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Principles of Plant Genetics and Breeding

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510 CHAPTER 30<br />

short, day-neutral genotypes for use in temperate regions,<br />

starting with 1,500 accessions. A large part <strong>of</strong> this material<br />

is also maintained in long-term storage at the National<br />

Seed Storage Laboratory, Fort Collins, Colorado.<br />

Germplasm resources<br />

In the USA, sorghum germplasm is maintained at<br />

the Regional <strong>Plant</strong> Introduction Station, Experiment,<br />

Georgia, with duplicate samples stored at the National<br />

Seed Storage Laboratory, Fort Collins. A more comprehensive<br />

collection with accessions in excess <strong>of</strong> 25,000 is<br />

maintained at the International Crops Research Institute<br />

for the Semi-Arid Tropics (ICRISAT) in India.<br />

Cytogenetics<br />

Sorghum comprises three known ploidy levels: x = 5,<br />

x = 10, <strong>and</strong> x = 20. Important sorghum species are:<br />

S. bicolor (L.) Moench (2n = 2x = 20) S. propinquun<br />

(Kunth) Hitche (2n = 2x = 20), <strong>and</strong> S. halepense (L.)<br />

Pers. (2n = 4x = 40)<br />

Of these three, the most important to crop production<br />

is S. bicolor. Grassy species such as S. arundinuceuin,<br />

S. verticillijlorum, <strong>and</strong> S. aethiopicum have the<br />

same chromosome number (2n = 2x = 20), <strong>and</strong> can be<br />

crossed with S. bicolor. Sorghum has also been successfully<br />

crossed with sugarcane <strong>and</strong> corn.<br />

Cultivated races are readily crossable with each other<br />

to produce fertile hybrids. In sorghum production,<br />

natural interspecific crossing between S. bicolor <strong>and</strong><br />

S. halepense is a common source <strong>of</strong> <strong>of</strong>f-types in the field.<br />

Two types <strong>of</strong> products arise from this type <strong>of</strong> cross –<br />

sterile plants with 30 chromosomes <strong>and</strong> fertile plants<br />

with 40 chromosomes, the former <strong>of</strong>ten persisting in<br />

the field as perennial weeds.<br />

Cytogenetic male sterility (CMS) <strong>and</strong> fertilityrestoration<br />

systems have been discovered in sorghum<br />

<strong>and</strong> are used in hybrid sorghum production. The CMS<br />

in sorghum genotypes was developed by backcrossing<br />

chromosomes <strong>of</strong> kafir into the cytoplasm <strong>of</strong> milo.<br />

Similarly, genetic male sterility (ms) has been discovered<br />

in sorghum male-sterile plants (Msms).<br />

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

Midrib color is controlled by a single dominant gene<br />

(D), whereas resistance <strong>of</strong> hydrogen cyanide (HCN)<br />

appears to be controlled by more than one factor.<br />

Researchers have determined that grain color in sorghum<br />

is influenced by pericarp color, pericarp thickness,<br />

presence <strong>of</strong> testa, color <strong>of</strong> testa, endosperm color,<br />

glume color, <strong>and</strong> plant color. Each <strong>of</strong> these characteristics<br />

is determined by one or a few genes. For example,<br />

two genes, R <strong>and</strong> Y, determine whether the pericarp<br />

will be red (RY ), colorless or white (Ry, ry), or lemon<br />

yellow (rY ). The kernel starch is conditioned by a dominant<br />

allele (Wx); wxwx conditions a waxy endosperm.<br />

Similarly, sugary endosperm is controlled by a single<br />

locus, susu.<br />

Genetic male-sterility genes occur in sorghum, the<br />

most commonly used being the ms 3 . It has stable expression<br />

over different environments. CMS also occurs in<br />

sorghum, conditioned by an interaction between two<br />

major genes, msc 1 <strong>and</strong> msc 2 .<br />

Important genes that have impacted sorghum breeding<br />

are those affecting maturity <strong>and</strong> plant height.<br />

Sorghum is a short-day plant. Maturity is influenced<br />

by photoperiod <strong>and</strong> temperature. Genes that influence<br />

sorghum maturity are designated Ma 1 , Ma 2 , Ma 3 , <strong>and</strong><br />

Ma 4 ; tropical cultivars being dominant at the Ma 1 locus.<br />

A genotype <strong>of</strong> Ma 1 Ma 2 Ma 3 Ma 4 takes about 90.5 days<br />

to anthesis, while ma 1 ma 2 ma 3 Ma 4 takes 55.3 days to<br />

anthesis. Similarly, four independent recessive genes,<br />

dw 1 , dw 2 , dw 3 , <strong>and</strong> dw 4 , reduce internode length <strong>of</strong> the<br />

sorghum stalk without affecting time <strong>of</strong> blooming <strong>and</strong><br />

leaf size. A genotype Dw 1 Dw 2 Dw 3 Dw 4 has a flag leaf<br />

at 127 cm, while dw 1 dw 2 dw 3 dw 4 has a flag leaf located<br />

43 cm above ground on the stalk. Sorghum cultivars<br />

containing dwarf genes are identified by the number <strong>of</strong><br />

specific dwarf genes they contain as 1-dwarf (contains<br />

one dwarf gene, e.g., dw 2 ), 2-dwarf (contains two<br />

dwarf genes, e.g., dw 2 dw 3 ), <strong>and</strong> so on. US commercial<br />

sorghum hybrids are 3-dwarf.<br />

General botany<br />

Sorghum (Sorghum bicolor (L.) Moench) is known by<br />

common names such as milo, kafir, <strong>and</strong> guinea corn.<br />

The annual sorghums have n = 10 <strong>and</strong> include grain<br />

sorghum, sorgo, broomcorn, <strong>and</strong> sudangrass. Sorghum<br />

halepense (johnsongrass) is a perennial sorghum with<br />

2n = 20. The sorghum plant has culms that may st<strong>and</strong><br />

0.6–4.5 m tall, depending on the type <strong>and</strong> variety. It<br />

may produce two or more tillers. The stalk is solid. The<br />

center <strong>of</strong> the stem can be dry or juicy, insipid or sweet<br />

to taste. A dry stalked variety has leaves with a white or<br />

yellow midrib, while a juicy stalked variety has a dull

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