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Principles of Plant Genetics and Breeding

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First cell division<br />

Second cell division<br />

Interphase<br />

Anaphase 2<br />

Centromeres divide; sister<br />

chromatids move to opposite poles<br />

Figure 3.3 Diagrammatic presentation <strong>of</strong> meiosis in a cell<br />

with a diploid number <strong>of</strong> 4. The process has two distinct<br />

cell divisions. Prophase I consists <strong>of</strong> five distinguishable<br />

stages; the most genetically significant event <strong>of</strong> crossing<br />

over occurs in the fourth stage, diplonema.<br />

PLANT CELLULAR ORGANIZATION AND GENETIC STRUCTURE 39<br />

Prophase 1<br />

Divisible into 5 substages;<br />

leptonema, zygonema,<br />

pachynema, diplonema,<br />

diakinesis<br />

Homologous chromosomes<br />

pair to form bivalents; each<br />

cromosome duplicates; sister<br />

chromatids pair; chiasmata<br />

form; crossing over occurs<br />

Metaphase 1<br />

Homologous chromosomes<br />

align at the equitorial plate;<br />

nuclear membrane<br />

disappears<br />

Anaphase 1<br />

Homologous chromosome<br />

pairs separate, sister<br />

chromatids remain<br />

together<br />

Telophase 1<br />

Two daughter cells form;<br />

nuclear membrane forms;<br />

each cell contains only<br />

one chromosome <strong>of</strong> the<br />

homologous pair<br />

Prophase 2<br />

Short stage; DNA does not<br />

duplicate<br />

Metaphase 2<br />

Nuclear membrane<br />

disappears; chromosomes<br />

align at the equitorial plate<br />

Telophase 2<br />

Nuclear membrane<br />

forms; four haploid<br />

daughter cells are<br />

produced following<br />

cytokinesis<br />

the ploidy level <strong>of</strong> the species. By reducing the diploid<br />

number to a haploid number before fertilization, the<br />

diploid number is restored thereafter.<br />

Mendelian concepts in plant breeding<br />

As previously stated, genetics is the principal science<br />

that underlies plant breeding. Gregor Mendel made<br />

significant contributions to the development <strong>of</strong> the<br />

discipline <strong>of</strong> genetics, albeit in absentia. He derived<br />

several postulates or principles <strong>of</strong> inheritance, which are<br />

<strong>of</strong>ten couched as Mendel’s laws <strong>of</strong> inheritance.<br />

Mendelian postulates<br />

Because plant breeders transfer genes from one source<br />

to another, an underst<strong>and</strong>ing <strong>of</strong> transmission genetics<br />

is crucial to a successful breeding effort. The method <strong>of</strong><br />

breeding used depends upon the heredity <strong>of</strong> the trait<br />

being manipulated, among other factors. According to<br />

Mendel’s results from his hybridization studies in pea,<br />

traits are controlled by heritable factors that are passed<br />

from parents to <strong>of</strong>fspring, through the reproductive<br />

cells. Each <strong>of</strong> these unit factors occurs in pairs in each<br />

cell (except reproductive cells or gametes).<br />

In his experiments, Mendel discovered that in a cross<br />

between parents displaying two contrasting traits, the<br />

hybrid (F1 ) expressed one <strong>of</strong> the traits to the exclusion<br />

<strong>of</strong> the other. He called the expressed trait dominant<br />

<strong>and</strong> the suppressed trait recessive. This is the phenomenon<br />

<strong>of</strong> dominance <strong>and</strong> recessivity. When the<br />

hybrid seed was planted <strong>and</strong> self-pollinated, he observed<br />

that both traits appeared in the second generation (F2 )<br />

(i.e., the recessive trait reappeared), in a ratio <strong>of</strong> 3 : 1<br />

dominant : recessive individuals (Figure 3.4). Mendel<br />

concluded that the two factors that control each trait do<br />

not blend but remain distant throughout the life <strong>of</strong> the<br />

individual <strong>and</strong> segregate in the formation <strong>of</strong> gametes.<br />

This is called the law <strong>of</strong> segregation. In further studies<br />

in which he considered two characters simultaneously,<br />

he observed that the genes for different characters are<br />

inherited independently <strong>of</strong> each other. This is called the<br />

law <strong>of</strong> independent assortment. In summary, the two<br />

key laws are as follows:<br />

Law I Law <strong>of</strong> segregation: paired factors segregate<br />

during the formation <strong>of</strong> gametes in a r<strong>and</strong>om<br />

fashion such that each gamete receives one<br />

form or the other.<br />

Law II Law <strong>of</strong> independent assortment: when two<br />

or more pairs <strong>of</strong> traits are considered simultaneously,<br />

the factors for each pair <strong>of</strong> traits<br />

assort independently to the gametes.<br />

Mendel’s pairs <strong>of</strong> factors are now known as genes,<br />

while each factor <strong>of</strong> a pair (e.g., HH or hh) is called an<br />

allele (i.e., the alternative form <strong>of</strong> a gene: H or h). The<br />

specific location on the chromosome where a gene<br />

resides is called a gene locus or simply a locus (loci for<br />

plural).

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