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Available data suggest that the variation in mutation rate among protein loci follows the gamma distribution. Thus, taking into account this variation, formulae are developed for the distribution of allele frequencies, mean and variance of heterozygosity, expected number of alleles, proportion of polymorphic loci, and genetic distance. These formulae should be more appropriate for the analysis of gene frequency data for protein loci than equivalent formulae with constant mutation rate.
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Nei et al. (1976) studied this question.
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