Saturday 17 December 2016

evolution - Model for fluctuating selection


Is there any mathematical model to predict the behaviour and long-term consequence of counter-acting selection at different time scale?


For example, let's consider the bi-allelic gene A, with alleles A1 and A2. During a long period of n1 generations A1 is slightly beneficial (differential of selection: s1). After this period, follows a short period of n2 generations when A2 is highly beneficial (differential of selection: s2).


What mathematical model describes the frequency fluctuations of alleles and which allele will get fixed at the long term given the initial frequency ( f0 ), assuming infinite population size and random mating.



Answer



The frequency fluctuations will be determined by a standard model of selection as found in any basic population genetics text. In this scenario they take a very basic form: during each long period $i$ the frequency of $A_1$ increases from $f_i$ to $f_i\cdot (1+s_1)^{n_1}$ and during each short period $j$ the frequency of $A_1$ decreases from $f_j$ to $f_j\cdot (1/(1+s_2))^{n_2}.$ Thus over each pair of periods the frequency of $A_1$ changes by $(1+s_1)^{n_1}/(1+s_2)^{n_2}$. If this quantity exceeds 1, $A_1$ goes to fixation; if it is less than one $A_2$ goes to fixation.


More generally, for an infinite population in a fluctuating environment, the allele with the higher geometric mean fitness will go to fixation. Early discussions of these results are due to Dempster (1955; Cold Spring Harbor Symp. Quant. Biol.), Haldane and Jayakar (1963; J. Genetics), and Lewontin and Cohen (1969; PNAS).



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