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Evolution__3rd_Edition

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..<br />

Log fitness of organisms<br />

Number of mutations in organism<br />

. . . that may or may not confirm the<br />

mutational theory of sex<br />

Work on a second prediction is also<br />

inconclusive<br />

3<br />

2<br />

1<br />

CHAPTER 12 / Adaptations in Sexual Reproduction 323<br />

Figure 12.6<br />

Three different relations between the fitness of an organism<br />

and the number of deleterious mutations it carries. The y-axis is<br />

logarithmic. 1, Synergistic epistasis: multiple mutations have an<br />

increasingly damaging effect on the organism. 2, Independent,<br />

or multiplicative, fitness effects. 3, Multiple mutations have a<br />

decreasingly damaging effect. Deleterious mutations are purged<br />

more (slope 1), equally (slope 2), and less (slope 3), efficiently<br />

with sexual, than asexual, reproduction. The relative positions of<br />

lines 1, 2, and 3 up the y-axis is irrelevant; only the slope matters.<br />

If slope 1 was drawn below 2 it would still be synergistic epistasis.<br />

deleterious mutation rates, U is less than one in bacteria and more than one in great<br />

apes, including humans. Kondrashov’s theory correctly predicts the absence of sex in<br />

bacteria. The problematic area is around flies and worms. They reproduce sexually<br />

and Kondrashov’s theory predicts U > 1. On the high estimate, the prediction is<br />

upheld; on the low estimate, it is falsified. Thus, further research is needed on the fraction<br />

of mutations that are deleterious. However, it is worth noting that U is > 1 in<br />

humans whether the correct figure is 2 or 30 deleterious mutations per generation. If<br />

Kondrashov’s theory turns out to be wrong, and sex does not help the selective purge of<br />

deleterious mutations, we shall be left with a paradox a how can humans exist, given<br />

their high deleterious mutation rate?<br />

The second prediction of Kondrashov’s theory concerns the relation between the<br />

fitness of an organism and the number of deleterious mutations it contains. Three sorts<br />

of relation are theoretically possible (Figure 12.6). Kondrashov’s theory only works if<br />

the graph slopes down a a condition called synergistic epistasis. Experimenters are also<br />

trying to test this prediction, but no conclusive results are yet available.<br />

In conclusion, the mutational theory suggests that sex exists to help life cope with<br />

its load of deleterious mutations. The theory has been worked out, and is internally<br />

consistent. It makes two predictions about real sexual creatures: they should have<br />

deleterious mutation rates of one or more, and their fitness relations should show<br />

synergistic epitasis. These predictions have inspired a major research programme a<br />

one of the most active and important in modern evolutionary biology a but it is<br />

currently inconclusive. The next few years of work should tell us whether U is > 1 in<br />

fruitflies, but we do not yet know.<br />

12.2.3 Coevolution of parasites and hosts may produce rapid<br />

environmental change<br />

The second theory we shall look at ignores the effect of deleterious mutations and concentrates<br />

on external environmental change. Sex is more likely to be advantageous if<br />

environments change rapidly: the problem is to work out how environments could

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