Advances in Potato Chemistry and Technology

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Advances in Potato Chemistry and Technology

Second Edition

Editors

Jaspreet Singh

Lovedeep Kaur

Riddet Institute and Massey Institute of Food Science and Technology, Massey University, Palmerston North, New Zealand

Table of Contents

Cover image

Title page

Copyright

List of Contributors

Foreword

Chemistry, Processing, and Nutritional Attributes of Potatoes—An Introduction

Chapter 1. Potato Origin and Production

1. Introduction

2. Origin, Domestication, and Diversity

3. Production in the Center of Origin

4. Spread and Global Production Trends Outside the Center of Origin

5. Conservation in the Center of Origin

6. Uses of Potato in the Center of Origin

7. Trend for Sustainable Conservation and Use

Chapter 2. Cell Wall Polysaccharides of Potato

1. Introduction

2. Isolation of Potato Cell Walls

3. Cell Wall Polysaccharides

4. Effects of Heating on Potato Cell Wall Polysaccharides

5. Distribution of Pectin in Potato Tuber and Function in Cell Walls

Chapter 3. Structure of Potato Starch

1. Introduction

2. Polysaccharide Components of Potato Starch

3. Starch Granules in Potato

4. Phosphorylated Potato Starch

5. Potato Starch Synthesis

6. Conclusions

Chapter 4. Potato Proteins: Functional Food Ingredients

1. Introduction

2. Potato Proteins

3. Conclusion

Chapter 5. Potato Lipids

1. Introduction

2. Lipids of Potato

3. Lipids of Transgenic Potato Cultivars

Chapter 6. Vitamins, Phytonutrients, and Minerals in Potato

1. Introduction

2. Potatoes, Nutrition, and the Food Debates

3. Basic Potato Nutritional Content

4. A Survey of Vitamins in Potatoes

5. Glycoalkaloids

6. Potato Minerals

7. Potato Phenylpropanoids

8. Effect of Development on Tuber Phytonutrients

9. Carotenoids

10. Effect of Cooking on Phytonutrient Content

11. The Role of Potatoes in Global Food Security

12. Conclusion

Chapter 7. Glycoalkaloids and Calystegine Alkaloids in Potatoes

1. Introduction

2. Glycoalkaloids

3. Calystegine Alkaloids

4. Conclusions

Chapter 8. Potato Starch and Its Modification

1. Introduction

2. Potato Starch versus Cereal Starches

3. Potato Starch Modification

4. Nutritional and Toxicological Aspects

5. Conclusions

Chapter 9. Colored Potatoes

1. Introduction

2. Potatoes with Red and Purple Flesh

3. Potato Antioxidants

4. Colored Potato AOA

5. Factors Influencing Levels of Beneficial Phytochemicals, AOA, and Antinutrients in Colored Potatoes

6. Health Benefits, Nutritional Aspects, and Use of Colored Potatoes

7. Conclusions and Future Trends

Chapter 10. Postharvest Storage of Potatoes

1. Introduction

2. Maturity of Tubers

3. Growing Conditions Affecting Postharvest Storage

4. Harvesting and Handling Factors Affecting Postharvest Storage

5. Pests and Diseases

6. Biochemical Changes of Tubers During Storage

7. Storage Preparations and Conditions

8. Storage Process

9. Management of Storage Environment

10. Effect of Postharvest Storage on Processing and Nutritional Quality of Potatoes

Chapter 11. Organic Potatoes

1. Introduction

2. Macroelements (N, P, K, Ca, Mg, S)

3. Phenolic Compounds

4. Carotenoids

5. Glycoalkaloids

6. Ascorbic Acid

7. Conclusions

Chapter 12. Potato Flavor

1. Introduction

2. Aroma and Flavor

3. Identifying Flavor Compounds

4. Taste

5. Texture

6. Influence of Growth and Storage Environment on Flavor

7. Combining Sensory Panels with Molecular and Metabolomics Approaches

8. Summary

Chapter 13. Microstructure, Starch Digestion, and Glycemic Index of Potatoes

1. Introduction

2. Starch Digestion and GI of Potatoes

3. Potato Microstructure and Starch Digestion

4. Rheology of Food Matrix and Starch Digestion

5. Formulated Foods and Starch Digestion

6. Conclusions

Chapter 14. Thermal Processing of Potatoes

1. Introduction

2. Product versus Quality

3. Processing versus Quality

4. Softening Kinetics

5. Quality Optimization

6. Novel or Emerging Methods and Outlook for the Future

Chapter 15. Fried and Dehydrated Potato Products

1. Introduction

2. Importance of Chemical Composition in Potato Processing

3. Importance of Microstructure in Potato Processing

4. Potato Processing: Important Derived Products

5. Conclusions

Chapter 16. Textural Characteristics of Raw and Cooked Potatoes

1. Introduction

2. Cooking and Sensory Characteristics

3. Textural Characteristics

4. Rheological and Thermal Characteristics

5. Conclusions

Chapter 17. Mechanisms of Oil Uptake in French Fries

1. Introduction

2. Physics of Cooling and Oil Uptake During Cooling

3. Typology of Defects Provoking Oil Uptake

4. Multiscale Modeling of Oil Uptake

5. Deviations to Generally Accepted Oil Uptake Mechanisms

6. Conclusions

Chapter 18. Acrylamide in Potato Products

1. Introduction

2. Aspects Affecting Acrylamide Formation in Fried Potato Products and Possible Mitigation Strategies

3. Additives or Processing Aids—from Lab Tests to Industrial Scale

4. Evolution of Risk Management

5. Future Outlook

Chapter 19. Advanced Analytical Techniques for Quality Evaluation of Potato and Its Products

1. Introduction

2. Analytical Techniques

3. Quality Evaluation

4. Advanced Techniques

5. Final Remarks

Chapter 20. The Role of Potatoes in Biomedical/Pharmaceutical and Fermentation Applications

1. Introduction

2. Biomedical Applications

3. Pharmaceutical Applications

4. Fermentation Applications

5. Other Applications

Chapter 21. Novel Applications of Potatoes

1. Introduction

2. Biodegradable Packaging

3. Fiber-Reinforced Biodegradable Composites for Constructive Parts in Aerospace, Automotive, and Other Areas

4. Edible Films

5. Textiles and Paper

6. Starch Spherulites and Nanocrystals

7. Potato Waste Utilization and Other Miscellaneous Uses

Chapter 22. Potato Proteomics: A New Approach for the Potato Processing Industry

1. Introduction

2. Proteomics Techniques—An Overview

3. Proteomics in Food Nutrition and Processing

4. Potato Proteomics and Quality Improvement

5. Getting the Best of Potato—Perspectives of Proteomics Applied to Product Quality

6. Conclusions

Chapter 23. Potatoes and Human Health

1. Introduction

2. Nutrient Contributions

3. Dietary Guidance

4. Obesity

5. Diabetes

6. Immune Function and Inflammation

7. Cardiovascular Disease

8. Cancer

9. Gastrointestinal Health

10. Chronic Kidney Disease

11. Consumer Benefits

12. Conclusions

Index

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Copyright

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Notices

Knowledge and best practice in this field are constantly changing. As new research and experience broaden our understanding, changes in research methods, professional practices, or medical treatment may become necessary.

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ISBN: 978-0-12-800002-1

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List of Contributors

María Dolores Álvarez Torres,     Department of Characterization, Quality, and Safety, Institute of Food Science, Technology and Nutrition (ICTAN-CSIC), Madrid, Spain

Silvia Arazuri,     Department of Agricultural Projects and Engineering, Universidad Pública de Navarra, Pamplona, Navarra, Spain

Cristina Barsan

Université de Toulouse, INP-ENSA Toulouse, Génomique et Biotechnologie des Fruits, Castanet-Tolosan, France

INRA, Génomique et Biotechnologie des Fruits, Chemin de Borde Rouge, Castanet-Tolosan, France

Led Academy, Toulon, France

Eric Bertoft,     Department of Food Science and Nutrition, University of Minnesota, St Paul, MN, USA

Andreas Blennow,     Department of Plant and Environmental Sciences, University of Copenhagen, Frederiksberg C, Denmark

Vaiva Bražinskienė,     Faculty of Business and Technologies, Utena University of Applied Sciences, Utena, Lithuania

Fanny Buffetto,     INRA, UR1268 Biopolymères, Interactions et Assemblages, Nantes, France

Mary E. Camire,     School of Food and Agriculture, University of Maine, Orono, ME, USA

Isabelle Capron,     INRA, UR1268 Biopolymères, Interactions et Assemblages, Nantes, France

Marie-Christine Ralet,     INRA, UR1268 Biopolymères, Interactions et Assemblages, Nantes, France

Rosana Colussi

Riddet Institute and Massey Institute of Food Science and Technology, Massey University, Palmerston North, New Zealand

Departamento de Ciência e Tecnologia Agroindustrial, Universidade Federal de Pelotas, Pelotas, Brazil

Virginia Corrigan,     The New Zealand Institute for Plant & Food Research Ltd, Palmerston North, New Zealand

Pablo Cortés,     Department of Chemical Engineering and Bioprocesses, Pontificia Universidad Católica de Chile, Santiago, Chile

Stef de Haan,     International Center for Tropical Agriculture (CIAT), Tu Liem, Hanoi, Vietnam

Bruno De Meulenaer,     NutriFOODchem Unit, Department of Food Safety and Food Quality, Faculty of Bioscience Engineering, Ghent University, Ghent, Belgium

Mendel Friedman,     United States Department of Agriculture, Agricultural Research Service, Western Regional Research Center, Albany, CA, USA

Kristina Gaivelytė,     Department of Pharmacognosy, Lithuanian University of Health Science, Kaunas, Lithuania

Fabienne Guillon,     INRA, UR1268 Biopolymères, Interactions et Assemblages, Nantes, France

Hanjo Hellmann,     School of Biological Sciences, Washington State University, Pullman, WA, USA

Carmen Jarén,     Department of Agricultural Projects and Engineering, Universidad Pública de Navarra, Pamplona, Navarra, Spain

Lachman Jaromír,     Department of Chemistry, Faculty of Agrobiology, Food and Natural Resources, Czech University of Life Sciences, Prague, Czech Republic

Salwa Karboune,     Department of Food Science and Agricultural Chemistry, McGill University, Ste-Anne de Bellevue, Quebec, Canada

Hamouz Karel,     Department of Plant Production, Faculty of Agrobiology, Food and Natural Resources, Czech University of Life Sciences, Prague, Czech Republic

Lovedeep Kaur,     Riddet Institute and Massey Institute of Food Science and Technology, Massey University, Palmerston North, New Zealand

Régis Kesteloot,     Régis Kesteloot Conseil, Lambersart, France

Carol E. Levin,     United States Department of Agriculture, Agricultural Research Service, Western Regional Research Center, Albany, CA, USA

Ainara López,     Department of Agricultural Projects and Engineering, Universidad Pública de Navarra, Pamplona, Navarra, Spain

María Salomé Mariotti,     Department of Chemical Engineering and Bioprocesses, Pontificia Universidad Católica de Chile, Santiago, Chile

Orsák Matyáš,     Department of Chemistry, Faculty of Agrobiology, Food and Natural Resources, Czech University of Life Sciences, Prague, Czech Republic

Owen J. McCarthy,     Massey Institute of Food Science and Technology, Massey University, Palmerston North, New Zealand

Marian McKenzie,     The New Zealand Institute for Plant & Food Research Ltd, Palmerston North, New Zealand

Raquel Medeiros,     NutriFOODchem Unit, Department of Food Safety and Food Quality, Faculty of Bioscience Engineering, Ghent University, Ghent, Belgium

Frédéric Mestdagh,     NutriFOODchem Unit, Department of Food Safety and Food Quality, Faculty of Bioscience Engineering, Ghent University, Ghent, Belgium

Duroy A. Navarre,     USDA-ARS, Washington State University, Prosser, WA, USA

Wenceslao Canet Parreño,     Department of Characterization, Quality, and Safety, Institute of Food Science, Technology and Nutrition (ICTAN-CSIC), Madrid, Spain

Anna Patsioura

INRA, UMR 1145 Ingénierie Procédés Alimentaires, Group Interaction between Materials and Media in Contact, Massy, France

AgroParisTech, UMR 1145 Ingénierie Procédés Alimentaires, Massy, France

Franco Pedreschi,     Department of Chemical Engineering and Bioprocesses, Pontificia Universidad Católica de Chile, Santiago, Chile

Reena Grittle Pinhero,     Department of Food Science, University of Guelph, Guelph, Ontario, Canada

Mohamed Fawzy Ramadan,     Agricultural Biochemistry Department, Faculty of Agriculture, Zagazig University, Zagazig, Egypt

M.A. Rao,     Cornell University, Geneva, NY, USA

Flor Rodriguez,     International Potato Center (CIP), La Molina, Lima, Peru

Roshani Shakya,     USDA-ARS, Washington State University, Prosser, WA, USA

Jaspreet Singh,     Riddet Institute and Massey Institute of Food Science and Technology, Massey University, Palmerston North, New Zealand

Rekha S. Singhal,     Food Engineering and Technology Department, Institute of Chemical Technology, Mumbai, Maharashtra, India

Paul Smith,     Cargill R&D Centre Europe, Vilvoorde, Belgium

Shrikant A. Survase,     Food Engineering and Technology Department, Institute of Chemical Technology, Mumbai, Maharashtra, India

Gilles Trystram,     AgroParisTech, UMR 1145 Ingénierie Procédés Alimentaires, Group Interaction between Materials and Media in Contact, Massy, France

Jean-Michaël Vauvre

INRA, UMR 1145 Ingénierie Procédés Alimentaires, Group Interaction between Materials and Media in Contact, Massy, France

AgroParisTech, UMR 1145 Ingénierie Procédés Alimentaires, Massy, France

McCain Alimentaire S.A.S., Parc d’entreprises de la Motte du Bois, Harnes, France

Olivier Vitrac

INRA, UMR 1145 Ingénierie Procédés Alimentaires, Group Interaction between Materials and Media in Contact, Massy, France

AgroParisTech, UMR 1145 Ingénierie Procédés Alimentaires, Massy, France

Amanda Waglay,     Department of Food Science and Agricultural Chemistry, McGill University, Ste-Anne de Bellevue, Quebec, Canada

Rickey Y. Yada

Department of Food Science, University of Guelph, Guelph, Ontario, Canada

Food, Nutrition, and Health Program, Faculty of Land and Food Systems, University of British Columbia, Vancouver, British Columbia, Canada

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Foreword

Advances in Potato Chemistry & Technology, Second Edition

Originating in the Andes, where there are over 4000 varieties of native potatoes, the humble potato has been cultivated since at least 5000 BC as a food source for humans.

Today, it is the fourth most important crop in the world after rice, wheat, and maize, with more than a billion people globally eating potatoes as part of their diet on a regular basis. Annual production exceeds 300  million  metric  tonnes in over 100 countries.

Globally, trends in potato growing and consumption are changing. Traditionally in the domain of Europe and the Americas, there has been a dramatic increase in potato production and consumption in Asia and Africa since 1990.

A global population increasing in both size and wealth is predicted to require the production of 70% more food than today by 2050. In this context, the potato’s role in global nutrition will become increasingly critical. To ensure the most efficient use of increasingly scarce natural resources, we must ensure that we understand the functional components within a potato and how to apply them to both food and nonfood applications.

Since its publication in June 2009, the first edition of Advances in Potato Chemistry and Technology has been embraced as a, if not the only, comprehensive and convenient single reference of current developments in the field of potato chemistry.

With advances in the field developing rapidly, the second edition is a comprehensive review that keeps this book at the cutting edge by adding the latest information on:

• Potato flavor: the effects of cooking, texture, and metabolites

• Potato proteins: their extraction, nutritional, and health-promoting properties and their application in the biogeneration of peptides

• Colored potatoes: their chemistry and health-related attributes

• Acrylamide in potato products

• Microstructure, starch digestion, and the glycemic index of potatoes

• Composition differences in organically and conventionally grown potatoes

• Mechanisms of oil uptake in french fries

• Potato proteomics: a new approach for the potato processing industry

• Advanced analytical techniques for quality evaluation in potato and its products

• Novel food and nonfood uses of potato and its by-products

• Potatoes and human health

• Novel methods of potato starch modification

• The role of potatoes in global food security

• Lipids of transgenic potato cultivars

We at Potatoes NZ, Inc., are very proud to be associated with the authors and congratulate them on this very comprehensive extensive and cohesive update to what is already a celebrated contribution to the field.

I have no hesitation in recommending this book to scientists, researchers, academics, and graduate students working in the fields of food chemistry, agronomy, genetics, horticulture, and nutrition.

I’m sure that they will find it an indispensable companion as they seek to deepen their understanding of the potato at the molecular level and develop new applications that will ensure that the humble potato remains a cornerstone of human nutrition into the next millennium.

Champak Mehta,     CEO, Potatoes NZ, Inc.

September 2015

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Chemistry, Processing, and Nutritional Attributes of Potatoes—An Introduction

Jaspreet Singh,  and Lovedeep Kaur,     Riddet Institute and Massey Institute of Food Science and Technology, Massey University, Palmerston North, New Zealand

The potato (Solanum tuberosum L.) has an annual world production exceeding 376  million  metric  tonnes (2013), with China being the top producer (FAOSTAT, 2015). Higher yield per unit area and nutritional value have led to an increase in potato production over past years compared with other tuber crops. In fact, the potato production from developing countries exceeds that of the developed countries (FAO, 2010). The potato plant, a perennial herb belonging to the family Solanaceae, bears white to purple flowers with yellow stamens, and some cultivars bear small green fruits, each containing up to 300 seeds. The potato tuber develops as an underground stem (swollen part of a subterranean rhizome or stolon) bearing auxiliary buds and scars of scale leaves and is rich in starch and storage proteins. Potatoes can be grown from the botanical seeds or propagated vegetatively by planting pieces of tubers. The eyes on the potato tuber surface, which are actually dormant buds, give rise to new shoots (sprouts) when grown under suitable conditions. A sprouted potato is not acceptable for consumption and processing. But optimum sprouting is a desired attribute when the tubers are used for propagation. The production potential of potatoes is quite high, as nearly 80% of the potato plant biomass constitutes economic yield (Osaki et al., 1996).

New cultivars of potatoes with better yield, disease resistance, and desirable end use are being developed with the help of breeding techniques. In the past many years, several potato cultivars with desired yield, dry matter, cooking texture (such as waxy, floury), flesh color, and disease resistance have been developed with the help of breeding. Following the rational development of genetic engineering, many genetically modified potatoes with very high amylose/amylopectin content, antioxidant levels, and tuber yield have also been developed. However, these transgenic varieties of potatoes are not permitted for food use in many countries because of the concerns related to consumer health and the environment. Until these genetically modified potatoes have been given proper clearance by the food authorities and acceptance by the consumers, they may have a good scope for their use in nonfood or other industrial applications.

Morphologically, a potato tuber is usually oval to round in shape, with white flesh and a pale brown skin, although variations in size, shape, and flesh/skin color are also frequently encountered, depending on the genetics of the cultivar. The color, size, and texture of potatoes are the main quality attributes assessed by the consumer for acceptability. Good-quality potatoes are considered to be relatively smooth, firm, and free from sprouts or any other disorders. In a potato tuber, about 20% is dry matter and the rest is water. The yield, the dry matter, and the composition of the dry matter vary among potato cultivars, soil type and temperature, location, cultural practices, maturity, postharvest storage conditions, and other factors (Burton, 1989). Starch is the major component of the dry matter, accounting for approximately 70% of the total solids. The major part of the fresh potato tuber comprises storage parenchyma in which the starch granules are stored as a reserve material. Potatoes are a rich source of high-value protein, essential vitamins, minerals, and trace elements. The average range of raw material composition of a potato tuber is as follows: starch (10–18%) having 22–30% amylose content, total sugars (1–7%), protein (1–2%), fiber (0.5%), lipids (0.1–0.5%), vitamin A (trace/100  g fresh weight, FW), vitamin C (30  mg/100  g FW), minerals (trace), and glycoalkaloids (1–3  mg/100  g FW). The average composition of a potato tuber is presented in Table 1.

In past years, potato breeding programs have targeted only the crop yield and disease resistance; therefore significant gaps exist in the knowledge of the nutrient range, processability, and health-related attributes of the new germplasm. The less well-known constituents of potato tuber are carotenoids and phenolics, which are potent antioxidants. Carotenoid content of potatoes ranges from 50 to 100  μg/100  g FW in white-fleshed cultivars to 2000  μg/100  g FW in deeply yellow- to orange-fleshed cultivars. Potatoes also contain phenolic compounds, predominantly chlorogenic acid, and up to 30  μg/100  g FW of flavonoids in white-fleshed potatoes and roughly 60  μg/100  g FW in red- and purple-fleshed potatoes. The total anthocyanin content of whole unpeeled red- and purple-fleshed potatoes may be around 40  mg/100  g FW (Brown, 2005). The colored potatoes, if processed in a way that does not destroy their anthocyanins and carotenoids, may help in lowering the incidence of several chronic diseases in humans.

Table 1

Average Composition of a Potato Tuber, per 100  g, After Boiling in Skin and Peeling Before Consumption

USDA, National Nutrient Database.

Potato is generally processed through boiling, mashing, frying, etc., before consumption. The influence of the chemical composition of potatoes during processing is of significance to maintain the quality of processed potato products. As an example, the texture of potato crisps is dependent mainly on the starch content of the raw potato tubers. The nonstarch polysaccharides (cell wall) also play a crucial role in determining the quality of the crisps while contributing to the tuber fiber content, also. Potatoes with closely packed small and irregular parenchymatous cells have been observed to be relatively hard and cohesive. In contrast, potatoes with large, loosely packed cells are generally less hard. The cell wall characteristics of cooked potato also play an important role in the release of glucose during starch digestion in our body (Singh et al., 2013).

Starch is the major component of potato dry matter and consists of amylose and amylopectin. The structural characteristics and amylose-to-amylopectin ratio of potato starch vary among cultivars. The nutritional and processing quality of potatoes and potato products (frozen and dry) are greatly affected by their starch characteristics. Several chemical, physical, and enzymatic modifications are performed to improve the processing performance of potato starch. Most of these modifications are listed as generally recognized as safe by the safety authorities. Several modified potato starches with slow digestibility are being developed that may provide nutritional benefits for humans. These starches have the potential to be used for the treatment of certain medical conditions (e.g., glycogen storage disease and diabetes mellitus).

The total sugars in potato tuber range from 1 to 7  g/kg. The reducing sugars (glucose, fructose) are at the highest levels in young tubers and decrease considerably toward the end of the growing season. The starch-to-sugar conversion during postharvest storage also causes variation in the sugar content of tubers, which is an important consideration in the potato crisp industry. In the past few years, advanced analytical and instrumentation techniques have been introduced to evaluate the quality of the potato and its products. These techniques provide in-depth information about the structure and functionality of potato components, which helps to tailor potato products for desirable attributes.

Potato is an essential and safe source of energy and dietary fiber for children and pregnant women. The nutritional characteristics such as starch digestibility, glycemic index, and relative glycemic impact are important in human health. The microstructure of potatoes, whether natural or created during processing/storage, plays an important role during digestion of starch in the gastrointestinal tract and affects the glycemic index of potatoes. The relationships between the composition of potato tubers and its impact on the release of glucose in the blood have been studied by various researchers through in vitro and in vivo methods. The careful selection of suitable potato cultivars, processing techniques, and storage conditions can prove helpful in getting better nutritional benefits from potatoes. New formats of processed potato products suitable for the nutritional needs and taste of various population groups may help to further increase the consumption of potatoes. Apart from food use, potato products are being used for nonfood applications such as biodegradable packaging, fermentation, vaccines, and pharmaceuticals. New applications are being developed for the utilization of potato by-products and waste, which are otherwise an expensive waste management challenge.

References

Brown C.R. Antioxidants in potato. American Journal of Potato Research. 2005;82:163–172.

Burton W.G. The Potato. New York: John Wiley and Sons, Inc.; 1989.

FAOSTAT. Food and Agriculture Organization of the United Nations. 2015. http://www.faostat3.fao.org (accessed 17.09.15.).

FAO. Strengthening Potato Value Chains: Technical and Policy Options for Developing Countries. Rome, Italy: Food and Agricultural Organization of United Nations; 2010.

Osaki M, Matsumoto M, Shinano T, Tadano T. A root-shoot interaction hypothesis for high productivity of root crops. Soil Science and Plant Nutrition. 1996;42(2):289–301.

Singh J, Kaur L, Singh H. Food microstructure and starch digestion. Advances in Food and Nutrition Research. 2013;70:137–179.

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Chapter 1

Potato Origin and Production

Stef de Haan¹,  and Flor Rodriguez²     ¹International Center for Tropical Agriculture (CIAT), Tu Liem, Hanoi, Vietnam     ²International Potato Center (CIP), La Molina, Lima, Peru

Abstract

The extent of the potato gene pool, with its abundant landrace diversity and numerous wild relatives, offers a wide range of options for prospecting, prebreeding, and niche market development. Landraces are still widely produced by smallholder farmers in the center of origin and are a key component of complex production systems that cover multiple agroecologies and production objectives. Sustainable future conservation and use will require attention to systematic monitoring of potato genetic diversity, gap analysis, screening, and prebreeding. Modern and traditional uses are both main drivers behind the rational use of potato biodiversity. Both offer promise for the scientific advancement of potato chemistry and technology.

Keywords

Genebanks; Genetic resources; On-farm conservation; Use systems

1. Introduction

Western South America is the primary center of the origin and diversity of the potato crop and its wild relatives. Contemporary landrace gene pools occur from 45° south in Chile to 12° northern latitude in Colombia (Hawkes, 1990). Wild relatives of the potato (Solanum section Petota; Solanaceae) have a much wider distribution range and occur from northern Patagonia to the southern US and the western Atacama desert to eastern South America (Hijmans et al., 2002; Spooner et al., 2004). The genetic diversity of landraces and wild relatives has been and continues to be an extremely valuable source of variation for genetic enhancement, crop improvement, and understanding of chemical variability (Brown et al., 2007; Hanneman, 1989; Jansky et al., 2013; Osman et al., 1978; Väänänen, 2007). At the same time, ongoing evolution of potato diversity in farmers’ hands is anticipated to allow for adaptation to climate change and continued food security in extreme agroecologies (Johns and Keen, 1986; Zimmerer, 2014).

The case of the potato in its center of origin is special because landraces are still widely grown by smallholder farmers in semitraditional and market-oriented production systems. Predictions regarding landrace loss, genetic erosion, and full-fledged extinction of genetic diversity (e.g., Fowler and Mooney, 1990; Hawkes, 1973; Ochoa, 1975) have not materialized in the Andes, because autonomous farming rationales and multiple innovations such as revaluation of local cuisines, farmers’ markets, and biodiversity seed fairs have renewed local interest in conservation (de Haan et al., 2010b; Monteros, 2011).

Opportunities to screen and evaluate potato genetic resources to explore chemical profiles, determine concentration ranges of (anti)nutritional compounds, and test technological innovations are abundant. The extent of the potato gene pool, with its abundant landrace diversity and numerous wild relatives, offers a wide range of options for prospecting, prebreeding, and niche market development. At the same time, advancements in biotechnology, genomics, analytical techniques, and postharvest technologies open up many new possibilities for the enhanced use of genetic resources (Jo et al., 2014; Singh and Kaur, 2009; Slattery et al., 2010).

2. Origin, Domestication, and Diversity

2.1. Biosystematics and Evolution

The evolutionary origin of the cultivated potato has not yet been conclusively unraveled, and geneticists, archaeobotanists, and taxonomists alike have explored different hypotheses for nearly 9 decades (Spooner et al., 2014). However, at a cultivated species level it is well documented that different species (Solanum tuberosum, Solanum curtilobum, Solanum ajanhuiri, and Solanum juzepczukii) and groups (S. tuberosum Chilotanum and Andigenum groups) are the result of unique evolutionary pathways and have different biogeographical distribution patterns.

Of all cultivated species, the S. tuberosum Chilotanum group (2n  =  4x  =  48) has contributed most to the founder effect of the European and North American gene pool and global crop improvement (Figure 1). According to van der Berg and Groendijk-Wilders (2014), over 99% of European extant modern cultivars possess Chilean cytoplasm. Yet, the level of intraspecific diversity within the S. tuberosum Chilotanum group is modest compared with the Andigenum group. Its contemporary distribution range is basically restricted to the Chiloe Island of south-central Chile (Contreras and Castro, 2008; Manzur, 2012). Two hypotheses are commonly put forth regarding the origin of Chilotanum landraces (Spooner et al., 2012). The first suggests that they originated independently in southern Chile, possibly involving the putative wild ancestor Solanum maglia (Dillehay, 1997; Ugent et al., 1987) or hybrids of Solanum tarijense (Solanum berthaultii) (Hosaka, 2003; Spooner et al., 2014). The second sustains an Andean origin with early introduction into Chile and a gradual adaptation of Andigenum landraces into long-day adapted Chilotanum landraces (Hawkes, 1990, 1999; Salaman, 1946; Simmonds, 1964, 1966).

The S. tuberosum Andigenum group as proposed in the taxonomic treatments of Ovchinnikova et al. (2011) and Spooner et al. (2005a, 2014) includes diploid, triploid, and tetraploid subgroups previously considered separate species by Hawkes (1990) and Ochoa (1990, 1999). The biogeographical distribution of the group spans from northern Chile and Argentina to Colombia. Spooner et al. (2005a, 2014) proposed a single origin of Andigenum landraces from the Solanum brevicaule complex (S. brevicaule and Solanum candolleanum) involving hybridization, natural variation, polyploidization events, and anthropogenic selection. On the other hand, Hawkes (1990) and Ochoa (1990, 1999) previously proposed separate origins based on taxonomies that recognized multiple likely ancestral species such as Solanum ambosinum, Solanum bukasovii, Solanum canasense, Solanum leptophyes, and Solanum sparsipilum within what is now considered the S. brevicaule complex.

Figure 1  Sample of mixed Chilotanum landraces from the island of Castro, Chile. Photo: S. de Haan.

The S. tuberosum Andigenum group contains more intraspecific diversity than all other species and the S. tuberosum Chilotanum group combined (Figure 2). Based on ploidy, ethnobiological classification, altitudinal distribution, and hotspot concentration, several cultivar groups can be distinguished. A clear example of such a group is the Phureja cultivar group, which is vernacularly classified as Phureja (northwestern Bolivia) or Chaucha (Peru), which has differential agronomic characters (early bulking, lack of dormancy, and up to three cropping cycles a year) and is spatially separated by altitude in its distribution range from other cultivar groups within the S. tuberosum Andigenum group (Ghislain et al., 2006; Zimmerer, 1991a,b).

The three species, S. ajanhuiri (2n  =  2x  =  24), S. juzepczukii (2n  =  3x  =  36), and S. curtilobum (2n  =  5x  =  60), each evolved through unique pathways. The distribution of S. ajanhuiri is restricted to the Bolivian-Peruvian altiplano region around Lake Titicaca and evolved through hybridization events between the cultivated diploid Solanum stenotomum (S. tuberosum Andigenum group) and the wild Solanum megistacrolobum (Solanum boliviense) (Huamán et al., 1982; Johns, 1985; Johns and Keen, 1986). Solanum juzepczukii and S. curtilobum are so-called bitter species containing high levels of glycoalkaloids. They are commonly used by Andean farmers for traditional freeze-drying (Figure 3). The biogeographical distribution range of both species spans from southern Bolivia to central Peru. Solanum juzepczukii has commonly been proposed to originate from hybridizations between S. stenotomum and tetraploid Solanum acaule (Hawkes, 1962; Schmiediche et al., 1980). Different possible origins for the pentaploid S. curtilobum have been proposed, involving the tetraploid Andigenum group  ×  S. juzepczukii (Hawkes, 1962, 1990) or the triploid Andigenum group  ×  S. acaule (Gavrilenko et al., 2013; Spooner et al., 2014).

Figure 2  Sample of mixed Andigenum landraces from central Peruvian Andes. Photo: S. de Haan.

2.2. Wild Relatives

The taxonomy of Solanum section Petota is complicated by sexual compatibility among species, interspecific hybridization, auto- and allopolyploidy, a mixture of sexual and asexual reproduction, possible species divergence, and phenotypic plasticity. The resulting complexity causes difficulty in defining and distinguishing species (Camadro et al., 2012; Huamán and Spooner, 2002; Knapp, 2008; Masuelli et al., 2009; Rodríguez et al., 2010; Spooner, 2009; Spooner and van den Berg, 1992). The potato has prezygotic and postzygotic hybridization barriers. Solanum species have been assigned endosperm balance numbers (EBNs) based on their ability to hybridize with each other (Hanneman, 1994; Johnston et al., 1980; Ortiz and Ehlenfeldt, 1992). The EBN is a strong postzygotic crossing barrier. Excluding other crossing barriers, successful hybridization is expected when male and female gametes have matching EBN values, regardless of ploidy. The other mechanism is unilateral incompatibility, the prezygotic hybridization barrier, in which pollen tube elongation is inhibited by stylar tissue but the reciprocal cross, when the self-compatible species is used as the female, is successful (Spooner et al., 2014).

The first modern comprehensive taxonomic treatment of the Petota section was provided by Hawkes (1956) and was followed by treatments prepared by Correll (1962), Hawkes (1963, 1990), Bukasov (1978), Gorbatenko (2006), and Spooner et al. (2014). Until 1990, only morphology was used to define species. A wide range of molecular markers and deoxyribonucleic acid sequences were progressively combined with morphology to obtain better insights into species boundaries. Morphological studies throughout the range of the Petota section showed wide variation of character states within and overlap among closely related species (Spooner et al., 2014). A reclassification by Spooner et al. (2014) recognizes 107 wild species instead of the 228 previously proposed by Hawkes (1990). The latest taxonomic treatments use morphology and molecular markers to reinvestigate species boundaries, combined with the practical ability to distinguish species, following a phylogenetic species concept. The complete list of wild potato species recognized by Spooner et al. (2014) is listed in Table 1.

Figure 3  Farmer from southern Andes of Peru preparing tubers from bitter potatoes for freeze-drying. Photo: C. Fonseca.

The degree of relatedness can also be classified based on Harlan and de Wet’s (1971) gene pool concept. It is based on the degree of hybridization among species and recognizes three gene pools: primary (GP-1), secondary (GP-2), and tertiary (GP-3). The primary gene pool consists of biological species, and crossing within this gene pool is easy. Hybrids are vigorous, exhibit normal meiotic chromosome pairing, and possess total fertility. In the GP-2 pool are wild relatives crossable with some manipulations, and in GP-3 are those species that need radical techniques to allow gene transfer. Bradeen and Haynes (2011), Jansky et al. (2013), and Veilleux and De Jong (2007) have attempted to determine the three potato gene pools, but EBN and other pre- and postzygotic barriers to hybridization have made it difficult to apply the gene pool concept to potatoes (Chen et al., 2004; Jansky et al., 2013; Masuelli and Camadro, 1997). Spooner et al. (2014) proposed five crossability groups based on EBN and self-compatible/self-incompatible systems. They predicted most possible successful crosses and pointed out that whereas hybridization across groups is less likely to be successful than hybridization within groups, no barrier is complete.

2.3. Landraces

Intraspecific diversity of potato is high and the clonal nature of the crop contributes to consistent folk taxonomic classifications of cultivar groups and individual cultivars (Brush, 1980; de Haan et al., 2007; La Barre, 1947). The basic unit of management and consequent conservation for Andean and Chiloe island farmers is the landrace. It is commonly recognized and named. The International Potato Center’s gene bank and most gene banks in the center of origin maintain clonal landraces as well as botanical seed. At the level of use, three landrace groups can be distinguished: (1) commercial or cosmopolitan floury landraces, (2) noncommercial floury landraces, and (3) bitter landraces.

Commercial or cosmopolitan landraces are a selected group of cultivars that enjoy market demand, consumer recognition, or relatively large crop areas (Brush, 2004). In the countries within the center of crop origin, these are among the most abundant landraces (Table 2). Some landraces such as the diploid Peruanita in Peru and Criollo Amarilla in Colombia are grown extensively and are widely available in regular markets and supermarkets alike. Others, such as the tetraploid Mechuñe Roja in Chile or Tuni in Argentina, are available only in specialty restaurants and fairs. Since the early 2000s several value chain projects in Bolivia, Ecuador, and Peru have brought about shifts in the market demand of landraces that were previously noncommercial.

Table 1

Solanum Section Petota Wild Species with Three-Letter Standard Abbreviations, Countries of Occurrence, Ploidy (and EBN), and Gene Pool Assignment.

a Spooner et al. (2014).

b Castañeda-Álvarez et al. (2015).

Table 2

Contemporary Crop Area, Areal Proportion, and Diversity of Landraces in the Center of Origin and Diversity.

Coca Morante (2002), Contreras and Castro (2008), FAO (2013), Iriarte et al. (2009), Monteros (2011), Monteros et al. (2010), Pumisacho and Sherwood (2002), Terrazas et al. (2008), and Ugarte and Iriarte (2000); expert opinion validation by Universidad Nacional de Colombia, CORPOICA, INIAP Ecuador, INIA Peru, INIAF Bolivia.

The bulk of landraces managed by smallholder farmers in the Andes and Chiloe islands are noncommercial and floury (nonbitter). This group consists of thousands of cultivars that are grown almost exclusively for home consumption. Farmers’ reasons for cultivating this diversity consist of multiple complementary livelihood rationales including risk mitigation and yield stability, management options, preference traits, and superior quality in local cuisines, and prestige and cultural identity, among other factors (Brush, 2004; Zimmerer, 1996). Some regions can be considered contemporary landrace hotpots with high levels of diversity, such as Huancavelica (Peru) (CIP, 2006; de Haan et al., 2013), Paucartambo (Peru) (Pérez Baca, 1996; Zimmerer, 1996), northern La Paz (Bolivia) (Iriarte et al., 2009), and northern Potosí (Bolivia) (Terrazas et al., 2008). Yet, whereas other regions may harbor only modest levels of total diversity, this same diversity could consist of unique or endemic landraces.

Native bitter landraces are grown by smallholder farmers in central southern Peru and Bolivia. It concerns a group of landraces that are robust and commonly frost resistant (Christiansen, 1977; Condori et al., 2014). They are almost exclusively used to freeze tubers into chuño, moraya, or tunta (de Haan et al., 2010a). Intraspecific diversity of S. juzepczukii and S. curtilobum consists exclusively of bitter landraces. However, a common misconception is that intraspecific diversity of S. ajanhuiri also consists exclusively of bitter landraces and that tetraploid Andigenum landraces are always nonbitter. Solanum ajanhuiri landraces are commonly nonbitter whereas Andigenum landraces are sometimes bitter and destined for freeze-drying. The diversity of bitter landraces is modest compared with floury landraces, but between Peru and Bolivia they amount to at least 100 distinct cultivars.

3. Production in the Center of Origin

Cropping systems of the potato crop in the center of origin are extremely diverse. They can be differentiated according to their main orientation and agroecological setting. Orientation ranges from small-scale, nonmechanized family farming for household food security to medium- to large-scale mechanized production for the market, and to medium-scale mixed production systems that pursue both self-sufficiency and income generation. Macrolevel potato agroecologies in the region are the central dry Andes (Peru and Bolivia), northern wet Andes (Ecuador to Venezuela), and Southern Cone.

Small-scale family farming is predominant in the region in terms of the number of households that depend on this activity. In Peru alone, more than 600,000 households depend on potato production to support their livelihoods (Table 3). Small-scale family farming (0.1–2  ha/household) is often characterized by the use of animal traction or foot plows, field scattering, relatively high landrace diversity, limited use of external inputs, and the use of semitraditional practices such as zero-tillage cropping (de Haan and Juarez, 2010; Oswald et al., 2009). Medium- to large-scale mechanized production (5–80  ha/household) is frequently intensive and orientated toward the production of bred varieties or cosmopolitan landraces. Although the demand for processing varieties has been growing steadily, the bulk of commercial production is oriented toward the supply of ware potatoes (Scott, 2011a). Medium-scale mixed production systems (2–5  ha/household) that pursue both self-sufficiency and income generation are common in the Andes and southern Chile. Depending on the final destination of the production, either for home consumption or markets, the use of external inputs is often differential.

Table 3

Number of Production Units, Average Yield, Production, and Consumption of Potato at Center of Origin.

a Private sector in Argentina and Chile.

Devaux et al. (2010), FAO (2013), INIA and Red LatinPapa (2012), and Scott (2011b); expert opinion validation by Universidad Nacional de Colombia, CORPOICA Colombia, INIA Venezuela, INTA, and McCain Argentina.

Potato production in the central dry Andes of Peru and Bolivia follows two cropping calendars, the big campaign or qatun tarpuy (November to May), which coincides with the rainy season, and the small or early campaign (miska or maway, June to December). Planting dates are generally the same for bred varieties and landraces, although the harvest of bred varieties tends to be earlier. Potatoes are produced under short-day conditions at altitudes up to 4300  m. Production in the northern wet Andes of Ecuador to Venezuela occurs up to 3300  m altitude under short-day conditions and follows several cropping calendars that are less defined compared with the dry Andes. In southern Chile, potatoes are grown during summer under long-day conditions with main planting for the papa guarda season starting in October and harvests lasting until April.

Seed tubers are predominantly supplied through farmer or informal seed systems (Thiele, 1999). The exception concerns the supply of certified tuber seeds of bred varieties in Chile and Argentina. Climate change and the consequent pressures of alterations in pest and diseases are a threat to seed quality and ware potato production alike. In particular, the altitudinal expansion of potato late blight (Phytophthora infestans) and potato tuber moth (Phthorimaea operculella) pose a threat to the potato in the high Andes (Giraldo et al., 2010; Kroschel et al., 2013).

4. Spread and Global Production Trends Outside the Center of Origin

4.1. Spread of a New World Crop

Botanists, historians, economists, anthropologists, and novelists alike have been interested in the fascinating story involving the spread of the potato from its center of origin to the rest of the world (Figure 4). At best, historical records of the early introduction of potatoes to Europe are sparse (Glendinning, 1983; Oliemans, 1988; Salaman, 1949). Initial fourteenth-century introductions have been shown to be of Andean origin; the introduction from Chile occurred as early as 1811, 34  years before the European late blight (P. infestans) epidemics (Ames and Spooner, 2008). The first written record of potatoes in Europe dates from 1567. The actual document reports a consignment of tubers from Gran Canary Island destined for Belgium (Lobo-Cabrera, 1988; Ríos et al., 2007; Spooner et al., 2005b). Hawkes and Francisco-Ortega (1993) speculated that potatoes were introduced into the Canary Islands as early as 1562. Eleven years later, in 1573, potatoes were reported in continental Spain (Hamilton, 1934; Hawkes and Francisco-Ortega, 1992; Salaman, 1937). The second record of a shipment of potatoes from the Canary Islands destined for France is from 1574 (Hawkes and Francisco-Ortega, 1993; Ríos et al., 2007).

Figure 4  Chronological time series map with some of major events of global potato diffusion.

According to Salaman (1949) potatoes were introduced to England by Sir Francis Drake around 1586, to India by Portuguese sailors in the early seventeenth century, and around the same time to Sri Lanka by the British (Graves, 2001; Nunn and Qian, 2011). The British took potatoes to Bermuda around 1613 and from there to Virginia (United States) in 1621 (Graves, 2001). The potato reached Ireland in the late sixteenth century (Salaman, 1949). Unquestionably, the potato transformed European society because farmers could produce more food in less time and in small plots. This jump boosted population growth and enabled the Industrial Revolution (Reader, 2008). In 1603, potatoes were reportedly taken by Dutch settlers to the Penghu Islands in the Taiwan Strait. Around that same period, toward the end of the Ming dynasty, cultivation also quickly spread throughout China. It is generally accepted that potatoes were first introduced to New Zealand in the late eighteenth century by Captain James Cook and the French explorer Marion du Fresne (Harris and Niha, 1999). Potatoes soon became both a staple crop, with various cultivars receiving Maori names. Introduction of the potato into Africa occurred late; German and English settlers and missionaries introduced it into East Africa in the nineteenth century (Kiple and Ornelas, 2000).

4.2. Production Trends

With a total cropping area of about 20  million hectares globally, the potato is currently the fourth most important staple crop after rice, wheat, and maize. Since the early 1960s, growth in potato production has rapidly overtaken all other food crops in Africa and Asia. In 2005, for the first time, the combined potato production of Africa, Asia, and South America exceeded that of Europe and the United States (Birch et al., 2012; FAO, 2008). Nowadays, roughly half of the global potato cropping area is concentrated in Asia (96  million hectares).

China, India, and Russia are by far the largest producers in the world, with a national production output of 88, 45, and 30  million tons, respectively, registered in 2013, compared with 52, 19, and 9  million tons for the 28 member countries of the European Union, the United States, and the center of crop origin, respectively. Over 3 decades (1981–2011), the total potato cropping area in the Americas and Oceania has remained more or less the stable. However, during that period, Africa and Asia have seen staggering growth, with 300% and 237% increases in total area. In contract, the cropping area in Europe halved during that period.

The global average potato yield is 18.9  metric  tons  per  hectare  (t/ha). The yield gap is particularly high in Africa, where average yields are 14.2  t/ha, compared with 18.3, 21.1, and 25.9  t/ha for Asia, Europe, and the Americas, respectively (FOASTAT, 2013). International and national breeding programs in low- to middle-income countries are focusing on bridging the yield gap, solving classical constraints such as viruses (Potato Virus Y, Potato Leafroll Virus) and late blight (P. infestans) through resistance breeding, as well new challenges. The latter increasingly include an emphasis on abiotic stress (heat, salt, and drought tolerance), quality aspects (micronutrient density and taste profiles), and early bulking and storability. Of course, improved and sustainable management options will have to accompany genetic gains to bridge the yield gap.

A clear trend in selected Asian, African, and Latin American countries concerns the increased investment of governments in potato research, development, and capacity building. Unfortunately, this positive trend is not observed in all low- and middle-income countries where potato is an important staple. Therefore, the divide between strong and weak potato research programs within the same continents remains notably sharp. Yet, potato research in China, India, Rwanda, Brazil, Ecuador, and Chile, among other countries, has benefitted tremendously from consistent government support and targeted research agendas in past decades. This has resulted in the development of nationally adapted expertise and the release of a diverse portfolio of technologies that have been nationally adopted, including bred varieties and seed potato technologies.

In Europe and the United States, more potatoes are processed to meet the rising demand from the fast food, snack, and convenience food industries. Food and Agriculture Organization of the United Nations data reveal that from 2003 to 2007, average shares of exports in total production quantities reached around 7% for the world as a whole (FAO, 2013). The major drivers behind this development include growing urban populations, rising incomes, and the diversification of diets and lifestyles that leave less time for preparing fresh tubers for consumption (FAO, 2008). Walker et al. (1999) highlight that per capita consumption of potato tends to steadily decline as per capita income rises. Yet, the global demand for fries continues to grow and exports from the top 10 exporting countries increased by 71.5% between 2000 and 2010. Demand in Africa, Asia, and South America also continues to expand but internal supply is relatively low, thus fueling imports from Europe, Oceania, and the United States.

International trade is dominated by french fries, starch, and seed potatoes. Developing countries are still net importers in the international potato trade. However, multinational and national processing companies have been expanding their operations and processing capacity in Asia and Africa. Thus, it can be foreseen that the internal supply of potato for processing and convenience food industries will increase in countries where the crop is currently a staple food (Bradshaw and Ramsay, 2009; Kirkman, 2007). China has been the world’s largest potato producer since 1993 and has accounted for more than 80% of the increase in global potato production from 1990 to 2002 (Wang and Zhang, 2004). The growth in China’s potato industry has yet not changed global markets and the country remains a modest exporter. China’s annual exports of fresh potatoes has held stable at 300,000–350,000  metric  tons since 2005, with Malaysia, Vietnam, and Russia being the most important importing countries.

Despite its importance as a staple food and in combating hunger and poverty, the potato has been largely neglected in agricultural development policies for food crops. This may be partly related to the image of the potato as a fast food contributing to rising levels of obesity in high-income countries. In this sense, an important new trend relates to the increased effort of the global potato community, including industry, to highlight the multiple health benefits of the potato when it is not fried. The Potato goodness unearthed campaign by the US Potato Board and the Potatoes are healthy message by HZPC in Holland are two examples of this important effort. Behind it is a growing body of research that provides an evidence base for the positive role that potatoes have in providing nonfat-derived energy, vitamin C, micronutrients, carotenoids, and antioxidants (Burgos et al., 2007, 2009a,b; Graham et al., 2007; King and Slavin, 2013; Nesterenko and Sink, 2003; Singh and Kaur, 2009; Vinson et al., 2012).

5. Conservation in the Center of Origin

5.1. Ex Situ Conservation

Ex situ conservation involves conserving components of biological diversity outside their natural habitat (UNCED, 1992) in botanical gardens, gene banks, or captive breeding programs. Ex situ conservation targets collections of cultivars of important food crops in gene banks, including old landraces and modern varieties, interspecific hybrids, breeding clones, and crop wild relatives (GCDT, 2006). Germplasm conserved ex situ is ideally demanded by users who need ready access for genetic enhancement, crop improvement, varietal selection, or rehabilitation of in situ reserves (Cohen et al., 1991).

In the center of the origin of the potato, at least eight important potato collections can be identified (Table 4). Together, these maintain approximately 19,777 accessions (GCDT, 2006; personal communication with curators). Potato genetic resources are largely conserved as botanical seeds (wild species) or vegetatively as tubers and in vitro plantlets (cultivated species and breeding stocks). Landraces form the largest group of accessions, followed by wild species. There is expected to be a high level of duplication among gene banks in the center of origin.

The International Potato Center (CIP) (Peru) is the custodian of the world’s largest in vitro field and true seed collection of potato, with approximately 4461 landrace accessions from all cultivated species and 2520 accessions of wild relatives of cultivated potato (Castañeda-Álvarez et al., 2015). The genetic diversity conserved at CIP, and related information about each accession, are globally available through the Multilateral System designed by the International Treaty on Plant Genetic Resources for Food and Agriculture.

Table 4

Composition and Size of Potato Collections in Latin America.

a GCDT (2006).

b CORPOICA (personal communication).

c INIAF Bolivia (personal communication).

d INIA Chile (personal communication).

e Breeding lines, hybrids, etc.

Important potato germplasm collections in Latin America are maintained at national gene banks. These include the Instituto Nacional de Tecnología Agropecuaria (INTA) (Argentina), Corporacion Colombiana de Investigacion Agropecuaria (CORPOICA) (Colombia), Universidad Nacional de Colombia (UNC) (Colombia), Instituto Nacional de Innovación Agropecuaria y Forestal (INIAF) (Bolivia), Instituto de Investigaciones Agropecuarias (INIA) (Chile), Universidad Austral de Chile (UACH) (Chile), Instituto Nacional de Investigaciones Agropecuarias (INIAP) (Ecuador), Centro Nacional de Recursos Genéticos–Instituto Nacional de Investigaciones Forestales, Agrícolas y