Additional evidence for the genomic imprinting model of sex determination in the haplodiploid wasp Nasonia vitripennis: isolation of biparental diploid males after X-ray mutagenesis

The primary sex-determining signal in the haplodiploid wasp Nasonia vitripennis is not known. In haplodiploid reproduction, unfertilized eggs typically develop into uniparental haploid males and fertilized eggs into biparental diploid females. Although this reproductive strategy is common to all Hymenoptera, sex-determination is not strictly specified by the number of genome copies inherited. Furthermore, primary sex-determining signals differ among haplodiploid species. In the honeybee, for example, the primary signal is the genotype at a single, polymorphic locus: diploid animals that are homozygous develop into males while heterozygotes develop into females. Sex determination in Nasonia cannot be explained by this mechanism. Various lines of evidence show that the inheritance of a paternal genome is required for female sexual development and suggest a genomic imprinting mechanism involving an imprinted gene, expressed only from a paternal copy, that triggers female sexual development. In this model, haploid or diploid uniparental embryos develop into males due to a maternal imprint that silences this locus. The genomic imprinting model predicts that a loss-of-function mutation in the paternal copy of the imprinted gene would result in male sexual development in a biparental diploid embryo. In support of this model, we have identified rare biparental diploid males in the F1 progeny of X-ray mutagenized haploid males. Although uniparental diploid male progeny of virgin triploid females have been previously described, this is the first report of biparental diploid males in Nasonia. Our work provides a new, independent line of evidence for the genomic imprinting model of Nasonia sex determination.