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doi:10.1016/j.tpb.2005.03.002
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4 S. Karlin / Theoretical Population Biology 68 (2005) 35
initial state is entirely ab individuals, then in subsequent generations the frequency of the double mutant is always larger without recombination than with it (see also Eshel and Feldman (1970). Karlin (1973) studied the foregoing two-locus model for all circumstances of the selection coefcients producing variable conclusions indicating sensitivity of the model in its dependence on initial population composition and inequalities among the selection coefcients. In summary, for the deterministic model with all mutation events favorable recombination is advantageous in accelerating the incorporation of the most favored double mutant provided that the initial population state exhibits negative linkage disequilibrium. The deterministic case is with favorable double mutations but singly deleterious mutants. It is possible for the recombination mechanism to establish a polymorphism and the nature of this equilibrium state is such that the wild type is most abundant while the other haplotypes are represented with positive but small frequency. In a haploid two-locus selection model under the condition where the forms of Ab and aB have tness disadvantage to ab (the wild type) and AB is the most t type, the pure population consisting only of the ab gamete presents a stable state provided recombination is large enough. If unidirectional mutation a ! A; b ! B is introduced into the system, then a mutation selection balance is developed but a sufciently large recombination force is essential for the stability of the polymorphism. The foregoing result testies to the importance of recombination as a mechanism for maintaining potential and actual variability rather than merely for expressing advantages for the efcient incorporation of favored double mutants. It is possible that the notion of the advantage of recombination should be phrased in terms not of the speed of incorporation of mutants but in terms of the variability it permits. In taking account of nite population size in these models, Bodmer (1970) regards the problem of the time until formation of the rst double mutant type as the important criterion by which to gauge the advantage of recombination. Intuitively, if a state exists where the single mutants Ab and aB are present together in the population, the formation of the rst double mutant will be expected earlier if recombination exists as against no recombination. Maynard Smith (1971) emphasized certain ecological factors. He envisages frequent situations where two genetically different populations migrate into a new environment such that the favored genotype is now a mixture of the genes from the invading populations, and sexuality (rather than recombination) will promote the mixing. There are two basic random variables or expectations: (i) the expected time until rst formation of a double mutant labeled F(i) and the expected time
until total xation of AB denoted by E(i) where i i1 ; i2 ; i3 ; i4 indicates the initial state of the population. Concentrating on the initial state w (0, 0, 0, N) such that the population consists initially of the haplotype ab only, we can infer the following contrasting facts: (i) For xed population size N, F(w) is a decreasing function of r (the recombination rate) while (ii) E(w) is an increasing function of r. Thus, in a nite population, without selection effects, increasing recombination is on an average advantageous in speeding the time of rst formation of the double mutant but has the opposite effect of impeding the time of total incorporation. In other words, sex (recombination) is advantageous in speeding the time of rst formation of the double mutant; but it later breaks apart the favored gamete types tending to stretch the xation time beyond that which would be obtained if no recombination mechanism was operating. These assertions are perhaps surprising since in the innite population model where no selection differences operate, recombination exerts no effect on the rate of incorporation. A proper understanding of the role of recombination in the evolutionary process cannot be discussed only in the context of the problem of incorporation of favorable mutations. There is a large literature underlining the importance and effects of recombination for explaining multi-locus equilibria theory, characterizations of modier genes, the nature of polygenic inheritance, the structure of special mating systems, problems related to sex determination and other factors. Obviously, Maynard Smith was highly innovative and simulating in all these matters. References
Bell, G., 1982. The Masterpiece of Nature: The Evolution and Genetics of Sexuality. University of California Press, Berkeley and Los Angeles. Bodmer, W.F., 1970. The evolutionary signicance of recombination in prokaryotes. Symp. Soc. Gen. Microbiol. 20, 279294. Charnov, E.L., 1982. The Theory of Sex Allocation. Princeton University Press, Princeton. Crew, F.A.E., 1965. Sex Determination, fourth ed. Dover, New York. Eshel, I., Feldman, M.W., 1970. On the evolutionary effect of recombination. Theor. Popul. Biol. 1, 88100. Karlin, S., 1973. Sex and innity: a mathematical analysis of the advantages and disadvantages of genetic recombination. In: Bartlett, M.S., Hiorns, R.W. (Eds.), The Mathematical Theory of the Dynamics of Biological Populations. Academic Press, London, pp. 155194. Karlin, S., Lessard, S., 1986. Theoretical Studies on Sex Ratio Evolution. Princeton University Press, Princeton. Maynard Smith, J., 1968. Evolution in sexual and asexual populations. Am. Nat. 102, 469473. Maynard Smith, J., 1971. What use is sex? J. Theor. Biol. 30, 319335. Maynard Smith, J., 1978. The Evolution of Sex. Cambridge University Press, Cambridge.
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S. Karlin / Theoretical Population Biology 68 (2005) 35 Maynard Smith, J., Price, G.R., 1973. The logic of animal conict. Nature 246, 1518. Maynard Smith, J., Stenseth, N.C., 1978. On the evolutionary stability of the female-biased sex ratio in the wood lemming (Myopus schisticolor): The effect of inbreeding. Heredity 41, 205214. 5
Samuel Karlin Department of Mathematics, Stanford University, Stanford, CA 94305-2125, USA E-mail address: karlin@math.stanford.edu 24 March 2005