Continuous populations Structured populations Low density of individuals Mainland Stepping Island Stone Some density of individuals model Metapopulation High density of individuals Low density of individuals Subpopulation Isolation by distance: the genetic distance between individuals is correlated with geographic distance Population (Genetics) A population is a group of individuals that interbreed and that is somewhat isolated from other populations. Population versus Deme Species Population Population Population A deme is a group of individuals that interbreed and that is somewhat isolated from other demes. Assessing structure Species Population Deme Deme Deme F-statistics HI = average heterozygosity in individuals over all populations F-Statistics (Wright 1922+) H S = average of expected heterozygosity over all subpopulations HT = expected heterozygosity ignoring structure Individuals I Subpopulations (Demes) S Total Population T FIS H S − HI = HS measures whether there are fewer or more heterozygous individuals than expected in subpopulation FIS > 0 → Fewer heterozygotes than average FIS < 0 → More heterozygotes than average Sewall Wright 1924 F-statistics HI = average heterozygosity in individuals over all populations Contribution to difference between subpopulations H S = average of expected heterozygosity over all subpopulations HT = expected heterozygosity ignoring structure Mutations Genetic drift Local adaptations Degree of genetic differentiation among subpopulations. FST = HT − H S HT FST = ! 0 1 Subpopulations are not differentiated They do not share alleles with each other Migration (Gene flow) reduces differentiation and equalizes allele frequencies among subpopulation Connection between migration and FST 0.44 0.2 FST 0.15 1 = 1 + 4Ne m 10080 Ne m 0.1 60 40 0 .2 0.2 20 0.3 1 0.75 0.1 0.05 Immigration rate 20 FST 40 60 80 100 m 0.5 FST 0.25 0 Ne FST Summary of genetic effects Low population size (N ): reduction of heterozygosity H due to e genetic drift, inbreeding High migration (Nm): increase of H in subpopulation (it brings foreign alleles), counteracts genetic drift and inbreeding, homogenizes populations. Low migration leads to independent evolutionary histories of the subpopulations (high F ), local adaptations ST