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Evolution__3rd_Edition

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Populations may be subdivided<br />

CHAPTER 5 / The Theory of Natural Selection 129<br />

higher at high frequencies, where the local birds are well educated about the dangers of<br />

eating the warningly colored forms.<br />

The purpose of Sections 5.11–5.13 has been to illustrate the different mechanisms by<br />

which natural selection can maintain polymorphism. In Chapter 6 we look at another<br />

mechanism that can maintain polymorphism a genetic drift. Then, in Chapter 7, we<br />

tackle the question of how important the mechanisms are in nature.<br />

5.14 Subdivided populations require special population<br />

genetic principles<br />

5.14.1 A subdivided set of populations have a higher proportion of<br />

homozygotes than an equivalent fused population: this is<br />

the Wahlund effect<br />

So far we have considered population genetics within a single, uniform population. In<br />

practice, a species may consist of a number of separate populations, each more or less<br />

isolated from the others. The members of a species might, for example, inhabit a number<br />

of islands, with each island population being separated by the sea from the others.<br />

Individuals might migrate between islands from time to time, but each island population<br />

would evolve to some extent independently. A species with a number of more or<br />

less independent subpopulations is said to have population subdivision.<br />

Let us see first what effect population subdivision has on the Hardy–Weinberg<br />

principle. Consider a simple case in which there are two populations (we can call them<br />

population 1 and population 2), and we concentrate on one genetic locus with two<br />

alleles, A and a. Suppose allele A has frequency 0.3 in population 1 and 0.7 in population<br />

2. If the genotypes have Hardy–Weinberg ratios they will have the frequencies, and<br />

average frequencies, in the two populations shown in Table 5.10. The average genotype<br />

frequencies are 0.29 for AA, 0.42 for Aa, and 0.29 for aa. Now suppose that the two<br />

Table 5.10<br />

The frequency of genotypes AA, Aa, and aa in two populations when A has frequency 0.3 in<br />

population 1 and 0.7 in population 2. The average genotypes are calculated assuming the two<br />

populations are of equal size.<br />

Genotype<br />

AA Aa aa<br />

Frequency (0.3) 2 = 0.09 2(0.3)(0.7) = 0.42 (0.7) 2 = 0.49 population 1<br />

(0.7) 2 = 0.49 2(0.7)(0.3) = 0.42 (0.3) 2 = 0.09 population 2<br />

Average 0.58/2 = 0.29 0.84/2 = 0.42 0.58/2 = 0.29

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