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Principles of Plant Genetics and Breeding

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62 CHAPTER 4<br />

homozygous <strong>and</strong> therefore become expressed. However,<br />

such depression is reversed upon cross-pollination.<br />

Hybrid vigor (the increase in vigor <strong>of</strong> the hybrid over<br />

its parents resulting from crossing unlike parents) is<br />

exploited in hybrid seed production (see Chapter 18).<br />

In addition to hybrid breeding, population-based<br />

improvement methods (e.g., mass selection, recurrent<br />

selection, synthetic cultivars) are common methods <strong>of</strong><br />

breeding cross-pollinated species.<br />

Asexual reproduction<br />

Asexual reproduction may be categorized into two –<br />

vegetative propagation <strong>and</strong> apomixis. Asexual reproduction<br />

is also called clonal propagation because the<br />

products are genetically identical to the propagules.<br />

Vegetative propagation<br />

As previously indicated, certain species may be reproduced<br />

by using various vegetative parts including bulbs,<br />

corms, rhizomes, stems, <strong>and</strong> buds. Vegetative propagation<br />

is widely used in the horticultural industry. Pieces<br />

<strong>of</strong> vegetative materials called cuttings are obtained from<br />

parts <strong>of</strong> the plant (e.g., root, stem, leaf ) for planting.<br />

Potato, cassava, sugarcane, rose, grape, <strong>and</strong> some perennial<br />

grasses are frequently propagated by stem cuttings.<br />

Methods such as grafting <strong>and</strong> budding are used for<br />

propagating tree crops, where two different plant parts<br />

are united by attaching one to the other <strong>and</strong> securing<br />

with a tape. Healing <strong>of</strong> the graft junction permanently<br />

unites the two parts into one plant.<br />

A variety <strong>of</strong> sophisticated techniques are used to vegetatively<br />

propagate high value plants. Numerous plantlets<br />

may be generated from a small piece <strong>of</strong> vegetative material<br />

(e.g., a segment <strong>of</strong> a leaf) by the technique <strong>of</strong> micropropagation.<br />

The tissue culture technique is used to<br />

rapidly multiply planting material under aseptic conditions<br />

(see Chapter 11 for more details). Perennial horticultural<br />

species tend to be clonally propagated, whereas<br />

annuals <strong>and</strong> biennials tend to be propagated by seed.<br />

Clonally propagated crops may be divided into two<br />

broad categories on the basis <strong>of</strong> economic use:<br />

1 Those cultivated for a vegetative product. Important<br />

species vegetatively cultivated for a vegetative product<br />

include sweet potato, yam, cassava, sugarcane,<br />

<strong>and</strong> Irish potato. These species tend to exhibit certain<br />

reproductive abnormalities. For example, flowering<br />

is reduced, <strong>and</strong> so is fertility. Some species such as<br />

potatoes have cytoplasmic male sterility. Sometimes,<br />

flowering is retarded (e.g., by chemicals) in production<br />

(e.g., in sugarcane).<br />

2 Those cultivated for a fruit or reproductive product.<br />

<strong>Plant</strong>s in this category include fruit trees, shrubs,<br />

<strong>and</strong> cane fruits. Examples include apple, pear, grape,<br />

strawberry, <strong>and</strong> banana.<br />

<strong>Breeding</strong> implications <strong>of</strong> vegetative propagation<br />

There are certain characteristics <strong>of</strong> clonal propagation<br />

that have breeding implications.<br />

1 Clonal species with viable seed <strong>and</strong> high pollen<br />

fertility can be improved by hybridization.<br />

2 Unlike hybridization <strong>of</strong> sexual species, which <strong>of</strong>ten<br />

requires additional steps to fix the genetic variability<br />

in a genotype for release as a cultivar (except for<br />

hybrid cultivars), clonal cultivars can be released<br />

immediately following a cross, provided a desirable<br />

genotype combination has been achieved. Clonal<br />

breeding is hence quick.<br />

3 When improving species whose economic parts are<br />

vegetative products, it is not important for the<br />

hybrid to be fertile.<br />

4 Because <strong>of</strong> the capacity to multiply from vegetative<br />

material (through methods such as cuttings or<br />

micropropagation), the breeder only needs to<br />

obtain a single desirable plant to be used as stock.<br />

5 Heterosis (hybrid vigor), if it occurs, is fixed in<br />

the hybrid product. That is, unlike hybrid cultivars<br />

in seed-producing species, there is no need to<br />

reconstitute the hybrid. Once bred, heterozygosity<br />

is maintained indefinitely.<br />

6 It is more difficult to obtain large quantities <strong>of</strong><br />

planting material from clones in the short term.<br />

7 <strong>Plant</strong> species that are vegetatively parthenocarpic (e.g.,<br />

banana) cannot be improved by hybrid methodology.<br />

8 Species such as mango <strong>and</strong> citrus produce polyembryonic<br />

seedlings. This reproductive irregularity<br />

complicates breeding because clones <strong>of</strong> the parent<br />

are mixed with hybrid progeny.<br />

9 Clonal crops are perennial outcrossers <strong>and</strong> intolerant<br />

<strong>of</strong> inbreeding. They are highly heterozygous.<br />

10 Unlike sexual crop breeding in which the genotype<br />

<strong>of</strong> the cultivar is determined at the end <strong>of</strong> the breeding<br />

process (because it changes with inbreeding),<br />

the genotype <strong>of</strong> a family is fixed <strong>and</strong> determined at<br />

the outset.<br />

11 Both general combining ability (GCA) <strong>and</strong> specific<br />

combining ability (SCA) (that is, performance in<br />

crosses) can be fully exploited with appropriate<br />

breeding method <strong>and</strong> population sizes.

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