycin-
potential depends on their chain length, which affects their
namic acids (Figure 8). Hydroxybenzoic acids of significance
amphiphilic behavior and incorporation into cell membranes;
in bran, although levels are relatively low (Figure 5), include
studies have shown that the shorter the chain length, the higher
vanillic acid, synergic acid, and r-hydroxybenzoic acid. They
the inhibitory level of lipid oxidation. Also, membrane incorpo-
are typically components of complex compounds like lignins
ration of alkylresorcinols could accommodate a biological effect
and tannins. The concentration of hydroxybenzoic acids in
given their high concentration in wheat and bran, but limited
wheat and bran is low compared to hydroxycinnamic acids,
data on alkylresorcinols effective dose levels require further
which includes r-coumaric, and ferulic acid, which form links
studies on risk modulation of cancer and cardiovascular disease.
to cell wall structural components such as arabinoxylans,
Alkylresorcinols are absorbed through the lymphatic system and
cellulose, and lignins through ester bonds.
have been recovered from rat adipose tissue, human lipoproteins
By far, the predominant phenolic acid in wheat is (trans)
(the main transporter), human erythrocyte membranes, and
ferulic acid, which comprises more than 90% of total phenolic
human plasma. Animal studies have shown that alkylresorcinol
acids in wheat bran. Wheat bran is one of the richest sources of
is highly bioavailable, which indicates that apparently, the
ferulic acid with levels as high as 1400 mg/100g being
indigestible bran matrix does not greatly inhibit absorption.
reported. Ferulic acid is predominant in wheat aleurone,
pericarp, and embryo cells, while only trace amounts can be
found in starchy endosperm. Aleurone cells are associated with
the highest concentration of ferulic acid (Figure 3). The seed
Phytic Acid
coat layer of wheat bran contains significant quantities of
ferulic acid, higher than those in the pericarp. Total ferulic
Perhaps the most notable phytochemical constituent of wheat
acid concentration in wheat bran is typically about four times
bran is phytic acid, which is composed of a simple sugar (inosi-
higher than that in whole wheat grain (Figure 5).
tol) with phosphate groups attached to each of the six carbons in
There is considerable scientific evidence that ferulic acid, like
the sugar molecule (Figure 8). It is typically found in food sources
other phenolic compounds, has antioxidant, anticarcinogenic,
high in fiber content, although wheat bran may be the richest
and anti-inflammatory properties. The great majority of these
source in nature. Phytic acid represents the main storage form of
studies are based on animal or cell models. The types of antiox-
phosphorus in wheat and other cereal grains. Phytic acid repre-
idant activities possessed by phenolic acids include radical scav-
sents the most highly concentrated phytochemical in wheat and
enging, metal ion chelation, and inhibition of prooxidant
wheat bran in particular with average levels of 0.7% and 4%,
enzymes. Ferulic acid has been extensively studied for its
respectively (Figure 7). By contrast, endosperm tissue of wheat
in vitro antioxidant effects and has been shown to be effective.
contains very little phytic acid. Phytic acid has traditionally been
However, in vivo studies of phenolic acids from cereals like
considered to be an antinutrient associated with its ability to bind
wheat and fractions like wheat bran are few. Cereal grains such
minerals with positively charged ions (i.e., cations) such as iron,
as wheat that have significant quantities of ferulic acid and other
as well as magnesium and calcium. This binding of minerals
phenolics require in vivo studies to determine if high enough
therefore reduces their bioavailability, but only for poorly nour-
levels of these bioactive compounds become bioavailable when
ished people consuming predominantly whole grains in their
consumed in food to provide health benefits.
diet. Many animal studies have shown phytic acid to be capable
of antineoplastic activities on multiple types of cancers of the
Alkylresorcinols breast, colon, liver, prostate, skin, and skeletal muscle. Potential
mechanisms suggested for phytic acids anticarcinogenic proper-
Alkyresorcinols are phenolic lipids that are present exclusively in ties include cell cycle inhibition, antioxidant ability (metal chela-
bran and specifically in the outer layer of the seed coat. tion), gene alteration, and increased natural killer cell activity.
276 BIOACTIVES AND TOXINS | Bioactive Compounds in Wheat Bran
Summary and Conclusions for food use and how much for feed? How do these pro-
portions vary in major wheat producing countries?
The nature of wheats value for human health derives from its Are there wheat genotypes or varieties that have bran with
structure, biochemical composition, and functionality of its enhanced levels of bioactive compounds?
components. Because wheat consumption as bread and other How does the wheat growing environment affect the con-
foods is so significant, whole wheat and the bran it contains tent of bioactive compounds in bran?
have the potential to deliver to the diet considerable amounts
of fiber and constituent bioactive compounds whose function-
ality extends well beyond the role of nutrients, which are Exercises for Readers to Explore the Topic Further
essential for growth and maintenance of life. There is now
ample scientific evidence that consumption of adequate How do bioactive compounds in the bran or whole grain of
amounts of bran in whole wheat is associated with lowering other cereals compare to wheat, especially oats, given its
the risk of many chronic diseases, although the specific actions reputation for having dietary benefits?
of individual biologically active compounds in bran have been Select a few fruits or vegetables that are commonly associ-
only studied using animal and cell models. A more thorough ated with health benefits. How similar or different are the
discussion of the potential health efficacies of the major bio- bioactives in those foods compared to wheat bran?
active compounds of wheat bran is well beyond the scope of Apart from eating whole wheat bread or pasta, think of
this article. Many excellent books or review papers have been other ways to increase your consumption of wheat bran
published and a few are offered at the end of this article as that would appeal to you.
suggestions for further reading. These articles are based on a
foundation of hundreds of research studies, and the science in
this area continues to expand in scope and knowledge. Most See also: Bioactives and Toxins: Bioactives: Antioxidants;
exciting perhaps is the microbiome-related research on the Carbohydrates: Cereals: Chemistry and Physicochemistry of Non-
fermentation of fiber and bran that is gaining momentum. starchy Polysaccharides; Food Grains and the Consumer: Grains
Wheat bran is generally considered to be one of the best, if and Health; Food Grains and Well-being: Functional Foods:
not the best, sources of fiber for fecal bulking and laxation. This Dietary Fibers, Prebiotics, Probiotics, and Synbiotics; Functional
sort of bioactivity represents the traditional health benefit Foods: Overview; Grain Composition and Analysis: The
ascribed to bran, but it also suggests that wheat bran interacts Composition of Food Grains and Grain-Based Products; The
strongly with the microbiome and that this area of research has Basics: Grain: Morphology of Internal Structure.
enormous potential for greater understanding of mechanisms
underlying the more compelling health efficacies generally
associated with the consumption of whole wheat and bran Further Reading
and its constituent bioactive compounds.
Anson NM, Hemery YM, Bast A, and Haenen GR (2012) Optimizing the bioactive
potential of wheat bran by processing. Food and Function 3: 362375.
Fardet A (2010) New hypotheses for the health-protective mechanisms of
Exercises for Revision whole-grain cereals: What is beyond fibre? Nutrition Research Reviews
23: 65134.
Marquart L, Jacobs Jr. DR Jr., McIntosh GH, Poutanen K, and Reicks M (eds.) (2007)
Are there wheat milling procedures that can generate bran Whole Grains and Health. Oxford: Blackwell.
of higher purity that is more concentrated in bioactive Okarter N and Liu R (2010) Health benefits of whole grain phytochemicals. Critical
compounds? Reviews in Food Science and Nutrition 50: 193208.
Shewry PR, Charmet G, Branlard G, et al. (2012) Developing new types of
What is the function of the bioactive compounds in wheat
wheat with enhanced health benefits. Trends in Food Science and Technology
bran from the perspective of the wheat seed? 25: 7077.
How much wheat bran is typically produced in a large Slavin J (2013) Fiber and prebiotics: Mechanisms and health benefits. Nutrients
milling operation and how much of that bran is destined 5: 14171435.
Bioactives: Antioxidants
T Beta, University of Manitoba, Winnipeg, Manitoba, Canada
KG Duodu, University of Pretoria, Pretoria, South Africa
2016 Elsevier Ltd. All rights reserved.
Source: Fardet A (2010) New hypothesis for the health-protective mechanisms of whole-grain cereals: What is beyond fibre? Nutrition Research Reviews 23: 65134.
diseases including cancer, cardiovascular disease, type 2 structure and mechanism of action, which can be direct or
diabetes, and even cataractogenesis. indirect. Mechanisms may involve direct free radical scaveng-
It is hypothesized that bioactives from cereal grains could ing or Fe chelation. Indirect antioxidants such as Fe, Zn, Cu,
offer potential health benefits by protecting against diseases and Se act as cofactors of antioxidant enzymes. Very few studies
associated with oxidative stress. A lot of in vitro evidence has have examined the antioxidant hypothesis; hence, in vivo inves-
emerged that shows the ability of cereal bioactives to protect tigations are needed to determine the complex and synergistic
cell constituents against oxidative stress and oxidative damage mechanisms taking place in human subjects.
of biological tissues. This observed protective effect has been
attributed to various mechanisms. These include the suppres-
sion of the formation of reactive oxygen species by either Sulfur Compounds
inhibiting enzymes or chelating trace elements involved in
free radical production, scavenging reactive oxygen species, or Total methionine and cystine contents of whole-grain wheat,
upregulating or protecting antioxidant defense systems. wheat bran, and wheat germ are 0.5%, 0.6%, and 1.0%, respec-
tively (Table 1). Other cereal varieties may contain higher levels.
Methionine and cystine contents of commercially prepared
whole-wheat bread based on USDA database are, respectively,
Antioxidants from Whole-Grain Cereals 155 and 214 mg/100 g, contributing an average 47 and 64 mg
day1 when a 30 g slice of bread is consumed. Significant
Whole-grain cereals serve as sources of antioxidants that amounts of at least 280 mg methionine and 380 mg cystine
include sulfur compounds, minerals, B vitamins, phytic acid, per day can be obtained by following USDA food guide pyramid
tocols, carotenoids, phenolic compounds, lignins, lignans, that recommends between 6 and 11 daily servings of whole-
alkylresorcinols, betaine, choline, policosanol, melatonin, grain cereal products. The two sulfur-containing amino acids are
g-oryzanol in rice bran, avenanthramides, and saponins in both precursors of reduced glutathione (GSH), an intracellular
oats as reviewed by Fardet. These antioxidants differ in their antioxidant that controls cell oxidative status by participating in
BIOACTIVES AND TOXINS | Bioactives: Antioxidants 279
gene expression through the modification of the thiol redox ADPFeoxygen complexes that initiate lipid peroxidation.
status. GSH is also present at levels of 1.045.74, 1.719.4, Phytic acid may play a role in the prevention of various cancers
and 19.4245.7 mg/100 g in whole-grain wheat, wheat bran, including colon cancer.
and wheat germ, respectively, although its absorption in
humans is deemed negligible in its free form.
Lignins and Lignans
Lignins range from 0.9% to 2.8% in whole-grain wheat, 1.9% to
Minerals
5.6% in wheat bran, and 1.5% in wheat germ. They have been
The contents of Fe, Zn, Mn, Mg, Cu, and Se in whole-grain shown to have antioxidant effects on ex vivo fresh lymphocytes.
wheat, wheat bran, and wheat germ are shown in Table 1 as They account for 2632% of the enterolactone (a mammalian
reviewed by Fardet. Low levels of 3.8% Fe are absorbed in lignan) formed from cereal bran. Mammalian lignans are anti-
humans following the consumption of wheat bran rolls. oxidants in vitro at concentrations of 10100 mM achievable
About 1720% and 1995% of zinc is absorbed in humans in vivo, particularly in the colon. Lignans make up 0.206,
and rats, respectively, following the consumption of whole- 2.86.7, and 0.49 mg/100 g of whole-grain wheat, wheat bran,
grain wheat. There is limited literature on apparent absorption and wheat germ, respectively (Table 1). Lignins are neither
of Mn in humans, although levels are known to reach absorbed nor fermented in the human GIT. Only 04% of
1418 mg/100 g in wheat bran and wheat germ compared to lignins are apparently absorbed in rats following the consump-
0.0780.079 mg/100 g for selenium in respective fractions. tion of raw and processed wheat bran. Their antioxidant activity
Studies have shown a 6085% absorption of selenium in rats due to the phenolic hydroxyl group may contribute to the
following the consumption of wheat and sodium selenite. prevention of colon cancer.
Fe, Zn, Cu, and Se act as cofactors of antioxidant enzymes
and therefore serve as indirect antioxidants. Cu exerts its anti-
Lipotropes and Methyl Donors (Betaine, Choline, and Folates)
oxidant effect mainly as superoxide dismutase cofactor. Mg
displays antioxidant effects by acting against lipid peroxida- Whole-grain wheat, wheat bran, and wheat germ contain about
tion. Se protects cell membranes from damage caused by lipid 0.28%, 1.04%, and 1.09% of betaine and choline, respectively,
oxidation. and about 51, 231, and 420 mg folates/100 g. Betaine, choline,
and folates are lipotropic compounds together with methionine
and myoinositol that prevent excess fat deposition in the liver by
Thiamin and Riboflavin
hastening fat removal or by limiting lipid synthesis. Whole-grain
Both thiamine (B1) and riboflavin (B2) are present in grains wheat, wheat bran, and wheat germ can supply 0.51, 1.31, and
including wheat (Table 1). The two B vitamins exert antioxi- 1.59 g lipotropes/100 g, respectively. Betaine is likely to be more
dant effects in addition to their several well-documented phys- bioavailable as it is not bound to cell wall constituents as is the
iological functions. case with ferulic acid. Data on apparent absorption of betaine
and choline are not available.
Phytic Acid
g-Oryzanols
Phytic acid (myoinositol hexakisphosphate) can reach up to 6%
in the bran of certain wheat varieties. It displays strong in vitro g-Oryzanols or phytosteryl ferulates mainly found in rice bran
antioxidant activity; however, it has been considered an antinu- have been reported to have antioxidant activity. They are con-
trient due to its ability to chelate minerals such as Zn, Fe, Ca, and sidered more effective than tocopherols and tocotrienols in the
Mg, reducing their bioavailability. Processing procedures such as prevention of cholesterol oxidation. They have the ability to
the preparation of sourdough to produce bread can activate scavenge radicals in multiphase lipid systems and inhibit UV
endogenous phytases and lower pH, thereby reducing the che- irradiation-initiated linoleic acid oxidation.
lation of minerals by phytic acid. Phytic acid can also chelate Fe
and limit the oxidative damage caused by the Fenton reaction,
Avenanthramides in Oats
which would otherwise lead to the production of the very
l
reactive free radical OH . Strategies are required that pre- Avenanthramides are found only in oats, particularly in the
hydrolyze phytic acid so as to maximize mineral bioavailability outer layers. Avenanthramides or hydroxycinnamoyl anthrani-
while simultaneously enhancing its antioxidant effect against linate alkaloids are amides of various cinnamic acids with dif-
free radicals in the gastrointestinal tract (GIT). ferent anthranilic acids. Avenanthramides 2p, 2f, and 2c are the
Phytic acid is poorly absorbed in humans. In the absence of most abundant among over 35 forms identified to date. Oat
the enzyme phytase, phytate is not likely to be absorbed at the grain contains 413 mg avenanthramides 1/100 g (the major
intestinal level in humans. About 5860% of phytate is avenanthramide), while oat bran contains 1.312.5 mg/100 g
degraded into lower myoinositol phosphates in ileostomates according to the type of avenanthramide considered.
following the consumption of raw wheat bran and only 5% The inclusion of avenanthramide-2c (N-(30 ,40 -dihydroxy-
following the consumption of phytase-deactivated wheat bran. cinnamoyl)-5-hydroxyanthranilic acid) in rat diets altered
As an antioxidant, phytic acid chelates various metals includ- oxidantantioxidant balance in various tissues. It resulted in
ing the suppression of damaging Fe-catalyzed redox reactions, selective reduction in the production of exercise-induced reac-
inhibits xanthine oxidase, suppresses oxidant damage to the tive oxygen species and lipid peroxidation in the rats. This is
intestinal epithelium, and interferes with the formation of related to the ability of the avenanthramides to modulate
280 BIOACTIVES AND TOXINS | Bioactives: Antioxidants
Future Prospects
Aruoma OI (1998) Free radicals, oxidative stress, and antioxidants in human health and Ndolo VU and Beta T (2014) Comparative studies on composition and
disease. Journal of the American Oil Chemists Society 75: 199212. distribution of phenolic acids in cereal grain botanical fractions. Cereal Chemistry
Belodrajdic DP and Bird AR (2013) The potential role of phytochemicals in wholegrain 91: 522530.
cereals for the prevention of type-2 diabetes. Nutrition Journal 12: 62. Neveu V, Perez-Jimenez J, Vos F, et al. (2010) Phenol-Explorer: An online
Fardet A (2010) New hypothesis for the health-protective mechanisms of comprehensive database on polyphenol contents in foods. Database. doi:10.1093/
whole-grain cereals: What is beyond fibre? Nutrition Research Reviews database/bap024. Version 1.5.2. http://www.phenol-explorer.eu.
23: 65134. Okarter N and Liu RH (2010) Health benefits of whole grain phytochemicals. Critical
Grassi D, Desideri G, and Ferri C (2010) Flavonoids: Antioxidants against Reviews in Food Science and Nutrition 50: 193208.
atherosclerosis. Nutrients 2: 889902. Perez-Jimenez J, Neveu V, Vos F, and Scalbert A (2010) Systematic analysis of the
Li W and Beta T (2013) Food sources of phenolics compounds. In: Ramawat KG and content of 502 polyphenols in 452 foods and beverages: An application of the
Merillon JM (eds.) Natural Products: Phytochemistry, Botany and Metabolism Phenol-Explorer database. Journal of Agricultural and Food Chemistry
of Alkaloids, Phenolics and Terpenes, pp. 25272558. Berlin, Heidelberg: 58: 49594969.
Springer-Verlag. Ryan D, Kendall M, and Robards K (2007) Bioactivity of oats as it relates to
Ndolo VU and Beta T (2013) Distribution of carotenoids in endosperm, germ, and cardiovascular disease. Nutrition Research Reviews 20: 147162.
aleurone fractions of cereal grain kernels. Food Chemistry 139: 663671.
The Antinutritional Components of Grains
CK Madsen and H Brinch-Pedersen, Aarhus University, Slagelse, Denmark
2016 Elsevier Ltd. All rights reserved.
Topic Highlights negatively charged phosphate groups (Figure 1). The strength
of the interactions depends on the charge of the cations
Antinutritional compounds interfere with the utilization of (Me3 > Me2 > Me) and to a lesser extent other parameters
nutrients and energy. such as ionic radius. At acidic pH, the metal cations compete
The major antinutritionals of grains are phytate, proteina- with H and interactions with most metal cations are insignif-
ceous inhibitors of digestive enzymes, tannins, and non- icant below pH 5.
starch polysaccharides. In grains, IP6 typically accounts for 2/3 of the total P
Antinutritionals may also offer nutritional benefits that content (see Table 1 for exact values). This IP6 is found as an
have to be balanced against their negative effects. insoluble mixed salt with cations called phytin. In many grains
The significance of antinutritionals depends on species and including cereals and legumes, phytin forms spherical inclu-
nutritional status of the human or animal consuming it. sions called globoids inside protein storage vacuoles. Not only
are the globoids the principal site of P, K, and Mg in the mature
cereal grain, but also they contain Ca, Fe, Zn, Cu, Mn, and Na.
Learning Objectives Globoids are predominantly found in the aleurone layer of
small grain cereals and in the embryo of maize, tissues that
are the main contributors to millers bran fraction. Accord-
To achieve an understanding of the chemistry, occurrence,
and antinutritional mechanism of the major antinutritional ingly, the IP6 contents in bran and white flour are higher and
compounds in grains. lower, respectively, than those in whole grain. IP6 is resistant to
most phosphatases, whereas the lower inositol phosphates are
To gain an overview of strategies to counter
antinutritionals. substrates of a wider range of phosphatases. Phosphatases that
can initiate dephosphorylation of IP6 are known as phytases.
All plants express phytases to assist in their IP6 metabolism but
the phytase activity in mature seed varies several orders of
Introduction magnitude, even in closely related species. This is in sharp
contrast to the modest variation in total and proportional
Antinutritionals are compounds that reduce the bioavailability content of IP6 (Table 1). The Triticeae tribe of cereals (such
of essential nutrients or energy in the diet. This leads to dis- as wheat, barley, rye, and triticale) stand out with exceptionally
crepancies between the formal and actual nutritional values of high mature grain phytase activity. This is caused by the rela-
a food or feed. It is therefore most important that both nutri- tively recent duplication of a purple acid phosphatase phytase
ents and antinutritionals are part of the equation when the gene whose archaic function was to be expressed during ger-
nutritional value of a food item is evaluated. Moreover, it is mination. Following the gene duplication, one copy has chan-
important to notice that some antinutritionals may also pos- ged expression pattern so it is expressed during grain filling.
sess positive dietary properties and that nutritional goals in The reported intake of IP6 for adults ranges from 180 to
human and animal nutrition can be very different, that is, 5577 mg day1. Omnivorous Westerners are in the low range
with respect to weight gain. Antinutritionals that are deleteri- of the interval (usually less than 1000 mg day1). Vegetarians
ous for the calorie yield of a diet may be used to counter and people in developing countries are in the high range
obesity in a human diet, whereas they result in poor resource (>1000 mg day1). The highest value (5577 mg day1) has
use efficiency in animal husbandry. been reported for a group of Trappist monks on a lacto-ovo
The use of the term antinutritionals is inconsistent in the vegetarian diet. The utilization of IP6 by humans and most
scientific literature and may sometimes refer to compounds nonruminants is negligible unless the meal has sufficient phy-
that are considered overall unhealthy (i.e., allergens, irritants, tase activity. Fast-growing animals on a high phytate contain-
and toxins). Such compounds are beyond the scope of this ing feed without P supplements may suffer from P deficiency.
article. The low retention of IP6 affects not only the molecule per se
but also its chelation partners. Because of the chemical prop-
erties discussed earlier in the text, IP6 will release chelated
Phytate metals in the low pH of the stomach but recruit ions again
from the food matrix as pH reaches neutral during the duode-
Phytic acid or myo-inositol-(1,2,3,4,5,6)hexakisphosphate nal passage. Therefore, IP6 from one source exerts an antinu-
(IP6) is the major phosphorus (P) storage compound in tritional effect on the entire meal. In vivo, this has been shown
plants. IP6 is hexaphosphorylated myo-inositol. At physiolog- to affect zinc, iron, calcium, magnesium, copper, and manga-
ical pH, it adopts a conformation with two phosphates in the ne