Linkage disequilibrium among multiple neutral alleles produced by mutation in finite population.

An analysis is undertaken for a finite random mating population of the linkage disequilibrium between two loci, at both of which all alleles are neutral, all mutant alleles differ from existing ones and several may be segregating at any time. Formulae are derived for the expected total squared disequilibrium, measured as the sum of squares of disequilibria between all pairs of alleles. The ratio of this quantity to the expected value of the product of the heterozygosities at the two loci is similar to that obtained previously by Ohta and Kimura for two nucleotide sites at each of which not more than two mutant types can segregate at any time.