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Evolution__3rd_Edition

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Figure 12.2<br />

Non-reproductive sex in<br />

Paramecium. Paramecium<br />

normally contains one<br />

micronucleus and one<br />

macronucleus. When it<br />

prepares for sex, the<br />

macronucleus dissolves and the<br />

micronucleus is duplicated.<br />

Two such cells can then<br />

conjugate, swapping one of<br />

their micronuclei. Meiosis then<br />

occurs within each cell. The<br />

sex act is non-reproductive.<br />

Paramecium cells reproduce by<br />

binary fission. Thus, sex and<br />

reproduction are not associated<br />

in Paramecium. The same is<br />

true in many single-celled life<br />

forms.<br />

Some kinds of sex are cost-free<br />

But a big unsolved problem remains<br />

CHAPTER 12 / Adaptations in Sexual Reproduction 315<br />

conjugate (Figure 12.2). The two cells swap copies of their DNA and then separate.<br />

Meiosis then occurs within each cell. Sex is non-reproductive: there were two cells<br />

before conjugation and there are two after it. Sex has no cost as in Figure 12.1. Sex<br />

acquired its cost as sex became associated with reproduction, perhaps around the<br />

time of the evolution of multicellular life. Sex probably originated in single-celled life,<br />

and had little cost at that time. The origin of sex therefore poses no deep evolutionary<br />

problem. But in many life forms today, sex does have a 50% cost and its existence is a<br />

problem.<br />

Fifty percent is a large cost. The problem of explaining sex is to find a compensating<br />

advantage of sexual reproduction that is large enough to make up for its cost. We are on<br />

the look out for an extraordinarily large selective advantage. Typical evolutionary<br />

events are thought to involve selective advantages of a few percent at most, and more<br />

often 1% or less. Consider this: a female who has survived to adulthood and is about to<br />

reproduce must be fairly well adapted to her environment. If she were to reproduce<br />

asexually, she would just make a copy of herself and produce a daughter as well adapted<br />

to the conditions of the next generation as she would be herself. If she reproduces sexually<br />

instead, she discards half her genes and produces an offspring by mixing the<br />

remaining half with other genes drawn from a stranger. If sex is to outweigh its twofold<br />

cost, the sexual female must by this procedure expect to produce a daughter who will be<br />

twice as fit as a simple copy of herself. The problem, therefore, is not trivial. Indeed,<br />

G. C. Williams has described it as “the outstanding puzzle in evolutionary biology.” The<br />

puzzle is still a puzzle, but we can look at some possible solutions to it. Box 12.1 discusses<br />

how the puzzle has taken on a practical importance, with the rise of cloning technologies.<br />

12.1.2 Sex is unlikely to be explained by genetic constraint<br />

One possibility is that life uses sexual reproduction because it is “stuck with it.” That<br />

is, the mutations to produce asexual reproduction have not occurred. (In terms of

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