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Proceedings of an International Symposium

H.E. Vivar and A. McNab, Editors

INTERNATIONAL MAIZE AND WHEAT IMPROVEMENT C ENTER


83

Breeding Barley
in the New Millenium:

Proceedings of an International
Symposium

held on
13-14 March 2000

in
Ciudad Obregon, Sonora, Mexico

H.E. Vivar and A. McNab, Editors

NTERNATIONAL CENTER FOR AGRICULTURAL RESEARCH IN THE DRY AREAS

INTERNATIONAL MAIZE AND WHEAT IMPROVEMENT CENTER


CIMMYT® (www.cimmyt.org) is an internationally funded, nonprofit, scientific research and
training organization. Headquartered in Mexico, CIMMYT works with agricultural research
institutions worldwide to improve the productivity, profitability, and sustainability of maize
and wheat systems for poor farmers in developing countries. It is one of 16 food and
environmental organizations known as the Future Harvest Centers. Located around the world,
the Future Harvest Centers conduct research in partnership with farmers, scientists, and
policymakers to help alleviate poverty and increase food security while protecting natural
resources. The centers are supported by the Consultative Group on International Agricultural
Research (CGIAR) (www.cgiar.org), whose members include nearly 60 countries, private
foundations, and regional and international organizations. Financial support for CIMMYT’s
research agenda also comes from many other sources, including foundations, development
banks, and public and private agencies.

Future Harvest® builds awareness and support for food and environmental research for a
world with less poverty, a healthier human family, well-nourished children, and a better
environment. It supports research, promotes partnerships, and sponsors projects that bring the
results of research to rural communities, farmers, and families in Africa, Asia, and Latin
America (www.futureharvest.org).

 International Maize and Wheat Improvement Center (CIMMYT) 2001. All rights reserved.
The opinions expressed in this publication are the sole responsibility of the authors. The
designations employed in the presentation of materials in this publication do not imply the
expression of any opinion whatsoever on the part of CIMMYT or its contributory organizations
concerning the legal status of any country, territory, city, or area, or of its authorities, or
concerning the delimitation of its frontiers or boundaries. CIMMYT encourages fair use of this
material. Proper citation is requested.

Correct citation: Vivar, H.E., and A. McNab (eds.). 2001. Breeding Barley in the New Millenium:
Proceedings of an International Symposium. Mexico, D.F.: CIMMYT.

AGROVOC Descriptors: Plant breeding; Breeding methods; Barley; Germplasm; Plant


diseases; Disease resistance; Environmental factors; Research projects

Additional Keywords: CIMMYT

AGRIS Category Codes: F30 Plant Genetics and Breeding


A50 Agricultural Research

Dewey Decimal Classif.: 633.16

ISBN: 970-648-078-1

Printed in Mexico.

Design and layout: Miguel Mellado E.

Front and back cover photos: H.E. Vivar.


iii

Table of Contents
iv Preface

1 Learning about Barley Breeding—D.C. Rasmusson

7 Hull-less Barley: An Overview—J.H. Helm

10 Contributions of Breeding and Crop Management to Increasing Barley


Yield and Grain Quality in Chile—E. Beratto M.

18 Importance of Identifying Stripe Rust Resistance in Barley—W.M.


Brown, J.P. Hill, and V.R. Velasco

25 Impact of ICARDA/CIMMYT Barley Germplasm on Barley Breeding in


Ecuador—O. Chicaiza

28 Malting Barley for the New Millenium—L. Wright

34 Barley Breeding in Peru—M. Romero Loli and L. Gomez Pando

39 Accumulating Genes for Disease Resistance in Two-Rowed Barley for


North Dakota—J.D. Franckowiak

47 Collaborative Stripe Rust Resistance Gene Mapping and Deployment


Efforts—P.M. Hayes, A. Castro, A.E. Corey, T. Filuchkin, C. Rossi, J.S.
Sandoval, I. Vales, H.E. Vivar, and J. Von Zitzewitz

61 Perspectives on Fusarium Head Blight Resistance in Barley—L.


Gilchrist

72 Resistance and/or Tolerance to BYDV: Recent Advances in Barley at


CIMMYT—M. Henry and H.E. Vivar

77 Two Decades of Barley Breeding—H.E. Vivar


iv

Foreword

One aspect of CIMMYT’s mandate is to barley yellow dwarf (BYD), into a high
develop improved barley varieties for Latin yielding background. Resistance to scald
America, a task we carry out in and leaf rust was incorporated in the early
collaboration with one of our CGIAR sister stages, followed by stripe rust and other
centers, the International Center for diseases. Genes conferring resistance to net
Agricultural Research in the Dry Areas blotch, spot blotch, and stem rust were
(ICARDA). Our collaborative efforts are added later. By applying this method, the
embodied in the ICARDA/CIMMYT Program succeeded in developing high
Barley Breeding Program for Latin yielding barleys with resistance to multiple
America, an excellent example of diseases that effectively safeguard
partnership within the CGIAR that has smallholder food security in the Andean
operated successfully since the early 1980s. Region. In essence, the Program provided
farmers with an exceptional resource—
Barley can grow in extremely inhospitable
greatly improved seed—that enabled them
environments where other crops would
to move beyond the limitations imposed by
have a difficult time surviving—for
their harsh farming environment.
example in the high valleys (2,700-3,500
meters above sea level) of the Andean The success of the Program has been such
Region in South America, where it is very that it has produced significant spillover
cold and soils are rocky and infertile. In this benefits in regions beyond Latin America. A
region, which includes Bolivia, Ecuador, case in point is its very impressive impact
Peru, and southern Colombia, barley does in China, where barleys developed by the
well, though farmers’ circumstances do not Program cover 40% of the one million
allow them to invest much in it: they do hectares planted to the crop. This impact is
very little land preparation and do not due mainly to the high yield potential of
control diseases or insect pests, despite the ICARDA/CIMMYT barley germplasm and
fact that barley provides both animal feed its resistance to fusarium head blight and
and food for farm families. barley yellow mosaic virus.
Regional farmers traditionally planted local Perhaps the most noteworthy impact of the
barleys that were low yielding and Program is that thanks to its efforts, there is
susceptible to diseases; as a result they now a large pool of barley germplasm
often suffered disastrous yield losses that possessing good agronomic traits, high
seriously threatened household food yield potential, and resistance to numerous
supplies. Breeding to incorporate durable diseases that is freely available to breeding
resistance to multiple diseases programs all over the world. It is our hope
consequently became one of the main that the achievements of this collaborative
goals—and outstanding achievements—of Program will be multiplied a hundred-fold
the ICARDA/CIMMYT Breeding Program. by the use barley breeders—particularly in
the developing world—will make of these
The approach utilized during the 20-year
resources.
breeding process was to create templates
for incorporating resistance to diseases
endemic in the region, such as stripe and
leaf rusts, scald, fusarium head blight, and Timothy G. Reeves
Director General, CIMMYT
v

Preface

These are the proceedings of an the three rusts, BYDV, fusarium head
international barley symposium held on blight (FHB), scald, and net blotch. It
13-14 March 2000 in Ciudad Obregon, should be noted that CIMMYT took the
Sonora, Mexico. The primary reason for lead in introducing FHB resistant
holding this special seminar was to varieties into China through the variety
honor Dr. Hugo E. Vivar, who retired in Gobernadora.
February 2000 after a long and
There is no doubt great progress has
extraordinarily fruitful career. Barley
been achieved in improving barley for
researchers from Latin America, the
food, feed, and forage. The crop has
USA, and Syria who were closely
been endowed with traits such as high
associated with Dr. Vivar were invited to
yield potential, multiple disease
make presentations at the symposium.
resistance, and good grain quality.
Dr. Vivar became the head of the However, in the future research will also
ICARDA/CIMMYT Barley Breeding have to focus on improving the quality
Program for Latin America in 1984, after of malting barley, a cash crop that would
working for CIMMYT for nine years. In provide barley producers in developing
the course of his long career, Hugo countries with a promising option for
worked on different types of barley for earning their living.
diverse environments and uses, but
In breeding one always builds on other
made a special effort to develop barleys
people’s work, and the exchange of
for marginal environments, such as
germplasm and information is crucial to
those in the Andean Region of his native
developing new varieties. We take this
South America, where subsistence
opportunity to acknowledge the
farmers use barley for food. The higher
openness and cooperation of the
yields produced by new, disease
international community of barley
resistant barleys have significantly
breeders, especially North American
improved farm families’ food security all
barley breeding programs, whose
year round.
collaboration over many years has been
The main focus of the ICARDA/ invaluable to ICARDA/CIMMYT barley
CIMMYT barley breeding program, improvement efforts. We hope these
under CIMMYT leadership, is on Latin proceedings will stand as a small
America, but the barleys it has memorial to the accomplishments not
developed are also sown in other parts only of Dr. Vivar, a dedicated breeder,
of the world, such as China, Pakistan, but also of a generous group of barley
and Kenya. One of the reasons they are scientists all over the world.
so widely used is that they possess
resistance to multiple diseases such as

Sanjaya Rajaram
Director, CIMMYT Wheat Program
1

Learning about Barley


Breeding
D.C. RASMUSSON1

Plant breeding programs are dynamic • The challenging and innovative years,
and always require change because when we adopted new strategies and
pests, target area, markets, the plant our breeding program became
breeder’s toolbox (e.g., plot equipment, productive.
computers, and molecular-based • The still challenging years, i.e., ensuring
methods), and available resources are gains in a mature program.
constantly changing. Thus there is need
for learning on a continuing basis. It is
interesting to speculate about learning The Formative Years
and to ponder how closely breeding Early in my career, I had the good fortune
success is related to learning and of associating with a large number of
adopting new philosophies and new interesting and innovative plant breeders.
strategies. In this context I would like to There are too many to mention, but I
share some of my memorable learning would like to comment on a few. Among
experiences with you. them were barley breeders Rollo
Woodward, at Logan, Utah, and Charles
In thinking about this presentation and
Schaller, at Davis, California. I learned
my title “Learning about Barley
pedigree breeding from Rollo Woodward,
Breeding,” it occurred to me that many
my master’s degree adviser. It was a
of you have had experiences similar to
small but productive barley program
those I will describe. Plant breeders tend
where hand cycling of plots was the
to move forward collectively in how we
norm. Charles Schaller, my Ph.D. adviser,
think about and how we do plant
did mainly backcross breeding. Together
breeding. So those of you that are older
we found the YD2 gene that provides
will likely conclude that you have had a
resistance to the barley yellow dwarf
learning experience similar to my own.
virus. He said that YD2 would be the
I have divided the presentation into largest contribution we would make in
three parts: our lifetime. Coit Suneson, also a barley
breeder at Davis, was a proponent of
• The formative years, when my
utilizing mixtures and allowing natural
thinking about barley breeding was
selection to improve populations. His
shaped by advisors and more
experienced mentors. hallmark paper was “An Evolutionary
Plant Breeding Method.” For an aspiring
barley breeder, Davis, California, in the

1 Agronomy and Plant Genetics, University of Minnesota, 411 Borlaug Hall, 1991 Buford Circle,
St. Paul, MN 55108, USA.
2 D.C. Rasmusson

1950s provided a first rate learning Adopting New Strategies


experience. Hardly a week passed
When I reflect on my early years in
without a thought-provoking discussion
barley breeding, a few items come to
about breeding methods.
mind that have contributed in a major
The most enriching experience of my way to our breeding effort. I am referring
career, in terms of learning plant to four practices or innovations that we
breeding, occurred in Ciudad Obregon, adopted between 1960 and 1970:
Mexico. For more than 20 years I had the germplasm sharing, arriving at
good fortune of having an annual appropriate breeding objectives, reducing
learning experience with CIMMYT staff. development time, and the paramount
When I came to Ciudad Obregon for the role of parental germplasm.
barley harvest each year, I had the
opportunity to tour the plots, view the Sharing germplasm
large impressive program and, more I learned about the value of shared
importantly, to explore breeding germplasm when we benefited
philosophies and breeding methods. We immensely from parent germplasm given
tried to resolve how many crosses to to us by colleagues located at North
make, which types of parents to use, Dakota, USA, and Brandon, Manitoba,
whether or not to select on the Canada. Their germplasm was ideal since
individual plant basis and how best to they are neighbors and they were leaders
breed for various mega-environments. in barley improvement in the USA and
Because it was recognized worldwide as Canada. Their germplasm contributed
a leading center for small grains improved agronomic performance,
improvement, Obregon was a gathering disease resistance, and malting quality.
place for small grain researchers from all Subsequently, we were able to return the
over the world. In addition to gaining favor when they used our varieties
valuable insights about plant breeding, I Morex and Robust, and their progeny, as
learned the importance of teamwork and parents in their programs. Morex was a
of sharing ideas and germplasm. hallmark variety quality-wise.

In reflecting on my career and about


Arriving at appropriate
learning how to do barley breeding, I breeding objectives
always come back to colleagues whose Upon arriving in Minnesota, I found
ideas, research methods, and germplasm many opportunities including the need
were available for the asking. One such for a sizeable breeding effort for
colleague was Hugo Vivar. I followed his resistance to diseases. Our germplasm
barley program with special interest. was relatively low yielding, tended to
When I came to Obregon, I would visit lodge, and kernel plumpness was not
the barley breeding plots with Hugo, adequate. So I concluded that it was
and every year I obtained useful appropriate to adopt a broad set of
information and a valuable gift of objectives focusing on field performance.
germplasm from one sub-program or
another. I was favorably impressed by But, after some time, I observed that
the many breeding objectives and successful varieties in the Midwest six-
overall quality of his nurseries. row malting barley area had one feature
Learning about Barley Breeding 3

in common. They had the malting and Germplasm is paramount


brewing industry stamp of approval and I learned about value of parent
brought a premium at the local elevator. germplasm slowly, and only after many
Subsequently, our breeding efforts came years did I come to realize that parental
to revolve around malting and brewing germplasm is paramount, especially
quality. A part of this learning experience when malting and brewing quality are
was recognizing that malting and high priority breeding goals. Two factors
brewing quality is determined by a were of primary importance in arriving
complex set of approximately 35 traits at this conclusion. The first was
that are evaluated in three major recognizing that Minnesota barley
groupings: 1) barley and malt variety candidates did not fare well in
performance (N ~ 14), 2) brewhouse competition with barley lines from other
wort/fermentation characteristics (N ~ breeding programs in the region.
12), and 3) beer characteristics (N ~ 9). Secondly, the notion that parental
Since mid-generation and even late- germplasm is paramount was solidly
generation selection cannot be practiced reinforced when I reaped the benefits
for some of these 30+ traits, it was from using elite North Dakota and
essential to limit parent choices to elite Canadian germplasm. Their germplasm
quality germplasm on both sides of the provided a large step forward for several
pedigree. Hence, parent building has traits including lodging resistance,
played an increasingly important role as disease resistance, grain yield, and
we attempted to incorporate desired traits malting/brewing quality.
into a malting barley genetic background.
Additionally, I had numerous
opportunities to observe a wide array of
Reducing development time
parental germplasm in our ideotype
In most breeding programs, a barley
research effort. Time and time again, I
breeder is challenged by evolving pest
observed that inferior and mediocre
populations, varying environments and
barley lines do not make good parents.
in some cases changing markets. These
We had to be content to do parent
factors may mandate a rapid response by
building for two to three cycles to obtain
the breeding program. Another reason for
parents that might lead to new varieties.
shortening development time is
We needed good performing parents on
competition among breeding programs.
both sides of the pedigree.
Acceptance of a new variety may depend
on whether it is the first variety to
provide new resistance or a trait of special
interest to a grower or an end-point user. Ensuring Progress in a
Yet another reason for reducing Mature Breeding Program
development time is the growing need for
parent building, which frequently In the early years of a barley breeding
requires two to three cycles of crossing program (perhaps a couple of decades),
and selection. In this situation, reducing there are usually opportunities for
the time per cycle becomes critical. improving several traits including
resistance to pests, lodging, shattering,
yield, and, sometimes, malting quality.
4 D.C. Rasmusson

The germplasm is genetically diverse and whose ultimate goal was to enhance
crossing leads to populations with wide grain yield. In the first case, we worked
segregation that lend themselves to with individual traits that we trusted
effective selection for several traits. But, would be yield-promoting. This came
over time, perhaps 50 years, for under the heading “ideotype breeding.”
discussion purposes, traits receiving high The second program was introgressing
priority improve substantially and the genes from elite two-row germplasm to
program germplasm is called elite. At this obtain useful diversity for yield and
time a breeder may observe that lines in malting quality. We call this program
field trials have similar phenotypes and “open-parent cyclic selection.” We used
to a degree similar pedigrees. three breeding cycles. It was an open-
parent strategy in that each cycle may
At this point, the program can be called
employ different parents, i.e., the most
mature and the breeder faces a new
promising lines or varieties at the time.
challenge that was not present in the
younger program. For future progress to
Ideotype breeding
occur, the breeder has two distinct
The concept of an ideal plant or an
challenges. Making gains for traits of
ideotype, as Donald (1968) called it, was
choice, as in a younger program, remains
an effective elaboration on an older idea
a primary goal. However, there is a
accompanied by a new terminology. The
second major goal, that is to preserve
idea was that yield could be increased by
gains that have been made over decades.
modifying individual traits like height,
One can visualize favorable combinations
leaf angle, and head number. The entire
of individual genes scattered across the
package of desired traits would be called
genome acting in an interactive way to
an ideotype or ideal plant. Rasmusson
produce elite phenotypes. The breeder is
(1987) described an ideotype for six-row
obliged to find a strategy that adds new
spring barley and elaborated on the
alleles for desired traits, while protecting
ideotype concept and its strengths and
and preserving accumulated favorable
limitations.
alleles and gene combinations. It is not an
easy task. The Minnesota ideotype breeding effort
began shortly after Donald’s hallmark
A new terminology describes the new
paper appeared in 1968 and occupied a
type of breeding designed to build future
significant proportion of the barley
gains upon the base of the already elite
breeding and genetics effort for more
germplasm. Parent building or
than 30 years. A brief consideration of
germplasm enhancement, terms used
two so-called ideotype traits will
interchangeably, refer to introgressing
illustrate important aspects of the
desired traits/genes into an elite genetic
program.
background. In mature programs, parent
building may utilize more than one-half Semidwarf barley. Jean Lambert, a barley
of the crossing/selection resources in a breeder at Minnesota from 1948 to 1962,
breeding effort. obtained short-stature mutants in the
‘Jotun’ background from Norway and
Faced with the challenge of ensuring
began breeding with them in 1956. The
gains in a mature program, we have
aspiration was to duplicate gains
gained experience with two programs
Learning about Barley Breeding 5

obtained in the Vogel short-stature were not realized because the traditional
wheats. The basic Minnesota germplasm breeding effort raised the bar, so to speak.
was a three-way cross, Jotun / Kindred / The traditional program provided the
2 / Vantage. The cross to Vantage was entire slate of traits needed in a new
critical as it provided the large-diameter variety, while the individual trait
sturdy culms that have been a ideotype effort too often came up short
distinguishing characteristic of the for one or more essential traits.
Minnesota short-stature lines.
Wide-leaf barley. The most interesting
Subsequently, nine cycles (40+ years) of
trait in our ideotype program at present is
recurrent breeding were done with the
a wide-leaf trait. The source of the
aspiration of achieving a yield
germplasm was Harlan’s Freak
breakthrough. It never came and,
Collection. The penultimate leaf width
furthermore, the Jotun germplasm lines
was about 2.5 cm at its widest point. We
appeared to preclude achieving
currently have 11 lines in advanced stages
favorable malting and brewing quality, at
of testing that have moderately wide
least in the Minnesota program and
leaves (much reduced from the source
target environment. Ultimately, only one
line) along with several positive
short-stature variety, Royal, was released
agronomic traits. The pedigree, which
in Minnesota. The most important
indicates the large breeding effort, is
contribution this program made was in
Robust / 63-8069 (source of wide leaf) / 2
sharing germplasm. The Lambert Jotun
/ M47 / 3 / Stander / 4 / Excel / 5 /
short-stature lines provided parental
M81. These lines have a large-diameter
germplasm to most barley breeders in
sturdy culm, large spike, high kernel
North America.
weight, and high grain yield. We
The contribution of the ideotype hypothesize that this attractive phenotype
approach to the barley program in is the result of a gene or genes which are
Minnesota can be viewed in different pleiotropic for several size-related traits.
ways. If ideotype-related research, The ultimate yield level in comparison to
ideotype-related breeding gains, and lines from our traditional program is yet
graduate education are considered, it has to be determined. But we are optimistic
been a worthwhile venture. Thirty-three because of the combination of what
graduate students conducted thesis appear to be yield-promoting traits.
research on so-called ideotype traits
within the barley breeding program, and
22 papers were published. In this
context, it has been a productive and
Open-Parent Cyclic
worthwhile program. Selection
When breeding gains are measured by In response to our concern about lack of
new varieties and enhanced germplasm genetic variation, we initiated a program
(parental germplasm), the conclusion is to introduce genetic diversity for grain
different. The ideotype approach could yield and malting quality. The extreme
not compete with the traditional ongoing narrowness of the Minnesota malting
breeding effort. Time and again barley germplasm was described by
promising gains from individual traits Rasmusson and Phillips (1997). The
6 D.C. Rasmusson

introgression research using open-parent having the highest theoretical percentage


cyclic selection was part of a thesis effort of adapted recurrent parent germplasm,
by Michael Peel (1998). i.e., when six-row gene combinations
were predominantly left intact. Results
The objective of this research was to
from the three breeding cycles support
assess the effects of transferring genes
the proposition that a strategy that
from two-row barley cultivars to the
maintains favorable gene combinations
Minnesota six-row gene pool. Our
while introgressing relatively small
premise was that the gene combinations
amounts of donor germplasm can lead
of the six-row cultivars should be largely
to incremental yield gains.
maintained with a small introgression of
donor germplasm. We conducted three
cycles of recurrent-type breeding
beginning with crosses involving five Final Comments
two-row donor parents. The crossing
• Barley breeding was a good career
scheme led to progenies with theoretical choice. It is always interesting and
portions of two-row germplasm of 25- challenging and frequently rewarding.
50% in cycle 1, 12.5-25% in cycle 2, and • There have been many changes since I
6.25-12.5% in cycle 3. first harvested barley plots with a sickle
and we obtained one generation per
We observed the yield penalty that is
year. As a result, barley programs are
commonly associated with introgression
several-fold more efficient and more
of donor genes into elite germplasm in
productive today.
cycle 1. In this cycle, no putative cultivar • Learning remains a high priority as we
candidates were found. Furthermore, the wrestle with how to utilize the new
two-row progeny were decidedly molecular tools and (for some of us)
inferior in all populations, especially in how to obtain varieties resistant to
the backcross and three-way cross Fusarium head blight to save our
populations. Cycle 2 populations, which malting barley crop.
theoretically contained 12.5-25% of two-
row germplasm, were improved, and
mean grain yield of the populations References
increased to 98% of the check mean. Donald, C.M. 1968. The breeding of crop
ideotypes. Euphytica 17:385-403.
In cycle 3, sets of lines representing three Peel, M.D. 1996. Ph.D. Thesis. University of
populations yielded 112-119% of the Minnesota, St. Paul.
check mean in 1997 and 1998, and Rasmusson, D.C. 1987. An evaluation of ideotype
breeding. Crop Sci. 27:1140-1146.
individual lines surpassed ‘Stander’, the Rasmusson, D.C., and Phillips, R.L. !997. Plant
highest yielding check in both 1997 and breeding progress and genetic diversity from
1998. The highest yielding lines in each de novo variation and elevated epistasis. Crop
Sci. 37:303-310.
cycle were derived from populations
7

Hull-less Barley: An Overview


J.H. HELM1

In the early 1970s, the CIMMYT hull- While they were often disappointed with
less barley program and my own the results from the samples I sent to
program started sharing germplasm and them, I always learned a lot from their
ideas. It was a struggle in the early results.
years, because much of our focus was on
It was from these early results that I
high lysine, which brought with it poor
changed my selection approach to
agronomics, low yield, and
selection for pig digestibility from
environmentally unstable protein
chemical analysis. Dr. Frank Aherne from
quality. While nothing of commercial
the University of Alberta got involved
value came from this research approach,
with our first full feed tests coupled with
it did teach me a lot about protein/
nylon bag tests; it was from these results
environment interactions and protein
that we released the hull-less variety
digestibility in the rat and pig.
Condor in 1988. Condor is still the
By the end of the 1970s and early 1980s, predominant hull-less variety in some
most breeders had quit chasing lysine areas of western Canada, though I am
genetics, and I changed my approach to convinced that other, newer varieties are
looking at environmental stability and better for all characteristics. It was
protein and energy digestibility. Our Condor, however, that began the increase
methods of determining these were not of the hull-less barley acreage in western
very scientific at that time, as we had Canada. The primary reason was that
very little animal data. We pieced feed quality was superior in all
together some rough NIRS calibrations environments to the hulled check
for protein, energy, starch, and lysine varieties.
and began screening large numbers of
While hull-less barley acreage took off
lines. While these screenings were only
with the release of Condor, we must
very rough, we did notice that we were
acknowledge that it was Dr. Brian
able to make improvements from
Rossnagel at the Crop Development
breeding cycle to breeding cycle. I found
Centre, University of Saskatchewan, who
two animal nutritionists who would
paved the way with the release of Scout
work with me, Dr. Richard Beames from
(1982) and Tupper (1984). Without that
the University of British Columbia and
push it is doubtful that I would have put
Dr. B.O. Eggum from the National
Condor forward.
Institute of Animal Science, Denmark.

1 Head, Field Crop Development Centre, Alberta Agriculture, Food & Rural Development, 5030 -
50 Street, Lacombe, AB T4L 1W8, Canada. Phone: (403) 782-4641. Fax: (403) 782-5514. Email:
james.helm@agric.gov.ab.ca
8 J.H. Helm

Since this beginning, the last decade has also has 5% higher protein and energy
seen hull-less barley acreage rise from digestibility in pigs than Condor.
near zero commercial acreage to nearly
one million acres. More than 16 varieties The future
are now in production, with the pig What is the future for hull-less barley? In
industry as the major market. New my opinion, we will continue to make
varieties are agronomically superior, and agronomic gains equal to what is
quality is now stable with the happening in the hulled barley breeding
development of Falcon (1992) and CDC area. The ICARDA/CIMMYT program is
Dawn (1995) as the best of the best rapidly broadening the hull-less barley
(Table 1). germplasm. Our germplasm base, thanks
to the work of Hugo, is now more than 700
The ICARDA/CIMMYT role in this, at
entries from many backgrounds. I think
least from my point of view, has been the
the pig industry will remain the primary
development of a diverse germplasm
market; to this end we will be able to
base that has also introduced new sources
increase digestible energy by 200 Kcal
of disease resistance, early maturity, and
(Figure 1) and protein digestibility by 5%
straw strength into the program. In 1999
(Figure 2).
the predominant hull-less variety grown
was Falcon. Falcon is a six-row There is a significant potential in the food
semidwarf variety selected from one of industry but these are not progressing as
CIMMYT’s F2 populations, about 50 rapidly as many think they should. I think
seeds sent to me in 1977. I was intrigued some of the work Hugo is doing in
by its short stature and large head size; I subsistence agriculture is interesting and
played around with it for a long time this is an area where we will not see the
before I put it into the variety registration food processing industry or large
system in 1989. The important companies put any effort. In my opinion,
characteristics of Falcon are its strong barley makes good tortillas, pancakes,
straw and lodging resistance under high muffins, soups, rice replacer, cookies,
yielding conditions. It also threshes noodles, breads, and of course beer. It has
easily; this characteristic is now the prime potential as a source of food additives for
quality factor in hull-less barley. Falcon fiber, tocols, and other fractions.

It has been an interesting crop to work on


for the last 32 years when I made my first
Table 1. Average feed quality analysis for three hull-less barley cross, to now when we see
hull-less barleys grown at 11 locations in
western Canada, 1993, 1994, and 1995.
an expanding interest.

Characteristics Falcon CDC Dawn Condor Breeding methodology


I would like to complete my presentation
Protein (%) 13.90 12.95 13.85
with a quick description of how we handle
Protein digestibility (%) 79.62 80.28 78.54
Digestible energy (kcal/kg) 3507 3442 3478 the breeding material in our program.
B-glucan 3.8 3.5 4.5 Crosses are made and F1s grown out in
Soluble fiber (%) 4.4 4.0 5.0 growth rooms or in California. The F2s
Pentosans (%) 4.2 4.0 4.1
from our crosses are grown out in a solid-
Dietary fiber (%) 14.9 13.8 15.1
Insoluble fiber (%) 10.5 9.7 10.1 seeded plot of approximately 4000 plants
Hull-less Barley: An Overview 9

kcal/kg the Mexican population. This same


3,900 procedure is used in the F3 to F5. Poor
populations are dropped along the way.
3,700
These large bulk populations are
3,500
inoculated with scald from straw each
year. The rationale is that diseased plants
3,300
will produce smaller, lighter seed. By
3,100 sieving to take out the small seed and
Hulled
using the gravity table to take out the
1990 2000 barley
light seed, we move the populations
Figure 1. Range of digestible energy in hull-less toward better disease resistance and
barley germplasm.
larger kernels and test weights. This
%
selection in the F2, F4, and F5 has shown
95 to be effective for the kernel
characteristics, and we have been able to
90 select our new semi-dwarf varieties with
85 kernel weights 30% higher than their
parents. We are also seeing an increase in
80 the level of field resistance to leaf
diseases.
75
Hulled The F5 populations are selected for plant
70
1990 2000 barley type by selecting a single head from up to
200 plants from the bulk population.
Figure 2. Range of protein digestibility in hull-
less barley germplasm. These are grown in head/row plots and
inoculated for scald. Rows with poor
agronomic type or moderate to heavy
in 15.25 square meters. The F2 from disease are discarded prior to harvest.
Mexico is grown in a 3.5-meter row The harvested rows are weighed and
(average 65 seeds). Good populations scored for seed characteristics and
are harvested in bulk. Seed from the analyzed in the research laboratory for
large plots are screened for plump seed quality. We use NIRS as a method to
(6/64 in slotted screen) then run over a screen for all quality characteristics.
gravity table. The top 10% of plump, Those rows that meet all agronomic,
heavy seed is advanced to an F3 disease, and quality standards are
population grown in Southern entered into yield trials. We have two
California. We select for plant type based years of preliminary yield testing and
on pedigree in California, selecting three years of advanced yield testing.
approximately 500 heads out of a Then the best lines are entered into a
population of 4000 F3 plants. The seeds cooperative testing system for two years
from these are bulked and 4000 random before receiving registration as a variety
seeds planted in Alberta in a 15.25 and released to seed growers.
square meter plot as are the F3 bulk from
10

Contributions of Breeding
and Crop Management to
Increasing Barley Yields and
Grain Quality in Chile
E. BERATTO M.1

The first barley trials in Chile were “Genetic Improvement and Production of
conducted by the National Agriculture Malting Barleys” was signed between
Society during the first quarter of the 20th INIA and the United Brewers Company
century and later by the Ministry of (Compañía Cervecerías Unidas, CCU); the
Agriculture’s Department of Genetics agreement is still operative 23 years later.
and Plant Improvement (1940-1947) and
the School of Agriculture (1955-1964) of
the Univeridad de Chile. The trials
focused on sowing date, seed rate,
Initial Status
adaptability, and yield of varieties
Released varieties
introduced mainly from Europe.
In 1978, when the agreement was signed,
Research efforts in Chile up to 1976 were CCU was sowing the varieties Breun’s
typically isolated trials conducted by Wisa, Carina, and Firlsbeck Union. Breun’s
public and private organizations; they Wisa was sown until 1979; Carina was
were geographically fragmented, withdrawn from the market in 1983, and
discontinuous, had virtually no Firlsbeck Union in 1984. Commercial
integrated research teams, and did not cropping of the latter two varieties
persevere in pursuing established stopped mainly because of their
objectives. These efforts produced susceptibility to yellow rust of barley,
irrelevant results that had little impact on which greatly reduced yields (Caglevic
domestic barley production. and Herrera, 1984), grain size, test weight,
and grain weight in both the northern and
The National Agricultural and Livestock central parts of Chile (Caglevic and
Research Institute (Instituto Nacional de Herrera, 1982; 1983).
Investigaciones Agropecuarias, INIA),
through the Carillanca Regional Research Barley yields
Center, officially incorporated barley into The national average barley yield in 1978
its research agenda in 1976. Two years was 1.9 t/ha, but the average yields
later, in 1978, a research agreement on produced by commercial farmers and seed

1 Instituto de Investigaciones Agropecuarias, Centro Regional de Investigación Carillanca, Casilla


58-D, Temuco, Chile. Tel.: 56-45-215706; Fax: 56-45-216112; Email: eberatto@carillanca.inia.cl.
Contributions of Breeding and Crop Management to Increasing Barley Yields and Grain Quality in Chile 11

producers under contract to CCU were 1.7 was demonstrated by their impact on
and 2.2 t/ha, respectively. yield (Figures 1, 2, and 3) and grain size,
and by how often they were a part of the
Diseases sowing agreements established by the
When the agreement was implemented, CCU (Table 1).
not much was known about barley
diseases. Damage to yield, grain size, and t/ha
test weight had not been evaluated, and
no chemical control methods had been
developed.

Grain size
Barley producers under contract to CCU
received premium prices because of good
grain size, i.e., when at least 80% of the
grain was retained on 2.5-mm sieves; this
cost the company 5%.
35
38
41
44
47

19 0
19 3
19 6
19 9
19 2
65
19 8
19 1
74
19 7
19 0
19 3
19 6
19 9
19 2
95
5
5
5
5
6

6
7

7
8
8
8
8
9
19
19
19
19
19
19

19

19
Figure 1. Progress in barley yields, 1935-97.
Crop management practices
Crop management practices (sowing date, t/ha
seeding rate, fertilization, and weed
control) used on barley at that time were
similar to those recommended for wheat.
This was based, with no scientific
evidence, on the principle that
“everything that was good for wheat was
good for barley.”

Advances in Barley
19 39
19 44
19 -49
19 -54
19 -59
19 64
19 -69
19 -74
19 79
19 -84
19 89
19 94
-97
-
-

-
-
35
40
45
50
55
60
65
70
75
80
85
90
95
19

Improvement Figure 2. Progress in barley yields by five-year


periods, 1935-97.
New varieties
Research activities conducted by INIA t/ha
under the agreement from 1978 to 1999
led to the release of five new barley
varieties with good malting quality.
Aramir, property of CEBECO, was
introduced from Holland; Leo INIA-CCU Acuario
originated from advanced lines in the Leo
Libra
ICARDA/CIMMYT nurseries. The other Granifen
Aramir
three, Granifen INIA-CCU, Libra INIA- B.Wisa Carina
F. Union
CCU, and Acuario INIA-CCU, were
developed by INIA’s Barley Improvement
79

81

83

85

87

89

91

93

95

97
19

19

19

19

19

19

19

19

19

19

Project. The significance of these varieties Figure 3. Progress in barley yields, 1978-97.
12 E. Beratto M.

Table 1. Distribution of barley varieties sown by United Brewers Company (Compañía Cervecerías
Unidas, CCU), 1977-99.

Area (ha)
Percentage of seed used by CCU† Under Total Percentage of
contract malting total area under
Year BW FU CA AR GA LI LE AC to CCU barley contract

1977-78 40 60 16,541 54,238 30.5


1978-79 16 84 11,163 50,804 22.0
1979-80 6 94 9,636 41,327 23.3
1980-81 83 17 13,447 39,066 34.4
1981-82 60 40 15,847 48,858 32.4
1982-83 90 10 10,984 32,436 33.9
1983-84 32 49 19 10,137 28,177 36.0
1984-85 7 76 17 12,195 29,784 49.9
1985-86 62 38 10,101 19,312 52.3
1986-87 64 36 7,889 13,914 56.7
1987-88 65 35 12,810 20,459 62.6
1988-89 51 49 14,854 20,901 71.1
1989-90 38 36 26 16,753 22,336 75.0
1990-91 38 62 12,173 26,953 45.2
1991-92 38 61 1 11,085 24,131 45.9
1992-93 35 57 8 11,080 18,946 58.5
1993-94 39 12 49 15,543 23,953 64.9
1994-95 68 32 11,083 21,399 51.8
1995-96 30 70 10,264 19,695 52.1
1996-97 30 70 11,374 18,634 61.0
1997-98 30 70 8,955 22,637 39.6
1998-99 15 85 8,028 22,527 35.6

† BW: Breun´s Wisa; FU: Firlsbeck Union; CA: Carina; AR: Aramir; GA: Granifen INIA-CCU; LI: Libra INIA-CCU; LE: Leo
INIA-CCU; AC: Acuario INIA-CCU.

Barley varieties sown by the Granifen, 38%, and Firlsbeck Union was
CCU in 1977-99 discontinued. In 1989-90, 38% was sown
When the agreement was implemented to Aramir, 36% to Granifen, and 26% to
(1977-78), the varieties Breun’s Wisa and Libra INIA-CCU. Aramir was
Firlsbeck Union covered 40 and 60%, withdrawn from the market in 1990-91
respectively, of the area sown to barley and was replaced by Libra, which
by the CCU. Two years later (1980-81), occupied 62% of the barley area; the
Bruen’s Wisa was discontinued, and 83% remaining 38% was sown to Granifen. In
of the cultivated area was sown to 1991-92, Granifen and Libra still covered
Firlsbeck Union and 17% to Carina. In 38 and 61%, respectively, and Leo was
1983-84, Carina was withdrawn from the sown on 1%. In 1995-99, Granifen and
market, and Firlsbeck Union, Aramir, Leo were no longer sown, and 70-85%
and Granifen INIA-CCU covered 32, 49, was covered by Acuario INIA-CCU; the
and 19%, respectively, of the barley area. remaining 15-30% was sown to Libra
In 1985-86, the area sown to barley was (Table 1).
distributed as follows: Aramir, 62%,
Contributions of Breeding and Crop Management to Increasing Barley Yields and Grain Quality in Chile 13

The length of time varieties introduced or The advance in annual average barley
developed by INIA were sown is as yields from 1935 to 1997 is shown in
follows: Aramir, 7 years (1983-89); Figure 1.
Granifen INIA-CCU, 12 years (1983-94);
Leo INIA-CCU, 4 years (1991-94); Libra Gains in barley area and
INIA-CCU, 10 years. Acuario INIA-CCU production
began to be sown 4 years ago; by 1999, Over a period of 63 years, barley area
100% of the barley area was sown to it. and production in Chile decreased by
67.8% and 20.8%, respectively (Table 3).
Progress in barley yields The reasons for these reductions are as
National average barley yields in Chile follows: about 80% of the total barley
increased from 1.5 t/ha (in 1935-39) to 3.6 area in Chile is sown to malting barley
t/ha (in 1995-97) within a 63-year period. under contract to domestic malting
This is equal to a yield gain of 149.3%, or companies and breweries. The former
an annual increase of 34.6 kg/ha (Table have significantly reduced the area
2). Upon comparing yields during the
first period (characterized by an almost
complete lack of systematic barley Table 3. Progress in national average barley
research) (from 1935-39 to 1970-74), one yields by five-year periods, 1935-97.
comes to the conclusion that in 40 years Five-year Area Production Yield
yields increased only 31.8%, or 11.5 kg/ period (ha) (000 t/ha) (t/ha)
ha annually. In contrast, between the
second (1975-79) (when barley research 1935 – 1939 74,320 1,097 1.5
1940 – 1944 48,910 752 1.5
under the CCU agreement started) and 1945 – 1949 49,024 798 1.6
last (1995-97) periods, yield rose by 1950 – 1954 53,578 797 1.5
85.7%, or 73.1 kg/ha per year, over a 23- 1955 – 1959 53,056 914 1.7
year period (Table 2). During that same 1960 – 1964 41,329 744 1.8
1965 – 1969 50,540 1,081 2.1
period, average commercial yields
1970 – 1974 65,904 1,260 1.9
obtained by the CCU increased from 1.7 1975 – 1979 58,360 1,149 2.0
t/ha in 1978 to 4.3 t/ha in 1997, a yield 1980 – 1984 41,958 882 2.1
gain of 160.8%. Average yields of CCU 1985 – 1989 22,814 749 3.3
1990 – 1994 27,149 831 3.5
seed producers rose from 2.2 t/ha in 1978
1995 – 1997 23,909 869 3.6
to 5.7 t/ha in 1997, an increase of 132.4%.

Table 2. Progress in national average barley yields


comparing different five-year periods, 1935-97.

Yield (t/ha) Yield Annual


1935- 1970- 1975- 1995- increase increase
Period 1939 1974 1979 1997 (%) (kg/ha)

Total period 1.5 x x 3.6 149.3 34.6


(1935-1997)
First period 1.5 1.9 x x 31.8 11.5
(1935-1974)
Second period x x 2.0 3.6 85.7 73.1
(1975-1997)
14 E. Beratto M.

under contract due to the drastic development of better agronomic


reduction in the amount of Chilean malt practices for managing the barley crop.
exported to countries such as Bolivia,
Brazil, Venezuela, and Peru. This Disease management
reduction in exports is the result of Research on disease control measures
strong competition from the European began in 1978, when disease
Common Market, which sells malt to identification surveys in Chile’s barley-
those countries at a lower price. Also, producing regions were conducted by
local brewers such as the CCU have Gilchrist (1979), Caglevic and Herrera
reduced the area they sow to barley as a (1984), and Andrade (1986). Later studies
result of the signing and implementation evaluated the impact of diseases on
of the Free Trade Agreement between barley yields and grain size, and
Chile and Canada, which allows the identified the most economically
tariff-free importation of Canadian barley important diseases nationally (scald, net
grain and malt into Chile. Thus it is blotch, yellow and leaf rusts, and
actually cheaper to import barley grain BYDV). They also determined the most
and malt from Canada than to produce it effective chemical treatments
in Chile. (fungicides) for controlling scald
(Rhynchosporium secalis) in particular,
Improving grain size since it reduces yield by more than 40%
The basis (in place since 1978) for paying in early sown barley in years with a
a premium for grain size was changed humid spring (Andrade, 1989). Yellow
for the first time in 1984, from 80% of the rust (Puccinia striiformis f.sp. hordei)
grain remaining on a 2.5-mm sieve to reduced yield by 48-56% in 1980-82, and
85%, as a result of genetic improvement caused grain size, test weight, and kernel
research. In 1996, it was modified for the weight to drop dramatically in the
second time to 90%, bringing Chilean northern and central parts of Chile
requirements to the same level as those (Caglevic and Herrera, 1984). At the
stipulated in the European Brewery same time, the breeding program
Convention. identified genetic resistance sources,
especially to scald. Most of these sources
Improving beer production are from the ICARDA/CIMMYT
efficiency nurseries (Beratto, 1983; CIMMYT, 1983).
In 1978, 18.5 kg of barley were needed to
produce 100 L of beer; today 16-17 kg are Fertilization
necessary to produce 100 L of beer The importance of applying nitrogen
(Devilat, 1988). and phosphorus for raising barley yields
and improving grain quality, especially
in volcanic soils, whether Andisols
Advances in Crop (volcanic soils) or Ultisols (red clay
soils), in southern Chile has been amply
Management demonstrated in studies conducted by
Gains in barley yields both in Chile and Peyrelongue (1983a, b; 1990; 1992) and
at the global level have been the result of Peyrelongue et al. (1984) (Figure 4).
improved barley varieties and the
Contributions of Breeding and Crop Management to Increasing Barley Yields and Grain Quality in Chile 15

a
Fertilization and weed and a

Number of grains/m2
a
disease control ab
In a study conducted at the Carillanca
Regional Research Center, Thomas (1997)
concluded that fertilizer application and c b
c
disease control were essential for d
producing high yields and improving
barley grain quality. He also found that V E E F
(1) E+CM )V+C V+CM F+C F+CM 2)V+ E+CM
of these techniques, combined nitrogen/ + C (4 (3) )V+ V+ (
F+
C
(7)
V (5 (6) V+
phosphorus fertilization has the greatest Treatment (8)
impact on yield (Figure 4), grain weight Figure 6. Effect of eight treatments on the
(Figure 5), number of grains per m2 number of grain per meter2 of Acuario INIA/CCU.
(Figure 6), test weight (Figure 7), grain a
ab ab
abc
Test weight (kg/hl)
size >2.8 mm (Figure 8), and grain size c c c bc
>2.8 + 2.5 mm (Figure 9).

a
+C
E CM CM )V+F V+CE +CM +CM (1)
V
b )V+F V+F+ +CE+ (2 (4) +CE (3)V
(5 )
(6 )V+F V
b
8 Treatment (7)
ce (
Yield (t/ha)

Figure 7. Effect of eight treatments on test


weight of Acuario INIA/CCU.
d a a
d d a
Grain size (>2.8 mm)

e ab

bc
c c c
V E E F
(1) E+CM )V+C V+CM F+C F+CM 2)V+ E+CM
+C (4 (3) (5)V+ 6)V+ ( C
)V ( F+
(7 V+
Treatment (8)

Figure 4. Effect of eight treatments on grain


(1)
V CM CE CM CE +F CM CM
yield of Acuario INIA/CCU. V+ (4)V+ CE+ V+F+ (2)V CE+ +F+
(3) )V+ (5) F+ 6)V
(7 )V+ (
Treatment (8
Figure 8. Effect of eight treatments on grain
size of Acuario INIA/CCU.
a
Grain weight (mg)

a a a a
a a
ab
Grain size (>2.8+2.5)

bc
c c c b b
b b

V E F E
(1) V+CM )V+C E+CM 2)V+ F+C F+CM E+CM V E E F
(3) (4 +C ( )V+ 6)V+ F+C (1) )V+C V+CM E+CM F+C 2)V+ E+CM F+CM
)V (5 (
(7 (
(8)
V+ (4 (3) )V+C (5)V+ C
F+ (6)V
+
Treatment (7 V+
Treatment (8)
Figure 5. Effect of eight treatments on grain Figure 9. Effect of eight treatments on grain
weight of Acuario INIA/CCU. size of Acuario INIA/CCU.
16 E. Beratto M.

Economic Quantification met with good farmer acceptance. The


commercial release of facultative hull-less
of Impacts Achieved
varieties such as Alteza-INIA (Beratto and
under the Agreement Rivas, 1999), with high yield potential,
• The amount of barley grain needed for resistance to economically important
producing 100 L of beer diminished diseases, good agronomic traits, and very
from 18.5 kg in 1984 to 17.0 kg in 1988. good grain and nutritional quality has
This has been estimated to generate a paved the way for barley production in
cost savings of US$ 420,000 for the CCU southern Chile, where it could displace
(Devilat, 1988). Today only 16.0 kg are maize, given that maize cropping in that
needed to produce 100 L of beer, but
region is very risky due to climatic
this has not yet been economically
conditions. Based on these two new
quantified.
alternatives, we predict that the area
• Changes in the premium paid for bigger
sown to barley for feeding livestock (both
grain size in 1978 and 1988 have
translated into a higher yearly income dairy and meat) will continue to rise as it
for the CCU of US$ 105,000 (Devilat, has in the past two years (1998 and 1999).
1988). In 1996, the basis for paying a The development of facultative hull-less
premium price was changed to 90% of and covered barley varieties means that
the grain retained on a 2.5-mm sieve,
farmers now have very promising genetic
but this has not yet been economically
materials that could be sown
quantified.
commercially in regions with high rainfall
• Campos, Ortiz, and Beratto (1989)
estimated that the internal rate of return
in autumn and winter, and severe
(IRR) for the 1978-87 period of the drought stress in spring and summer.
INIA/CCU Agreement was 48-50%. In Sowing spring habit barleys is not
a recent and as yet unpublished study, recommended in these regions (Beratto,
Campos and Beratto (2000) found that 1990). This type of barley, besides having
the IRR for the 1978-99 period of the high yield potential, gives farmers a
INIA/CCU Agreement was 52%, which practical advantage: it has a longer
produced a social benefit valued at sowing period (Salinas, 1996). As a result,
5,350 million Chilean pesos (US$1 = barley cultivation can extend to regions,
516-520 Chilean pesos). The distribution or subregions, where until recently it
of these benefits was 65% for producers could not be sown due to the prevailing
and 53% for consumers. climatic conditions.

Future prospects for improving yield and


Future Prospects malting and nutritional quality of both
spring and facultative barley continue to
About 80% of Chile’s barley area is
be extremely good. There is still a
currently dedicated to the production of
considerable gap between the national
grain for malt and beer, 15% to feed
average yield and the average yields of
grain, and 5% to seed. In the last four
innovative farmers, and between the
years, the area sown to barley for silage
average yield achieved in research centers
production has increased, especially in
with the leading commercial variety and
the cattle-raising region in southern
maximum yield obtained with the best
Chile (Goic and Ponce, 1999); this has
advanced line (Beratto, 1999).
Contributions of Breeding and Crop Management to Increasing Barley Yields and Grain Quality in Chile 17

References Devilat B., J. 1988. Avances y logros del Convenio


de Mejoramiento Genético de Cebadas con
Andrade V., O. 1986. Enfermedades que afectan a Calidad Maltera entre Instituto de
la cebada en la IX Región. Investigación y Investigaciones Agropecuarias y Compañía
Progreso Agropecuario Carillanca (Chile) Cervecerías Unidas. Informe Interno de
5(2):2-4. Compañía del Departamento Agrícola de
Andrade V., O. 1989. Rincosporiosis o escaldadura Compañía Cervecerías Unidas. 11 p.
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Caglevic D., M., and Herrera M., G. 1984. La roya de Investigaciones Agropecuarias, Centro
amarilla de la cebada en Chile (Puccinia Regional de Investigación La Platina
striiformis f.sp. hordei). Estación Experimental La (Santiago, Chile). Trabajo presentado en las
Platina (Santiago, Chile). Trabajo publicado en XXXV Jornadas Agronómicas.
las XXXV Jornadas Agronómicas. Salinas G., A. 1996. Influencia de la época de
Campos M., A; Ortiz R., C., and Beratto M., E. 1989. siembra sobre el rendimiento y calidad física
Análisis del impacto económico del Convenio del grano de tres líneas avanzadas invernales
INIA-CCU para mejoramiento genético de y alternativas. Universidad de la Frontera,
cebada cervecera. Boletín N°6, Programa de Facultad de Ciencias Agropecuarias y
Economía, Instituto de Investigaciones Forestales (Temuco, Chile). Tesis para optar al
Agropecuarias (Santiago, Chile). 55 p. título de Ingeniero Agrónomo. 49 p.
Campos M., A., and Beratto M., E. 2000. Análisis Thomas A., E. 1997. Influencia de la fertilización,
del impacto económico del Convenio INIA- control de malezas y enfermedades sobre el
CCU en Mejoramiento Genético de Cebada rendimiento y calidad de grano de una cebada
Cervecera. Trabajo en etapa de revisión por el (Hordeum vulgare L.) de primavera con calidad
Comité Editor. maltera. Universidad de la Frontera, Facultad
CIMMYT. 1983. Materials resistant to scald in de Ciencias Agropecuarias y Forestales
Mexico and Chile and to stripe rust in the (Temuco, Chile). Tesis para optar al título de
Andean countries in 1983. In: CIMMYT Report Ingeniero Agrónomo. 79 p.
on Wheat Improvement 1983. 73 p.
18

Importance of Identifying
Stripe Rust Resistance in
Barley
W.M. BROWN, J.P. HILL, AND V.R. VELASCO1

Stripe rust (Puccinia striiformis f. sp. stripe rust fungus could pose a major
hordei) of barley is a new disease of threat to the industry in those areas.
barley in the American hemisphere. It While there are no commercial resistant
was recognized as a problem in Mexico malting varieties readily available at this
and the United States only in the last 10 time, sources of resistance are becoming
years (Brown, 1992; Brown et al., 1993a; available.
Calhoun et al., 1988; Line and Chen,
In 1990, Dr. Darrell Wesenberg, Director
1999). In South America, race 24 of the
of the USDA/ARS Small Grain
fungus was recorded near Bogota,
Germplasm Collection (NSGC) Research
Colombia, in 1975. Within five years,
Laboratory at Aberdeen, ID, contacted
barley stripe rust was found throughout
the Colorado State University barley
western South America causing losses of
disease research team to initiate stripe
30-70% (Dubin and Stubbs, 1984; 1986).
rust field trials with cooperators in South
The fungus in recent years has spread America. The authors and their Bolivian
north to Mexico (Calhoun et al., 1988), associates were familiar with the fungus
and in 1991, Marshall and Sutton (1995) and had initiated some of the first
reported it in Uvalde, Texas. It was studies of the disease in the late 1970s in
confirmed in Arizona,1993; Bolivia (Velasco and Brown, 1982;
California,1994; Colorado,1992; Velasco et al., 1993).
Idaho,1993; Montana,1993; New Mexico,
Field work was established in
1992; Oklahoma, 1992; Oregon, 1995;
Cochabamba, Bolivia, South America, in
Washington, 1995; and Utah, 1994
1991 (Brown, 1992). The Cochabamba
(Brown et al., 1996; Brown et al., 1993a;
area is a high Andean valley
Chen et al., 1995; Line and Chen, 1996;
approximately 7,500 ft above sea level.
1999). Barley stripe rust is also present in
This valley shows a high endemic
western Canada. Since 1995 it has
occurrence of barley stripe rust race 24
become established and on occasion
(BSR-24) and heavy infections are a
causes considerable damage and concern
yearly occurrence. Field germplasm and
in the northwest states of the USA. The

1 Department of Bioagricultural Sciences and Pest Management, Colorado State University, Fort
Collins, CO 80523-1177, USA.
Importance of Identifying Stripe Rust Resistance in Barley 19

associated studies were initiated in the Colorado State University, Idaho State
1991 growing season (January-May) and University, and Oregon State University.
continued through 1996. USDA/ARS
NSGC staff prepared germplasm for Race variation studies
planting. CSU and Bolivian staff carried Differential nurseries were developed in
out the field trials. cooperation with Dr. Mareike Johnston at
Montana State University. The differential
Germplasm screening trials nurseries were planted in Bolivia,
Trials were composed of all entries from Colorado, Ecuador, and Germany (Brown
the National Small Grain Collection and et al., 1996; Hill et al., 1995). Considerable
selections developed by cooperators variation was observed both between
from: field nurseries and also under controlled
conditions in greenhouse trials, where
USDA/ARS (Aberdeen, ID) University of California-
Davis comparisons were made by Dr. Johnston
North Dakota State University Montana State University (Tables 1 and 2).
Oregon State University Utah State University
Washington State University Adolf Coors
Busch Agricultural Resources Western Plant Breeders

Table 1. Variation of BSR-24 susceptibility in


Selections showing some level of Bolivia, Ecuador, and Colorado, 1994 (field test).†
resistance were field tested a second year Differentials Bolivia Ecuador Colorado
in Bolivia and also tested in Colorado, Emir S MS MS
Ecuador, Germany, and eventually Mazurca MS MS S
Mexico at different times over the period Zephyr S S S
from 1993 to 2000. In 1995, Ecuador was I5 MS S S
BBA 2890 MS MS MS
deleted in favor of adding the Toluca
Trumpf MS 0 S
Valley of Central Mexico. Field testing is Hor 4020 R MS MS
now carried out at Davis, California, by Bigo MS R MS
University of California staff, at the Abed Binder 12 S S MS
Cambrinus S S S
Mexico site by the authors in cooperation
Luttichauer Landgerste MS S MS
with Dr. Hugo Vivar, and in Idaho by Heils Franken MS MR S
USDA/ARS staff. S 3170 (Hor 3209) S MR S
S 3192 (Hor 2926) MS R S
During the period from 1991-1999, over Grannelose Zweizeilige R 0 S
40,000 selections of barley were field Weisse Von Fong Tien S S S
evaluated for stripe rust resistance. Varunda MS R S
Stauffers Obersulzer S MS S
Numerous sources of resistance were Hor 1428 MS S MS
identified and are being incorporated Morex S S S
into commercial lines by various barley Steptoe S S S
breeding programs throughout the Larker S S S
Abyssinian 14 MS 0 MS
western U.S. All results are available on
Topper S MS S
the USDA/ARS GRIN data base system. BBA 809 MR R MS
In 1999 a resistant variety, Bancroft (P.I. Hokkaido Chevalier MS R S
605474), was developed and released Hiproly MS S S
cooperatively by the USDA/ARS, † R = resistant, MR = moderately resistant,
MS = moderately susceptible, S = susceptible.
20 W.M. Brown, J.P. Hill, and V.R. Velasco

Table 2. Selected stripe rust differential Dr. Johnston also tested isolates from
reactions in greenhouse inoculations.† Colorado, Montana, and Utah against
GER GER WPD the differentials under controlled
Differentials 24 23 93 PF94 Utah Colo conditions. When these were compared
to known races 24 and 23 in tests carried
Mazurka R S R R R out by Dr. Walters in Germany,
HOR 4020 R R,S I R S
considerable variation was again seen
Cambrinus S R S S R R
Heils Franken S R R I R I (Tables 3 and 4).
HOR 1428 R S R R R R

† R = resistant, MR = moderately resistant,


MS = moderately susceptible, S = susceptible.

Table 3. Stripe rust barley differential reactions Table 4. Effect of fungicide seed treatment on
in Montana, Germany, and Colorado, 1994 stripe rust race 24, at different planting times,
(greenhouse test).† 1994.†
Montana Germany Germany Colorado Treatment January† April
Differentials R - 24 R-24 R-23 R-24 (?)
Emir R R-S R R CBG‡3.00 45 a 23 a
Mazurca R R S R CBG 2.00 45 a 22 a
Zephyr S S I S CBG 1.50 43 a 22 a
I5 R R R R CBG 1.00 43 a 23 a
BBA 2890 R R - R CBG 0.75 43 a 23 a
Trumpf R R R R Captan/Baytan 40 b 19 b
HOR 4020 R,S R - S Thiram/Raxil 39 b 14 c
Bigo R R R R Vitavax extra 39 b 13 c
Abed Binder 12 S MS - S Vitavax 200 34 c 13 c
Cambrinus S S R R Untreated 34 c 13 c
Luttichauer S S - S
Landgers † Days after emergence when first pustules observed.
Heils Franken R S R S ‡ Captan/Baytan/Gaucho.
S 3170 (Hor 3209) R R R R
S 3192 (Hor 2926) R R R R
Grannenlose R R - R
Zweizeil
Weisse S S S S Summary of germplasm
Von Fong Tie screening
Varunda R R R R
Stauffers R R - R
Germplasm evaluation and screening in
Obersulzer Bolivia and elsewhere continue to
Hor 1428 R R S R identify a wide range of barley stripe
Morex S S S S rust resistance sources. Of note is that
Steptoe S MS - S
our work and that of our cooperators
Larker S I - S
Abyssinian 14 R - - MS show considerable variation in what has
Topper S S - S been called “race 24.”
BBA 809 R - - R
Hokkaido R - - S Variation in race 24 is also supported by
Chevalier observations reported in 1994 by
Hiproly R R - R Marshall and Sutton (1995) and Line
† R = resistant, MS = moderately susceptible, (Chen et al., 1995; Line and Chen, 1996;
I = intermediate, S = susceptible.
Importance of Identifying Stripe Rust Resistance in Barley 21

1999). Thus it is clear that barley stripe Integrated trials using seed treatments
rust in North America is a very and foliar spray combinations were
heterogenous population. carried out in Bolivia from January
through April, 1994 and 1995. These
trials demonstrated that foliar
applications of Tilt, Folicur, or Bayleton
Fungicide Trials for applied at first sign of stripe rust were
Control of Barley Stripe effective against stripe rust (Brown et al.,
Rust Race 24 1996; Velasco et al., 1993). Other trials
have demonstrated effective control with
Early results in 19792 in Bolivia
additional fungicides (Calhoun et al.,
indicating that seed treatments might be
1988; Line 1998, 1999; Navarro and
effective led to reevaluating this
Zamora, 1990).
approach (Velasco et al., 1995). Seed
treatment field trials were hand planted One trial of interest in Bolivia was
twice in Bolivia (January and April) in planted with a local barley landrace and
1998 (Table 4). Seed was commercially had no seed treatment other than noted
treated by Gustafson Corporation. The (Table 5). A companion trial was planted
first planting in January was at the with the variety Russell. The seed for
normal time for the area, a period prior this trial had been supplied by the ARS/
to the development of high stripe rust USDA Small Grains Laboratory in
occurrence. The second planting was Aberdeen, ID, and unknown to the
much later, in April, when the fungus authors had been treated with Vitavax.
was very active, and under conditions of All plots were hand-planted and sprays
extremely high stripe rust inoculum applied with a manual backpack sprayer
pressure. at 33 psi at label rates. No adjuvants
were used.
In all cases Baytan (triadimenol) gave
significantly enhanced stripe rust
protection after emergence (Table 5).
Table 5. Barley stripe rust race 24 fungicide trial,
cv. Criolla, 1994.
Combination seed treatment and
foliar fungicide trials No. of Disease
Fungicide trials were initiated in 1978, Treat- applica- Rate severity (%) Yield
ment tions (kg/ha) at milk stage (kg/ha)
when the disease was first found in
Bolivia. Unfortunately at least two B/T 2 † 0.55 a 3,537.66 a
fungicide applications of Bayleton are B/T 1 † 7.35 a 3,177.50 ab
required to keep the disease below a Baytan ‡ 12.72 a 3,140.00 ab
Tilt 2 .6 13.22 a 2,532.00 ab
potentially damaging level (Vealsco and
Tilt 1 .6 66.77 b 2,499.16 b
Brown, 1982; Velasco et al., 1980). It is Control - - 91.05 c 1,094.16 c
unlikely that under U.S. conditions two
field applications of any fungicide † Baytan/Tilt at 0.6kg/ha and 200g/100kg seed.
would be economically feasible for ‡ As seed treatment 200g/100kg seed.
barley stripe rust control.

2 Velasco and Brown, unpublished field observations, Cochabamba, Bolivia.


22 W.M. Brown, J.P. Hill, and V.R. Velasco

All combinations of Baytan and foliar The importance of stripe rust resistance
fungicides enhanced disease depends on all or any of several
suppression and resultant yield. Of characteristics:
considerable interest, the second trial
• location and environment
(Table 6) with a combination of Vitavax
• other pest and diseases present
and Baytan gave better protection than
• total context of the cropping system
two foliar sprays without seed • economics
treatment. When compared to the
efficacy of Baytan alone in the previous Each of the above impacts the potential
trial, it indicated a possible synergistic usefulness of developing stripe rust
effect between Vitavax and Baytan. resistance in barley. Barley stripe rust is
favored by humid and cooler climates.
These do not necessarily prevail in many
Table 6. Barley stripe rust race 24 fungicide trial, of the locations where malting and feed
cv. Russell, 1994. barleys are produced.

No. of Disease Other pests and diseases impacting the


Treat- applica- Rate severity (%) Yield crop, such as Russian wheat aphid and
ment tions (kg/ha) at milk stage (kg/ha)
scab, may be more important. Putting a
Baytan † 0.82a 3,352.66a high priority on stripe rust resistance
Tilt 2 0.600 7.90ab 3,075.83ab may not be appropriate or the best use of
Bayleton 2 1.000 8.20ab 2,715.83 bc limited resources.
Folicur 2 0.750 25.45 bc 2,553.33 bcd
Manzate 2 2.000 27.47 c 2,240.83 cd Looking at the whole cropping system is
Control - 73.77 d 2,002.50 d also important. In some areas such as
† Seed treatment 200 g/100 kg seed. All seed pretreated
Colorado, barley stripe rust is only a
with Vitavax. problem if barley is planted late and
even then only in some years. Use of
early planting provides a period when
the fungus is not present and allows
Discussion and major development of the crop in the
Conclusions absence of inoculum.

Considerable progress has been made in The economics of fungicide use is also
the identification of sources of resistance important. Effective fungicides are
to barley stripe rust race 24 (Brown et al., available for stripe rust, but in less
1996; Brown et al., 1993b; Hill et al., 1995; affluent societies, are beyond the
Velasco et al., 1991). The observed economic resources of the growers. Even
variability and the conclusion reached in more affluent agriculture, the cost
by ourselves and others that the race 24 benefit ratio of the new generation
designation is not just one race, but strobulin-based fungicides may not
potentially many, brings in to question justify fungicide use.
the value of searching for gene specific
Additionally, other pest or disease
“vertical resistance.”
problems may not be as amenable to
pesticide treatment. Therefore,
development and deployment of
Importance of Identifying Stripe Rust Resistance in Barley 23

resistance to those problems are more Therefore a stripe rust program is


important in the total management recommended that uses the following
context. integrated management tactics:

Use of a fungicide to manage stripe rust • Use a slow-rusting (TMS-5MS) line.


will not be necessary on a yearly basis • Treat seed with an appropriate
due to the sporadic development of the fungicide.
disease in many areas. In those years • Plant early.
when the fungus does develop early • Scout and apply appropriate fungicide
enough to become a threat, timely if 5% BSR prior to boot.
fungicide application may be economic
and outweigh the expense and priority
of developing resistance to stripe rust. Acknowledgments
The authors would like to acknowledge
Associated knowledge gained from
and thank our cooperators: Dr. Ursula
barley screening trials over 10 years,
Walter (Germany), Ing. Juan Cardova
from field observations of the
(Bolivia), Ing. Miguel Rivadeneira
epidemiology and work with both seed
(Ecuador), and Dr. Hugo Vivar (Mexico).
and foliar fungicide trials, suggests a
We also thank Dr. Mareike Johnston,
long-range integrated approach is best.
Montana State University, for
Such an approach would emphasize the
developing and providing the
use of trace to moderately susceptible
differential nursery lines for field
barley lines and other management
planting and race comparison studies,
techniques. Even in using such an
and Dr. Darrell Wesenberg and Dr.
integrated approach it is still critical to
Harrold Bockelman at the USDA/ARS
manage host plant resistance (Brown
Small Grains Laboratory, Aberdeen, ID,
1976; Brown 1993) by the use of:
for providing and preparing barley lines
• gene rotation, where different sources used in this program. Also we would
of resistance are moved in and out of like to thank Dr. Rollie Line, USDA/ARS
the host plant population, at Washington State University for
• gene diversity, where a range of identifying multiple races of barely
resistance genes are present in different stripe rust from Colorado and other U.S.
cultivars within an area at any one collections. This project is supported by
time, the USDA/ARS Small Grains
• gene rotation, where continued and
Laboratory, Aberdeen, ID, and the
careful monitoring of fields are
American Malting Barley Association
routinely carried out and initial
(AMBA).
outbreaks are detected early and spot
treated with appropriate fungicides
early, before the disease can spread.
24 W.M. Brown, J.P. Hill, and V.R. Velasco

References Hill, J.P., Johnston, M.R., Velasco, V.R., and Brown,


W.M. 1995. Response of selected barley lines to
Brown, W.M. 1976. Emerging disease problems in barley stripe rust in Bolivia, Ecuador,
Korean rice and their origins. Korean Journal Colorado, and Germany. Phytophath. 85:1040.
of Plant Protection 15:39-46. Line, R.E. 1998. Control of stripe rust and stem
Brown, W.M. 1992. New diseases of significance rust of spring barley with seed treatments and
to Colorado. Proc. of the Colorado Crop foliar fungicides, 1997. F & N Tests 53:383-386.
Protection Institute, Ft. Collins, CO. pp. 72-80. Line, R.E. 1999. Control of stripe rust of spring
Brown, W.M. 1993. The role of IPM in barley with foliar fungicides, 1998. F & N Tests
contemporary agriculture and environmental 54:299-303.
issues. In: Successful Implementation of Line, R.F., and Chen, X.M. 1996. Wheat and barley
Integrated Pest Management for Agricultural stripe rust in North America. Proc. of the
Crops. A.R. Leslie and G.W. Cuperus (eds.). Europe Mediterranean Cereal Rust Powdery
Lewis Publishers, CRC Press, Boca Raton, FL. Mildew Conference. 9:2-6.
pp. 171-79. Line, R.F., and Chen, X.M. 1999. Epidemiology
Brown, W.M., Velasco, V.R., and Hill, J.P. 1996. and control of barley stripe rust in North
Integrated management of barley stripe rust America. Proc. 16th American Barley
disease: A case study in successful Researchers’ Workshop. Idaho Falls, ID. p. 24.
international cooperation. Technical Quarterly Marshall, D., and Sutton, R.L. 1995. Epidemiology
33:241-245. of stripe rust virulence of Puccinia striiformis
Brown, W.M., Velasco, V.R., Hill, J.P., and Dillon, f.sp. hordei, and yield loss in barley. Plant Dis.
M.A. 1993. Barley stripe rust, Puccinia 79:732-737.
striiformis, found in Colorado. Phytopath. Navarro, F.M., and Zamora, M.D. 1990. Control
83:690 (abstr.). quimico de la roya lineal amarilla de la cebada
Brown, W.M., Velasco, V.R., Hill, J.P., Marshall, D., en la Mesa Central. SARH. INIFAP-CIFAEM.
Wesnberg, D.M., and Bockelman, H.C. 1993. Mexico. 10 pp.
Responses of national small grain collection Velasco, V.R., and Brown, W.M. 1982. Stripe rust
and other elite barley germplasm to stripe rust occurrence and control in Bolivia. Phytopath.
race 24. 15th North America Barley Research 72:975 (Abstr.).
Workshop, Guelph, Canada. 1992. Barley Velasco, V.R., Brown, W.M., and Hill, J.P. 1995.
Newsletter. 36:183 (abstr.). Evaluation of barley stripe rust control with
Calhoun, D.S., Arhana, K., and Vivar, H.E. 1988. seed treatments and/or foliar fungicides.
Chemical control of barley stripe rust, a new Phytopath. 85:1042.
disease for North America. Barley Newsletter Velasco, V.R., Brown, W.M., Hill, J.P., Bockelman,
32:109-12. H.E., and Wesenberg, D.M. 1991. Barley stripe
Chen, X.M., Line, R.F., and Leung, H. 1995. rust race 24 resistance identified in field
Virulence and polymorphic DNA relationships screenng trials in Bolivia, SA. Phytopath.
of Puccinia striiformis f.sp. hordei to other rusts. 81:1347 (Abstr.).
Phytopath. 85:1335-1342. Velasco, V.R., Hill, J.P., and Brown, W.M. 1993.
Dubin, H.J., and Stubbs, R.W. 1984. Puccinia Barley stripe rust-race 24 control with triazole
striiformis f.sp. hordei, cause of barley yellow fungicides. Phytopath. 83:1335 (Abstr.).
rust epidemic in South America. Phytopath. Velasco, V.R., and Brown, W.M. 1980. Incidencia y
74:821 (abstr.). control de la raza nueva de Puccinia striiformis
Dubin, H.J., and Stubbs, R.W. 1986. Epidemic en cebada. III Seminario Nacional de
spread of barley stripe rust in South America. Investigaciones Agropecuarias y Forestales.
Plant Dis. 70:141-144. Santa Cruz de la Sierra, Bolivia.
25

Impact of ICARDA/CIMMYT
Barley Germplasm on Barley
Breeding in Ecuador
O. CHICAIZA1

Since 1984, the barley breeding program Development of barley cultivars


in Ecuador started a very close with multiple disease resistance
collaboration with the ICARDA/ The main goal of Ecuador’s barley
CIMMYT Barley Program to develop breeding program has been to develop
varieties suitable for the Ecuadorian new barley cultivars with multiple
highlands. In Ecuador, barley is disease resistance. Diseases or disease
cultivated under rainfed conditions on complexes determine the type of
mountain slopes at altitudes of 2400 to germplasm that needs to be developed
3600 m. The highlands of Ecuador feature for the Ecuadorian highlands. Since
small subsistence farms where 1984, a breeding methology for the
mechanization is limited because of the incorporation of genetic resistance to
terrain, and most threshing is done by major diseases was adopted by the
animals. ICARDA/CIMMYT Barley Breeding
Program, because most available
Statistical data show that the area planted resistance sources, most of them in poor
to barley dropped from 100,231 ha in agronomic backgrounds, were useful
1969 to 26,878 ha in 1978, due mainly to only for a single disease.
the presence of the pathogen Puccinia
striiformis f.sp. hordei, which causes The incorporation of two or more
yellow rust. The new disease wiped out disease resistance genes into well
all the barley varieties in Ecuador. By the adapted varieties or advanced lines,
late 1970s, the barley program had followed by rounds of crossing and
released three resistant varieties (Dorada, selection, allowed the combination of
Duchicela, and Teran) selected from resistance genes for yellow rust, leaf
materials introduced from the world rust, scald (Rhynchosporium secalis), and
collection. Yet the rapid adoption of these BYD.
varieties by farmers was not enough to
Because most sources of resistance were
return barley to its previous area. Even
available at the ICARDA/CIMMYT
though the varieties Dorada and Teran
Program, initial crosses, at least at the
are still resistant to yellow rust, they have
very beginning of the joint program,
been eliminated from the barley growing
were done in Mexico. As the gene(s) was
area because of their susceptibility to leaf
incorporated into local germplasm,
rust of barley, P. hordei.

1 Leader, Cereals Program, Instituto Nacional de Investigaciones Agropecuarias (INIAP), Ecuador.


26 O. Chicaiza

crosses were also done by the national barley area, which increased to 80,000 ha
breeding program. In both cases, by 1998.
segregating populations were evaluated
The most recent release is the variety
in both Mexico and Ecuador to speed up
INIAP-SHYRI 2000, generated for the
the selection process.
national program using as parents the
well adapted variety INIAP-SHYRI 89
New barley varieties
and GRIT, introduced by the ICARDA/
released in Ecuador
INIAP-SHYRI 89 was the first variety CIMMYT Program in 1990. The new
released in Ecuador in 1989. This variety variety will replace INIAP-SHYRI 89.
originated from the cross LIGNEE 640/ The origin and characteristics of barley
KOBER//TERAN 78; the original cross varieties released in Ecuador are
was made in Mexico and introduced to presented in Table 1.
the national program as an F5 line in
1985. The new variety had high levels of New barley germplasm
available in the national
resistance to yellow rust, scald, leaf rust,
program
net blotch, and BYD, but was susceptible
The origin and yield performance of the
to loose smut. It has been the most
best lines in Santa Catalina in 1999 are
widely adopted barley cultivar in
presented in Table 2. Yields of lines
Ecuador. In 1992, the national program
derived from crosses made by the
released another two varieties originated
national program are very similar to
in the ICARDA/CIMMYT Program:
those of lines derived from crosses made
INIAP-CALICUCHIMA 92 and INIAP-
by the ICARDA/CIMMYT Program. The
ATAHUALPA 92. Together these three
best five lines yielded 42-75% more than
varieties cover almost 90% of all the
INIAP-SHYRI 89. This yield increase

Table 1. Origin and characteristics of barley varieties released in Ecuador.

Year of Days to Plant Yellow Leaf Yield


Variety release harvest height (cm) rust rust (kg/ha)
DORADA (selection from C.I. 9650) 1973 170 110 0 80S 3636
DUCHICELA 1978 180 120 30S 5MS 3700
TERAN 78 (selection from Abyssinian 669) 1978 145 105 0 60MS 2715
INIAP-SHYRI 89 1989 154 105 0 5MS 4937
LIGNEE640/KOBER//TERAN 78
CMB83A-2561-A-2M-1Y-1LAG-1E-OGH
INIAP-CALICUCHIMA 92 1992 160 110 15MS 60S 3958
LB IRAN/UNA8271//GLORIA”S”/COME”S”
CMB84A-1127-D-2B-1Y-6M-0Y
INIAP-ATAHUALPA 92 1992 155 100 5MS 0 3542
SUTTER/GLORIA”S”/COME”S”/3/PI6384/
CAPUCHONA
CM86-767-C-2Y-168GH-2M-2Y
INIAP-SHYRI 2000 2000 160 110 0 TR 8056
SHYRI/GRIT
E-II-93-8891-2E-1E-4E
Impact of ICARDA/CIMMYT Barley Germplasm on Barley Breeding in Ecuador 27

may be due to: 1) incorporation of yellow introduce the variety INIAP-SHYRI 89.
rust, leaf rust, and scald resistance genes The high yield potential shown by this
into a single genotype, and 2) selection variety gave farmers enough confidence
for high yield per se. Of these new lines, to start applying inputs such as fertilizers
INIAP-SHYRI 2000 was released as a and herbicides in the next cycles. The
variety in 1999; the line CARDO “S” was results of four years of work are
released in the year 2000. summarized in Table 3. The rapid increase
in the number of families participating
Performance of barley varieties and the number of hectares planted to the
in Saraguro, Loja, Ecuador new varieties is a clear demonstration that
Until 1995, the average yield obtained by small-scale farmers have adopted new
small poor farmers in Saraguro, Loja, was varieties released by the national
700 kg/ha. This low yield was due to the program. Average yields now range from
susceptibility of the local variety Clipper 1.9 to 3.5 t/ha, with some farmers
to yellow rust, scald, and leaf rust; and obtaining nearly 5.0 t/ha.
the lack of new, improved varieties. In
1995, the breeding program decided to Conclusions
• Barley germplasm developed by the
ICARDA/CIMMYT Program has
Table 2. Origin and yield performance of the
allowed the release in Ecuador of new
best barley advanced lines in Santa Catalina, varieties with multiple disease resistance
1999. and high yield potential.
Yield % yield
• New barley germplasm that will allow
Advanced line (kg/ha) increase continued progress is available in the
INIAP-SHYRI 89 (check) 4954
national program.
• Seed of the new high yielding barley
ROLAND/EH11//ESC-II-72-83-3E-7E- 8704 75
5E-1E/3/SEN”S”/4/ALELI
varieties has to be available for small-
CMB90A-871-C-1M-1Y-1M-0Y-0E scale farmers to give them the
INIAP-SHYRI 2000 8056 62
opportunity of benefiting from the new
SHYRI89/GRIT varieties.
E-II-93-8891-2E-1E-4E-0E-0E-0E- • Better varieties have helped small-scale
0E-0E-0E
farmers to develop enough confidence to
GAL/PI6384//CN48/CI8985/3/ 7569 52 apply inputs such as fertilizers and
GLORIA”S”/COPAL”S”
E-II-89-8889-1E-4E-1E-3E-0E
herbicides.

CARDO “S” 7153 44


CMB85A-1300-E-15B-5E-0E
GLORIA”S”/COPAL”S”//ABN/3/SHYRI 7037 42
CMB87-E-2Y-3B-2Y-3M-1Y-0M-0E

Table 3. Performance of barley varieties in Saraguro, Loja, Ecuador.


Participating families (no.) Hectares (no.) Mean yield (t/ha)
Varieties 1996 1997 1998 1999 1996 1997 1998 1999 1996 1997 1998 1999
SHYRI 89 13 240 334 600 8 80 141 190 3.16 2.05 1.94 2.1
SHYRI 2000 16 6 3.5
28

Malting Barley for the


New Millennium
L. WRIGHT1

A brief look at the history of By the 16th century certain European


malting barley regions, mainly the French and Danish
What will malting barley look like in the areas, were prized as having better
next thousand years? What will the malting barley than others. Why it was
malting quality be? To successfully better was not specifically recorded, but
answer these questions we need to look a statement from a 18th century brewer
at what has happened in the last 1000 summed it up best: “You can’t make a
years and perhaps from the beginning of good beer out of poor malt”. At that time
recorded time. Around 6000 years ago, plump barley, with low protein and
the Sumarians wrote the first references bright hull color, was a very rough “rule
about barley and beer. During the next of thumb” for malt quality.
5000 years the Sumarian beer recipes
were refined by the Babylonians, and The Industrial Revolution of the 19th
then by the Egyptians, by the Greeks century also revolutionized brewing
and then the Romans. science. Scientific instruments were
invented to help measure the various
With the spread of the Roman Empire, stages of beer making. Instruments like
beer making spread throughout Europe hydrometers, attemperators, and
and especially to the northern part of the improvements in thermometers and
Empire to the Gauls, the Teutons and refrigeration made it possible to make
into Scandinavia. By the end of the first better, more consistent beer, and in
millennium AD the art and science of larger volumes.
malting barley and beer making were
firmly established in the monasteries; Another historical event took place in
each monastery had its own special 1842 in Plzen, in what is now the Czech
sources of barleys used for malting and Republic: the creation of a clear, golden-
its special way of brewing beer. After the colored lager beer. Until that time lager
Reformation, brewing shifted from the beers were dark colored and cloudy. This
monasteries to small family breweries in clear, golden-colored lager beer became
the towns and cities. a much-sought-after standard in the
making of lager beer, the most popular
style of beer in the world. This beer was

1 Busch Agricultural Resources, Inc. The opinions outlined in this talk are the author’s and do not
necessarily reflect those of Anheuser-Busch or Busch Agricultural Resource.
Malting Barley for the New Millennium 29

made from locally grown, low-protein In the early 20th century, many local
barley, malted using a new British malting varieties evolved into regionally
malting system that applied indirect heat adapted malting varieties. In the latter
and used very soft water having a very part of the same century those regional
low soluble minerals content. malting varieties moved globally. This
was driven for the most part by malting
In North America, beer making
companies buying suitable malting
developed in the 18th century and
barley for their brewing customers, who
flourished during the 19th century.
in turn had gone from being local
Malting barley varieties developed in the
breweries to regional breweries. These
United States were higher in protein for
regional breweries needed larger
several reasons. First, the original barley
amounts of high quality malt as they
areas in North America were better
increased their size to meet regional
suited for growing six-rowed malting
demands. They knew which malting
barley than two-rowed barley. Second,
varieties gave them good brewing
six-rowed barleys were inherently higher
results, so they favored these varieties
in protein than two-rowed malt barley.
over several lesser known malting
And third, the use of supplemental starch
varieties. Thus, locally adapted varieties
additives in the mashing process, called
gave way to regionally grown varieties.
adjunct, required a higher enzyme level
and thus a higher protein level in the These regional varieties are now giving
malt. All of these events, plus many way to even fewer malting varieties
more, gave us the beer and the malting grown in wider and more diverse
barleys we have today. regions. As malting and brewing
companies source malting barley from
Trends in current and future wider and more diverse malting growing
malting varieties areas, malting barley varieties in the
What will the trends in malting barley future will need to be widely adapted in
quality be in the 21st century? Since terms of yield, disease resistance, and
malting barley and the brewing industry malting quality. If a malting variety is
have changed so rapidly in the past 200 not widely adapted, it may not make it
years, it is impossible to accurately on the list of malting varieties that
predict too far in the future. Also brewing breweries give to their malt suppliers.
techniques in the future may change Therefore, future malting varieties will
some of the parameters for barley quality be tested in more than one growing
we have today. Yet, based on what has region and will have wide adaptation. If
happened in the past 200 years, certain there is a slogan for the malting barley
predictions can be made on where breeder in the future, it is: “Develop
malting barley could be in the next 100 malting barley varieties locally, but test
years. Much of the information used in globally.”
this talk is from the American Malting
Barley Association (AMBA), which Malting barley must keep pace as an
recommends malting barleys for the economically viable crop for farmers to
United States on behalf of its member grow. The return per acre that a crop
maltsters and breweries, Anheuser-Busch generates will remain the factor that
among them. determines which crop the farmer will
30 L. Wright

grow in any given year. Since commodity This is just a sampling of the many
prices have remained fairly constant, the important agronomic traits that the
increase in the value of the crop must malting barley of the future will need.
come from increased grain yield. Yet it gives an idea of what is needed for
Increased grain yield comes from developers of new malting varieties. The
improved cultural practices and newer, bottom line is that future malting barley
higher yielding varieties. breeders need to think globally when
they develop new malting varieties.
In the US, new malting varieties are
accepted, on average, every 10 years.
Quality requirements
This rate of acceptance has not kept pace
What will be needed in malting quality
with the introduction of varieties of
in the future? Basically, brewers want
competing crops. The lag in the
large quantities of grain of known
introduction of new varieties is putting
varieties with consistent quality.
increasing pressure on the brewing
Remember that malting companies are
industry to accept newer varieties at a
buying malting barley for their brewing
faster rate. In the future, economically
customers. The brewing industry, as a
viable malting varieties will probably
whole, is very conservative and changes
have a shorter market life and will make
slowly. Breweries need consistent
way for higher yielding malting varieties
malting quality and approve only those
with very similar malting quality.
new varieties that meet their needs. Yet
In conjunction with grain yield and wide there are some trends in malting quality
adaptation, disease resistance also needs that may be predicted.
to be thought of globally. Disease affects
Extract is the measure of everything that
not only yield but also malting quality
is put in solution from the malt at the
through reduced plumpness, increased
start of the brewing process. It is a
protein, and reduced malt extract. Future
measure of soluble sugars, dextrins,
malting barleys will be grown in many
soluble proteins, beta-glucans, and other
areas and under different disease
compounds. Just as grain yield is money
pressure. The developers of future
to the farmer, extract yield is money to
malting varieties must be aware of which
the brewer. Generally speaking, the more
genes for resistance to minor as well as
extract there is in the wort, the more food
major diseases their varieties possess.
for the yeast in the fermentation process.
This is important to determine whether a
The amount of fermentable sugars in the
variety can be introduced into other
extract is particularly important because
regions.
the yeast converts these sugars into
The successful malting barley of the alcohol and CO2.
future will have to have a set bank of
In the last 40 years the extract of US
disease resistance genes to give it the
varieties has increased from around 75%
extra flexibility to be sown in diverse
to roughly 80%. Will we see another 5%
growing areas. This is being greatly
increase in the next 40 years? Perhaps,
facilitated today with the advent of
but we should point out that there is an
molecular marker assisted selection and
upper limit for extract in the barley
technologies to screen for several
kernel, and we are close to reaching it. In
markers at the same time.
Malting Barley for the New Millennium 31

determining the maximum extract important because it controls and


potential, we have to take into account influences many aspects of malt quality,
not only the soluble compounds in including enzymatic activity, foam
barley malt, but also all the insoluble properties, and beer flavor. Therefore, a
compounds. The hull component, the balance needs to be maintained between
insoluble proteins, and other insoluble these factors. Higher malt protein will
components have been estimated at increase the enzymatic activity of the
around 15%; thus the maximum malt, but will also lower the fermentable
extract potential in the barley kernel is sugar content in the extract. Lower malt
around 85%. protein will increase the fermentable
sugar content in the extract, but could
There are only a few ways that extract
lower enzymatic activity below the level
could increase beyond this limit. The
needed to convert the adjunct starch. A
author of a recent publication states that
proper balance has to be maintained to
the percent extract would increase by
accomplish both high fermentable sugars
more than 5% through the use of a hull-
and high enzymatic activity.
less malting barley. This increase in
extract is very interesting to the Soluble protein, or wort protein, is the
brewers, who consider a 0.1% increase amount of malt protein that becomes
to be substantial. Will the brewing soluble in the mashing process. Brewing
industry shift to using hull-less malting yeast needs a certain level of soluble
barleys? Not without changing the protein to provide the nutritional amino
brewing process. Barley hulls are an acids needed for proper yeast growth.
important component of the wort- Too little or too much protein will cause
filtering process and contribute essential problems in the fermentation process and
flavor to the beer. However, due to the may lead to undesirable beer flavor
significant jump in extract that hull-less compounds and beer color. Soluble
malting barley could contribute, its use protein has increased over the last 40
will no doubt be studied closely by the years. For example, the high level of
brewers. If hull-less malting barley is soluble protein in the six-rowed variety
used in brewing, the smaller breweries “Stander” has caused problems in the
will start using it first; the larger and brewing process. As a result, the AMBA
more conservative breweries will has implemented new guidelines for the
probably not use it until all processing maximum desired level of soluble
and flavor concerns are satisfactorily protein.
answered.
These new guidelines have brought
Another way to increase extract is by about a progression toward lower
reducing protein. There is a reverse soluble protein malting varieties.
correlation between malt extract and Breeding guidelines are 11.5% to 12.5%
barley protein. Less protein is related to for malt protein and 4.7% to 5.5% for
more starch in the grain. From a soluble protein. These levels may be
brewer’s standpoint, a high starch lowered again some time in the future to
content in malting barley is desirable as 11.0% to 12.0% for malt protein and 0.0%
a source of fermentable sugars. Grain to 5.0% for soluble protein. This
protein content, on the other hand, is reduction should maintain favorable
32 L. Wright

brewing components while minimizing diastatic power and alpha-amylase


unsatisfactory processing conditions, levels must be similar to either
beer color, and/or beer flavor. Even Harrington, a two-rowed malting
lower levels may be possible if brewing variety, or Robust or Morex for six-
processes and yeast nutrition are not rowed malting barley. These varieties are
adversely impacted. generally lower in these enzymes than
Stander. Desired levels of diastatic
Enzymatic activity of malting barley is
power are now 120 to 150, while alpha-
needed to break down the starch in the
amylase levels are 45 to 55. These new
kernel into sugars that in turn are
levels should remain fairly stable or
converted into alcohol by the yeast. In
perhaps be slightly lower in the future,
North America, a higher level of
unless there are significant changes in
enzymatic activity is needed to convert
the brewing process.
the starch of the adjunct as well.
Enzymatic activity is generally
Application of new technologies
measured using two parameters:
New technology will be developed to
diastatic power and alpha-amylase.
assist in breeding better malting barley
Diastatic power is the measure of
varieties. Molecular marker assisted
amylolytic enzymes levels in the malt,
selection (MAS) will continue to be a
including alpha- and beta-amylase, limit
tool to determine the genetic make-up of
dextrinase, and the alpha-glucosidases.
new experimental barleys. As locations
This is a general measurement of the
of genes conferring important traits and
ability of the malt to rapidly break down
markers for them are discovered, a
the starch into fermentable sugars.
library of markers will be built to enable
Alpha-amylase is the major enzyme in
the breeder first to test experimental
starch conversion and is measured
malting lines in the lab and then to
separately.
verify the selected lines in the field. New
North American malting barleys, testing procedures and instrumentation
especially the two-rowed malting will make it possible to screen for several
varieties, have increased both their hundred markers at the same time. This
diastatic power and alpha-amylase will greatly improve the effectiveness of
levels in the past 40 years. Consequently, the breeder in selecting successful
there is concern that these higher malting lines.
enzymatic varieties have more spouting
Since we are speculating on future
problems under wet harvest conditions.
developments, there is a possibility that
In addition, there are indications that the
improved brewing chemistry could help
upper limit for both diastatic power and
the breeder by finding appropriate and
alpha-amylase may have been reached
inappropriate flavor compounds and the
when looking at brewhouse
genes behind them. This would allow
performance. Again, the six-rowed
the breeder to include additional
variety “Stander” is in the center of the
markers for these flavor genes in his
debate due to its higher levels of both
library of markers. It should be stressed
diastatic power and alpha-amylase.
that the malting and brewing industry
After the release of Stander, breeding
will still require barley breeders to grow
guidelines were changed to state that
Malting Barley for the New Millennium 33

out their malting lines to determine their using GMO raw materials in the
malting and brewing potential. brewing process. However, I believe this
type of research needs to be encouraged
One new technology that will take more
in barley while the GMO issue is being
time to be accepted for use in approved
reviewed by the brewing industry.
malting barley varieties are genetically
modified organisms (GMOs), which This is a little of what may be waiting in
mean that non-barley genes have been the future of malting barley. Some
incorporated into malting barley. GMOs researchers can already see the
are a controversial subject yet to be beginnings of new tools that may help
settled in Europe, Japan, and the U.S. It select malting barley varieties in the
is a mixed blessing that barley has not future. Others researchers may already
been easily tissue cultured. By the time a be implementing some of the ideas
truly genetically modified malting mentioned here. There are other new
barley is ready to be tested by the and exciting improvements in both the
brewing community, the issue of GMOs barley field and in the brewhouse that
may be resolved. In the meantime, the have not been mentioned here but we
brewing community is not ready to start await the report of these improvements
at a later date.
34

Barley Breeding in Peru


M. ROMERO LOLI AND L. GOMEZ PANDO1

Barley is a very important crop in the contributes at least 20% of the total
highlands of Peru, especially at altitudes caloric intake of rural families.
above 3000 m (Table 1). Few crops, aside According to national statistics, there are
from barley, are suitable for growing 3 million poor people living in highland
under the adverse environmental rural areas who subsist mainly thanks to
conditions (such as frost and drought) their farming activities.
that predominate in this region. Barley
Table 2 shows the barley area,
production, and yield per hectare from
1977 to 1998. Barley is produced mainly
Table 1. Main crops, area, production, and yield by subsistence farmers as a staple food
per unit area in Peru, 1998.
crop in relatively small plots (Table 3).
Food Area Production Yield
Barley grows mostly in low-fertility
crops (ha) (t) (kg/ha)
soils. In many cases, the use of
Rice 269080 1548778 5756 machinary is not possible due to the
Potato 268847 2589338 9631 steeply sloping terrain. There is virtually
Maize 214590 230450 1074
no flat, well-watered agricultural land.
Barley 146698 165831 1130
Wheat 125894 146285 1162 Plots are perched one above another up
Banana 117792 1321890 11222 mountain sides rising thousand of
Cassava 80708 884118 10955 meters. Frost, drought, hail, and other
Bean 75164 67563 899 climatic factors reduce barley production
Faba bean 34184 38129 1115
Pea 31415 33365 1062 at that altitude.
Quinoa 30720 28614 931
Olluco 23523 116897 4969
The rusts are the most aggressive of the
Oca 21626 101639 4700 diseases present in the region. Leaf rust
Sweet potato 17397 221609 12738 causes damage all the way from sea level
Onion 14317 315622 22045 (the coast) to 2800 masl. Stem rust can be
Mashua 7244 32859 4536
found at the same altitudes, but in recent
Industrial years there have been very few
Corn 229114 702479 3066 outbreaks of the disease. Stripe rust is
Cotton 73629 95262 1294 the main disease above 2600 masl,
Sugar cane 52614 5705340 108438 affecting 90% of the cereal area. Since
Marigold 5507 96070 17445
1978, it has been the most serious
Source: OIA. Producción Agrícola 1998.

1 Programa de Cereales, Universidad Nacional Agraria La Molina, Apdo. Postal 456, Lima 100.
Email: pcereal@lamolina.edu.pe.
Barley Breeding in Peru 35

Table 2. Barley area, yield, and production in In 1968, Universidad Nacional Agraria
Peru, 1977–98. La Molina started a cereal breeding
Malt and program that continues to this day. The
Harvest barley
program was developed in close
area Yield Production imports
Year (000 ha) (kg/ha) (000 MT) (000 MT) collaboration with Malteria Lima S.A.
(Backus Corporation), ICARDA/
1977 169.72 861 146.20 43.10
1978 151.54 855 129.51 38.20 CIMMYT, Nebraska State University,
1979 152.59 861 131.44 51.00 and the International Atomic Energy
1980 109.93 890 97.87 74.10 Agency (IAEA). It focuses on cereals, but
1981 118.13 969 114.50 82.80 especially on barley production in the
1982 114.82 960 110.24 61.30
1983 105.76 830 87.81 51.30
highlands.
1984 108.67 946 102.80 64.70
1985 117.08 1060 124.11 95.60 Objectives
1986 104.05 1130 118.14 83.90 • To develop improved varieties with
1987 110.57 997 110.23 108.40 high yield potential, resistance or
1988 123.70 1040 128.59 92.40
tolerance to abiotic and biotic stress,
1989 120.22 1045 125.60 46.20
1990 75.10 954 71.64 84.20 and good quality.
1991 111.08 1028 115.22 77.70 • To develop production technologies
1992† 81.76 842 68.81 80.55 adapted to different barley production
1993† 101.17 1112 112.49 86.40 areas.
1994 112.50 1154 129.84 • To transfer research results to
1995 107.03 1143 131.19
agronomy students, agronomists, and
1996 108.93 1190 152.94 39.65
1997 129.91 1062 138.03 29.55 farmers.
1998 149.89 1130 165.83 29.95

† Jan.-Nov. 1993/1992 (preliminary data).


Breeding
Source: Oficina de Información Agraria. The breeding program has an excellent
balance between conventional plant
breeding, ICARDA/CIMMYT
germplasm augmentation, induced
limiting factor for barley production. mutation, and biotechnology.
There are many other diseases such as
powdery mildew, BYDV, and foliar Germplasm introduction and
diseases caused by Helminthosporium spp. evaluation
and Rhynchosporium secalis, which are The United States Department of
important in the highland region during Agriculture and CIMMYT have
warm and wet years. provided more than 10,000 barley
accessions. This entire collection was
In general, very little farming technology screened for adaptability in several parts
is applied to barley production. The of the Andean Region. Only a few
average barley yield was 0.8 t/ha some foreign accessions were adapted to such
years ago. However, the average yield extreme environmental conditions.
has now increased to 1.5 t/ha in some
areas thanks to new varieties developed National collections of local barley were
by Universidad Nacional Agraria La put together for the highlands of Peru
Molina (Pasco; Table 4). and Bolivia. These materials have been
36 M. Romero Loli and L. Gomez Pando

Table 3. Size and number of farm units and uses of barley grain in Peru.

Grain uses
Sold at the Sold on the Home Sold as
Total units farmgate market consumption seed
Below 0.5 ha
Number of units 22657 74 433 22137 73
Area 1589.75 7.93 42.28 1537.43 2.12
0.5 to 4.9 ha
Number of units 162166 1204 7832 154624 281
Area 59279.45 545.45 3756.54 54891.77 85.69
5.0 to 9.9 ha
Number of units 37151 499 2738 34722 89
Area 29512.47 490.68 2888.86 26069.66 63.27
10 to 19.9 ha
Number of units 16505 250 1299 15309 53
Area 17790.38 385.7 2120.2 15206.68 77.8
20 to 49.9 ha
Number of units 7635 131 624 7024 34
Area 10455.31 285.4 1519.99 8585.76 64.15
Above 50 ha
Number of units 5319 89 406 3076 19
Area 7251.42 240.47 1728.94 5241.09 40.91
Total no. of units 249633 2247 13332 236892 549
Total area 125878.78 1955.63 12056.81 111532.39 333.94

Source: INEI. III Censo Nacional Agropecuario, Resultados definitivos, 1996.

used to develop and improve new Table 4. Barley area, yield per hectare, and
varieties. The continuous introduction of number of farm units in different departments
in the Peruvian highlands.
new accessions through the ICARDA/
CIMMYT Barley Program and other Barley Yield Farm
institutions is very useful to our Department area (ha) (t/ha) units
breeding program.
Ancash 6657.65 939 11005
Apurimac 3128.00 1026 7892
Methods Ayacucho 8764.96 738 18941
Hybridization. In this method, one Cajamarca 7011.36 986 11679
parent is usually a line locally developed Cusco 8905.66 1250 23143
by the cereals program; the other is Huancavelica 20632.97 1319 31801
Huanuco 3803.27 1300 8474
generally a selected introduction.
Junin 8724.07 1422 16796
Selection is done using bulk or La Libertad 11475.12 1209 10278
individual selection. Pasco 29.26 1519 191
Puno 43970.57 1003 191

Source: INEI. III Censo Nacional Agropecuario,


Resultados definitivos.
Barley Breeding in Peru 37

Mutation induction. Mutation is used to performance (Table 5). It should be


improve the adaptability of modern, noted that UNA La Molina 95 is an early
high yielding cereal varieties to make maturing hull-less variety developed by
them suitable for cultivation in stress- inducing mutation in the variety
prone areas of the Peruvian Highlands Buenavista using gamma rays.
and to improve grain quality
characteristics.

Doubled haploid production. Plants Increase in National


selected from F1, F3, or M1 are used each Average Barley Yields
cycle to produce doubled haploids using
anther culture. According to statistics of the Peruvian
Ministry of Agriculture, the national
Improved crop management average barley yield increased from 859
practices kg/ha in 1978 to 1130 kg/ha in 1998.
Studies on chemical fertilization, seeding This considerable increase in yield was
density, seeding methods, time of due to the use of fertilizers and other
seeding, and chemical weed control are cultural practices in the highland region,
conducted to develop or improve which is not usual among small farmers
agronomic practices specifically for the in Peru. The increase in barley yields
region. Each year in the highlands was of great significance for small
demonstration plots using new varieties landholders in the Peruvian highlands.
are established with the cooperation of
many institutions to show farmers new
crop management practices. Socioeconomic Impact
New improved varieties The value of the annual barley
Yellow rust resistant Zapata 588 played production in Peru is estimated at US$
an important role during the severe 12 million. Barley varieties released by
epidemic that occurred from 1976 to Universidad Nacional Agraria La Molina
1978. Almost all barley varieties account for 80% of this production. From
cultivated in Peru were killed by the 1977 to 1998 barley production totalled
yellow rust pathogen introduced into US$ 228 million, to the direct benefit of
South America at that time. the highland population.

The varieties UNA 80, UNA 8270, National statistics indicate that
Yanamuclo 87, Buenavista, UNA La aproximately 49.6% of Peru’s total
Molina 94, UNA La Molina 95, and UNA population lives in poverty or extreme
La Molina 96 were released between poverty; three million of those people
1980 and 1996. These varieties show live in rural highland areas. Barley is
higher yield potential, improved yellow mainly used for food and feed in Peru.
rust resistance, and better grain quality. In the highland region, about 70% of
Highland farmers adopted improved barley grain is used for human
varieties due mainly to their good consumption.
38 M. Romero Loli and L. Gomez Pando

Table 5. Improved barley varieties released by Universidad Nacional Agraria La Molina from 1978 to
1996.

Yield Adaptation
Genealogy (t/ha) (masl) Disease reaction

Zapata 588 2–6 3000 –3800 Tolerant to yellow rust


B112(F7-1962)D2hsIII/Compuesto XXI-51Cz

UNA 8270 2 –5 3000 - 3800 Resistant to yellow rust


CNC 203131/Compuesto XXI Tolerant to scald

UNA 80 2.2 – 6 3000 – 3800 Resistant to yellow rust


CNC 203346/CXXI

Yanamuclo 87 2.4 – 5 3000 - 3800 Resistant to yellow rust


UNA 8309 x Itintec 77

Buenavista 2.5 –3 3000 – 3800 Resistant to yellow rust


P71318-Row134.78 (ICARDA/CIMMYT: F3 Generation)

UNA La Molina 95 2.5 –5 3000 – 3800 Resistant to yellow rust


Parent Material Buenavista (Gamma ray 300 Gy) Tolerant to Pyrenophora teres

UNA La Molina 96 2.5 -5 3000 – 3600 Resistant to yellow rust, leaf


Gloria”s”/Celo”s”/ESCII-77-83-3E-7E-5E/1E/3/Lignee 527 rust and powdery mildew
(ICARDA/CIMMYT: F3 Generation)
39

Accumulating Genes for Disease


Resistance in Two-Rowed Barley
for North Dakota
J.D. FRANCKOWIAK1

A large portion of the barley (Hordeum Many of the six-rowed barley cultivars
vulgare) crop in North Dakota (ND) is released for production in Manitoba,
grown for the malting and brewing Minnesota, North Dakota, and South
industry. Most cultivars have a six-rowed Dakota have similar agronomic traits and
spike type and are used as malting barley are often referred to as Midwest six-
based on recommendations made by the rowed barley. Historically, spot blotch,
American Malting Barley Association, incited by Cochliobolus sativus, and wheat
Inc. (AMBA). Eastern ND is generally stem rust, incited by Puccinia graminis f.
more favorable for producing malting sp. tritici, are the barley diseases that have
barley than western ND, where low caused the greatest production losses in
yields and thin kernels are frequently eastern and central ND. Barley diseases
production problems. During relatively rarely cause losses in western ND.
dry growing seasons in western ND,
two-rowed barley often yields more than Development of two-rowed
six-rowed barley. Therefore, the ND barley for ND
Agricultural Experiment Station funded In 1970, Dr. Glenn A. Peterson made
an improvement program for two-rowed crosses designed for improvement of
barley in the 1970s. two-rowed barley. He assumed that some
traits of Midwest six-rowed barley were
Barley research at North Dakota State needed in two-rowed barley for ND. The
University (NDSU) is partially supported best two-rowed cultivars from the
by grants from AMBA and its precursory western USA and prairie provinces of
agencies. Support for two-rowed barley Canada were susceptible to spot blotch,
improvement was based on the wheat stem rust, and net blotch, incited
assumptions that two-rowed cultivars by Pyrenophora teres f. teres. Crosses were
will show greater yield stability in made between the two types of barley
western ND and will be suitable for and the F1 plants were crossed to another
production of malting barley in eastern two-rowed cultivar to increase the
ND. The two-rowed program frequency of two-rowed plants in the
complements the six-rowed program, progenies. In 1974, Dr. Melvern K.
which was started in the early 1940s.

1 Department of Plant Sciences, North Dakota State University, Fargo, North Dakota, 58105 USA.
40 J.D. Franckowiak

Anderson was employed as the first two- Bowman was widely grown in
rowed barley breeder at NDSU. southwestern ND because it yielded well
and often had much higher test weight
Two-rowed lines selected from the initial
values than other barley cultivars.
three-way crosses were tall and lacked
Bowman was utilized in breeding
adequate disease resistance. To generate
subsequent two-rowed cultivars for ND
better material, good selections were
because of its unique combination of
crossed to six-rowed cultivars and the F1
plant height and maturity genes. Other
plants crossed to a two-rowed cultivar. In
two-rowed barley cultivars released for
1978, when selections from the second
ND include Stark in 1991, Logan in 1995,
cycle of crosses were in preliminary yield
and Conlon in 1996. Conlon was
trials, this author was employed as the
recommended as a malting barley
two-rowed barley breeder. Research on
cultivar by AMBA in 2000 and as such is
barley is conducted at NDSU by a team
the first two-rowed malting barley
of scientists including a two-rowed
recommended for production in ND.
barley breeder, a six-rowed barley
These two-rowed cultivars have similar
breeder, a cereal chemist, a plant
agronomic characteristics and are best
pathologist, a geneticist, a virologist, and
adapted for production in western ND
agronomists at the ND Research
and adjacent areas of Montana and South
Extension Centers.
Dakota. They can be referred to as
‘Midwest’ two-rowed barley.
Development of ‘Midwest’ two-
rowed barley cultivars
Barley diseases in eastern ND
As two-rowed lines suitable for
When released, Bowman was resistant to
production in western ND were
wheat stem rust and moderately resistant
identified, phenotypic variability among
to net and spot blotch. However, its levels
the lines derived from the two-rowed by
of net and spot blotch resistance were not
six-rowed crosses was rapidly restricted.
adequate to recommend Bowman for
In 1979, the line ND4994 was observed to
production in eastern ND. Bowman is
be earlier, shorter, and stiffer than most
susceptible to number of pathogens of
other selections. ND4994 had large,
minor importance in ND. They include
plump kernels; acceptable levels of malt
leaf rust, Puccinia hordei; barley yellow
extract; and moderate values for grain
dwarf virus (BYDV); scald,
protein, diastatic power, and alpha-
Rhynchosporium secalis; bacterial blight,
amylase; but its yields were below
Xanthomonas campestris pv. translucens;
average. In 1980, however, when the
powdery mildew, Blumeria graminis f. sp.
growing season was relatively hot and
hordei; loose smut, Ustilago nuda; and
dry, ND4994 was the only early line with
covered smut, Ustilago hordei. Bowman is
high yields. A reselection was released in
resistant to barley stripe mosaic virus
1984 under the name Bowman (PI483237)
(BSMV) and shows some resistance to
and recommended for production in
black point and common root rot, both
western ND (Franckowiak et al., 1985).
incited by C. sativus.
Bowman was classified by AMBA as a
non-malting barley cultivar. After the release of Bowman, changes
occurred in the relative importance of
Accumulating Genes for Disease Resistance in Two-Rowed Barley for North Dakota 41

several barley pathogens. In 1989, a new Development of disease-


race of Puccinia graminis f. sp. tritici resistant, Midwest two-rowed
attacked cultivars previously classified as barley
resistant (Jin et al., 1994). In 1990, a new Because barley is a low value crop and
pathotype of C. sativus was found to disease epidemics are often sporadic,
attack Bowman and many of its many ND farmers do not apply
derivatives (Fetch and Steffenson, 1994). fungicides. A few barley diseases can be
This pathogenicity change greatly controlled by clean seed programs or by
lowered the yields of Bowman in trials at seed treatments. Genetic resistance to
Fargo and Langdon in eastern ND (Table barley pathogens is considered the best
1). In the early 1990s, speckled leaf means to minimize losses caused by
blotch, incited by Septoria passerinii and diseases. Thus, incorporation of genetic
Stagonospora avenae f. sp. triticea, returned resistance to barley pathogens was
as production problems in northeastern established as a breeding goal when
ND (Toubia-Rhame and Steffenson, breeding of six-rowed barley was
1999). In the 1990s, Bowman showed a initiated.
susceptible reaction to new isolates of P.
The two most important diseases, wheat
teres. However, the major event in the
stem rust and spot blotch, received more
Upper Midwest barley production area
attention than other diseases. Genes for
was epidemics of Fusarium head blight
resistance to loose smut, speckled leaf
(FHB), incited primarily by Fusarium
blotch, net blotch, and BSMV were
graminearum, which started in 1993
incorporated into a few six-rowed
(Schwarz et al., 1995a). The presence of
cultivars. When other diseases occurred
the toxin deoxynivalenol (DON) reduced
in field plots, highly susceptible lines and
the value of crop and products made
selections were discarded. When
from infected grain (Schwarz et al.,
improvement of two-rowed barley was
1995b).
undertaken, disease testing focused on
wheat stem rust, spot blotch, and net
blotch.
Table 1. Grain yields (t/ha) of selected barley
cultivars in trials conducted in North Dakota
from 1995 to 1998. Resistance to spot blotch
Controlling losses caused by spot blotch
Trial locations in North Dakota has been challenging. Shortly after the
Row Williston Carrington Fargo Langdon Average barley improvement program was
Cultivar type 4† 4 4 4 28 started, the six-rowed selection ND B112
Bowman 2 3.72 3.51 2.98 3.86 3.32 (CIho 11531) was shown to be highly
resistant to spot blotch (Wilcoxson et al.,
Logan 2 3.87 4.42 3.99 4.93 4.23
1990). Six-rowed cultivars developed
Conlon 2 3.57 4.12 3.52 4.53 3.84
from crosses to ND B112 were released in
Morex 6 3.71 3.90 3.56 4.29 3.71 the 1960s and 1970s. Bowman has a
Stander 6 3.79 4.60 3.75 4.67 4.18 moderate level of spot blotch resistance,
which is probably controlled by two
† Number of trials.
genes from its six-rowed parents
(Steffenson et al., 1996).
42 J.D. Franckowiak

Although this level of resistance is lower susceptible field reactions to net blotch in
than that of ND B112, spot blotch eastern ND while many two-rowed lines
epidemics were not observed on were resistant. In the mid 1990s, Bowman
Bowman during the 1980s. During the and Stark showed susceptible reactions
1990 growing season, however, over 80% to net blotch in western ND, but six-
of the two-rowed lines in nurseries near rowed cultivars were resistant in those
Fargo were prematurely defoliated by a trials. These observations suggest that
spot blotch epidemic. This and host-pathogen interactions for net blotch
subsequent epidemics on two-rowed in ND are similar to those reported in
barley were caused by a new pathotype other barley growing areas (Khan, 1982;
of C. sativus (Fetch and Steffenson, 1994). Steffenson and Webster, 1992; Tekauz,
Six-rowed cultivars are resistant to the 1990). As new genes for net blotch
new isolate, which was found only in resistance are incorporated into cultivars,
ND (Valjavec-Gratian and Steffenson, isolates of P. teres with different virulence
1997). This isolate continued to cause patterns are identified.
epidemics in two-rowed breeding
Some lines selected from crosses between
nurseries until susceptible lines were
Bowman-derived cultivars and the two-
discarded.
rowed cultivar Norbert (PI 452125), bred
Finding two-rowed lines that are highly in Manitoba and released by Agriculture
resistant to the ‘old’ isolate of C. sativus Canada, have continued to show low
has been more difficult. Two recent reactions to net blotch in greenhouse and
observations have been helpful in in field tests. The origin of these Rpt
selecting lines having better spot blotch genes for net blotch resistance is
resistance. First, six-rowed cultivars unknown, but the cultivar Norbert,
retain green leaves and culms longer which was derived from a complex cross
than two-rowed cultivars in trials grown to CIho 5791 (Metcalfe and Bendelow,
in eastern ND. Second, lines with the 1981), is one possible source. Some lines
lowest spot blotch readings in seedling derived from crosses to accessions from
tests often have the lowest spot reactions the ICARDA-CIMMYT barley
in field tests. Using these selection improvement program in Mexico also
criteria, two-rowed lines with better spot have low net blotch scores. Hopefully,
blotch resistance have been identified presence of Rpt genes from more than
recently. one source will permit their rapid
deployment when future changes in
Resistance to net blotch virulence occur.
Two-rowed breeding lines with a
moderately resistant reaction to P. teres f. Resistance to BYDV
teres were easy to identify with seedling Barley yellow dwarf symptoms occur
tests; however, a portion of them showed sporadically on barley grown in eastern
susceptible reactions in subsequent field ND, but production losses are often, but
tests. These changes in disease reactions not always, low (Gill, 1970). Severe
illustrate the variability associated with BYDV epidemics do occur frequently in
genetic resistance to net blotch. In the late-planted breeding nurseries. Crosses
late 1980s, Midwest six-rowed barley were made in the early 1980s to
cultivars were found to exhibit introduce the Ryd2 gene for BYDV
Accumulating Genes for Disease Resistance in Two-Rowed Barley for North Dakota 43

resistance from CIho 2376 (Rasmusson accessions from the ICARDA-CIMMYT


and Schaller, 1959), but BYDV resistant barley program may have Rph genes that
selections were late and susceptible to are different from those reported in the
lodging. Additional cycles of crossing literature.
and selection did not improve greatly the
agronomic traits of BYDV resistant lines. Resistance to speckled leaf blotch
Thus, ICARDA-CIMMYT lines were In the 1950s and 1960s, speckled leaf
used as an alternative source of the Ryd2 blotch was a problem in northeastern
gene. Lines with BYDV resistance and ND; however, severe losses were not
good agronomic traits have been observed again until the early 1990s. The
recovered, but their malting quality is disease develops late during the growing
not acceptable. Malt extract values need season and causes premature death of
improvement and the low protein gene leaves and post-maturation straw
from the six-rowed cultivar Karl needs to breakage. Several genes for resistance to
be added. S. passerinii have been identified
(Rasmusson and Rogers, 1963), but they
Resistance to leaf rust are not present in Midwest six-rowed
Yield and quality losses caused by leaf cultivars. In 1993, several six-rowed lines
rust of barley in ND are generally low from the barley ICARDA-CIMMYT
because the crop is seldom infected program were observed to show resistant
before heading. The inoculum is blown reactions in field plots at Langdon, ND.
northward from over-wintering sites in These lines maintained green leaves and
the southeastern USA. Rapid changes in stems longer than susceptible cultivars.
leaf rust races are not expected because In trials at Langdon, delayed loss of
resistant cultivars are rarely grown in the green leaves is a criteria used to select
over-wintering sites. Since Midwest speckled leaf blotch resistant lines.
barley cultivars do not have genes for Subsequent seedling tests confirmed that
leaf rust resistance, crosses were made to selections from crosses to ICARDA-
introduce the Rph3 and Rph7 genes from CIMMYT lines are resistant to an isolate
Estate (CIho 3410) and Cebada Capa of S. passerinii (Toubia-Rhame and
(CIho 6193), respectively. Experimental Steffenson, 1999). Septoria resistant lines
lines with leaf rust resistance were from the second cycle of three-way
selected, but they are not suitable for crosses have been identified.
release as cultivars. Because the Rph3 and
Rph7 genes are no longer effective Resistance to Fusarium head
against all leaf rust isolates worldwide blight
(Jin et al., 1996), accessions of wild Fusarium head blight (FHB) or scab is a
barley, H. vulgare subsp. spontaneum, frequent barley production problem in
were evaluated as a source of new leaf many humid and subhumid climates.
rust resistance genes. The Rph15 gene Yield losses are often low, but the
from PI 355447 was isolated in a accumulation of toxins reduces the value
Bowman backcross-derived line and is of the grain. Until 1993, FHB was
being incorporated into Midwest two- considered a minor disease problem in
rowed breeding material (Chicaiza et al., ND. Above normal rainfall during
1996). Preliminary tests indicate heading is believed to be partially
44 J.D. Franckowiak

responsible for the FHB epidemics in some resistance to FHB based on tests in
1993 and subsequent years. The best eastern China, but that resistance is
resistance to FHB has been found in poorly expressed in ND (Urrea-Flórez,
accessions closely related to the two- 2000), where these photoperiod sensitive
rowed cultivar Svanhals and the six- cultivars head extremely early.
rowed cultivar Chevron (Urrea-Flórez,
2000). Although resistance to FHB is Multiple disease resistance in
inherited in a quantitative manner Midwest two-rowed barley
(Takeda, 1992), resistance is frequently New two-rowed cultivars for ND should
associated with the two-rowed spike ideally contain a large number of disease
trait, controlled by the Vrs1.b allele at the resistance genes. however, accumulation
vrs1 locus (Prom et al., 1997; Takeda, of these genes in elite breeding material
1990). Molecular mapping of FHB is slow because the donor parents are
resistance in Chevron identified poorly adapted to ND. Thus,
several QTLs of which one is associated development of locally adapted material
with the vrs1 locus in the proximal with multiple disease resistance genes
region of chromosome 2HL (de la Peña consumes a large portion of the resources
et al., 1999). available for two-rowed barley
improvement. The pattern using donor
Most FHB resistant selections from parents in three-way crosses, which was
crosses between Midwest six-rowed employed by Dr. Glenn Peterson, is still
cultivars and accessions related to followed. Greenhouses and off-season
Svanhals have a two-rowed spike (Urrea- nurseries have facilitated crossing and
Flórez, 2000). FHB resistant six-rowed generation advance. A modified pedigree
selections are very rare, very tall, and scheme is used to select lines with
late. This phenomenon has also been desirable agronomic traits. Seedling tests
observed in crosses to Midwest two- and off-season nurseries are used to
rowed cultivars. A close linkage between identify ones that are disease resistant.
the vrs1 locus and one of the genes for The best lines become parents for another
FHB resistance appears to cause the cycle of three-way crosses.
problem. The Vrs1.b and vrs1.a alleles in
Midwest two-rowed and six-rowed Two cycles of crossing and selection are
barley, respectively, are also closely generally adequate to recover disease
linked to a short culm gene (hcm1.a) resistant lines with acceptable agronomic
(Swenson and Wells, 1944) and one or traits. Disease resistance genes from
more early maturity genes. The FHB different sources are combined in
resistance locus is likely positioned in the subsequent breeding cycles. However,
middle of this chromosome 2HL linkage this scheme for breeding locally adapted
group. Thus, a double crossover is lines with multiple disease resistance has
required to obtain FHB resistant lines not been highly successful because it
adapted to ND. Identification of takes a long time. For acceptance by the
desirable recombinants is difficult malting and brewing industry, new
because several plant height and cultivars must also have better malt
maturity genes are segregating in crosses quality than that of the recurrent parents.
to FHB resistant accessions. Barley
cultivars from Japan and China have
Accumulating Genes for Disease Resistance in Two-Rowed Barley for North Dakota 45

Sources of disease resistant Summary


barley germplasm Since most barley grown for North
The transfer of disease resistance genes Dakota (ND) has a six-rowed spike type,
from several landraces to elite breeding two-rowed spring barley can be
materials takes a long time. Utilization of considered a new crop in ND. Drought
donor parents having multiple disease and high temperatures in western ND
resistance could shorten this time and barley diseases in eastern ND are the
requirement. As an added benefit, primary factors limiting barley
accessions with multiple disease production. A large portion of the ND
resistance might contain resistance to barley crop is used by the malting and
some minor diseases. Minor diseases brewing industry; therefore, desirable
become important because genetic malt quality parameters are needed in
control of major diseases alters both crop elite breeding material. The two-rowed
physiology and pathogen interactions. cultivars and breeding materials
Predominant pathogens can no longer developed for ND have a unique
adequately suppress less aggressive ones combination of plant height and maturity
in disease resistant cultivars. Diseases genes and can be referred to as Midwest
such as bacterial blight, powdery mildew, two-rowed barley. Multiple disease
scald, black point, and root rots may resistance was established as a breeding
need attention in the future. goal because a large number of barley
pathogens can cause losses in ND.
Many barley lines bred by Dr. Hugo E.
Accessions from many countries were
Vivar for the ICARDA-CIMMYT barley
used as sources of disease resistance
program in Mexico have multiple disease
genes, but lines from the ICARDA-
resistance. Although they are not well
CIMMYT barley improvement program
adapted to ND, they have been utilized
in Mexico have been an extremely
in the development of Midwest two-
valuable source of disease resistance
rowed barley. This was, however, more
genes.
by accident than by design. In 1988,
several ICARDA-CIMMYT barley lines
References
were introduced because barley stripe Chicaiza, O., J.D. Franckowiak, and B.J. Steffenson.
rust, Puccinia striiformis f. sp. hordei, was 1996. New sources of resistance to leaf rust in
spreading into North America (Dubin barley. In: A.E. Slinkard, G.J. Scoles, and B.G.
Rossnagel (eds.). Proc. Fifth Int. Oat Conf. &
and Stubbs, 1986). In 1990, control of Seventh Int. Barley Genet. Symp. Saskatoon.
wheat stem rust provided by the Rpg1 Univ. of Saskatchewan, Saskatoon. pp. 706-708.
gene was found to be inadequate. New de la Peña, R.C., K.P. Smith, F. Capettini, G.J.
Muehlbauer, M. Gallo-Meagher, R. Dill-Macky,
genes for stem rust resistance were D.A. Sommers, and D.C. Rasmusson. 1999.
identified in lines from the ICARDA- Quantitative trait loci associated with resistance
CIMMYT barley program (Jin et al., to Fusarium head blight and kernel
discoloration in barley. Theor. Appl. Genet.
1994). Although stripe rust never became 99:561-569.
a problem in ND and the Rpg1 gene is Dubin, H.J., and R.W. Stubbs. 1986. Epidemic
effective against current pathotypes of spread of barley stripe rust in South America.
Plant Dis. 70:141-144.
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CIMMYT barley lines helped establish
multiple disease resistance as a barley
improvement goal.
46 J.D. Franckowiak

Fetch, T.G., Jr., and B.J. Steffenson. 1994. Schwarz, P.B., H.H. Casper, and S. Beattie. 1995b.
Identification of Cochliobolus sativus isolates The fate and development of naturally
expressing differential virulence on two-row occurring Fusarium mycotoxins during
barley genotypes from North Dakota. Can. J. malting and brewing. J. Am. Soc. Brew. Chem.
Plant Pathol. 16:202-206. 53:121-127.
Franckowiak, J.D., A.E. Foster, V.D. Pederson, and Steffenson, B.J., P.M. Hayes, and A. Kleinhofs.
R.E Pyler. 1985. Registration of ‘Bowman’ 1996. Genetics of seedling and adult plant
barley. Crop Sci. 25:883. resistance to net blotch (Pyrenophora teres f.
Gill, D.D. 1970. Epidemiology of barley yellow teres) and spot blotch (Cochliobolus sativus) in
dwarf in Manitoba and effect of the virus on barley. Theor. Appl. Genet. 92:552-558.
yield of cereals. Phytopathology 60:1826-1830. Steffenson, B.J., and R.K. Webster. 1992. Pathotype
Ho, K.M., A. Tekauz, T.M. Choo, and R.A. Martin. diversity of Pyrenophora teres f. teres on barley.
1996. Genetic studies on net blotch resistance Phytopathology 82:170-177.
in a barley cross. Can. J. Plant Sci. 76:715-719. Swenson, S.P., and D.G. Wells. 1944. The linkage
Jin, Y., G.H. Cui, B.J. Steffenson, and J.D. relation of four genes in chromosome 1 of
Franckowiak. 1996. New leaf rust resistance barley. J. Am. Soc. Agron. 36:429-435.
genes in barley and their allelic and linkage Takeda, K. 1990. Selection response and parent-
relationships with other Rph genes. offspring correlation of the resistance to
Phytopathology 86:887-890. Fusarium head blight in barley. Japan J. Breed.
Jin, Y., B.J. Steffenson, and J.D. Miller. 1994. 40:91-101.
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and MCC of Puccinia graminis f. sp. tritici in disease resistance in barley. Japan J. Toxic.
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84:452-455. of and sources of pathogenic variation in
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Drechslera teres relating to changes in barley from western Canada. Can. J. Plant Pathol.
cultivars grown in Western Australia. Plant 12:141-148.
Dis. 66:655-656. Toubia-Rhame, H., and B.J. Steffenson. 1999.
Metcalfe, D.K., and V.M. Bendelow. 1981. Norbert Sources of resistance to Septoria passerinii in
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Prom, L.K., B.J. Steffenson, B. Salas, T.G. Fetch, Jr., spontaneum. p. 156-158. In: M. van Ginkel, A.
and H.H. Casper. 1997. Barley accessions McNab, and J. Krupinsky (eds.). Septoria and
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yellow dwarf. Agron. J. 51:661-664. Miles. 1990. Development of barley resistant to
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University, Fargo.
47

Collaborative Stripe Rust


Resistance Gene Mapping and
Deployment Efforts
P.M. HAYES,1 A. CASTRO,1 A.E. COREY,1 T. FILICHKIN,1 C. ROSSI,1 J.S. SANDOVAL,2
I. VALES,1 H.E. VIVAR,3 AND J. VON ZITZEWITZ1

Our collaborative stripe rust resistance integrate gene discovery and deployment.
research projects have multiple objectives.
The objective of this report is to
The first and foremost is to provide
summarize multiple areas of endeavor
agronomically-competitive, disease-
that converge on stripe rust resistance
resistant varieties to our clients, who
gene mapping and deployment: 1) stripe
range from the Saraguro Indians of
rust epidemiology, 2) quantitative vs.
Ecuador to North American malting
qualitative resistance, and 3) and linkage
barley producers. A second objective is to
mapping and QTL analysis efforts in
broaden the genetic base of our barley
barley. The results of our collaborative
germplasm via the systematic
stripe rust resistance mapping and
characterization and introgression of
germplasm improvement efforts are
unique alleles. A third objective is to
summarized in an accompanying Figure 1
contribute to a better understanding of
and Table 1. Because this stripe rust
the genetic basis of durable disease
resistant germplasm development is an
resistance in crop plants.
ongoing process, updated summaries will
Three key components of this research be maintained on the Internet at http://
are: 1) phenotyping at the ICARDA/ www.css.orst.edu/barley/orbarley/
CIMMYT facilities in Toluca, Mexico, an collab.htm.
environment in which the heritability of
disease symptom expression is
maximized; 2) genotyping with molecular
markers; and 3) accelerated germplasm
Barley Stripe Rust
advance. Initially, we mapped resistance Barley stripe rust (Puccinia striiformis f.sp.
genes in ICARDA/CIMMYT germplasm hordei) has caused serious yield losses in
and then introgressed them into North Europe, the Asian subcontinent, and the
American germplasm via marker-assisted Americas (Dubin and Stubbs, 1985; Hayes
selection. More recently, we have et al., 1996c). The disease was first
pyramided quantitative and qualitative reported in South America in 1975 and in
resistance genes and attempted to

1 Dept. of Crop and Soil Science, Oregon State University, Corvallis, Oregon, USA.
2 Instituto de Fitosanidad, Colegio de Postgraduados, Montecillo, Texcoco, Mexico.
3 ICARDA/CIMMYT Barley Program, Apdo. Postal 6-641, Mexico, D.F., Mexico, 06600.
48 P.M. Hayes et al.

the U.S. in 1991 (Marshall and Sutton, Quantitative and qualitative


1995). By 1995, it had been reported in resistance
every state of the western U.S. There is an extensive literature on the
Commercial-scale epidemics have merits of different types of resistance, and
occurred annually in California and much of the debate is phrased in the
Oregon since 1995. Environments in the context of probable durability (Johnson,
Pacific Northwest favor the disease: 1981). The terminology of disease
wheat stripe rust (Puccinia striiformis f.sp. resistance genetics does justice to the
tritici) is the most important disease complexity of the subject. The terms
of wheat in the Pacific Northwest “quantitative,” “qualitative,” “vertical,”
(Line, 1993). “horizontal,” “partial,” and “tolerance”
have precise definitions (Browning et al.,
The two f.sp. of Puccinia striiformis are, 1977). Resistant phenotypes are a
however, highly specialized, and cross- continuum ranging from the
infection is not of economic importance. hypersensitive response (HR) to a modest
Initially, only race 24 of P. striiformis f.sp. reduction in the rate of epidemic
hordei was thought to be present in the development. Throughout the continuum,
Americas (Dubin and Stubbs, 1985). there are cases representing permutations
Recently, more extensive analysis has of locus number, allele effect, race-
revealed considerable variation in specificity, stage of expression, and
pathogen isolates collected in the U.S. durability (Browning et al., 1977).
(Chen et al., 1995b; Roelfs and Huerta- Therefore, locus number, allele effects,
Espino, 1994). stage of expression, and race-specificity
Genetic resistance is the most cost- need to be defined on a case-by-case basis.
effective and environmentally In the case of cereal rusts, a large body of
appropriate technique for crop disease theory has developed regarding the risks
management. Durability of resistance is a associated with race-specific resistance
key consideration in disease resistance genes (Johnson, 1981; Parlevliet, 1983;
breeding. Unfortunately, durability can Vanderplank, 1978). There is evidence that
only be demonstrated in hindsight. pathogen virulence can evolve more
Barley germplasm developed by the quickly than plant breeders can deploy
ICARDA/CIMMYT Barley Program in single resistance genes in new varieties
Mexico allows limited symptom (Parlevliet, 1977). Accordingly, a number
development when exposed to the of alternative disease resistance breeding
spectrum of virulence encountered in strategies have been proposed and, in
field tests in South America, Mexico, and some cases, implemented. One approach
the U.S. The fact that this germplasm has is to pyramid multiple race-specific genes
remained resistant over a 15-year period into a single genotype (Huang et al., 1997;
may be grounds for describing it as McIntosh and Brown, 1997; Mundt, 1991).
“durable.” Sandoval-Islas et al. (1998)
provided additional evidence for the Another approach is to use resistance
quantitative and durable nature of the genes that do not exhibit gene-for-gene
resistance of genotypes in the ICARDA/ relationships. Distinctions between
CIMMYT program. quantitative and qualitative resistance,
adult plant and seedling resistance, and
Collaborative Stripe Rust Resistance Gene Mapping and Deployment Efforts 49

partial and complete resistance are At the same time, QTL analysis
important considerations within this procedures have facilitated dissection of
general approach to disease quantitative disease resistance (see
management. At the risk of reviews by Michelmore, 1995; Young,
oversimplification, there is empirical 1996). In some cases, a significant
evidence that non-race-specific resistance proportion of the total variance in the
genes may be more durable than race- expression of quantitative traits may be
specific single genes (Parlevliet, 1983). attributable to one locus or a few loci
(Chen et al., 1994; Hayes et al., 1996;
Despite the existence of considerable
Michelmore, 1995; Young, 1995),
quantitative resistance theory, there is
confirming classic quantitative genetic
relatively little empirical data on the
studies (summarized by Vanderplank,
inheritance and mechanism of
1978). This may be an oversimplification
quantitative resistance. Molecular tools
due to overestimation of locus effects and
have revealed some unexpected results:
underestimation of locus numbers
unsuspected complexities in some gene-
(Beavis,1998; Jansen and Stam,1994;
for-gene resistance systems and
Kaeppler, 1997; Melchinger et al., 1998;
unsuspected large-effect determinants in
Utz et al., in press; Visscher et al, 1997;
some quantitative resistance systems (see
Zeng, 1994). However, the overall picture
reviews by Michelmore, 1995; Young,
is one of converging lines of evidence
1996). If quantitative resistance genes are
supporting complexity in some
to be useful, we need to understand their
qualitative models and simplicity in
effects, interactions, and relationships
some quantitative models.
with genes determining other
economically important, quantitatively Differentiation of qualitative versus
inherited phenotypes. quantitative resistance has long been a
source of controversy. At one extreme is
Characterization of plant resistance genes
the view that these classifications
at the molecular level has provided
represent genes with distinctly different
information upon which to develop
mechanisms and race specificity
models involving signal detection, signal
(Vanderplank, 1968, 1978). At the other
transduction, and response (Beynon,
extreme is the view that all resistance
1997). These studies (Buschges et al.,
genes are similar, but are merely
1997; Martin et al., 1993; Salmeron et al.,
expressed differently in different
1994; Schulze-Lefert, 1997; Zhou, 1995)
combinations and in different genetic
have provided molecular evidence
backgrounds (Nelson, 1978).
confirming hypotheses based on whole
Unfortunately, three decades of debate
plant data (summarized by Ellingboe,
have failed to resolve the issue.
1976; Gabriel and Rolfe, 1995) indicating
that “monogenic” gene-for-gene Wang et al. (1994) used recombinant
relationships are recognition processes inbred lines of rice (Oryza sativa) to show
that turn on multiple genes in a that a cultivar with durable resistance
resistance pathway. (sensu Johnson, 1981) to rice blast (caused
by Magnaporthe grisea) contains two race-
specific, qualitative genes for resistance
and ten QTLs contributing to partial
50 P.M. Hayes et al.

resistance (sensu Parlevliet, 1989). Qi et Research Institute (SCRI) has a very


al. (1999) have presented evidence for productive SSR development program
race-specific QTLs and argued that (http://www.scri.sari.ac.uk/SSR/). We
quantitative resistance to leaf rust is an are currently cooperating with the SCRI
example of minor gene-for-minor gene in an international barley SSR
interaction. However, the contribution of characterization effort and are
the qualitative vs. quantitative genes to systematically mapping the SCRI SSRs
resistance remains unclear. on NABGMP populations (Toojinda et
al., 2000).

Linkage maps are useful from the


Barley Linkage Maps standpoint of understanding genome
and QTLs organization, establishing synteny as a
platform for map-based cloning, and for
Barley is an excellent system for genome QTL detection. For a review of QTL
mapping and map-based analyses. This detected in barley with references and
diploid (2n = 14) species has seven links, see Hayes et al., 1996; GrainGenes
cytologically distinct chromosomes (http://wheat.pw.usda.gov/
containing approximately 5.3 x 109 bp graingenes.html); and the NABGMP
DNA (Bennett and Smith, 1976). home pages (http://www.css.orst.edu/
Although barley is an autogamous barley/nabgmp/nabgmp.htm; http://
species, there is sufficient DNA-level gnome.agrenv.mcgill.ca). In barley, as in
diversity for efficient linkage map other crop species, much of the activity
construction in populations derived from in QTL mapping has been descriptive.
crosses between related genotypes The experiments required for validation
(Graner et al., 1991; Kleinhofs et al., 1993; of estimates of QTL number, effect, and
Kasha et al., 1995; Becker et al., 1995; interaction are just coming to fruition.
Hayes et al., 1997). The North American
Barley Genome Mapping Project Larson et al. (1996), Romagosa et al.
(NABGMP) has focused on building (1996), Spaner et al. (1999) and Zhu et al.
maps in elite germplasm to facilitate the (1999) have conducted marker-assisted
direct application of these maps to plant selection experiments to verify QTLs for
breeding (reviewed by Hayes et al., agronomic traits in barley. Han et al.
1996). Several thousand loci have been (1997) and Marquez-Cedillo et al. (in
placed on these maps, providing a press) have conducted similar
comprehensive catalog of markers. experiments for malting quality traits. In
all cases, marker-assisted selection was
Higher throughput markers, such as effective for some, but not all QTLs. For
AFLPs, have been used for barley map stripe rust, we have successfully
construction (Becker et al., 1995; Hayes et introgressed resistance QTL alleles into a
al. l997). Microsatellite polymorphism susceptible genotype (Toojinda et al.
has been demonstrated (Saghai-Maroof 1998). The limited population sizes used
et al., 1994) and used for barley in many of the reported QTL detection
germplasm characterization and map experiments may have led to
construction (Powell et al., 1996; Russell underestimation of QTL number,
et al., 1997; Becker and Heun, 1995; overestimation of QTL effects, and a
Toojinda et al., 2000). The Scottish Crop
Collaborative Stripe Rust Resistance Gene Mapping and Deployment Efforts 51

failure to quantify QTL interactions QTL to the same region on chromosome


(Beavis, 1998; Jansen and Stam, 1994; 5 (1H) in the winter six-row variety Kold
Kaeppler, 1997; Melchinger et al. 1998; and a resistance QTL on chromosome 7
Utz et al., in press; Visscher et al, 1997; (5H) (at the same position as the
Zeng, 1994). chromosome 7 (5H) QTL in Cali-sib), in
the CIMMYT/ICARDA spring two-row
Mapping and deployment germplasm CMB643 (Hayes et al., 1999).
of stripe rust resistance These stripe rust resistance mapping
genes in barley efforts are summarized in the
Four qualitative resistance genes, Yr1 - accompanying Table 1.
Yr4, were described by Lehmann et al.
(1975). Of these genes, only the Yr4 Deployment of resistance genes has
locus has been mapped, and it is on the involved marker-assisted selection in
short arm of chromosome 5 (1H) various germplasm advanced
(vonWettstein-Knowles, 1992). We strategies—backcrossing in the case of
recently mapped a qualitative resistance the six-row variety “Tango” (Toojinda et
gene to the long arm of chromosome 1 al., 1998)—and, more recently,
(7H) in CI10587 (Hayes et al., 1999), but pyramiding. Pyramiding efforts, focused
the identity of this gene relative to the on two-row barley, are summarized in
Yr1 - Yr3 genes remains to be the accompanying Figure 1. The first step
established. We mapped resistance in construction of resistance gene
QTLs on chromosomes 4 (4H) and 7 pyramids involved positioning the
(5H) in Calicuchima-sib (Cali-sib), an resistance QTL alleles from Shyri and
ICARDA/CIMMYT germplasm line Cali-sib in a malting quality background
(Chen et al.1994; Hayes et al. 1996c). We contributed by Harrington and Galena.
also mapped a major adult plant stripe These resistance QTL allele pyramid lines
rust resistance QTL on the short arm of constitute the “BCD” population. The
chromosome 5 (1H) in the ICARDA/ acronym has local athletic allusions and
CIMMYT-derived variety Shyri and stands for “Beavers conquer Ducks”.
smaller-effect QTLs on chromosomes 2 This germplasm has been extensively
(2H), 3 (3H), and 6 (6H) (Toojinda et al., phenotyped, and its allelic structure at
2000). At the level of resolution afforded the target stripe rust resistance QTL is
by the available maps, the chromosome currently being defined (Castro et al.,
5 (1H) QTL coincides with the position 2000).
of the Yr4 locus. Yr4 is reported to The second step in the construction of the
confer resistance to race 23 (von resistance gene pyramids involved
Wettstein-Knowles, 1992) while the adding a qualitative resistance gene,
virulence spectrum in the Americas is contributed by CI10587. Because CI10587
described in terms of race 24 and its is agronomically disadvantaged, the
variants (Chen et al., 1995b). qualitative resistance gene was
Thomas et al. (1995) also mapped an transferred to Baronesse, the leading feed
adult plant resistance QTL in the same variety in the Pacific Northwest of the
region in the variety Blenheim and U.S. The quantitative/qualitative
hypothesized that it was an effect of an resistance gene pyramids are referred to
allele at the Yr4 locus. We mapped a as the “Ajo, Sal, Bu, and Ops”
52 P.M. Hayes et al.

Table 1. Summary of OSU/ICARDA/CIMMYT collaborative disease resistance mapping and


germplasm enhancement projects.†

Two-row populations. Phase I


Cross: CB. Calicuchima-sib/Bowman (Yd2). AKA: ‘BSR’
Parents: Cali-sib: Stripe rust, leaf rust, scald
Bowman (Yd2): Stress tolerance, BYDV
Population: F1-derived doubled haploid. N = 94. Stripe rust, leaf rust, scald, BYDV
Mapped genes: Stripe rust, leaf rust, scald, BYDV
Selections: BSR45 (2-row) = Orca. Stripe rust, scald, BYDV BSR41 (6-row). Stripe rust, scald
References: Chen et al., 1994; Hayes et al., 1996; Hayes et al., 2000
Comments: Also mapped malting quality QTL Orca released in 1998 as a feed variety.
Not recommended as a malting variety due to high protein and enzymes.

Cross: SG. Shyri/Galena. AKA: ‘D1’


Parents: Shyri: Stripe rust, leaf rust, scald, net blotch, BYDV, FHB
Galena: Malt quality, leaf rust, BYDV
Population: F1-derived doubled haploid. N = 94. Stripe rust, leaf rust, scald, net blotch, BYDV
Mapped genes: Stripe rust, leaf rust, BYDV
Selections: D1-72: Stripe rust
References: Toojinda et al., 2000
Comments: Scald and net blotch mapping in progress

Cross: CG. CI10587/Galena. AKA: ‘D3’


Parents: CI10587: Stripe rust, leaf rust, Russian wheat aphid
Galena: Malt quality, leaf rust, BYDV
Population: F1-derived doubled haploid. N = 94. Stripe rust, RWA
Mapped genes: Stripe rust
Selections: D3-6: Stripe rust, RWA
References: In progress
Comments: RWA mapping deferred until full map available

Cross: DB. D3-6/Baronesse. AKA: D3-6/B


Parents: D3-6: Stripe rust, RWA
Baronesse: Yield
Population: F1-derived doubled haploid. N = 100. Stripe rust, RWA
Mapped genes: NA
Selections: D3-6/B-23 Stripe rust, RWA
D3-6/B-45 Stripe rust, RWA
D3-6/B-61 Stripe rust, RWA
References: NA
Comments: CI10587 is agronomically challenged. D3-6 was the most prepossessing line.
We crossed it with Baronesse to improve agronomic type. Stripe rust and RWA resistance
phenotypes were confirmed in the D3-6/B progeny.

Cross: GA. Gobernadora/Azafran. AKA: Gobe x CMB643, Gobe


Parents: Gobernadora: Fusarium
Azafran: Fusarium, stripe rust, leaf rust
Population: F1-derived doubled haploid. N = 144. Fusarium (I,II,III), stripe rust
Mapped genes: Fusarium (I,II, III), stripe rust
Selections: Gobe 24: Fusarium, stripe rust
Gobe 96: Fusarium, stripe rust, scald
References: Zhu et al., 1999b
Comments: Stripe rust QTL not published. Maps to same position on chromosome 7 (5H) as QTL in
Cali-sib.
Collaborative Stripe Rust Resistance Gene Mapping and Deployment Efforts 53

Two-row populations: Phase II


Cross: BCD. Orca/1*Harrington//D1-72 AKA: ‘BCD’
Parents: Orca: Stripe rust, scald, BYDV
Harrington: Malting quality
D1-72: Stripe rust
Population: BC1-derived doubled haploid. N = 115. Stripe rust, scald, BYDV
Mapped genes: Stripe rust
Selections: BCD-12: Stripe rust, leaf rust, scald, BYDV
BCD-47: Stripe rust, leaf rust, scald
References: In progress
Comments: Objective of this population is to pyramid stripe rust resistance QTL alleles tracing to
Cali-sib and Shyri. BCD-47 is in regional testing and the AMBA pilot program.

Cross: RECLA. AKA: Red Cebada Latina


A complex cross involving:
Gobe-24; Gobe-96; AF9216; Orca; Kredit; and Azafran
Parents: Gobe-24: Fusarium
Gobe-96: Fusarium, stripe rust, leaf rust
AF9216: Leaf rust
Orca: Stripe rust, scald, BYDV
Kredit: Stripe rust, leaf rust
Azafran: Fusarium, stripe rust, leaf rust
Population: DH = 125
Mapped genes: NA
Selections: Lines advanced to breeding program
References: http://www.css.orst.edu/recla/la_red.htm
Comments: Objective of this population is to pyramid multiple resistance genes and to stimulate a
Latin American Barely initiative. Parents selected for adaptation to various environments
in Latin America.

Two-row populations: Phase III


Cross: Bu. BCD-47//D3-6/B-23
Parents: BCD-47: Stripe rust, leaf rust, scald
D3-6/B-23: Stripe rust, RWA
Population: F1-derived doubled haploid. N = 89. Stripe rust, scald, BYDV
Mapped genes: In progress
Selections: Lines advanced to breeding program
References: NA
Comments: Objective of this population is to pyramid stripe rust resistance QTL alleles tracing to
Cali-sib and Shyri with the single gene tracing to CI10587.

Cross: Sal. BCD-47//D3-6/B-45


Parents: BCD-47: Stripe rust, leaf rust, scald
D3-6/B-45: Stripe rust, RWA
Population: F1-derived doubled haploid. N = 11. Stripe rust, scald, BYDV
Mapped genes: In progress
Selections: Lines advanced to breeding program
References: NA
Comments: Objective of this population is to pyramid stripe rust resistance QTL alleles tracing to
Cali-sib and Shyri with the single gene tracing to CI10587.
54 P.M. Hayes et al.

Cross: Ajo BCD-47//D3-6/B-61


Parents: BCD-47: Stripe rust, leaf rust, scald
D3-6/B-61: Stripe rust, RWA
Population: F1-derived doubled haploid. N = 119. Stripe rust, scald, BYDV
Mapped genes: In progress
Selections: Lines advanced to breeding program
References: NA
Comments: Objective of this population is to pyramid stripe rust resistance QTL alleles tracing to Cali-sib
and Shyri with the single gene tracing to CI10587.

Cross: Ops. BCD-12//D3-6/B-61


Parents: BCD-47: Stripe rust, leaf rust, scald
D3-6/B-61: Stripe rust, RWA
Population: F1-derived doubled haploid. N = 108. Stripe rust, scald, BYDV
Mapped genes: In progress
Selections: Lines advanced to breeding program
References: NA
Comments: Objective of this population is to pyramid stripe rust resistance QTL alleles tracing to
Cali-sib and Shyri with the single gene tracing to CI10587.

Two-row populations: Phase IV


Cross: Bub. BCD-47//D3-6/B-23 F1///BCD-47
Parents: BCD-47: Stripe rust, leaf rust, scald
D3-6/B-23: Stripe rust, RWA
Population: SSD N = 130
Mapped genes: NA
Selections: NA
References: NA
Comments: Objective of this population is to pyramid stripe rust resistance QTL alleles tracing to Cali-sib
and Shyri with the single gene tracing to CI10587.

Cross: Buh. BCD-47/D3-6/B-23, F1///He6890


Parents: BCD-47: Stripe rust, leaf rust, scald
D3-6/B-23: Stripe rust, RWA
He6890: Stripe rust, Czech malting barley
Population: SSD N = 160
Mapped genes: NA
Selections: NA
References: NA
Comments: Objective of this population is to pyramid stripe rust resistance QTL alleles tracing to
Cali-sib and Shyri with the single gene tracing to CI10587 and the uncharacterized
resistance in He6890.

Cross: Opb. BCD-12/D3-6/B-61, F1///BCD-47


Parents: BCD-12: Stripe rust, leaf rust, scald, BYDV
D3-6/B-61: Stripe rust, RWA
BCD-47: Stripe rust, leaf rust, scald
Population: SSD N = 149
Mapped genes: NA
Selections: NA
References: NA
Comments: Objective of this population is to pyramid stripe rust resistance QTL alleles tracing to Cali-sib
and Shyri with the single gene tracing to CI10587.
Collaborative Stripe Rust Resistance Gene Mapping and Deployment Efforts 55

Cross: Oph. BCD-12/D3-6/B-61, F1///He6890


Parents: BCD-12: Stripe rust, leaf rust, scald, BYDV
D3-6/B-61: Stripe rust, RWA
He6890: Stripe rust, Czech malting barley
Population: SSD N = 130
Mapped genes: NA
Selections: NA
References: NA
Comments: Objective of this population is to pyramid stripe rust resistance QTL alleles tracing to
Cali-sib and Shyri with the single gene tracing to CI10587 CI10587 and the uncharacterized
resistance in He6890.

Cross: Ajb. BCD-47/D3-6/B-61, F1///BCD47


Parents: BCD-47: Stripe rust, leaf rust, scald
D3-6/B-61: Stripe rust, RWA
BCD-47: Stripe rust, leaf rust, scald
Population: SSD N = 149
Mapped genes: NA
Selections: NA
References: NA
Comments: Objective of this population is to pyramid stripe rust resistance QTL alleles tracing to Cali-sib
and Shyri with the single gene tracing to CI10587.

Cross: Ajh. BCD-47/D3-6/B61, F1///He6890


Parents: BCD-47: Stripe rust, lLeaf rust, scald
D3-6/B-61: Stripe rust, RWA
He6890: Stripe rust, Czech malting barley
Population: SSD N = 169
Mapped genes: NA
Selections: NA
References: NA
Comments: Objective of this population is to pyramid stripe rust resistance QTL alleles tracing to
Cali-sib and Shyri with the single gene tracing to CI10587 and the uncharacterized
` resistance in He6890.

Cross: Sah. BCD-47/D3-6/B-45, F1///He6890


Parents: BCD-47: Stripe rust, leaf rust, scald
D3-6/B-45: Stripe rust, RWA
He6890: Stripe rust, Czech malting barley
Population: SSD N = 94
Mapped genes: NA
Selections: NA
References: NA
Comments: Objective of this population is to pyramid stripe rust resistance QTL alleles tracing to
Cali-sib and Shyri with the single gene tracing to CI10587 and the uncharacterized
resistance in He6890.
56 P.M. Hayes et al.

Two-row populations: Phase V


Cross: BAR12. BCD-12/Baronesse
Parents: BCD12: Stripe rust, leaf rust, scald, BYDV
Baronesse: Yield
Population: In progress. Target n = 500 single seed descent
Mapped genes: NA
Selections: NA
References: NA
Comments: Objective of this population is to optimize estimates of resistance gene effects and measure
resistance gene interactions.

Cross: BAR47. BCD-47/Baronesse


Parents: BCD47: Stripe rust, leaf rust, scald
Baronesse: Yield
Population: In progress. Target n = 500 doubled haploids
Mapped genes: NA
Selections: NA
References: NA
Comments: Objective of this population is to optimize estimates of resistance gene effects and measure
resistance gene interactions.
† Cross: Abbreviation in bold corresponds to Figure.
Parents: Parents contribute favorable alleles for phenotypes listed.
Population: Population assayed for phenotypes listed.
Mapped genes: Phenotypes measured in the population for which genetic determinants have been mapped.
Selections: Correspond to designation in Figure. Selections contribute favorable alleles for phenotypes listed.

Two-row Six-row

Orca
CB SG CG GA GA

BSR-45 BSR-41 DI-72 D3-6 Gobe-24 Gobe-96

Harrington Baronesse Colter Steptoe

BCD DB Tango SR CR
BCD-12 BCD-47 D3-6/B-23 D3-6/B-45 D3-6/B-61 SR58-4 CR30-3
CRSR

OPS BU SAL AJO Excel Stander

He6890

OPH BUH BUB SAH AJH AJB SOE TSE SOT SOST

BAR-12 BAR-47 RECLA

Mapping population Selection from mapping population


Variety selected from Variety used as parent
mapping population

Figure 1. Construction of resistance gene pyramids in two- and six-row barley using marker-
assisted selection.
Collaborative Stripe Rust Resistance Gene Mapping and Deployment Efforts 57

germplasm. These germplasm descriptors Faced with a field nursery of


have gastronomic allusions; the agronomically promising germplasm, all
germplasm itself has been extensively of which is phenotypically resistant to
phenotyped. Genotyping is in progress. diseases, knowledge regarding the
genetic architecture of each germplasm
The third step in pyramid construction
accession can be invaluable in deciding
has involved He6890, an agronomically
which genotypes to advance as varieties,
attractive Czech selection with
and which genotypes to use as parents
uncharacterized stripe rust resistance.
in order to continue the allele
Our stripe rust resistance breeding effort
accumulation process. Knowledge
in six-row barley is not as advanced as
regarding the genetic architecture of
our effort in two-row barley. As shown in
exotic germplasm should increase its
the accompanying Figure, the six-row
utility, ensuring a broader germplasm
effort is currently directed deploying
base and providing an impetus for
resistance QTL alleles, tracing to Cali-sib,
germplasm conservation.
in a Midwestern malting quality
background.

Acknowledgments
Conclusions This collaboration has succeeded thanks
to the efforts of numerous individuals
In summary, the ICARDA/CIMMYT
and to support from a number of
program has been very successful in
organizations. Special thanks to all who
accumulating resistance to multiple
have contributed to this research over
diseases in single genotypes. Although
the years, including Fuqiang Chen,
this review has focused on stripe rust, this
Xianming Chen, Jeanine DeNoma,
germplasm is also rich in genes
Aihong Pan, Mareike Johnston, John
conferring resistance to a range of
Korte, Rollie Line, Vicente Morales,
diseases including barley yellow dwarf,
Chris Mundt, Doris Prehn, Bernardo
leaf rust, net blotch, scald, and spot
Ramirez, and Theerayut Toojinda. For
blotch. This accumulation of disease
the OSU participants, this research was
resistance alleles has been accomplished
made possible by the barley growers of
based on phenotypic data alone, and this
Oregon, Washington, and Idaho; the
success is testimony to Dr. Hugo Vivar’s
American Malting Barley Association;
sharp eye and powers of recollection, and
the Anheuser Busch Company; Busch
to the dedication of his staff. The
Agricultural Resource, Inc.; Great
genotypic characterization of this
Western Malting/ConAgra Malt; the
germplasm should be of assistance to all
North American Barley Genome
participants.
Mapping Project; and the Oregon
Agricultural Experiment Station. The
Colegio de Postgraduados and
ICARDA/CIMMYT provided support
for their participating staff.
58 P.M. Hayes et al.

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61

Perspectives on Fusarium Head


Blight Resistance in Barley
L. GILCHRIST1

Global Importance and North America


The USA and Canada. Since 1993, scab and
Distribution
vomitoxins have affected barley crops in
Fusarium head blight (FHB), or scab, has the Northern Great Plains, including
economic impact in barley-producing North Dakota, Minnesota, and Canada.
areas all over the world. It reduces yield From 1993 to 1997, North Dakota farmers
and affects quality due to the toxin it suffered cumulative losses due to scab and
produces in the grain. vomitoxin estimated at US$200 million
(Anonymous, 1999).
Asia
Surveys of US regional crops have shown
Among Asian countries, China has the
DON to be the primary mycotoxin
most extensive scab-affected area in its
associated with infected barley, although
barley growing regions. The disease
ZEA and other fusarium toxins have also
occurs mainly in the lower Yangtze River
been detected. It is estimated that 67 and
Basin (100,000 ha), where the humidity is
82% of the malting barley crop have been
high. Scab has also appeared in the
contaminated with DON (0.6-60mg/g)
Heilongjiang Province in recent years
during 1993 (Schwartz, 1995).
(Sun et al., 1999). Scab may cause 20-50%
yield losses in an epidemic year (Chen et Scab is very common in the highlands of
al., 1991). central Mexico (the states of Mexico,
Tlaxcala, Hidalgo, Puebla, and Jalisco). In
Barley is produced in four provinces of
a first survey conducted in the states of
Korea (Chonbuck, Chonnam,
Puebla and Tlaxcala in 1999, 33 samples of
Kyungbuck, and Kyunnam). The natural
malting barley were collected and the
occurrence of fusarium mycotoxins was
following Fusarium species detected: F.
surveyed in 39 barley samples collected
semitectum, F. avenaceum, F. sabucinum, F.
in those provinces. Five toxin compounds
poae, F. graminearum, F. moniliforme, F.
were detected: deoxynivalenol (DON), dimerum, F. equiseti, and F. subglutinans. Of
nivalenol (NIV), 4-acetylnivalenol (4- these, F. sabucinum and F. dimerum are not
ANIV), 3-acetildeoxynivalenol (3-DON), reported in the literature. However, in this
4,15-diacetynivalenol (4,15-DANIV), and study, these two species were isolated
zearalenone (ZEA). DON, NIV, and ZEA from grains, and from root and stem
were the major contaminants (Kim, 1993). lesions in blotter and germination tests
(Gutierrez, 2000).

1 CIMMYT Wheat Program, Apdo. Postal 6-641, Mexico, D.F., Mexico, 06600.
62 L. Gilchrist

South America Table 1. Pathogenic Fusarium species reported


Fusarium head blight is very important on barley by country.
in the Southern Cone and the Andean
Fusarium Country†
Region, including the Ecuadorian and species Canada USA Mexico Japan Poland
Peruvian highlands, where barley is used
for feed and food purposes. In Uruguay, F. graminearum + + + + +
F. culmorum + + + +
Brazil, and Argentina, barley is used
F. avenaceum + + + + +
mainly for malting and feed (Vivar, 1997). F. poae + + + +
Incipient damage has been detected in F. sporotrichum + + + +
southern Chile (regions IX and X) around F. equiseti + + + + +
lakes and rivers, and wherever maize has F. acuminatum + + +
F. moniliforme + +
been introduced into the crop rotation for F. semitectum + +
silage production (Mellado, 1999; von F. nivale +
Baer, 1999, pers. comm.). F. tricinctum +

In Uruguay barley was evaluated over a † Gordon, 1959; Clear et al., 1996.
Mihuta-Grimm, 1989; Salas et al., 1999.
period of four years (1993-97) for the Gutierrez, 2000.
natural occurrence of fusarium toxins. Koizumi et al., 1995; Takeda et al., 1995.
DON and Fumonosine B1 (FB1) were Perkowski et al., 1996.
predominant. Barley was second to
wheat in severity of infection. ZEA levels
were considerable only in barley and FHB symptoms and interaction
mixed feed, drawing attention to its use with other pathogens
for animal consumption and its potential The first symptom of scab infection is a
deleterious estrogenic effects. Locally, small, water-soaked, somewhat
barley feed is used extensively in animal brownish or pinkish brown spot at the
nutrition. The 1993-94 crop season was base or in the middle of the glumes or on
the first in the last decade to show severe the rachis. Water-soaking and
fusarium damage (Piñeiro and Silva, discoloration spread in all directions
1997). from the point of infection and can cover
the grain partially or completely. If
A comparative analysis of F. graminearum conditions are favorable, spread is
isolates from Canada, USA, Mexico, evident in adjacent grains. If the tissue
Argentina, and Uruguay confirmed that takes on a brown color, the symptoms
they belong to chemotype IB. There is a may be confused with those produced
regional relationship between the origin by other pathogens such as Bipolaris
of F. graminearum and the production of 3 sorokiniana, Rynchosporium secalis,
or 15 Ac DON as the major isomer Pyrenophora teres, and Alternaria spp.,
(Piñeiro et al., 1996). and saprophytes such as Diccocum spp.
Field diagnoses can be difficult to make,
and a laboratory analysis is
Causal Agents recommended in specific cases.
Most countries recorded F. graminearum
If plants are affected by other diseases
as the most important scab causing
like the rusts and BYDV before
pathogen. However, a broad range of
inoculation, inoculation is not
Fusarium species was reported to cause
the disease (Table 1).
Perspectives on Fusarium Head Blight Resistance in Barley 63

recommended, nor should the Today there is evidence that this model
germplasm even be evaluated. Resistant is also applicable to barley. Field data
germplasm may show a susceptible presented in Table 2 for Gobernadora
reaction and produce very brownish, doubled haploid populations and QTLs
shriveled grain. One clear example of this confirmed that Type I resistance is
is the interaction observed during the last independent of Type II, with Type II
two years in Toluca, Mexico (no stripe having the largest QTL detected near the
rust in 1998 and a heavy stripe rust centromeric region of chromosome 2
epidemic in 1999), where scab resistant (Zhu et al., 1999).
advanced lines from a breeding program
in Minnesota, which are very susceptible
to Puccinia striiformis f. sp. hordei (race 24), Table 2. Relative ranking of five doubled
showed a resistant reaction to fusarium in haploid lines (out of 100) and both parents for
resistance to head blight (Fusarium
1998 and a susceptible one in 1999.
graminearum) against fungal penetration (Type
I) and hyphae spread (Type II), or a combination
FHB resistance mechanisms of both. Atizapan Station, Toluca, Mexico, 1996.
Understanding the different types of
resistance mechanisms and their Cultivar Type I Type II
epidemiological role is necessary for DH-83 5 71
breeding head scab resistance. Key points DH-96 8 2
are to use specific inoculation, screening, DH-98 9 87
and evaluation methods for each DH-52 50 1
DH-89 98 25
resistance mechanism. This is important Gobernadora (Zhenmai 1) 1 6
when working with a virulent pathogen CMB 643 (Azafran) 45 5
population under extreme environmental
conditions. As Parry et al., (1995)
reported, “Differences in resistance Snijder and Perkowski (1990) found a
between cultivars could be masked, with negative correlation in wheat between
even the most resistant cultivars head blight and incubation period, given
becoming infected under extreme that the more resistant genotypes had
conditions.” longer incubation periods. This
correlation makes the incubation period
The model proposed by Schroeder and a potentially useful criterion for selecting
Christensen for wheat in 1963, which for resistance.
suggested the existence of two
components of resistance (Type I and Type III resistance was described and
Type II), has been accepted by most proved to occur in wheat by a team of
authors. Type I resistance operates Canadian researchers (Miller and
against initial infection and Type II Arnison, 1986). It was proposed that
against the spread of the pathogen within resistant cultivars possessed a factor that
the host. Type I and Type II resistances enabled them either to prevent DON
varied independently among cultivars. synthesis or to promote its degradation.
The resistant cultivar Frontana was able
to degrade 18% radioactive (C14) DON
into two breakdown products, while the
64 L. Gilchrist

susceptible cultivar Casavant degraded Influence of toxins on


only 5%. Snijder and Krechting (1992), pathogenesis and virulence
also working in wheat, indicated that factors
DON played a part in pathogenicity, as Inoculum composition. Today we know
they found it to be transported from that different F. graminearum isolates have
fusarium infected chaff to the young different toxigenic capacities. For this
kernel, which the pathogen later reason, it is absolutely necessary that
colonized. The phytotoxic effect of DON when inoculating for different types of
can be explained by the fact that it is a scab resistance, both inoculum
very potent inhibitor of eukaryotic composition and the conditions and
protein synthesis (Wang and Miller, timing of inoculation be controlled. For
1988). In a resistant wheat line, DON example, the phytotoxicity of some
transport was inhibited, so colonization trichothecenes has been demonstrated
was reduced. (Miller and Arnison,1986; Wong et al.,
1994). A USDA team created a genetically
To lower DON concentration in the grain modified strain of F. graminearum GzT40
is the aim of most barley breeders. that does not produce trichothecene
Steffenson (1998) mentions that there is a because it does not carry gene Tri 5-,
fairly positive correlation (r=0.64; which confers virulence (Proctor et al.,
P=0.0001) between FHB incidence and 1995). Eudes et al., (1998) inoculated 17
DON concentration in barley grain. wheat varieties with specific inocula of
Skadhauge et al., (1997) discovered a GzT40 and of its wild parent Gz3639 in a
proanthocyanidin-free mutant of barley controlled environment (Figure 1). The
that shows extreme resistance to strain possessing toxigenic capacity
fusarium in vitro. This resistance is due (Gz3639) was virulent and produced host
to the accumulation of dihydroquercitin, damage depending on the resistance
a potent inhibitor of fusarium growth. present in each variety. This suggests it is
essential that the composition of the
Type IV resistance was described as inoculum used as well as the conditions
tolerance to high DON concentrations by during inoculation be controlled within
Wang and Miller in 1988. They reported the same year and across years, to avoid
that some cultivars could tolerate high variation and incorrect interpretations.
mycotoxin concentrations with no
negative effects on growth. Ma et al., A barley example is presented in Table 3,
(1999) conducted a preliminary study where two varieties [Robust (MS) and
where they injected pure DON solution Chevron (MR)] were inoculated with
(100 ppm) 10 cm below the spike in three nine different fusarium isolates (Evans et
different varieties [Sumai 3 (R), Norm al., 1996). The reaction of the two
(S), and Pioneer 2375(MS)]. They found varieties differed depending on the
that tolerance to DON may be inoculated strain.
independent of Type II reaction since Inoculum concentration. According to
Pioneer 2375 (MS type II) had lower studies carried out in Toluca during
average dry kernel weight. This type of 1980-82 (G. Bekele, pers. comm.) on
research has not been done in barley. wheat, no differences were detected
using 30,000-70,000 spores per ml. This
Perspectives on Fusarium Head Blight Resistance in Barley 65

S Norseman
Mondor
Max
Messier
Laval-19 Concorde

Veerys

Katepwa
QW 550.13
Voyageur
Toropi
Frontana Strain Gz3639
Trichothecene+
Ning 8331
Key 94/28 Strain Gz3T40
Key 94/34 Nanjing 7840
Trichothecene+
Nyu-Bay
R

0.0 0.15 0.30 0.45 0.60 0.75 0.90


Number of florets showing symptoms in greenhouse

Figure 1. Virulence of the Fusarium graminearum genetically modified GzT40 strain (Tri 5-gene) and
the Gz3639 wild strain (Tri 5-gene) on 17 wheat cultivars.
Source: Eudes et al. (1998).

Table 3. Number of infected spikelets on barley Inoculation methods for


varieties Robust (moderately susceptible) and evaluating different types of
Chevron (moderately resistant). FHB resistance
Robust Chevron At CIMMYT different plots are used to
Isolate Rep I Rep II Rep I Rep II screen for each FHB resistance type.
2B-2A1 17 25 12 26 Type I. Wheat inoculation is carried out
3A-2A1 3 11 0 1
in the evening with a hand sprayer
3A-4A1 18 23 21 16
4B-5A2 10 1 15 3 (Bekele, 1984). A spore suspension is
5C-3A2 4 6 4 4 sprayed on 15 spikes that were selected
6A-2A1 13 3 6 5 and marked at the initial anthesis
Butte 86-ADA-11 24 11 5 12
growth stage. After 10-15 days spikes
M66-ADA-B1-A1 3 1 13 1
94RO/STE 4 9 17 25 21 are evaluated for penetration points.
Control 0 0 0 0 The precise timing of the evaluation
Source: Evans et al. (1996).
will depend on the reaction of resistant
and susceptible checks to the
environment that particular year.
was confirmed in barley at the same
location during the 1996 cycle. Based on Barley spikes can be infected by
this information, a suspension of 50,000 Fusarium species as soon as they emerge
spores per ml is recommended for use in from the flag leaf sheath, and they
inoculation. remain susceptible throughout the
66 L. Gilchrist

grain filling period. Spike growth stage of fungicide (Folicur plus) every 10 days.
resistant cultivars at inoculation affects At harvest, a 200-g sample is used for
the amount of damage observed in a DON analysis (Type III), and a
particular year (Tables 4 and 5). comparison of 300-kernel weight is
done between the two plots (Type IV).
Type II. Inoculation using the cotton
method (Bekele, 1984) is done on 20
barley spikes at the initial anthesis
growth stage. Spikes are evaluated 25-30 Resistance Sources from
days after inoculation. As with Type I All Over the World
resistance, the precise timing of the
evaluation will depend on the reaction of Nearly 40 barley genotypes have been
resistant and susceptible checks to the identified as having partial FHB
environment that particular year. resistance and low DON accumulation.
The genetic distance between
Types III and IV. A small plot (two 50-cm genotypes was calculated separately
rows) is sprayed with a spore suspension using morphological, agronomic, FHB
when 50% of barley spikes have reached severity and DON content, and foliar
anthesis. A similar plot is sprayed with disease data. Five clusters were

Table 4. Inoculation using two methods (spray Table 5. Inoculation using two methods (spray
and cotton) at three spike growth stages (pre- and cotton) at three spike growth stages (pre-
anthesis, initial anthesis, and post-anthesis) and anthesis, initial anthesis, and post-anthesis) and
the damage produced in barley varieties. the damage produced in barley cultivars.
Atizapan, Toluca, 1998. Atizapan, Toluca, 1999.

Spike Spike
growth Spray Cotton growth Spray Cotton
Variety stage† method method Variety stage† method method

Shyri PRA 6.3 (3.6-11.0)ai 39.6 (31-48.2) k Shyri PRA 5.0 (2.9-8.2) cl 9.3 (5.9-14.2) ag
IA 4.9 (2.6-9.1)af 26.1 (19.6-33.7)hk IA 7.0 (4.4-10.9) gn 10.5 (6.9-15.7) ag
PA 1.7 (0.6-4.9)ac 21.6 (15.7-29.0) fj PA 2.4 (1.2-4.7) bg 4.4 (2.4-8.0) a

Arupo/K8755// PRA 7.9 (4.7-13.3)cl 23.1 (16.3-31.8)fk Arupo/K8755// PRA 6.1 (3.8-9.6) fm 34.3 (27.8-41.6) ln
Mora (Selec 2) IA 6.6 (3.8-11.3)ak 11.7 (7.2-18.4)cg Mora (Selec 2) IA 3.7 (2.1-6.5) bk 38.7 (32.1-45.7) n
PA 9.0 (5.6-14.2)dm 3.9 (1.7-8.8)ac PA 0.4 (0.1-2.2) b 24.5 (18.9-31.2)lm

Gob/Humai 10 PRA 11.1 (7.3-16.5)fo 29.1 (22.1-37.4)ik Gob/Humai 10 PRA 1.6 (0.6-4.3) bf 14.7 (10.5-20.3) ei
IA 9.5 (6.1-14.6)dm 0.3 (6.3-16.3)cf IA 1.2 (0.4-3.2) be 6.5 (3.8-10.9) ae
PA 7.0 (4.1-11.6)ck 19.1 (13.4-26.5)ej PA 2.3 (1.2-4.7) bg 4.3 (2.3-7.7) a

Azafran PRA 5.1 (2.7-9.4)ah 23.5 (16.3-32.6)gk Azafran PRA 5.4 (3.2-8.9) cl 8.8 (5.7-13.2) af
IA 1.4 (0.4-4.3)ac 32.3 (25.0-40.6)jk IA 9.1 (6.1-13.3) jn 8.1 (5.1-12.8) af
PA 1.0 (0.3-3.9)a 28.0 (21.2-36.0)ik PA 9.4 (6.7-13.2) ln 4.8 (2.7-8.3) ab

LP/Shyri PRA 3.5 (1.6-7.2)ad 28.3 (21.7-35.9)ik LP/Shyri PRA 3.2 (1.6-6.0) bf 11.8 (8.0-17.1) bh
(Selec 1) IA 7.4 (4.1-13.0)bl 28.7 (21.9-36.7)ik (Selec 1) IA 2.8 (1.4-5.7) bh 8.3 (5.2-13.1) af
PA 10.2 (6.6-15.5)dn 22.1 (16.0-29.8)fj PA 3.2 (1.6-6.0) bi 7.0 (4.3-11.2) ae

† PRA = Pre-anthesis; IA = Initial anthesis; PA = Post anthesis. † PRA = Pre-anthesis; IA = Initial anthesis; PA = Post anthesis.
Perspectives on Fusarium Head Blight Resistance in Barley 67

identified (Figure 2), and three Information presented in Figure 2 is


genotypes did not fall into any cluster. useful for breeding programs, since it
Most two-rowed Chinese barley allows the identification of FHB-
genotypes are grouped in cluster 1, plus resistant cultivars whose resistance base
Svanhals, developed in Sweden. might be different, thereby increasing
Chevron and Chevron –derived the probability of accumulating levels of
genotype Ciho 16128 (comprising resistance when gene action for FHB
cluster 5) are the most resistant to FHB resistance is additive. However,
and have the lowest DON susceptibility to leaf blotches and rust
concentrations. diseases continues to be a problem that
could prevent the use of these
Most FHB-resistant, two-rowed barley
genotypes in some regions.
genotypes are susceptible to wheat stem
rust, net blotch, and powdery mildew,
and all are susceptible to leaf rust. All
FHB-resistant, six- rowed genotypes are QTLs for FHB Resistance
susceptible to leaf rust, wheat stem rust
Mapping QTLs for FHB resistance
(pathotype Pgt-QCC), and powdery
genes was carried out in doubled
mildew (Urrea et al., 1999).
haploid populations. In Gobernadora x
CMB 643 (Azafran), a QTL for type II

Zhedar 1
1 Svanhals
Zhedar 2

2 Gobernadora

Shyri
3

5 Chevron

0.0 0.23 0.45 0.68 0.90


Genetic distance

Figure 2. Dendrogram based on cluster analysis of the distance among 39 barley genotypes.
Source: Conci et al. (1999).
68 L. Gilchrist

resistance was found in chromosome 2 Conclusions


(Zhu et al., 1999). Chevron and Chevron
Fusarium graminearum resistance is
progeny have been used extensively in
difficult to evaluate under field
breeding for resistance to FHB and
conditions. Although producing
kernel discoloration (KD) in a
symptoms is easy, it is essential to control
Minnesota breeding program (Canci et
the precise composition of the inoculum
al., 1999). Four of the QTLs associated
used, the method of application, and the
with FHB in chromosomes 1, 2, and 4
circumstances in which inoculation is
were also associated with KD (De la
performed. Otherwise, great variability
Peña et al., 1999).
may be introduced that would make it
practically impossible to interpret the
observed results correctly.
Evaluating Barley The basis of FHB resistance is complex
Varieties and Advanced and not well understood today. The
Lines for FHB Resistance environment interacts with the host plant
In the last five years, barley varieties and the pathogen to produce a resistant
and advanced lines used in the reaction. The extent of the infected area in
CIMMYT/ICARDA Barley Program the host plant is the product of these three
were evaluated for FHB resistance at components (Figure 3). However,
Toluca, Mexico. The resulting data are evaluating FHB reaction in the field is not
presented in Tables 6, 7, and 8. enough for establishing resistance; kernel

Table 6. Characterization of fusarium head blight resistance (Types I, II, III, and IV) in nine barley
cultivars.

Type I Type II Type III Type IV Grain


Variety or % infec. % infec. DON ppm. % losses (1-5)†
line 98 99 98 99 98 99 98 99 99
Shyri 4.9 7.1 26.1 10.6 3.9 16.0 5.0 4.1 2
Atahualpa 92(H) 16.8 5.0 21.7 8.9 0.0 0.99 10.1 8.1 2
Arupo/K8755 6.6 11.7 3.7 38.7 1.0 2.9 13.6 4.0 2
//Mora (selec. 2)
Gob/Humai10 9.5 1.2 0.3 6.5 NA‡ 2.2 4.6 27.6 4
B89.3.1 (early)
Gob/Humai10 5.8 5.7 27.1 20.4 1.6 1.1 3.9 15.4 3
B89.3.2 (late)
Chevron 5.1 4.0 11.8 26.6 1.1 2.1 2.06 40.6 4
Gobernadora 5.3 9.2 20.7 14.6 1.9 12.0 13.4 28.5 2
Azafran 1.4 9.1 32.3 8.1 NA 120.0 11.3 6.28 3
LP/Shyri 12.5 2.1 23.4 10.6 NA 2.8 15.6 14.1 4
(Selec. 2)

† 1=good grain; 5=poor grain. ‡ NA=Not available.


Perspectives on Fusarium Head Blight Resistance in Barley 69

weight losses and toxin (DON)


concentration in the grain are also
important. Environment Pathogen

Mesterhazy (1997) made an important and


appropriate statement regarding this Host
issue: “Without reliable methods nothing Host

nment
can be said about the level of resistance,

Enviro
factors of resistance, about their

ent
Pathogen

ronm
inheritance and about an appropriate

Host
Envi
selection procedure. Without adequate Environment

methods we will not have appropriate Pathogen


Host
information, or what is even worse, Pathogen
incorrect information will be the result.”
Figure 3. Area of the disease.

Table 7. Characterization of fusarium head blight resistance (Types I, II, III, and IV) in a group of
advanced hull-less barley lines.

Type I Type II Type III Type IV Grain


Variety or % infec. % infec. DON ppm. % losses (1-5)‡
line† 98 99 98 99 98 99 98 99 99

Tocte//Gob/Humai 1.9 5.3 14.8 7.2 2.8 3.6 12.4 6.4 1


10/3/Atah 92/Aleli
CBSS96M00766D-B-2M-3Y

Tocte//Gob/Humai 2.2 3.7 14.4 15.3 1.1 12.0 10.2 3.0 2


10/3/Atah 92/Aleli
CBSS96M00766D-B-2M-4Y

Penco/Chevron-Bar 10.2 1.5 38.1 20.5 NA 9.2 2.0 6.1 1


CBSS96Y00341s-1Y-1M-10Y

Penco/Chevron-Bar – 2.6 16.9 13.2 NA 11.0 4.5 4.1 1


CBSS96Y00341s-1Y-1M-16Y

Ataco/Bermejo// 0.9 5.1 18.7 9.3 NA 14.0 0.3 3.8 1


Higo/3/CLNB/80.5138//
Gloria-Bar/Copal/4/
Chevron-Bar
CBSS96Y00653T-A-14Y-1M-16Y

Ataco/Bermejo// 0.9 3.7 7.3 5.1 2.3 5.3 4.7 3.9 2


Higo/3/CLNB/80.5138//
Gloria-Bar/Copal/4/
Chevron-Bar
CBSS96Y00653T-A-14Y-1M-17Y

Chamico/ Tocte// 6.8 4.6 53.1 12.6 2.0 34.0 14.8 8.8 3
Congona
CBSS95Y00352T-D-12Y-2Y-0M

† Resistance sources are in boldface. ‡ 1=good grain; 5=poor grain.


70 L. Gilchrist

Table 8. Characterization of fusarium head blight resistance (Types I, II, III, and IV) in seven advanced
covered barley lines.

Type I Type II Type III Type IV Grain


Variety or % infec % infec DON ppm % losses (1-5)‡
line† 98 99 98 99 98 99 98 99 99
Gob/Humai 10/3/ 1.0 9.7 16.3 9.4 38.0 12.9 4.3 1
Mpyt169.1Y/Laurel/
Olmo/4/ Canela
CBSS95M00804T-F-1M-7Y-0M
Escoba/Moradilla/3/ 3.8 11.0 7.2 8.4 24.0 7.1 3.3 2
Zhedar#2/NDB112//
Mora
CMB94A.595-A-2M-6Y-2M-3Y-0M
GOB 24 DH 4.0 14.6 6.0 8.0 27.0 13.0 5.9 2
(Gobernadora/Azafran)
GOB 45 DH 5.7 19.9 6.5 19.0 26.0 2.2 15.4 2
GOB 96 DH 9.7 25.0 7.4 33.1 3.6 4.3 4.9 1
Mosquera 7.3 18.2 8.2 18.1 27.0 16.3 16.4 1
(Fingal/F784.70/4/Zhedar/
3Hlla/Gob/Hlla/Shyri)
Zhedar #1/Shyri // 2.5 23.4 5.2 11.7 31.0 7.2 4.5 2
Olmo
CMB93.572-A-4Y-1Y-2M-1Y-0M

† Resistance sources in boldface. ‡ 1=good grain; 5=poor grain.

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72

Resistance and/or Tolerance


to BYDV: Recent Advances in
Barley at CIMMYT
M. HENRY1 AND H.E. VIVAR2

Barley yellow dwarf is the most concentration is affected. In some oats


important viral disease of barley. It is (Jedlinski et al., 1977) and barleys (Skaria
caused by a complex of luteoviruses et al., 1985, Ranieri et al., 1993), resistance
known as barley yellow dwarf viruses has been associated with field tolerance.
(BYDVs). They are transmitted in a
The most common and effective source
persistent manner by aphids to all
of tolerance to BYD in barley was
common cereals. The five serotypes PAV,
identified in Ethiopian barleys (Schaller
MAV, RPV, RMV, and SGV differ in
et al., 1963) and found to be associated to
severity, PAV being in general the most
the major semidominant gene Yd2
severe and most common, followed by
(Rasmusson and Schaller, 1959). As many
MAV and RPV. RMV and SGV are only
authors report, this gene is associated
found occasionally. Control of BYDV can
with field tolerance and with reduction
be achieved through elimination of its
in virus concentration (resistance). It is
insect vector by insecticide application,
located on chromosome 3 (Schaller et al.,
cultural practices, and the use of tolerant
1964) close to the centromere (Collins et
and/or resistant materials.
al., 1996). It is associated with reduction
According to Cooper and Jones (1983), in virus concentration with BYDV-PAV
tolerance in a plant is associated with an and MAV but not with BYDV-RPV
attenuation of symptoms without (Banks et al., 1992; Skaria et al., 1985;
reduction in virus multiplication. In a Herrera and Plumb, 1989).
resistant plant, virus multiplication or
Recently, Paltridge et al. (1998)
spread is affected. The term resistance has
developed an assay based on a protein
been broadly used to refer to any type of
showing allelic variation that correlates
interaction to the disease development;
with Yd2. Later, Ford et al. (1998)
most field resistance reported in the
developed a PCR marker for the Yd2-
literature was assessed through symptom
associated allele of Ylp using allele-
expression in the field and is therefore
specific primer pair. These molecular
tolerance. In this paper, we will
markers are useful for identifying the
differentiate the two mechanisms and
presence of Yd2 in barley cultivars.
refer to true resistance only when virus

1 CIMMYT Wheat Program, Apdo. Postal 6-641, Mexico, D.F., Mexico, 06600.
2 Head, CIMMYT-ICARDA Barley Program, Apdo. Postal 6-641, Mexico, D.F., Mexico, 06600.
Resistance and/or Tolerance to BYDV: Recent Advances in Barley at CIMMYT 73

The CIMMYT-ICARDA Barley Program approximately 10 aphids were deposited


for Latin America holds a number of lines at the base of each seedling using a
with high field tolerance to BYDV isolates calibrated mechanical dispenser. Aphid
PAV, MAV and RPV combined with movement was controlled through
resistance to foliar diseases. The objectives fortnightly insecticide application
of the present study were to evaluate the (Metasystox) on the entire trial, starting
type of resistance present in the CIMMYT one week after inoculation.
barley lines, distinguish between true
resistance and tolerance, and measure to Plant materials
what extent the gene Yd2 was present in Ninety-one lines from the CIMMYT-
those lines. Some of the results presented ICARDA Barley Program were tested in
here are still preliminary. this study. These lines were selected
because they showed field tolerance to
one of the BYDV isolates tested. The
controls were non-Yd2 lines Atlas 57,
Materials and Methods Centinela, and Calicuchima “S” and Yd2
lines Atlas 68, Sutter, Sutter /2*Numar.
Virus isolates and inoculation
The BYDV PAV-Mex, MAV-Mex, and RPV-
Field evaluation of BYDV
Mex isolates used in the experiment were
tolerance and/or resistance
collected in Mexico in 1993 and
Lines were tested under BYDV infection
maintained in CIMMYT’s greenhouse
in Toluca and El Batan, Mexico, during
through transmission by aphids. The
the summers of 1997, 1998, and 1999. For
aphid species Rhopalosiphum padi (PAV
each line, four 1-m double plots (20
and RPV) and Metopolophium dirhodum
plants) were sown adjacent to each other,
(MAV) were used for transmission.
to constitute the four treatments: non-
Inoculation was performed in the inoculated, inoculated with PAV, MAV, or
greenhouse by infesting five 6-day old RPV.
seedlings per line with 10 viruliferous
Symptoms were evaluated at flowering
aphids that had acquired BYDV by
using a 1-9 scale as described by
feeding on infected plants for 48 hours.
Berstchinger (1994), 1 being the most
Seedlings were isolated from each other
tolerant and 9 the most susceptible. The
by transparent plastic tubes. After a two-
three parameters measured were intensity
day inoculation period, aphids were killed
of yellowing, dwarfism, and reduction in
with the insecticide Metasystox (Bayer). In
tillering. In 1999, biomass was evaluated
all experiments, one or two plants were
instead of tillering because it seems to
kept free of aphids to serve as the non-
better represent the effect of BYD on plant
inoculated controls.
growth.
In the field, plants were infested at the
three-leaf stage with aphids reared in the Evaluation of resistance to BYDV
greenhouse on BYDV infected plants. To using ELISA
avoid contamination, the non-inoculated Flag leaf-1 and roots were collected seven
treatments were sprayed soon after days after inoculation to evaluate virus
emergence with the insecticide titers by ELISA (enzyme-linked
Metasystox; in the inoculated treatments, immunosorbent assay). Double Antibody
74 M. Henry and H.E. Vivar

Sandwich ELISA (DAS ELISA) was used OD


1.4
as described in Ayala et al. (2000). The
Roots
coating polyclonal antibodies against the 1.2
Leaves
US PAV, MAV, or RPV isolates were 1.0
provided by K. Perry (Purdue
University). Optical density (OD) was 0.8
measured at 410 hm using and MR 700 0.6
Microplate reader (Dynatech
0.4
Laboratories). A plant was considered
infected when the OD obtained in ELISA 0.2
was higher than twice the one obtained
0.0 Sutter Sutter*2/Numar Atlas 68
with the non-infected control. The higher
Yd2+ Yd2+ Yd2+
the OD, the higher the concentration and Centinela No Calicuchima “S” Atlas 57 No
the lower the resistance. Yd2+ No Yd2 Yd2

Figure 1. Virus titers in roots and leaves of


Detection of the presence of Yd2 barley genotypes seven days after infection
using molecular markers with BYDV-PAV.
The marker pair (Ylp PCR MF and Ylp
PCR MR) described by Ford et al. (1998)
was used in this study following the Effect of Yd2 on BYDV
procedure described by the same concentration
authors. A 311 base pair product was In the lines with Yd2, Sutter and Sutter/
obtained after PCR of DNA from both 2*Numar, virus titers were low in both
Yd2 and non-Yd2 plants. After digestion roots and leaves (Figure 1). However, in
of the PCR product with the restriction Atlas 68, the virus titers reached
enzyme NlaIII, two products of 253 and moderate levels in the leaves. According
58 base pairs were obtained with the to this, differentiation between Yd2 and
DNA from plants with the Yd2 gene, non-Yd2 lines is easier by comparing
while a single product of 311 base pairs virus titers in roots.
was obtained with plants not containing
As shown in Table 1, virus titers were
Yd2. The products were visualized on a
lower when the Yd2-associated allele was
3% agarose gel.
present than when it was absent,
confirming that Yd2 reduces
multiplication of BYDV-PAV in barley.
Results
As expected, a single product of 311bp
Table 1. Mean virus titers in ELISA measured in
was obtained with the non-Yd2 lines leaves and roots of barley genotypes with and
(Atlas 57, Centinela and Calicuchima without Yd2, seven days after inoculation with
“S”) while a 253bp product could be BYDV-PAV.
visualize with the Yd2 lines (Atlas 68,
Yd2-associated Number Average OD
Sutter, Sutter /2*Numar) (data not allele of lines Leaves Roots
shown), confirming the usefulness of the
marker for the detection of Yd2. The Yd2- Present 46 0.244 0.195
associated allele (Ylp) was detected in Absent 16 0.682 0.998
82% of the CIMMYT barleys tested (73).
Resistance and/or Tolerance to BYDV: Recent Advances in Barley at CIMMYT 75

Effect of Yd2 on field tolerance Symptoms were more severe in the absence
to BYDV of the Yd2-associated allele after BYDV PAV
Most lines carrying the Yd2-associated and MAV infection (Table 3). The difference
allele had a tolerant or intermediate was not so marked with BYDV-RPV
response to BYDV-PAV and MAV (Table infection.
2). In contrast, most lines not carrying
the gene were sensitive to infection by Germplasm with field
the two BYDV serotypes. The response tolerance to BYDV
to BYDV-RPV did not differ greatly Table 4 shows a list of lines with field
between lines with and without the tolerance to BYDV. The following lines
Yd2-associated allele. combined field tolerance to the three BYDVs
tested as well as resistance to BYDV-PAV:
LIMON, BOLDO/MJA, GUAYABA,
JACAPA, CANTUA, and CLAVO.

Table 2. Response of CIMMYT barleys to BYDV Table 4. Selected CIMMYT barley lines with
field infection. combined tolerance to the three BYDV Mexican
isolates PAV, MAV, and RPV.
BYDV Yd2 Non-Yd2
response PAV MAV RPV PAV MAV RPV Name Selection history

Tolerant 57.8† 69.6 34.8 0.0 6.7 14.3 PETUNIA 1 CMB93.855-G-15Y-1M-0Y


Intermediate 22.2 13.0 30.4 7.1 6.7 28.6 CHAMICO/TOCTE//CONGONA CBSS95Y00352T-H-7Y-2M-0Y
Sensitive 20.0 17.4 34.8 92.9 86.7 57.1 DUMARI CMB93.755-C-1Y-1M-0Y
FAIQUE CMB93.885-A-7Y-2M-1Y-0M
† Percentage of lines in each category according to the MADRE SELVA CMB92A.790-H-4M-1Y-2B-0Y
presence of Yd2 as identified by the marker pair. LIMON CMB92A.655-B-7M-1Y-1B-0Y
INCIENSO CMB92A.1264-P-1M-1Y-1B-0Y
PALTON CMB92.391-B-3Y-1M-1Y-1B-0Y
LIMON CMB92A.655-C-9M-1Y-1B-0Y
ARAVISCO CMB91A.629-B-0M-1Y-1M-1Y-2B-0Y
Table 3. Severity of BYDV response after BOLDO/MJA CMB91A.60-1M-1Y-1M-1Y-0B
artificial inoculation in El Batan, Mexico.† TINCTORIA CMB91A.210-23M-2Y-1M-1Y-2B-0Y
BOLDO/MJA CMB91A.60-1M-1Y-3M-1Y-1B-0Y
Yd2 GRAMALOTE CMB92A.893-A-1M-2Y-1B-0Y
associated PAV MAV RPV CANTUA CMB92.546-X-2Y-4M-0Y
allele Y D B Y D B Y D B CANTUA CMB92.546-I-1Y-1M-0Y
CANTUA CMB92.546-I-1Y-4M-0Y
Present 1.2 0.7 0.7 1.0 0.5 0.4 1.3 0.8 0.9 PAPELILLO CMB92A.920-A-2M-1Y-0B
Absent 3.3 3.8 5.1 1.8 1.4 1.3 2.3 0.4 0.8 CALENDULA CMB93.947-D-5Y-1M-0Y
INCIENSO CMB92A.1264-A-3M-1Y-0M
† Y= yellowing, D= dwarfing, B= reduction in biomass, JACINTO CMB92.490-E-6Y-1M-1Y-2B-0Y
measured on a 1-9 scale. MJA/BRB2//QUINA CMB93.987-C-1Y-1M-0Y
BBSC/CONGONA CBSS95M00144S-3M-3Y-0M
ABN-B/KC-B//RAISA/3/ALELI CMB92.338-A-6Y-1M-2Y-1B-0Y
QUINN/ALOE//CARDO CMB92A.1439-I-6M-1Y-1B-0Y
COMINO/3/MATICO/JET// CMB92.494-C-2Y-1M-2Y-1B-0Y
SHYRI/4/ALELI
ATACO/COMINO//ALELI CMB92.367-E-7Y-1M-1Y-1B-0Y
ATACO/COMINO//ALELI CMB92.367-Q-6Y-1M-0Y
CANTUA CMB92.546-I-1Y-5M-1Y-1B-0Y
76 M. Henry and H.E. Vivar

Conclusions Berstchinger, L. 1994. New procedures for the


effective screening of cereals for symptomatic
The CIMMYT-ICARDA Barley Program tolerance to barley yellow dwarf luteoviruses.
BYD Newsletter 5:14.
for Latin America holds germplasm with Burnett, P.A., Comeau, A., and Qualset, C.O. 1995.
combined field tolerance to the three Host plant tolerance or resistance for control of
major BYDVs (PAV, MAV and RPV) and barley yellow dwarf. In: Barley Yellow Dwarf,
40 Years of Progress. D’Arcy, C.J. and Burnett,
to other major diseases. Furthermore, P.A. (eds.). APS Press, St Paul.
some of the material is truly resistant Collins, N.C., Paltridge, N.G., Ford, C.M., and
(low virus concentration) at least to Symons, R.H. 1996. The Yd2 gene for barley
yellow dwarf virus resistance maps close to
BYDV-PAV. Most lines possesses the centromere on the long arm of barley
gene Yd2; however, it is still possible that chromosome 3. Theoretical and Applied
other genes are present, because of the Genetics 92:858-864.
Cooper, J.I., and Jones, A.T. 1983. Responses of
extremely high level of tolerance is some
plants to viruses: proposals for the use of
lines. This work is still in progress and terms. Phytopathology 73:127-128.
will be completed by studying true Ford, C.M., Paltridge, N.G., Rathjen, J.P., Moritz,
R.L., Simpson, R.J., and Symons, R.H. 1998.
resistance to MAV and RPV.
Rapid and informative assays for Yd2, the
barley yellow dwarf virus resistance gene,
based on the nucleotide sequence of a closely
linked gene. Molecular Breeding 4:23-31.
Acknowledgments Herrera, M.G.F. 1989. Interactions between host
plants and British isolates of barley yellow
We thank Dr. Mireille Khairallah for her dwarf virus. Thesis, University of London.
assistance in setting up the PCR assay, Jedlinski, H., Rochow, W.F., and Brown, C.M.
1977. Tolerance to barley yellow dwarf virus in
Javier Segura and Gabriel Posadas for oats. Phytopathology 67:1408-1411.
their technical assistance, and Alma Paltridge, N.G., Collins, N.C., Bendahmane, A.,
McNab for her editorial input. and Symons, R.H. 1998. Development of YLM,
a codominant PCR marker closely linked to
the Yd2 gene for resistance to barley yellow
dwarf disease. Theoretical and Applied
Genetics 96:1170-1177.
References Ranieri, R., Lister, R.M., and Burnett, P.A. 1993.
Relationships between barley yellow dwarf
Ayala, L., Khairallah, M., González-de-Leon, D., titer and symptom expression in barley. Crop
van Ginkel, M., Mujeeb-Kazi, A., Keller, B., Sci. 33:968-973.
and Henry, M. 2000. Identification and use of Rasmusson, D.C., and Schaller, C.W. 1959. The
molecular markers to detect barley yellow inheritance of resistance in barley to the
dwarf virus resistance derived from Th. yellow dwarf virus. Agron. J. 51:661-664.
intermedium in bread wheat. Submitted to Schaller, C.W., Qualset, C.O., and Rutger, J.N.
Theoretical and Applied Genetics. 1964. Inheritance and linkage of the Yd2 gene
Banks, P.M., Waterhouse, P.M., and Larkin, P.J. conditioning resistance to the barley yellow
1992. Pathogenicity of three RPV isolates of dwarf disease in barley. Crop Sci. 4:544-548.
Barley Yellow Dwarf Virus on barley, wheat Schaller, C.W., Rasmusson, D.C., and Qualset,
and wheat alien addition lines. Annals of C.O. 1963. Sources of resistance to the yellow
Applied Biology 121:305-314. dwarf virus in barley. Crop Sci. 3:342-344.
Skaria, M., Lister R.M., Foster J.E., and Schanner,
G. 1985. Virus content as an index of
symptomatic resistance to barley yellow dwarf
virus in cereals. Phytopathology 75:212-216.
77

Two Decades of Barley Breeding


H.E. VIVAR1

Plant breeders seeking to improve a of accessions in Mexico. Our work was to


particular trait, such as disease field test 273 resistant accessions against
resistance, search for sources of that trait races of R. secalis present in central
in collections stored in germplasm Mexico. Most accessions were still
banks. Most germplasm bank accessions resistant to US and Mexican races, and
have been screened for different traits only 13% proved to be scald susceptible
and the results published. The and discarded.
ICARDA/CIMMYT barley breeding
In 1983, accessions from the world barley
program relies on this type of
collection were screened against leaf rust
information for identifying cultivars to
at the CIANO Experiment Station located
be used as parents in crosses.
in the Yaqui Valley in northwestern
The Colombian National Research Mexico. Of 11,087 accessions screened
Institute (Anonymous, 1984) screened against races 8, 19, and 30 of P. hordei,
8,650 barley accessions from the world only 285 were found to be resistant to leaf
collection against race 24 of stripe rust rust (Vivar, 1986).
caused by Puccinia striiformis f.sp. hordei,
In Japan, Takeda and Heta (1989)
a disease recently introduced to the
screened 5,000 barley accessions against
Andean Region of South America. In
Fusarium graminearum and found 23
1982, accessions found to be stripe rust
entries with good levels of scab
resistant in Colombia were brought to
resistance.
Mexico, where they were used to launch
extensive breeding efforts aimed at Breeders and pathologists who have to
developing germplasm resistant to the work with resistant accessions from the
disease. Stripe rust has since become world collection invariably find that
endemic in the Andean Region. resistance is in poor agronomic plant
types. Therefore, they have to devote a
In California, Webster et al. (1980) found
lot of effort to transferring disease
273 entries with no symptoms of
resistance genes into improved cultivars.
pathogen virulence of Rynchosporium
secalis present in the state. The extensive
Multiple disease resistance
effort devoted to finding scald resistant
Researchers at Montana State University
cultivars among 18,000 accessions from
advanced the concept of introducing
the world barley collection in the US
multiple disease resistance into barley
saved us from screening a large number
using male-sterile plants to facilitate

1 Head, ICARDA/CIMMYT Barley Program, Apdo. Postal 6-641, Mexico, D.F., Mexico, 06600.
78 H.E. Vivar

recurrent selection. The project required scab, in addition to scald and the three
distributing several barley populations rusts (leaf, stripe, and stem).
among cooperators around the world.
Several important factors were involved
The objective was achieved after several
in the incorporation of multiple disease
years of work. Plants with multiple
resistance, but a few were essential to the
disease resistance became available, but
success of the project. First, the use of
barley breeders were reluctant to use
large numbers of crosses (single and top)
resistant plants in crosses with their elite
made it possible to combine resistance to
cultivars because of their poor agronomic
several diseases in a single plant. Second,
plant type.
large segregating populations were
In the 1970s, Colombian breeders carried screened in the field at several
out activities aimed at developing stripe experiment stations. The Toluca Station
rust resistant varieties. Their efforts was used for screening scald, stripe rust,
culminated in the release of Quibenras, a and head scab; the CIANO Station for
variety characterized by its outstanding leaf and stem rust, El Batan for leaf rust,
stripe rust resistance. Despite this and Poza Rica for spot blotch. The use of
resistance, Quibenras had a short life in the stations helped us produce reliable
farmers’ fields due to its susceptibility to artificial epidemics and effective plant
leaf rust, which was predominant where selection. Third, international testing in
Quibenras was sown. Farmers who had cooperation with national program
been using expensive chemical scientists spread over five continents
treatments to control stripe rust were allowed us to identify plants resistant to
forced to continue using fungicides pathogenic races not present in Mexico.
(Bayleton) to control leaf rust.
The ability to create artificial large-scale
Both experiences pointed to the need for epidemics was key for selecting against
cultivars with multiple disease resistance scald and the three rusts. With diseases
to provide stability in barley production. such as barley yellow dwarf and head
Incorporating multiple disease resistance scab, developing artificial large-scale
into high yielding barley germplasm epidemics is cumbersome and expensive;
thus became the main objective of the for this reason, the identification of
ICARDA/CIMMYT barley program in potentially resistant plants was done by
1982. visual selection in advanced segregating
populations at the Toluca Station. The
This breeding strategy was implemented
preliminary identification of potentially
in several steps. The first template was
resistant plants required more detailed
achieved by crossing scald and leaf rust
screening by CIMMYT plant pathologists
resistant cultivars. Once advanced lines
Drs. L. Gilchrist and M. Henry, both of
with resistance to both diseases were
whom presented their results at this
obtained, they were used as parents to
meeting.
form a second template, to which stripe
rust resistance was added. Over 18 years Artificial inoculation at times creates
of breeding (two generations per year) friction between plant pathologists and
and a total of 36 generations, we have plant breeders. A frequent complaint is
built templates that contain resistance to that too much inoculum was applied,
BYD, net blotch, spot blotch, and head that the resistance reactions of the host
Two Decades of Barley Breeding 79

plants are blurred or that too little parents are resistant to scald.
inoculum was applied and no Calicuchima has remained scald
differences are visible. The high rainfall resistant for more than 10 years in
(800 to 1000 mm) recorded during the Mexico.
growing cycle at the Toluca Station
favors the development of severe Partial resistance
epidemics of scald, stripe rust, and head Studies of leaf rust virulence conducted
scab. Since in the barley program disease in Israel and Ecuador by Brodny and
epidemics develop without direct Rivadeneira (1996) showed that the
intervention by plant pathologists, virulence present in both countries was
complaints about the intensity of capable of rendering all major genes
epidemics are not uncommon. reported in the literature non effective.
This suggests that the resistance
In our experience, the use of one
achieved by pyramiding major gene
resistant parent in a single cross does not
combinations may not be durable.
provide an adequate level of resistance
under conditions favoring disease Parlevliet and Kuiper (1977), working
development. This is true not only with with leaf rust in barley in The
scald, but also rust and head scab. For Netherlands, proposed a partial
this reason, three-way crosses were resistance mechanism based on minor
adopted as a tool in the crossing genes that provide varying levels of
program. All F1s are now top-crossed to resistance against all virulences of the
a third parent so as to combine at least pathogen. Parlevliet developed barley
two resistance sources. In recent years, cultivars (Vada, LP) in which several
as the program matured, a combination minor resistance genes were
of three different sources of resistance accumulated; they showed better and
was attempted for scald, head scab, and more durable resistance against the
stripe rust. For example, advanced lines disease.
(F7) from the cross Svanhals/M.Selva//
The use in the ICARDA/CIMMYT
Azafran/Gob24 were screened for head
barley program of the sources of partial
scab resistance with good results.
resistance to leaf rust developed by
Savanhals, Azafran, and Gobernadora
Parlevliet was hampered by the presence
are three sources of head scab resistance.
of other diseases, such as scald, net
The combined use of three parents blotch, spot blotch, and BYD. Parental
resistant to the same disease has resulted material developed in Holland was
in cultivars showing durable resistance. excellent for leaf rust but very poor for
The variety Shyri (released in Ecuador) leaf blotches and BYD.
was the result of crossing three stripe
The program continues to breed for
rust resistance sources (Motan, Kober,
partial resistance against leaf rust.
and Teran). Shyri has remained resistant
Several barley lines carrying good levels
to stripe rust since 1989 in the US,
of partial resistance to leaf rust, stripe
Germany, and several Latin American
rust, and scald have been identified. Our
countries. The variety Calicuchima was
aim is to combine minor and major
selected from the cross L.B.Iran/Una
genes to achieve durable resistance.
8271//Gloria/Come, all of whose
80 H.E. Vivar

Head scab Over the last decade, fusarium toxin


In 1984 breeding for scab resistance by production in the grain has become a
the ICARDA/CIMMYT barley program major concern due to the implications for
was not popular with ICARDA human health. Toxins such as
management, who argued that the deoxynivalenol (DON) are present in
disease was not considered a problem in grain infected with Fusarium
the West Asia and North Africa (WANA) graminearum. In the developed world,
region, and that there were no reports of toxin levels in grain to be used as food or
its presence in the Americas. I learned feed must not exceed 2 ppm; however, in
that breeding against a disease that is not third world countries that have no
considered a problem is a difficult regulations on toxin content in the grain,
enterprise. Despite early doubts about the problem becomes more severe,
the project, breeding for scab resistance especially in areas such as the Andean
became one of the more successful Region, where barley is a staple food.
initiatives of the ICARDA/CIMMYT
Clear et al. (1997) reported that hull-less
barley program in Mexico.
barley showed a 59% DON reduction in
Two national programs released three grain toxin content immediately after
scab resistant varieties in two countries; being threshed in the field. Apparently
Ecuador (Atahualpa and Shyri) and the toxin is attached to the lemma and
China (Zhenmai-1). The variety palea, which are removed during
Zhenmai-1 is sown on more than 100,000 threshing. Clear and colleagues
hectares in several Chinese provinces concluded that hull-less barley may be a
located in the lower basin of the Yangtze good option for farmers in areas affected
River, and is showing yield increases of by head scab.
20-25% over the Chinese barley varieties
After several years of testing under
it replaced. Dr. He Zhonghu, CIMMYT
artificial inoculation at the Toluca Station,
representative in China, estimates that
several hull-less, six- and two-rowed
40% of one million hectares sown to
barley lines were identified as being
barley in China is planted to varieties
resistant to head scab. Seed of these lines
introduced from Mexico or derived from
will be distributed to national programs
crosses between local varieties and
for testing in areas prone to the disease.
Mexican introductions.
Molecular markers were used to map
Head scab caused by Fusarium
scab resistance in the doubled haploid
graminearum has become a major disease
population Gobernadora/Azafran in
in North America. Since 1993, the
cooperation with Oregon State
epidemic has produced more economic
University (Zhu et al., 1999). The
losses than any other disease in US
doubled haploid lines were phenotyped
history. In areas such as the upper
for scab resistance in Mexico, North
Midwest (the Dakotas, Minnesota), head
Dakota, and China. Type II resistance (to
scab has become a major problem, and
spread of the fungus within the spike)
extensive efforts are being invested in
was mapped near the centromeric region
breeding and pathology research.
of chromosome 2 with the largest R2
value. Lateral florets, a morphological
Two Decades of Barley Breeding 81

trait, was mapped in the same region, has received more attention. Two
opening the possibilities for indirect Ecuadorian varieties, Shyri and
selection for head scab resistance. Calicuchima, have been extensively
studied. They carry different quantitative
In Alberta, Canada, Dr. J. Helm selected
trait loci (QTL) for stripe rust resistance.
40 two-rowed advanced lines from a
Shyri’s stripe rust resistance was mapped
cross developed in a disease-free
on chromosome 5, while Calicuchima has
environment. The selected lines had
resistance QTLs on chromosomes 4 and 7
never been exposed to scab during their
(Toojinda et al., 2000). The diversity of
development. The main selection
the resistance found in these two
objective was to find spikes lacking
Ecuadorian varieties provides additional
lateral florets. The 40 lines were sent to
insurance against a sudden change in
Mexico for screening under artificial
pathogen virulence.
inoculation during the summer of 2000.
The ICARDA/CIMMYT Barley Program
Results of the screening done by Gilchrist
has been successful in developing high
are shown in Table 1. Five lines were
yielding barley with multiple disease
found to possess Type I and II resistance
resistance. Record yields (11 t/ha)
(to primary infection and fungal spread,
produced by introductions from Mexico
respectively). The percentage (12.5%) of
have been reported by breeders in Chile
head scab resistant lines selected
and Peru. However, when these high
provides a clear indication of the
yields (obtained in test plots) are
effectiveness of indirect head scab
compared to the national average barley
selection in two-rowed barley
yield in Peru (1.1 t/ha), it is evident that
populations.
the gap between experimental and
farmers’ yields has increased.

Table 1. Percent of grain infected with Fusarium In 1999, farmers were provided with
graminearum in five two-rowed lines introduced fertilizer, barley seed, and herbicides as
from Alberta, Canada, and screened at the
part of a project conducted in Saraguro,
Toluca Experiment Station, Mexico, summer of
2000. located in the highlands of southern
Ecuador. The average yield obtained by
Resistance (%) farmers growing the new variety Shyri-
Line Type I Type II 2000 was 3 t/ha, compared to 7 t/ha
H93123-003 5.35 5.39 harvested with the same variety in large
H93126-002 7.23 6.23 seed increase plots at the Chuquipata
H93125 6.88 6.83 Experiment Station. This shows that
H93123-008 5.98 7.00
H93126-011 4.44 8.30
closing the gap entails more that just
Susceptible check (Gob-89) 13.43 27.78 breeding activities.

I have described the successes of the


Ten years of cooperation with Oregon ICARDA/CIMMYT barley program, but
State University has led to a deeper I have not mentioned its weaknesses. To
understanding of the genetic basis of be fair, I would call your attention to a
resistance to several diseases. Stripe rust, problem exhibited by germplasm
present in Mexico and northwest USA, developed by the program: its
82 H.E. Vivar

susceptibility to loose smut (Ustilago Parleviet, J.E., and Kuiper, H.J. 1977. Resistance of
some barley cultivars to leaf rust, Puccinia
nuda), a disease easily controlled by hordei: Polygenic partial resistance hidden by
applying fungicides such as Vitavax. monogenic hypersensitivity. Neth. J. Plant
Unfortunately, subsistence farmers with Pathol. 83:85-889.
Takeda, K., and Heta, H. 1989. Establishing the
limited economic resources, who keep testing method and search for the resistant
their grain for use as seed the following varieties to Fusarium head blight in Japan.
year, could decide to skip the seed Japan J. Breed. 39:203-216.
Toojinda, T., Baird, E., Broers, L., Chen, X.M.,
treatment and run into a potentially
Hayes, P.M., Kleinhofs, A., Korte, J., Kudrna,
serious production problem. D., Leung, H., Line, R.F., Powell, W., and Vivar,
H. 2000. Mapping qualitative and quantitative
disease resistance genes in a doubled haploid
References
population of barley. Theor. Appl. Genet.
Anonymous. 1984. Control de la roya amarilla y
(in press).
parda en cebada. Colombia, 1975-84.
Vivar, H.E. 1986. Cereal Improvement Program.
Programa Nacional de Cereales Menores, 9.
ICARDA Annual Report.
Instituto Colombiano Agropecuario, Bogota,
Webster, R.K., Jackson, L.F., and Schaller, C.W.
Colombia.
1980. Sources of resistance in barley to
Brodny, U., and Rivadeneira, M. 1996. Physiologic
Rynchosporium secalis. Plant Dis. 64:88-90.
specialization of Puccinia hordei in Israel and
Zhu, H., Gilchrist, L., Hayes, P., Kleinhofs, A.,
Ecuador, 1992-1994. Can. J. Plant Pathol.
Kudrna, D., Liu, Z., Prom, L., Steffenson, B.,
18:375-378.
Toojinda, T., and Vivar, H. 1999. Does function
Clear, R.M., Patrick, S.K., Nowicki, T., Gaba, D.,
follow form? Principal QTLs for Fusarium
Edney, M., and Babb, J.C. 1997. The effect of
head blight (FHB) resistance are coincident
hull-removal and pearling on Fusarium species
with QTLs for inflorescence traits and plant
and trichocenes in hulless barley. Can. J. Plant
height in a doubled-haploid population of
Sci. 77:161-166.
barley. Theor. Appl. Genet. 99:1221-1232.