Evolution of intermediate selfing rates in plants: pollination ecology versus deleterious mutations

The evolutionarily stable rate of self-fertilzation is studied in phenotypic models that incorporate pollination ecology as well as the correlated evolution of inbreeding depression and the population mean selfing rate. Inbreeding depression is assumed to be caused by continual mutation to deleterious, partially recessive alleles. Several mutation rates and dominance levels are included. Two separate ecological cases are studied: how selfing rate affects proportion of ovules fertilized (pollination assurance, seed discounting) and how selfing rate affects male outcrossing success through pollen discounting. Evolutionarily stable rates are invariably zero or intermediate in two circumstances, namely when increased selfing causes (1) a decrease in the proportion of ovules fertilized or (2) an increase in pollen discounting and, therefore, a disproportionate decrease in male outcrossing success. Complete selfing is stable when selfing increases the proportion of ovules fertilized for all selfing rates. Stable selfing is zero or one in cases where the selfing rate has no effect on the proportion of ovules fertilized or when pollen discounting does not increase with selfing. Higher inbreeding depression tends to decrease the optimal selfing rate, and lower inbreeding depression (higher dominance coefficients and lower mutation rates) is more favorable to the existence of stable intermediate selfing rates. Approaches such as this that explicitly incorporate the interdependence of selfing, ovule fertilization, and male outcrossing may help explain the persistence of intermediate selfing rates in animal-pollinated plants.