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Page 137
C0153-09.gif is bounded below by C0153-10.gif (following Lemma 7.2). For a fraction Dt of a generation we can write
C0153-01.gif
If M(t) is the size of the population at time t (allowing the overall population size to be variable for the time being), then
C0153-02.gif
and
C0153-03.gif
using the fact that the population as a whole increases at a rate determined by the observed average fitness C0153-11.gif. It follows that
C0153-04.gif
where C0153-05.gif.
If we use a discrete time-scale t = 1, 2, 3, . . . then Dt = (t + 1) - t = 1 and
C0153-06.gif
If we take the limit as Dt ® 0, in effect going to a continuous time-scale, we have
C0153-07.gif
The equation C0153-08.gif, when restricted to alleles (schemata defined on one position), is just Fisher's (1930) classical result, relating the change in proportion of an allele to its average excess. We see however that the equation holds for arbitrary schemata. This gives us a way of predicting the rate of increase of a set of alleles with epistatic interactions from a sample average C0153-12.gif of the fitnesses of chromosomes carrying the set of alleles.

 
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