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Principles of Plant Genetics and Breeding

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The Ac element has an open reading frame. The activities<br />

<strong>of</strong> corn transposable elements are developmentally<br />

regulated. That is, the transposable elements transpose<br />

<strong>and</strong> promote genetic rearrangements at only certain<br />

specific times <strong>and</strong> frequencies during plant development.<br />

Transposition involving the Ac–Ds system is<br />

observed in corn as spots <strong>of</strong> colored aleurone. A gene<br />

required for the synthesis <strong>of</strong> anthocyanin pigment is<br />

inactivated in some cells whereas other cells have normal<br />

genes, resulting in spots <strong>of</strong> pigment in the kernel<br />

(genetic mosaicism).<br />

Biotechnology for creating genetic<br />

variability<br />

Gene transfer<br />

The rDNA technology is state-<strong>of</strong>-the-art in gene transfer<br />

to generate genetic variability for plant breeding.<br />

With minor exceptions, DNA is universal. Consequently,<br />

DNA from an animal may be transferred<br />

to a plant! The tools <strong>of</strong> biotechnology may be used to<br />

incorporate genes from distant sources into adapted<br />

cultivars. An increasing acreage <strong>of</strong> cotton, soybean, <strong>and</strong><br />

maize are being sown to genetically modified (GM) cultivars,<br />

indicating the importance <strong>of</strong> this technology for<br />

creating variability for plant breeding. Economic gene<br />

transfers have been made from bacteria to plants to confer<br />

disease <strong>and</strong> herbicide resistance to plants. The most<br />

common GM products on the market are Roundup<br />

Ready® cultivars (e.g., cotton, soybean) with herbicide<br />

tolerance, <strong>and</strong> Bt products (e.g., corn) with resistance<br />

to lepidopteran pests. The technique <strong>of</strong> site-directed<br />

mutagenesis allows scientists to introduce mutations<br />

into specified genes, primarily for the purpose <strong>of</strong> studying<br />

gene function, <strong>and</strong> not for generating variability for<br />

breeding per se. Other tissue-culture-based techniques<br />

include protoplast fusion, cybrid formation, <strong>and</strong> the use<br />

<strong>of</strong> transposons. Chapter 14 is devoted to the application<br />

<strong>of</strong> biotechnology in plant breeding.<br />

Somaclonal variation<br />

In vitro culture <strong>of</strong> plants is supposed to produce clones<br />

(genetically identical derivatives from the parent material).<br />

However, the tissue culture environment has been<br />

known to cause heritable variation called somaclonal<br />

variation. The causes cited for these changes include<br />

karyotypic changes, cryptic chromosomal rearrangements,<br />

somatic crossing over <strong>and</strong> sister chromatid<br />

exchange, transposable elements, <strong>and</strong> gene amplification.<br />

VARIATION 83<br />

Some <strong>of</strong> these variations have been stable <strong>and</strong> fertile<br />

enough to be included in breeding programs.<br />

Scale <strong>of</strong> variability<br />

As previously indicated, biological variation can be<br />

enormous <strong>and</strong> overwhelming to the user. Consequently,<br />

there is a need to classify it for effective <strong>and</strong><br />

efficient use. Some variability can be readily categorized<br />

by counting <strong>and</strong> arranging into distinct non-overlapping<br />

groups; this is said to be discrete or qualitative variation.<br />

Traits that exhibit this kind <strong>of</strong> variation are<br />

called qualitative traits. Other kinds <strong>of</strong> variability occur<br />

on a continuum <strong>and</strong> cannot be placed into discrete<br />

groups by counting. There are intermediates between<br />

the extreme expressions <strong>of</strong> such traits. They are best categorized<br />

by measuring or weighing <strong>and</strong> are described as<br />

exhibiting continuous or quantitative variation. Traits<br />

that exhibit this kind <strong>of</strong> variation are called quantitative<br />

traits.<br />

However, there are some plant characters that may be<br />

classified either way. Sometimes, for convenience, a<br />

quantitative trait may be classified as though it were<br />

qualitative. For example, an agronomic trait such as<br />

earliness or plant maturity is quantitative in nature.<br />

However, it is possible to categorize cultivars into maturity<br />

classes (e.g., in soybean, maturity classes range from<br />

000 (very early) to VIII (very late)). <strong>Plant</strong> height can be<br />

treated in a similar fashion, <strong>and</strong> so can seed coat color<br />

(expressed as shades <strong>of</strong> a particular color).<br />

Qualitative variation<br />

Qualitative variation is easy to classify, study, <strong>and</strong> utilize<br />

in breeding. It is simply inherited (controlled by one<br />

or a few genes) <strong>and</strong> amenable to Mendelian analysis<br />

(Figure 5.6). Examples <strong>of</strong> qualitative traits include diseases,<br />

seed characteristics, <strong>and</strong> compositional traits.<br />

Because they are amenable to Mendelian analysis, the<br />

chi-square statistical procedure may be used to determine<br />

the inheritance <strong>of</strong> qualitative genes. The success <strong>of</strong><br />

gene transfer using molecular technology so far has<br />

involved the transfer <strong>of</strong> single genes (or a few at best),<br />

such as the Bt <strong>and</strong> Ht (herbicide-tolerant) products.<br />

<strong>Breeding</strong> qualitative traits<br />

<strong>Breeding</strong> qualitative traits is relatively straightforward.<br />

They are readily identified <strong>and</strong> selected, although<br />

breeding recessive traits is a little different from breeding<br />

dominant traits (Figure 5.7). It is important to have

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