The cytonuclear effects of facultative apomixis. I. Disequilibrium dynamics in diploid populations.

We comprehensively analyze the cytonuclear effects of generalized mixed mating, including all combinations of selfing, outcrossing, and apomixis, the asexual production of seeds. After first deriving the time-dependent solutions for nonrandom associations (disequilibria) between a diallelic cytoplasmic marker and the alleles and genotypes at a diploid nuclear locus, we delimit all possible dynamical behaviors and the conditions under which each occurs. As in standard mixed mating systems, all disequilibria ultimately decay to zero except when outcrossing is absent, in which case permanent disequilibria result if the allelic association is initially nonzero. When at least some outcrossing is present, any initial allelic association decays at a constant geometric rate, whereas genotypic disequilibria may first increase in magnitude or change sign. Although selfing and apomixis tend to retard the decay of disequilibria (or approach to equilibrium) and often to the same extent, apomixis can have a stronger effect under some conditions. We also determine the dynamics of cytonuclear disequilibria in specific examples that may be of particular interest for empirical studies of hybrid zones. The results suggest several practical guidelines for experimental design and data analysis and show how the cytonuclear disequilibrium dynamics under mating system alone furnish a quantitative baseline for null hypotheses against which to test for the presence of other evolutionary forces.