Isolation allows populations to diverge and to fix different alleles. Deleterious alleles that reach locally high frequencies contribute to genetic load, especially in inbred or selfing populations, in which selection is relaxed. In the event of secondary contact, the recessive portion of the genetic load is masked in the hybrid offspring, producing heterosis. This advantage, only attainable through outcrossing, should favor evolution of greater outcrossing even if inbreeding depression has been purged from the contributing populations. Why, then, are selfing-to-outcrossing transitions not more common? To evaluate the evolutionary response of mating system to heterosis, we model two monomorphic populations of entirely selfing individuals, introduce a modifier allele that increases the rate of outcrossing, and investigate whether the heterosis among populations is sufficient for the modifier to invade and fix. We find that the outcrossing mutation invades for many parameter choices, but it rarely fixes unless populations harbor extremely large unique fixed genetic loads. Reversions to outcrossing become more likely as the load becomes more polygenic, or when the modifier appears on a rare background, such as by dispersal of an outcrossing genotype into a selfing population. More often, the outcrossing mutation instead rises to moderate frequency, which allows recombination in hybrids to produce superior haplotypes that can spread without the mutation's further assistance. The transience of heterosis can therefore explain why secondary contact does not commonly yield selfing-to-outcrossing transitions.
Focal scenario
These data are the simulated fixation proportions for the focal scenario. Rows represent the selection coefficient. Columns represent the number of fixed deleterious alleles per population.
focal.zip
Genotype frequency trajectories
These data represent the distribution of genotypes at the outcrossing modifier locus over time. The first column represents the frequency of mm, the second column represents the frequency of Mm, and the third column represents the frequency of MM. Each row represents a generation. NAs represent untracked generations after fixation has already occurred.
trajectories.zip
Unequal fitness
These data represent fixation proportions for scenarios in which the two parental populations have unequal fitnesses.
unfair_data.zip
Unequal size
These data represent fixation proportions when the two parental populations contribute unequally to the daughter population. Columns represent number of fixed deleterious alleles per population, and rows represent the initial frequency of the ancestral genotype on which the outcrossing mutation arose.
asymmetric_data_coefficient
Unpurged inbreeding depression
These data represent trials used to determine the effect of initial unpurged inbreeding depression in the parental populations. Each file contains a fixation proportion, designates whether there was initial inbreeding depression or not, and designates how many additional fixed sites per population there were.
initial_depression.zip
Trajectories with inbreeding depression tracked
Contains population trajectories in which average fitness, maximum fitness, inbreeding depression, and modifier genotype frequencies are tracked through time. Except for inbreeding depression, each trajectory is a three-column matrix in which each column represents a modifier-locus genotype (e.g., average fitness of individuals with genotype Mm). Each row is a generation. Inbreeding depression is vector of the population's inbreeding depression through time.
depression_trajectories.zip