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Principles of Plant Genetics and Breeding

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Table 13.4 <strong>Genetics</strong> <strong>of</strong> autoploids.<br />

Diploid Polyploidy Name<br />

Cross<br />

Aa × Aa AAaa × AAaa<br />

Products<br />

1/4 AA 1/36 AAAA Quadruplex<br />

2/4 Aa 8/36 AAAa Triplex<br />

1/4 aa 18/36 AAaa Duplex<br />

8/36 Aaaa Simplex<br />

1/36 aaaa Nulliplex<br />

Table 13.5 Genetic frequencies following chromosome<br />

segregation <strong>of</strong> an autotetraploid.<br />

Gametic frequency<br />

Genotype AA Aa aa<br />

AAAA 1 0 0<br />

AAAa 1/2 1/2 0<br />

AAaa 1/6 4/6 1/6<br />

Aaaa 0 1/2 1/2<br />

aaaa 0 0 1<br />

Note: chromatid segregation occurs less frequently than<br />

chromosomes segregation <strong>and</strong> produces alternative types <strong>of</strong><br />

segregation. For example the simplex (Aaaa) can produce<br />

gametes that are homozygous (AA) by the process called<br />

double reduction.<br />

Assuming complete dominance <strong>and</strong> chromosome segregation<br />

the following phenotypic ratios are observed. Certain<br />

segregation ratios are sometimes indicative <strong>of</strong> the nature <strong>of</strong><br />

autotetraploid inheritance.<br />

Cross Progeny (dominant : recessive)<br />

AAAA × AAAA 1 : 0<br />

AAAa × AAAa 1 : 0<br />

AAaa × AAaa 35 : 1<br />

AAaa × Aaaa 11 : 1<br />

AAaa × aaaa 5 : 1<br />

Aaaa × Aaaa 3 : 1<br />

Aaaa × aaaa 1 : 1<br />

aaaa × aaaa 0 : 1<br />

to allele a, there would be only two phenotypes. If<br />

dominance is incomplete or the effect <strong>of</strong> allele A is<br />

cumulative, there could be up to five phenotypes.<br />

Upon selfing, a dominant phenotype in a diploid (AA,<br />

Aa) would produce a progeny that is all dominant, or<br />

POLYPLOIDY IN PLANT BREEDING 219<br />

Table 13.6 Multiple allelelism in autotetraploids.<br />

Tetrasomic condition<br />

a1a1a1a1 a1a1a1a2 a1a1a2a2 a1a1a2a3 a1a2a3a4 All alleles are identical; monoallelic; balanced<br />

Two different alleles; diallelic; unbalanced<br />

Two different alleles; diallelic; balanced<br />

Three different alleles; triallelic<br />

Four different alleles; tetra-allelic<br />

Number <strong>of</strong> possible interactions are: (i) first order (e.g., a1a2 ,<br />

a1a3 ); (ii) second order (e.g., a1a2a3 , a1a3a4 ); <strong>and</strong> (iii) third<br />

order interaction (a1a2a3a4 ). This depends on the tetrasomic<br />

condition.<br />

Tetrasomic condition 1st 2nd 3rd Total<br />

a 1 a 2 a 3 a 4 6 4 1 11<br />

a 1 a 1 a 2 a 3 3 1 0 4<br />

a 1 a 1 a 2 a 2 1 0 0 1<br />

a 1 a 1 a 1 a 2 1 0 0 1<br />

a 1 a 1 a 1 a 1 0 0 0 0<br />

segregate in the 3 : 1 ratio. Selfing each <strong>of</strong> the five<br />

categories would produce many different outcomes in<br />

autotetraploids, assuming r<strong>and</strong>om chromosome segregation<br />

(Table 13.5).<br />

An autoploid individual can have up to four alleles<br />

(abcd) per locus. Five different genotype categories are<br />

similarly possible except that there may be only four<br />

nulliplex genotypes (aaaa, bbbb, cccc, dddd) <strong>and</strong> only<br />

one tetragenic genotype (abcd), but numerous combinations<br />

for the intermediates (Table 13.6). The possible<br />

gametic array is shown for each genotype. Interallelic<br />

<strong>and</strong> intra-allelic interactions may occur for as many as<br />

four alleles per locus in an autotetraploid. The degree<br />

to which intra-allelic interaction occurs determines the<br />

expression <strong>of</strong> heterosis <strong>and</strong> inbreeding depression in<br />

an autotetraploid. Because four identical alleles are<br />

required to achieve homozygosity in an autotetraploid<br />

compared with only two in a diploid, homozygosity<br />

is achieved at a less rapid rate in autotetraploids<br />

(Figure 13.4).<br />

Another aspect <strong>of</strong> autoploid genetics with a plant<br />

breeding implication, is the difficulty <strong>of</strong> distinguishing<br />

between a triplex <strong>and</strong> a quadruplex on the basis <strong>of</strong> a<br />

progeny test (assuming r<strong>and</strong>om chromosome segregation).<br />

Both genotypes (AAAA <strong>and</strong> AAAa) will breed<br />

true for the dominant allele. To identify a triplex plant,<br />

the breeder would have to advance the progeny one<br />

more generation to identify the duplex plants <strong>of</strong> the S 1 .<br />

Achieving genetic purity in autotetraploid stocks is

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