Reinforcement of genetic coherence in a two-locus model

In order to maintain populations as units of reproduction and thus enable anagenetic evolution, genetic factors must exist which prevent continuing reproductive separation or enhance reproductive contact. This evolutionary principle is called genetic coherence and it marks the often ignored counterpart of cladistic evolution. Possibilities of the evolution of genetic coherence are studied with the help of a two-locus model with two alleles at each locus. The locus at which viability selection takes place is also the one that controls the fusion of gametes. The second locus acts on the first by modifying the control of the fusion probabilities. It thus acts as a mating modifier whereas the first locus plays the role of the object of selection and mating. Genetic coherence is enhanced by modifications which confer higher probabilities of fusion to heterotypic gametic combinations (resulting in heterozygous zygotes) at the object locus.

It is shown that mutants at the mating modifier locus, which increase heterotypic fusions but do not lower the homotpyic fusions relative to the resident allele at the object locus, generally replace the resident allele. Since heterozygote advantage at the object locus is a necessary condition for this result to hold true, reinforcement of genetic coherence can be claimed for this case. If the homotypic fusions are lowered, complex situations may arise which may favor or disfavor the mutant depending on initial frequencies and recombination rates. To allow for a generalized analysis including alternative models of genetic coherence as well as the estimation of its degrees in real populations, an operational concept for the measurement of this degree is developed. The resulting index is applied to the interpretation of data from crossing experiments in Alnus species designed to detect incompatibility relations.