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Principles of Plant Genetics and Breeding

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218 CHAPTER 13<br />

1<br />

2<br />

3<br />

(a)<br />

(b)<br />

Figure 13.3 Cytology <strong>of</strong> polyploids: (a) triploidy <strong>and</strong> (b) autotetraploidy. Bivalents <strong>and</strong> quadrivalents usually produce<br />

functional gametes, while univalents <strong>and</strong> trivalents produce sterile gametes.<br />

as some trivalents <strong>and</strong> univalents. These meiotic abnormalities<br />

are implicated in sterility to some extent, more<br />

so in triploids. The microspores <strong>and</strong> megaspores with x<br />

or 2x genomes are usually viable.<br />

The amount <strong>and</strong> nature <strong>of</strong> chromosome pairing<br />

directly impacts the breeding behavior <strong>of</strong> autoploids.<br />

Autoploids are induced artificially by chromosome doubling<br />

using colchicine. Doubling a hybrid between two<br />

diploid cultivars would produce a tetraploid in which<br />

there may be a tendency for the doubled set <strong>of</strong> chromosomes<br />

from one parent to pair independently <strong>of</strong> the<br />

doubled set <strong>of</strong> chromosomes <strong>of</strong> the other parent. This<br />

propensity is called preferential or selective pairing, a<br />

phenomenon with genetic consequences. If preferential<br />

pairing is complete, there would be no new genetic<br />

recombination <strong>and</strong> hence the progeny would look<br />

like the doubled F 1 . Furthermore, the bivalent pairing<br />

would contribute to sterility originating from meiotic<br />

disorders, while preserving heterosis indefinitely, should<br />

there be any produced by the original cross. The concept<br />

<strong>of</strong> preferential pairing is applied in the modern<br />

breeding <strong>of</strong> polyploids whereby alloploids are stabilized<br />

Trivalent<br />

Trivalent<br />

Bivalent<br />

Univalent<br />

Bivalents<br />

Quadrivalents<br />

Univalent<br />

Possible functional<br />

gametes<br />

Non-functional<br />

gametes<br />

<strong>and</strong> made reliable as diploids, a process called<br />

diploidization.<br />

<strong>Genetics</strong> <strong>of</strong> autoploids<br />

The ploidy level may also be defined as the number <strong>of</strong><br />

different alleles that an individual can possess for a single<br />

locus on a chromosome. A diploid can have two alleles<br />

per locus, whereas an autotetraploid can have four different<br />

alleles. The genetics <strong>of</strong> autoploids is complicated<br />

by multiallelism <strong>and</strong> multivalent association <strong>of</strong> chromosomes<br />

during meiosis. Consider the segregation <strong>of</strong><br />

alleles <strong>of</strong> a single locus (A, a). In a diploid species,<br />

there would be three possible genotypes AA, Aa, <strong>and</strong><br />

aa. However, in an autotetraploid there would be five<br />

genotypes ranging from nulliplex (aaaa) to quadruplex<br />

(AAAA) (Table 13.4). The proportion <strong>of</strong> dominant<br />

(A) to recessive (a) genes is different in two <strong>of</strong> the<br />

five genotypes (AAAA <strong>and</strong> Aaaa) in autotetraploids<br />

from that which obtains in diploids. The number <strong>of</strong><br />

phenotypes observed depends on the dominance relationship<br />

<strong>of</strong> A <strong>and</strong> a. If allele A is completely dominant

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