Can females benefit from selfing avoidance? Genetic associations and the evolution of plant gender

Evolutionary stability of dioecy and nuclear gynodioecy in higher plants requires that females produce over twice as many successful seeds as hermaphrodites. This fitness differential is widely thought to derive primarily from the advantages of outcrossing caused by high selfing rates and inbreeding depression in the hermaphrodite. This study hypothesized that (i) extraordinarily high deleterious mutation rates are necessary to double female seed success due to outcrossing, and (ii) the large difference in outcrossing rates between sex morphs causes differential purging of these mutations, resulting in additional genetic selection on male sterility. Using genetically explicit models, I showed that the phenotypic outcrossing advantage requires at least one new highly recessive deleterious mutation per genome per generation, regardless of selection coefficient. However, under this mutational regime, differential purging created strong genetic selection against recessive male sterility that overwhelmed the phenotypic selection in favour of outcrossing. In very small populations and for dominant male sterility, this genetic selection was weaker or absent. This first genetically explicit study of the outcrossing advantage of unisexual females may shed new light on both the genetic and selective conditions for the evolution of gynodioecy and dioecy.