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Evolution__3rd_Edition

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..<br />

Figure 12.3<br />

<strong>Evolution</strong> in (a) asexual and<br />

(b) sexual populations. The<br />

mutations A, B, and C are all<br />

advantageous. In the asexual<br />

population, an AB individual<br />

can only arise if the B mutation<br />

arises in an individual that<br />

already has an A mutation<br />

(or vice versa). In the sexual<br />

population, an AB individual<br />

can be formed by the breeding<br />

of a B mutation-bearing<br />

individual with an A mutationbearing<br />

individual; the second<br />

mutation of B is not needed.<br />

(c) If favorable mutations are<br />

rare, each will have been fixed<br />

before the next arises, and<br />

sexual populations do not<br />

evolve faster. The relative rates<br />

of evolution in asexual and<br />

sexual populations depends<br />

on the rate at which favorable<br />

mutations arise.<br />

Sexual populations can evolve<br />

faster than asexual populations, ...<br />

. . . if the rate of favorable mutation<br />

is high<br />

(a) Asexual: high rate of favorable mutation<br />

C<br />

A<br />

Time<br />

B AC<br />

Time<br />

CHAPTER 12 / Adaptations in Sexual Reproduction 317<br />

AB ABC<br />

(c) Sexual or asexual: low rate of favorable mutation<br />

A AB<br />

(b) Sexual: high rate of favorable mutation<br />

single individual (Figure 12.3). Suppose, for example, that a sexual and an asexual<br />

population are both fixed for genes A′ and B′ at two loci. In the environment where the<br />

two populations are living, mutations A and B are advantageous. A and B mutations<br />

would be likely to arise initially in different individuals. The asexual population will<br />

then come to consist of A′B and AB′ individuals, because the A mutant cannot spread<br />

into the A′B clone or vice versa. AB individuals cannot appear until an A′ gene mutates<br />

to A within the A′B clone (or B′ to B in the AB′ clone).<br />

In the sexual population, evolution proceeds much faster. After A and B have arisen<br />

in different individuals, they can soon combine in a single individual by sex without<br />

waiting for the mutations to occur twice. Natural selection can therefore take the population<br />

from the state A′B′ to AB faster than under asexual reproduction. This argument<br />

was first put forward by Fisher and by Muller in the 1930s. They concluded that sexual<br />

populations have a more rapid rate of evolution than would an otherwise equivalent<br />

group of asexual organisms.<br />

However, subsequent research has shown that the rate of evolution in sexual populations<br />

is not necessarily faster than in equivalent asexual populations. For instance, the<br />

result depends on the rate of mutation. If favorable mutations are rare, each one will<br />

have been fixed in the population before the next one arises (Figure 12.3c). Sexual and<br />

asexual populations then evolve at the same rate. New favorable mutations will always<br />

arise in individuals that already carry the previous favorable mutation: they must,<br />

because the previous favorable mutation is already in every member of the population.<br />

In terms of the example, the B mutation will arise in an AB′ individual in both sexual<br />

and asexual populations. However, if favorable mutations arise more frequently, Fisher<br />

and Muller’s argument works: the sexual population evolves faster. Each new favorable<br />

mutation will usually arise in an individual that does not already possess other<br />

A<br />

C<br />

B<br />

AC<br />

AB<br />

Time<br />

ABC

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