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Principles of Plant Genetics and Breeding

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encloses the s<strong>of</strong>t starch at the center. The kernels are<br />

smooth, hard, <strong>and</strong> usually rounded at the top. This<br />

type <strong>of</strong> corn is grown widely in Europe, Asia, Central<br />

America, <strong>and</strong> South America. It is less widely grown<br />

in the US.<br />

3 Flour corn (Z. mays amylacea). As the name implies,<br />

flour corn consists almost entirely <strong>of</strong> s<strong>of</strong>t starch,<br />

making the kernels s<strong>of</strong>t. It has the shape <strong>of</strong> dent<br />

corn but shrinks uniformly upon drying. It is grown<br />

in the drier sections <strong>of</strong> the USA, mainly by American<br />

Indians, <strong>and</strong> also in the Andean region <strong>of</strong> Central<br />

<strong>and</strong> South America. Different kernel colors exist, the<br />

most common being white, blue, <strong>and</strong> variegated.<br />

4 Popcorn (Z. mays everta). Popcorn is an extreme<br />

form <strong>of</strong> flint corn. It has a very hard corneous<br />

endosperm with only a small portion <strong>of</strong> s<strong>of</strong>t starch.<br />

The kernels are characteristically small <strong>and</strong> may either<br />

be pointed or have a rounded tip. Different colors<br />

exist, most corneous varieties being yellow or white.<br />

The kernel pops upon heating as a result <strong>of</strong> the<br />

unique quality <strong>of</strong> the endosperm that makes it resist<br />

the steam pressure generated, until it reaches explosive<br />

proportions.<br />

5 Sweet corn (Z. mays saccharata). This corn is characterized<br />

by a translucent <strong>and</strong> wrinkled appearance<br />

upon drying, <strong>and</strong> a sweet taste when immature.<br />

St<strong>and</strong>ard sweet corn is a mutant <strong>of</strong> the dent corn with<br />

a mutation at the sugary (sy) locus. This mutation<br />

causes the endosperm to accumulate about two times<br />

more sugar than field corn. New mutants have been<br />

developed – sugary enhanced (se) <strong>and</strong> shrunken-2<br />

(sh2) or supersweet corn. Some sweet corn varieties<br />

are unable to convert sugar to starch. Sweet corn is<br />

grown as a winter crop in the southern US, especially<br />

Florida.<br />

6 Waxy corn. Waxy corn has a uniformly dull appearance.<br />

Instead <strong>of</strong> amylose, the starch <strong>of</strong> waxy corn<br />

consists <strong>of</strong> amylopectin, the result <strong>of</strong> waxy (wx)<br />

mutation. Ordinary corn consists <strong>of</strong> about 78% amylopectin<br />

(a high molecular weight branched chain)<br />

<strong>and</strong> 22% amylose (a low molecular weight straight<br />

chain).<br />

7 Pod corn (Z. mays tunicata). Pod corn has primitive<br />

features, each kernel being enclosed in a pod or husk,<br />

before the entire ear is enclosed in husks like other<br />

corns. Pod corn versions <strong>of</strong> the other types <strong>of</strong> corn<br />

(e.g., flint pod corn, dent pod corn) exist.<br />

Similarly, corn that is indigenous to the USA (excluding<br />

sweet corn <strong>and</strong> popcorn) may be classified in up<br />

to nine or 10 races. Of these, the most important are<br />

the Corn Belt dents, the southern dents, <strong>and</strong> the northern<br />

dents.<br />

BREEDING CORN 487<br />

Germplasm resources<br />

Over 13,000 accessions <strong>of</strong> corn are kept in storage at<br />

the Centro Internationale de Mejoramiento de Maiz y<br />

Trigo (CIMMYT) in Mexico, with duplicates <strong>of</strong> these<br />

accession held in the US (at the US National Seed<br />

Storage Laboratory, Fort Collins, Colorado), Columbia,<br />

<strong>and</strong> Peru. Many heterotic populations have been<br />

identified in maize, the most well known <strong>and</strong> exploited<br />

including “Reid” × “Lancaster” <strong>and</strong> “Lancaster” ×<br />

“Stiff Stalk”. Others are the European flint × dents,<br />

“Tuxpeno” × “ETO”, <strong>and</strong> “Pantap × Suwan 1”.<br />

Cytogenetics<br />

Corn (Zea mays L.) is a diploid (2n = 20) <strong>and</strong> a monocot<br />

<strong>of</strong> the family Poaceae (Gramineae), or grass family.<br />

The genus has four species: Z. mays (cultivated corn<br />

<strong>and</strong> teosinte), Z. diploperennis Iltis et al. (diploperennial<br />

teosinte), Z. luxurians, <strong>and</strong> Z. perennis (perennial<br />

teosinte). Of these four species, only Z. mays is widely<br />

grown commercially in the US. The closest generic<br />

relative <strong>of</strong> Zea is Tripsacum, which has seven species,<br />

three <strong>of</strong> which are known to occur in the US. Teosinte<br />

occurs in the wild in Mexico <strong>and</strong> Guatemala. Cultivated<br />

corn has 10 pairs <strong>of</strong> chromosomes (n = 10). However,<br />

plants with 1–8 sets <strong>of</strong> chromosomes have been developed<br />

for various purposes.<br />

In addition to the autosomes or normal or st<strong>and</strong>ard<br />

chromosomes (A-chromosomes), maize has supernumerary<br />

elements such as the B-chromosomes (also<br />

called supernumerary chromosomes). The role <strong>of</strong> the<br />

B-chromosomes in the cell varies from being practically<br />

a nuisance to having some definite function, depending<br />

on the organism. However, when the B-chromosomes<br />

number 10–15 or more, certain abnormalities may<br />

occur (e.g., reduced fertility, decreased vigor, aborted<br />

pollen, defective kernels). Seed is rarely produced with<br />

the presence <strong>of</strong> a B-chromosome number in excess <strong>of</strong><br />

25. Maize B-chromosomes are among those most widely<br />

studied in plants. They are suspected to influence the<br />

frequency <strong>of</strong> crossing over, among other roles. Thous<strong>and</strong>s<br />

<strong>of</strong> translocation events have been described in maize.<br />

They are used for locating genes on chromosomes.<br />

Monoploids (haploids) may arise spontaneously by<br />

pathogenesis (the unfertilized egg develops into a plant).<br />

Occasionally, paternal haploids develop by <strong>and</strong>rogenesis.<br />

Haploid generation via these systems has low frequency<br />

<strong>of</strong> occurrence. The average frequency <strong>of</strong> this event in<br />

corn is estimated at one per 1,000 kernels. These lines

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