& Hatfield, 2000). These have shown that the mutational load and inbreeding depression are generally increased by high migration rates and large local population sizes, while heterosis in betweenpopulation crosses is reduced. While the theoretical expectations are in qualitative agreement with the empirical results, no attempt has been made to determine whether the data quantitatively match theoretical predictions based on current estimates of mutation rates and the distribution of fitness effects of new deleterious mutations obtained from genomic data, reviewed by Keightley (2012) andCharlesworth (2015). Furthermore, the theoretical work has assumed that mutation rates per locus or nucleotide site are small, relative to the strength of selection against deleterious alleles. In addition, the forward mutation rate (from wild type to mutant) is usually assumed to be much greater than the backward mutation rate (from mutant to wild type), the latter often being ignored. The first assumption is usually well justified for mutations affecting protein sequences and strongly constrained functional non-coding sequences. The second assumption applies to the totality of mutations that affect a functional unit such as a coding sequence, provided that selection is sufficiently Abstract This paper examines the extent to which empirical estimates of inbreeding depression and interpopulation heterosis in subdivided populations, as well as the effects of local population size on mean fitness, can be explained in terms of current estimates of mutation rates, and the distribution of selection coefficients against deleterious mutations provided by population genomics data. Using population genetics models, numerical predictions of the genetic load, inbreeding depression and heterosis were obtained for a broad range of selection coefficients and mutation rates. The models allowed for the possibility of very high mutation rates per nucleotide site, as is sometimes observed for epiallelic mutations in plants. There was fairly good quantitative agreement between the theoretical predictions and empirical estimates of heterosis and the effects of population size on genetic load, on the assumption that the deleterious mutation rate per individual per generation is approximately one, but there was less good agreement for inbreeding depression. Weak selection, of the order of magnitude suggested by population genomic data, is required to explain the observed patterns. Possible caveats concerning the applicability of the models are discussed.
K E Y W O R D Sepiallelic mutations, genetic load, heterosis, inbreeding depression, mutation rate, subdivided population