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.

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 ha1 worldwide,
but varies widely by country, ranging from <40 000 hg ha1 in
some developing countries to >450 735 hg ha1 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

Brenners Encyclopedia of Genetics, 2nd edition, Volume 6

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.

changed considerably in recent years based on extensive field

work and studies on morphology and genetics of collections
obtained from genebanks. For 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 100110 wild species and four cultivated
species.
Chromosome numbers in wild species vary from diploid
(2n = 2x = 24), triploid (2n = 2x = 36), tetraploid (2n = 4x = 48),
pentaploid (2n = 5x = 60), to hexaploid (2n = 6x = 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 hens 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.

doi:10.1016/B978-0-12-374984-0.01442-X

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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 (2x), Solanum curtilobum (5x), Solanum juzepc
zukii (3x), Solanum phureja (2x), Solanum stenotomum (2x), and
Solanum tuberosum (4x) with the latter containing subspecies
andigenum and subsp. tuberosum. However, combined morpho
logical studies of Huamn 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 landraces 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 purpletop

wilt. Virus Calico; corky ring spot; leaf roll; moptop; 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 out to 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 germplasm, 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: 252257.
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)

Hanneman RE, Jr. (1989) The potato germplasm resource. American Potato Journal 66:
655667.
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: 21012112.
Huamn Z and Spooner DM (2002) Reclassification of landrace populations of cultivated
potatoes (Solanum sect. Petota). American Journal of Botany 89: 947965.
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 (19212008). Annals of Botany 104: 795807.
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

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

example in wild potatoes. American Journal of Botany 96: 11771189.
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. 47. 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:
1469414699.
Spooner DM, Nez 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: 1939819403.

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