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Solanum tuberosum (Potatoes)

DM Spooner, USDA Agricultural Research Service, University of Wisconsin, Madison, WI, USA

© 2013 Elsevier Inc. All rights reserved.

This article is a revision of the previous edition article by K Schuler, volume 4, pp 18481850, © 2001, Elsevier Inc.

Glossary

Glycoalkaloid A bitter chemical compound with a combination of a glycoside and an alkaloid, which is present in potatoes and some other plants, for example, solanine. Landrace Native varieties grown by indigenous peoples that are not improved by modern breeding methods. Pedigree The breeding history showing the parents of a cultivar. Ploidy The number of chromosome sets which in potato range from diploid (two sets of 12 chromosomes for a total of 24), triploid (three sets, 36), tetraploid (four sets, 48), pentaploid (five sets, 60), to hexaploid (six sets, 72). Polygenetic A character formed through the interaction of many genes.

Protoplast fusion A nonsexual method of joining genes of different organisms by fusing the protoplasts (cell constituents without the cell wall) from different cells. Rhizome An underground stem, which in potatoes have tubers positioned at the end. Sexual hybridization A method of joining genes of different organisms through the sexual process, which in potatoes and other flowering plants involve transfer of pollen (containing male gametes) to styles (containing female gametes). Somatic hybridization A method of joining genes of different organisms through any of the variety of nonsexual processes, such as protoplast fusion. Tuber A thickened and short underground stem having numerous buds or eyes and used for food storage.

Introduction

Potato is the fourth most important food crop worldwide. In 2009, the world potato area was 18 651 838 ha, and the amount produced was 325 302 445 tons. The major producing countries (as assessed by million tons grown; 2009 data) are China (73 281 890 million tons), India (34 391 000), Russian Federation (31 134 000), United States of America (19 569 100), and Germany (11 617 500). Potatoes grow under a wide variety of climates. The potato yield averages 176 702 hg ha 1 worldwide, but varies widely by country, ranging from <40 000 hg ha 1 in some developing countries to >450 735 hg ha 1 in developed countries. The potato is food for both humans and animals, and raw material for the food processing (e.g., potato chips, French fries, and dried potatoes) and starch industries. Substantial advantages of the potato are its high yield potential in a short growth time, the high edible dry matter content of its tubers, and its high dietary value as a staple food. For example, an average size raw potato contains only about 115 calories, 3.2 g of protein, 80 mg of phosphorus, 1 mg of iron, and 30 mg of vitamin C (about one-half the amount of an average-sized orange). Boiled potatoes have a similar nutrient composition but lose some vitamin C. Baked potatoes contain about 25% more solids and proportionately higher levels of all nutrients than raw or boiled potatoes. The yield from 0.4 ha (1 acre) of potatoes meets both the energy and protein requirements for over 10 people, with a better balance of nutrients than corn, rice, wheat, and soybeans. A good balance of essential nutri­ ents is met with supplements of foods high in calcium and vitamin A. Wild potatoes are native from the southwestern United States to south-central Chile, with centers of species diversity in central Mexico and in the central Andes of Peru and Bolivia. The taxonomy of wild and cultivated potatoes has

changed considerably in recent years based on extensive field work and studies on morphology and genetics of collections obtained from genebanks. Fo r example, in a manuscript from 1990 Hawkes states that there were over 220 wild potato species (tuber-bearing species of the huge genus Solanum ) and seven cultivated potato species, but recent estimates are 100 110 wild species and four cultivated species. Chromosome numbers in wild species vary from diploid (2 n = 2 x = 24), triploid (2 n = 2 x = 36), tetraploid (2 n = 4 x = 48), pentaploid (2 n = 5 x = 60), to hexaploid (2 n = 6 x = 72). The cul­ tivated potato has all these ploidy levels, except hexaploid. Cultivated potatoes can be classified as landraces that are native varieties still grown in South America today, or improved varieties that are grown around the world. Potato landraces originated from a group of closely related wild species in southern Peru, and were rapidly diffused into two areas, one group in a broad swath of the upland Andes from western Venezuela south to northern Argentina and a second group in the lowlands of south-central Chile. Our modern cultivars that are grown worldwide come from the Chilean landraces. The edible part of the potato plant is called a tuber, and is actually a modified underground stem, not a root. While tubers of most of the landraces are large, palatable, and grow near the plant, those of many of the wild species are often no larger than the size of a pea or hen s egg and can be quite bitter and poisonous due to the presence of chemicals called glycoalka­ loids. The tubers of the wild species can grow on thin underground stems (rhizomes) up to some distance away from the plant, but close to the plant in the cultivated species. While high concentrations of glycoalkaloids may confer protec­ tion against predation in the wild, they are a hindrance to potato breeders, who must ensure that they are greatly reduced in advanced cultivars.

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Solanum tuberosum (Potatoes)

History

Perhaps 6000 years ago in the Central Andes of what is today southern Peru and northern Bolivia, native peoples began to select wild potato species for human use. According to Hawkes, who outlined a taxonomic system still used by some today, there are seven cultivated species: Solanum ajanhuiri (2x ), Solanum chaucha (2 x), Solanum curtilobum (5x ), Solanum juzepc­ zukii (3x ), Solanum phureja (2x), Solanum stenotomum (2 x), and Solanum tuberosum (4x ) with the latter containing subspecies andigenum and subsp. tuberosum . However, combined morpho­ logical studies of Huamán and Spooner and molecular studies of Spooner et al. documented that not all of these seven are good species, and the current taxonomy by Spooner et al. recognizes only four species: S. tuberosum with Andean and Chilean landrace groups (Andigenum Group and Chilotanum Group), S. ajanhuiri, S. curtilobum , and S. juzepczukii . The most widely grown landrace is S. tuberosum Andigenum Group and the other landraces are not as commonly grown. Potato first appeared outside of South America in Europe in 1567 and rapidly diffused worldwide. Our modern cultivars of S. tuber­ osum originated from the Chilean landraces. The Irish potato famine caused by potato late blight disease, Phytophthora infes­ tans , caused widespread famine and migration in Europe beginning in 1845. Late blight remains one of the most serious potato diseases worldwide, but modern breeding and chemical controls afford protection against this disease.

Potato Breeding

The potato is threatened by numerous pests and pathogens, which makes resistance breeding so important. Both the land- races and wild potato species have been extensively collected and are maintained in a series of genebanks worldwide, and are used by potato breeders to incorporate a variety of disease resistances as well as traits to improve growth in harsh environ­ ments such as heat, cold, and drought, and quality traits such as yield or improved potato chipping qualities. Radcliffe men­ tions that more than 150 species of insects attack potatoes in North America alone, and Johnson et al. list the following potato diseases affecting potato in North America alone:

bacteria ’ – bacterial brown rot, Ralstonia solanacearum ; bacterial soft rot, Pectobacterium carotovorum var. carotovora ; blackleg, Pectobacterium carotovorum var. atrosepticum ; common scab, Streptomyces scabies, Streptomyces spp.; and ring rot, Clavibacter michiganensis subsp. sepedonicus. Chromista late blight, Phytophthora infestans ; leak, Pythium debaryanum, Pythium spp.; and pink rot, Phytophthora erythroseptica, Phytophthora spp. Fungus ’ – blackdot, Colletotrichum coccodes ; dry rot, Fusarium spp.; early blight, Alternaria solani, Alternaria alternata; Fusarium wilt, Fusarium solani var. eumartii, Fusarium oxysporum; Rhizoctonia, Rhizoctonia solani ; silver scurf, Helminthosporium solani; Verticillium wilt, Verticillium spp.; and white mold, Sclerotinia sclerotiorum . Nematode ’ – Columbia root knot, Meloidogyne chitwoodi ; golden, Globodera rostochiensis ; potato tuber rot, Ditylenchus destructor ; root knot, Meloidogyne spp.; and poot lesion or meadow, Pratylenchus penetrans. Plasmodium ’ – powdery scab, Spongospora subterranean , and wart, Synchytrium endobioticum. Phytoplasma ’ – purple top

wilt. Virus ’ – Calico; corky ring spot; leaf roll; mop top; PVA; PVM; PVY; PVS; PVX; and rugose mosaic (PVX + PVY). For a long period after the potato was introduced into North America, little effort was made to improve it or to introduce new kinds. The period from 1719 to 1850 was characterized by no marked or lasting improvement in the crop. Improvement of potatoes near the end of this period became imperative because the available cultivars ran outto the extent that yields decreased to low levels and production was uneconomical. Running out is caused by increasing levels of tuber-transmitted virus diseases in existing stocks and lack of proper seed (potato tuber planting stock) maintenance meth­ ods. In 1851, C. E. Goodrich, a clergyman of Utica, NY, introduced a small amount of potatoes received from the American consulate in Panama. One of them, Rough Purple Chili, was one of the most valuable plant introductions in history and continues to have a tremendous impact on the North American potato industry. It contributed 100% of the pedigree of the main US potato cultivar, Russet Burbank, and an average of about 25% of the pedigrees of the 10 most important cultivars grown in North America. Since 1925, plant explorations have been made in the United States, Mexico, Central America, and South America in search of new wild species and landrace cultivars of potato for use in improv­ ing commercial types. In addition, many named cultivars of potato have been introduced into genebanks worldwide for use in commercial breeding. Potato introductions and their exploitation are especially important because: (1) the potato

is the most economically important vegetable in the world,

(2) there is a broad array of easily accessible-related germplasm, (3) solutions to most of the present limitations

of the potato have been demonstrated to exist in related germ-

plasm, and (4) potato has a great potential to provide nutritious food in a diversity of environments for an increas­

ingly hungry world. The ability to make crosses between cultivated species and many of the wild species allows use of a vast germplasm resource useful for potato improvement. Although potato has

a series of mechanisms that inhibit crossing among many

cultivated and wild species, these can be overcome to various degrees by manipulation of chromosome numbers and other mechanisms that allow widespread germplasm transfer for

breeding programs. Prebreeding at the diploid level makes

the interpretation of genetic segregation and selection of poly­ genetic traits easier. Besides the classic sexual hybridization, there is the possibility of combining genomes asexually by protoplast fusion. This somatic hybridization can be applied

to species that are impossible or very difficult to cross sexually

(e.g., S. bulbocastanum S. tuberosum subsp. tuberosum ).

See also: Somatic Mutation.

Further Reading

Ames M and Spooner DM (2008) DNA from herbarium specimens settles a controversy about origins of the European potato. American Journal of Botany 95: 252 257. Emsinger AH, Emsinger ME, Konlande JE, and Robson JRK (1994) Foods & Nutrition Encyclopaedia , 2nd edn. Boca Raton, FL: CRC Press. Food and Agriculture Organization (FAO) (2009) FAOSTAT. http://faostat.fao.org (accessed 12 October 2011).

Solanum tuberosum (Potatoes)

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Hanneman RE, Jr. (1989) The potato germplasm resource. American Potato Journal 66:

655 667.

Hawkes JG (1990) The Potato: Evolution, Biodiversity and Genetic Resources . London:

Belhaven Press. Hijmans RJ and Spooner DM (2001) Geographic distribution of wild potato species. American Journal of Botany 88: 2101 2112. Huamán Z and Spooner DM (2002) Reclassification of landrace populations of cultivated potatoes ( Solanum sect. Petota ). American Journal of Botany 89: 947 965. Jansky SH (2000) Breeding for disease resistance in potato. Plant Breeding Reviews 19: 69155. Johnson SB, Stevenson W, and Miller J (2010) Disease control. In: Bohl WH and Johnson SB (eds.) Commercial Potato Production in North America , 2nd edn., pp. 6772. The Potato Association of America Handbook, Supplement Vol. 57 of USDA Handbook 267. Orono, ME: The Potato Association of America. http:// potatoassociation.org/documents/A_ProductionHandbook_Final.pdf Ortiz R, Simon P, Jansky S, and Stelly D (2009) Ploidy manipulation of the gametophyte, endosperm, and sporophyte in nature and for crop improvement: A tribute to Prof. Stanley J. Peloquin (1921 2008). Annals of Botany 104: 795 807. Radcliffe EB (2010) Insect control. In: Bohl WH and Johnson SB (eds.) Commercial Potato Production in North America , 2nd edn., pp. 6467. The Potato Association of America Handbook, Supplement Vol. 57 of USDA Handbook 267. Orono, ME: The Potato Association of America. http://potatoassociation.org/documents/ A_ProductionHandbook_Final.pdf

Spooner DM (2009) DNA barcoding will frequently fail in complicated groups: An

example in wild potatoes. American Journal of Botany 96: 1177 1189. Spooner DM (2010) Botany of the potato; morphology and anatomy; plant introduction and maintenance. In: Bohl WH and Johnson SB (eds.) Commercial Potato Production in North America , 2nd edn., pp. 4 7. The Potato Association of America Handbook, Supplement, Vol. 57 of USDA Handbook 267. Orono, ME:

The Potato Association of America. http://potatoassociation.org/documents/ A_ProductionHandbook_Final.pdf Spooner DM, McLean K, Ramsay G, Waugh R, and Bryan GJ (2005) A single domestication for potato based on multilocus AFLP genotyping. Proceedings of the National Academy of Sciences of the United States of America 102:

14694 14699.

Spooner DM, Núñez J, Trujillo G, et al. (2007) Extensive simple sequence repeat genotyping of potato landraces supports a major reevaluation of their gene pool structure and classification. Proceedings of the National Academy of Sciences of the United States of America 104: 19398 19403.

Relevant Websites

http://cipotato.org International Potato Center. http://www.ars-grin.gov NRSP-6: United States Potato Genebank.