Introgression of extranuclear genomes in finite populations: nucleo-cytoplasmic incompatibility

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Abstract
Summary: A ‘two locus two allele’ model is developed with special reference to the introgression of extranuclear genomes between two species of finite size. The model assumes that one locus, coded by a nuclear genome, causes the reproductive barrier while the other locus, coded by an extranuclear genome, causes nucleo-cytoplasmic incompatibility in particular genotypes. To fully study this model, simulations are conducted, and a diffusion equation is derived when introgression or extranuclear gene flow occurs in one direction. It is shown that although selection against the nuclear genome can reduce the levels of extranuclear gene flow and retard the introgression process, the dynamics are very similar to those without such selection. In contrast, the nucleo-cytoplasmic incompatibility directly affects the dynamics of introgressing extranuclear genomes: in large populations the ability of incompatibility to overcome extranuclear gene flow is conspicuous, but in small populations it is overshadowed by random sampling drift. Paternal leakage of extranuclear genomes, if present, is of evolutionary importance only when the male's migration rate is much larger than the female's. When the sizes of two populations are unequal, the introgression is most likely to occur from the larger population to the smaller one in the absence of mating preferences of backcross progeny. Recent observations on interspecific mitochondrial transfer in various species do not support the ubiquitousness of nucleo-cytoplasmic incompatibility as an efficient reproductive barrier.