HYBRID BREAKDOWN BETWEEN TWO HAPLODIPLOID SPECIES: THE ROLE OF NUCLEAR AND CYTOPLASMIC GENES.

A central question in evolutionary biology concerns the population and genetic processes by which new species arise. Here, the genetic basis of hybrid breakdown between two haplodiploid species, Nasonia vitripennis and N. giraulti is investigated. Hybridization between the two species is normally prevented by microorganisms that cause bidirectional incompatibility. However, after elimination of microorganisms, F hybrids females are readily produced (due to haplodiploidy, males develop from unfertilized eggs and are therefore not hybrids). F hybrid females are viable and fecund, but recombinant (haploid) F male offspring suffer from severe hybrid breakdown (larval and pupal mortality). This is typically interpreted as evidence for the existence of different coadapted gene complexes in the two species, which are broken up by recombination. F recombinant eggs were rescued by fertilization with the complete chromosome complement from either species, supporting the view that hybrid lethality genes tend to be recessive. Negative epistatic interactions occur between nuclear genes of the two species, and between cytoplasmically inherited factors (cytoplasmic genes) of giraulti and nuclear genes of vitripennis. Interactions between nuclear genes and cytoplasmic genes are asymmetric. Experiments clearly demonstrate that the latter incompatibility is not due to maternal-effect genes, but to cytoplasmically inherited elements. Nuclear-mitochondrial interactions are possibly involved.