 Individuals within the subpopulation are neighbors in the sense that their gametes may come together and inbreeding within the subpopulation increases homozygosity.Wright's statistical theory for isolation by distance looks at population genetic consequences measured by F-statistics where the correlation of randomly uniting gametes within a subpopulation relative to those of the total population is the FST value.formula_1

The equation takes into account the variance in the distribution (var), the allele frequency within the total population (qT), and the allele frequency within the subpopulation (qST).