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Principles of Plant Genetics and Breeding

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520 CHAPTER 31<br />

the US from China, India, Manchuria, Korea, Taiwan,<br />

<strong>and</strong> Japan. In 1852, J. J. Jackson is reported to have first<br />

planted soybean as an ornamental plant in Davenport,<br />

Iowa. Most <strong>of</strong> the production <strong>of</strong> soybean in the US<br />

prior to the 1920s occurred in the southern states,<br />

mostly for hay, <strong>and</strong> then spread to the Corn Belt after<br />

about 1924.<br />

Soybean is a subtropical plant. However, it is grown<br />

over a wide range <strong>of</strong> ecological zones, ranging from the<br />

tropics to 52°N. Its climatic requirements are similar to<br />

those <strong>of</strong> corn.<br />

History <strong>of</strong> breeding in the USA<br />

Soybean was introduced into the USA in about the late<br />

1700s. Initially, the crop was grown primarily as a forage<br />

species. The early introductions had weedy characteristics.<br />

They also shattered pr<strong>of</strong>usely, making them unsuitable<br />

for mechanized production. Producing soybean<br />

for seed started in the 1920s. Formal soybean research<br />

was initiated as a joint activity between the US Department<br />

<strong>of</strong> Agriculture (USDA) research wing <strong>and</strong> the<br />

Agricultural Experiment Stations <strong>of</strong> l<strong>and</strong> grant universities<br />

in the midwest <strong>and</strong> southern states. In 1936, a cooperative<br />

state <strong>and</strong> federal program was established at the<br />

University <strong>of</strong> Illinois, Champaign/Urbana, to conduct<br />

soybean research. This program is now known as the US<br />

Regional Soybean Laboratory.<br />

Seed companies became involved in soybean breeding<br />

as a result <strong>of</strong> the United States <strong>Plant</strong> Variety Act <strong>of</strong><br />

1970. With this protection against unauthorized use <strong>of</strong><br />

proprietary material, seed companies could make money<br />

developing <strong>and</strong> selling improved seed. Modern cultivars<br />

were developed for seed, shifting the emphasis away from<br />

soybean as a forage crop to soybean as an oilseed crop.<br />

Cultivars<br />

Soybean may be produced for forage, the varieties for<br />

this purpose generally being small seeded, finer stemmed,<br />

<strong>and</strong> very leafy. There are also cultivars for edible, dry,<br />

or green-shelled beans. These cultivars usually have<br />

straw-yellow or olive-yellow seed <strong>and</strong> a yellow, brown,<br />

or black hilum. Soybean grown for grain is grouped into<br />

13 maturity classes, ranging from 000 to X. The 000<br />

group consists <strong>of</strong> the earliest maturing cultivars while<br />

the X group consists <strong>of</strong> the latest maturing cultivars.<br />

Groups 000–IV are considered indeterminate, while<br />

groups V–X are determinate cultivars. Further, early<br />

maturing cultivars (000–IV) are adapted to the northern<br />

production regions, while those with high maturity<br />

class designation are adapted to southern production<br />

regions. The 000 cultivars are adapted to the short<br />

summer growing seasons <strong>of</strong> northwest USA <strong>and</strong> Canada.<br />

Cultivars in groups II <strong>and</strong> III are best adapted to the<br />

midwest growing area. Maturity groups VIII <strong>and</strong> higher<br />

are grown in the southern or coastal plain counties.<br />

Germplasm resources<br />

The USDA maintains about 15,000 accessions <strong>of</strong> G. max<br />

<strong>and</strong> a smaller number <strong>of</strong> other Glycine species at Urbana,<br />

Illinois, <strong>and</strong> Stoneville, Mississippi. Soybean accessions<br />

are also held by the Applied <strong>Genetics</strong> <strong>of</strong> Korea Atomic<br />

Energy Research Institute in Seoul, Korea, the National<br />

Institute <strong>of</strong> Agricultural Sciences at Hiratsuka, Japan,<br />

<strong>and</strong> the Asian Vegetable Research <strong>and</strong> Development<br />

Center, Taiwan.<br />

Cytogenetics<br />

The genus Glycine has two subgenera – Soja <strong>and</strong> Gycine.<br />

The subgenus Soja consists <strong>of</strong> two species: G. max (2n =<br />

2x = 40), the cultivated species, <strong>and</strong> G. soja (L.) Sieb<br />

or G. ussuriensis (2n = 2x = 40), a wild species. These<br />

two species are cross-fertile. There are 15 wild species <strong>of</strong><br />

soybean <strong>of</strong> which G. tabacinia <strong>and</strong> G. tomentella have<br />

polyploidy forms (including 2n = 4x = 80).<br />

<strong>Genetics</strong><br />

Several hundreds <strong>of</strong> genes for qualitative traits have<br />

been identified for soybean. Among them are four recessive<br />

genes for genetic male sterility, designated ms 1 , ms 2 ,<br />

ms 3 , <strong>and</strong> ms 4 . Linkage groups have been identified for<br />

13 <strong>of</strong> the 20 soybean chromosomes. Soybean has pigmentation<br />

on various parts <strong>of</strong> the plant. The black <strong>and</strong><br />

brown seed coat <strong>and</strong> hilum color are conditioned by two<br />

gene pairs, Tr <strong>and</strong> Rr. The expression <strong>of</strong> these genes is<br />

modified in some conditions by brow seed coat (TrO),<br />

which is dominant over reddish-brown color (Tro), <strong>and</strong><br />

green seed coat (G), which is dominant over yellow seed<br />

coat (g).<br />

Soybean may have either brown (tawny) or gray<br />

pubescence, the trait being conditioned by a single gene

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