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Encyclopedia of Evolution.pdf - Online Reading Center

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eliminated—that is, if sexual recombination occurs between<br />

individuals that are not genetically similar to one another.<br />

Consanguineous matings (between close relatives, who<br />

may share many <strong>of</strong> the same bad mutations) may produce<br />

a relatively large number <strong>of</strong> <strong>of</strong>fspring that are homozygous<br />

for those mutations, a result frequently called inbreeding<br />

depression. In humans it is well known that incest leads to<br />

a high incidence <strong>of</strong> deleterious mutations (see figure on page<br />

372). This would seem to be the case for all species, but it<br />

apparently is not. Some lineages <strong>of</strong> plants exhibit inbreeding<br />

depression, and some do not. Inbreeding depression in<br />

plants is most noticeable when individuals experience stressful<br />

conditions or competition. Depression in plants may show<br />

up more clearly in the second generation <strong>of</strong> inbreeding; the<br />

effects <strong>of</strong> inbreeding may have been hidden during the first<br />

year by maternal resources provided in the seeds. Sibling and<br />

other consanguineous matings occur routinely in some invertebrates.<br />

For example, there is a wasp (Scleroderma immigrans)<br />

in which the females not only mate with their own<br />

sons but their own grandsons. Their populations appear to<br />

have very few deleterious mutations. In the case <strong>of</strong> the wasps,<br />

this may be because the males are haploid, under which condition<br />

all deleterious genes are expressed and eliminated (see<br />

reproductive systems).<br />

It is possible that sexual reproduction has not been<br />

selected at all. Paleontologist Niles Eldredge explains that<br />

sexual species tend to produce new species more quickly than<br />

do asexual species, a process called species sorting. A lineage<br />

that produces many new lineages will persist over time, just<br />

by chance, more than a lineage that produces few or no new<br />

lineages. Random events <strong>of</strong> extinction can readily eliminate<br />

asexual lineages, which speciate little, but not sexual lineages.<br />

To Eldredge, then, the evolution <strong>of</strong> sex may represent species<br />

sorting rather than natural selection.<br />

Several plausible reasons for the evolutionary advantage<br />

<strong>of</strong> sexual reproduction have been proposed. The major difficulty,<br />

according to some evolutionary biologists, is not to<br />

explain why sexual reproduction exists but rather to explain<br />

why it is obligate in so many species.<br />

The evolution <strong>of</strong> sex provides the perfect illustration<br />

that natural selection favors individuals, rather than the<br />

whole species. Since male animals (and the male function <strong>of</strong><br />

plants) can produce many more <strong>of</strong>fspring than females, it<br />

would seem that a population would not require very many<br />

males. Indeed, in agricultural settings, farmers and ranchers<br />

keep only a few choice males as sources <strong>of</strong> pollen (in<br />

trees such as date palms) or sperm. The sex ratio <strong>of</strong> agricultural<br />

populations is therefore heavily weighted toward<br />

the females. However, in wild populations, the sex ratio is<br />

usually close to 1:1. Suppose that one begins with a wild<br />

population that consists, like an agricultural population,<br />

<strong>of</strong> many females and a few males. The few males would<br />

leave many <strong>of</strong>fspring. Any mutation that caused a female<br />

to produce more male <strong>of</strong>fspring would therefore be favored<br />

by natural selection. This process <strong>of</strong> frequency-dependent<br />

selection would continue until males constituted half <strong>of</strong> the<br />

population. Departures from a 1:1 sex ratio can occur as a<br />

sexual selection<br />

result <strong>of</strong> cytoplasmic sterility factors (see selfish genetic<br />

elements).<br />

There is little evidence regarding the origin <strong>of</strong> the process<br />

<strong>of</strong> meiosis, which produces sex cells, or <strong>of</strong> fertilization, which<br />

brings sex cells together. Lynn Margulis and Dorion Sagan<br />

point out that sexual fusion is not very different from other<br />

kinds <strong>of</strong> cellular fusion that merge the genetic components <strong>of</strong><br />

the two cells (see symbiogenesis).<br />

Natural selection favors the sexual production <strong>of</strong> diverse<br />

<strong>of</strong>fspring, mainly because it cleanses bad mutations from the<br />

population and produces a great diversity <strong>of</strong> genes that resist<br />

parasites; these benefits accrue to individuals, not just to<br />

populations.<br />

Further <strong>Reading</strong><br />

Bell, Graham. The Masterpiece <strong>of</strong> Nature: The <strong>Evolution</strong> and Genetics<br />

<strong>of</strong> Sexuality. Berkeley: University <strong>of</strong> California Press, 1982.<br />

Bittles, A. H., and J. V. Neel. “The costs <strong>of</strong> human inbreeding and<br />

their implications for variations at the DNA level.” Nature Genetics<br />

8 (1994): 117–121.<br />

Cosmides, Leda M., and John Tooby. “Cytoplasmic inheritance and<br />

intragenomic conflict.” Journal <strong>of</strong> Theoretical Biology 89 (1981):<br />

83–129.<br />

Hamilton, William D., R. Axelrod, and R. Tanese. “Sexual reproduction<br />

as an adaptation to resist parasites.” Proceedings <strong>of</strong> the<br />

National Academy <strong>of</strong> Sciences USA 87 (1990): 3,566–3,573.<br />

———. Narrow Roads <strong>of</strong> Gene Land, volume 2: <strong>Evolution</strong> <strong>of</strong> Sex.<br />

New York: Oxford University Press, 2002.<br />

Hurst, Laurence D. “Selfish genetic elements and their role in evolution:<br />

the evolution <strong>of</strong> sex and some <strong>of</strong> what that entails.” Philosophical<br />

Transactions <strong>of</strong> the Royal Society <strong>of</strong> London B 349<br />

(1995): 321–332.<br />

———. “Why are there only two sexes?” Proceedings <strong>of</strong> the Royal<br />

Society <strong>of</strong> London B 263 (1996): 415–422.<br />

Lively, Curtis M. “Evidence from a New Zealand snail for the maintenance<br />

<strong>of</strong> sex by parasitism.” Nature 328 (1987): 519–521.<br />

Margulis, Lynn, and Dorion Sagan. Mystery Dance: On the <strong>Evolution</strong><br />

<strong>of</strong> Human Sexuality. New York: Simon and Schuster, 1991.<br />

Otto, Sarah P., and Scott L. Nuismer. “Species interactions and the<br />

evolution <strong>of</strong> sex.” Science 304 (2004): 1,018–1,020.<br />

Paland, Susanne, and Michael Lynch. “Transitions to asexuality result<br />

in excess amino acid substitutions.” Science 311 (2006): 990–992.<br />

Ridley, Matt. The Red Queen: Sex and the <strong>Evolution</strong> <strong>of</strong> Human<br />

Nature. New York: HarperCollins, 1993.<br />

Takebayashi, Naoki, and Peter L. Morrell. “Is self-fertilization an<br />

evolutionary dead end? Revisiting an old hypothesis with genetic<br />

theories and a macrorevolutionary approach.” American Journal<br />

<strong>of</strong> Botany 88 (2001): 1,143–1,150.<br />

West, Stuart A., Curtis M. Lively, and Andrew F. Read. “A pluralist<br />

approach to the evolution <strong>of</strong> sex and recombination.” Journal <strong>of</strong><br />

<strong>Evolution</strong>ary Biology 12 (1999): 1,003–1,012.<br />

Williams, George C. Sex and <strong>Evolution</strong>. Princeton, N.J.: Princeton<br />

University Press, 1975.<br />

sexual selection Sexual selection is the selection <strong>of</strong> characters<br />

that enhance reproductive success rather than (even at<br />

the expense <strong>of</strong>) survival. The individuals in which the characteristic<br />

is well developed will reproduce more <strong>of</strong>ten, and leave

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