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Principles of Plant Genetics and Breeding

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18 CHAPTER 2<br />

discovered. As will be discussed in detail later in the<br />

book, scientists collect, process, <strong>and</strong> store this natural<br />

variation in germplasm banks for use by breeders in their<br />

breeding programs.<br />

The process <strong>of</strong> evolution has parallels in plant breeding.<br />

Darwin’s theory <strong>of</strong> evolution through natural selection<br />

can be summed up in three principles that are at the<br />

core <strong>of</strong> plant breeding. These are the principles <strong>of</strong>:<br />

1 Variation. Variation in morphology, physiology,<br />

<strong>and</strong> behavior exist among individuals in a natural<br />

population.<br />

2 Heredity. Offspring resemble their parents more<br />

than they resemble unrelated individuals.<br />

3 Selection. Some individuals in a group are more<br />

capable <strong>of</strong> surviving <strong>and</strong> reproducing than others<br />

(i.e., more fit).<br />

A key factor in evolution is time. The changes in<br />

evolution occur over extremely long periods <strong>of</strong> time.<br />

<strong>Plant</strong> breeders depend on biological variation as<br />

a source <strong>of</strong> desired alleles. Induced mutation <strong>and</strong><br />

hybridization for recombination are major sources <strong>of</strong><br />

variation. Once variation has been assembled, the<br />

breeder imposes a selection pressure (artificial selection<br />

in this case) to discriminate among the variation to<br />

advance only desired plants. <strong>Plant</strong> breeding may be<br />

described as directed or targeted <strong>and</strong> accelerated evolution,<br />

because the plant breeder, with a breeding objective<br />

in mind, deliberately <strong>and</strong> genetically manipulates<br />

plants (wild or domesticated) to achieve a stated goal,<br />

but in a very short time. Conceptually, breeding <strong>and</strong><br />

evolution are the same, a key difference being the<br />

duration <strong>of</strong> the processes. <strong>Plant</strong> breeding has been<br />

described as evolution directed by humans. Compared<br />

to evolution, a plant breeding process is completed in a<br />

twinkle <strong>of</strong> an eye! Also, unlike evolution, plant breeders<br />

do not deal with closed populations. They introgress<br />

new variability from different genotypes <strong>of</strong> interest, <strong>and</strong>,<br />

for practical <strong>and</strong> economic purposes, deal with limited<br />

population sizes.<br />

Domestication<br />

Domestication is the process by which genetic changes<br />

(or shifts) in wild plants are brought about through<br />

a selection process imposed by humans. It is an evolutionary<br />

process in which selection (both natural <strong>and</strong><br />

artificial) operates to change plants genetically, morphologically,<br />

<strong>and</strong> physiologically. The results <strong>of</strong> domestica-<br />

tion are plants that are adapted to supervised cultural<br />

conditions, <strong>and</strong> possessing characteristics that are preferred<br />

by producers <strong>and</strong> consumers. In some ways, a<br />

domesticated plant may be likened to a tamed wild<br />

animal that has become a pet.<br />

There are degrees <strong>of</strong> domestication. Species that<br />

become completely domesticated <strong>of</strong>ten are unable to<br />

survive when reintroduced into the wild. This is so<br />

because the selection process that drives domestication<br />

strips plants <strong>of</strong> natural adaptive features <strong>and</strong> mechanisms<br />

that are critical for survival in the wild, but undesirable<br />

according to the needs <strong>of</strong> humans.<br />

Like evolution, domestication is also a process <strong>of</strong><br />

genetic change in which a population <strong>of</strong> plants can experience<br />

a shift in its genetic structure in the direction <strong>of</strong><br />

selection imposed by the domesticator. New plant types<br />

are continually selected for as domesticates as new<br />

dem<strong>and</strong>s are imposed, thereby gradually moving the<br />

selected individuals farther away (genetically, morphologically,<br />

<strong>and</strong> physiologically) from their wild progenitors.<br />

Both wild <strong>and</strong> domesticated populations are<br />

subject to evolution.<br />

Patterns <strong>of</strong> plant domestication<br />

Domestication has been conducted for over 10,000<br />

years, <strong>and</strong> ever since agriculture was invented. Archeological<br />

<strong>and</strong> historical records provide some indications<br />

as to the period certain crops may have been domesticated,<br />

even though such data are not precise. Archeological<br />

records from arid regions are better preserved<br />

than those from the humid regions <strong>of</strong> the world.<br />

Concepts <strong>of</strong> domestication<br />

As G. Ladizinsky points out in discussing patterns <strong>of</strong><br />

domestication, the challenge is to determine whether<br />

the domesticate evolved under wild conditions, or was<br />

discovered <strong>and</strong> then cultivated by humans, or whether<br />

cultivation preceded the selection <strong>of</strong> domesticates. This<br />

is a subject <strong>of</strong> debate. For example, seed dormancy is a<br />

problem in wild legumes, <strong>and</strong> hence would have hindered<br />

their use in cultivation. It is likely that the domesticates<br />

evolved in the wild before being used in cultivation.<br />

However, in most cereal species, most experts believe<br />

that domestication occurred after cultivation. In wheat<br />

<strong>and</strong> barley, for example, a tough rachis, which is resistant<br />

to natural seed dispersal, <strong>and</strong> characterizes domesticates,<br />

would have been selected for during cultivation.<br />

Two categories <strong>of</strong> crop plants are identified, with<br />

respect to domestication, as primary crops or secondary

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