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Principles of Plant Genetics and Breeding

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Stigma<br />

Pin flower Thrum flower<br />

Figure 4.5 Heteromorphic incompatibility showing floral<br />

modifications in which anthers <strong>and</strong> pistils are <strong>of</strong> different<br />

lengths in different plants (heterostyly). This type <strong>of</strong><br />

incompatibility is believed to be always <strong>of</strong> the sporophytic<br />

type. Pin <strong>and</strong> thrum flowers occurs in flowers such as<br />

Primula, Forsythia, Oxalis, <strong>and</strong> Silia.<br />

(a) Gametophytic incompatibility. In gametophytic<br />

incompatibility (originally called the<br />

oppositional factor system), the ability <strong>of</strong> the<br />

pollen to function is determined by its own<br />

genotype <strong>and</strong> not the plant that produces it.<br />

Gametophytic incompatibility is more widespread<br />

than sporophytic incompatibility. Gametophytic<br />

incompatibility occurs in species such as red<br />

clover, white clover, <strong>and</strong> yellow sweet clover.<br />

Homomorphic incompatibility is controlled by<br />

a series <strong>of</strong> alleles at a single locus (S 1 , S 2 ,...S n )<br />

or alleles at two loci in some species. The system<br />

is called homomorphic because the flowering<br />

structures in both the seed-bearing (female)<br />

<strong>and</strong> pollen-bearing (male) plants are similar. The<br />

alleles <strong>of</strong> the incompatibility gene(s) act individually<br />

in the style. They exhibit no dominance.<br />

The incompatible pollen is inhibited in the style.<br />

The pistil is diploid <strong>and</strong> hence contains two incompatibility<br />

alleles (e.g., S 1 S 3 , S 3 S 4 ). Reactions<br />

occur if identical alleles in both pollen <strong>and</strong> style<br />

are encountered. Only heterozygotes for S alleles<br />

are produced in this system.<br />

(b) Sporophytic incompatibility. In sporophytic<br />

incompatibility, the incompatibility characteristics<br />

<strong>of</strong> the pollen are determined by the plant<br />

(sporophyte) that produces it. It occurs in<br />

species such as broccoli, radish, <strong>and</strong> kale. The<br />

sporophytic system differs from the gametophytic<br />

system in that the S allele exhibits dominance.<br />

Also, it may have individual action in both pollen<br />

<strong>and</strong> the style, making this incompatibility system<br />

PLANT REPRODUCTIVE SYSTEMS 69<br />

(a)<br />

(b)<br />

S 1<br />

Figure 4.6 Types <strong>of</strong> self-incompatibility: (a) sporophytic<br />

<strong>and</strong> (b) gametophytic. Sporophytic incompatibility occurs<br />

in families such as Compositae <strong>and</strong> Cruciferae. It is<br />

associated with pollen grains with two generative nuclei,<br />

whereas gametophytic incompatibility is associated with<br />

pollen with one generative nucleus in the pollen tube as<br />

occurs in various kinds <strong>of</strong> clover.<br />

complex. The dominance is determined by the<br />

pollen parent. Incompatible pollen may be inhibited<br />

on the stigma surface. For example, a plant<br />

with genotype S 1 S 2 where S 1 is dominant to S 2 ,<br />

will produce pollen that will function like S 1 .<br />

Furthermore, S 1 pollen will be rejected by an S 1<br />

style but received by an S 2 style. Hence, homozygotes<br />

<strong>of</strong> S alleles are possible.<br />

Incompatibility is expressed in one <strong>of</strong> three general<br />

ways, depending on the species. The germination <strong>of</strong> the<br />

pollen may be decreased (e.g., in broccoli). Sometimes,<br />

removing the stigma allows normal pollen germination.<br />

In the second way, pollen germination is normal, but<br />

pollen tube growth is inhibited in the style (e.g.,<br />

tobacco). In the third scenario, the incompatibility reaction<br />

occurs after fertilization (e.g., in Gesteria). This<br />

third mechanism is rare.<br />

Changing the incompatibility reaction<br />

Mutagens (agents <strong>of</strong> mutation) such as X-rays, radioactive<br />

sources such as P32 , <strong>and</strong> certain chemicals have been<br />

S 2<br />

1 2<br />

1<br />

S1 S3 1 3<br />

S2 S3 1<br />

4<br />

S 3<br />

3<br />

1<br />

2<br />

2<br />

S 2<br />

S 4<br />

4<br />

2

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