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Wolbachia  are maternally inherited bacteria that infect arthropod species worldwide and are deployed in vector control to curb arboviral spread using cytoplasmic incompatibility (CI). CI kills embryos when an infected male mates with an uninfected female, but the lethality is rescued if the female and her embryos are likewise infected. Two phage WO genes,  cifA   wMel   and  cifB   wMel   from the  w Mel  Wolbachia  deployed in vector control, transgenically recapitulate variably penetrant CI, and one of the same genes,  cifA   wMel  , rescues wild type CI. The proposed Two-by-One genetic model predicts that CI and rescue can be recapitulated by transgenic expression alone and that dual  cifA   wMel    and cifB   wMel   expression can recapitulate strong CI. Here, we use hatch rate and gene expression analyses in transgenic  Drosophila melanogaster  to demonstrate that CI and rescue can be synthetically recapitulated in full, and strong, transgenic CI comparable to wild type CI is achievable. These data explicitly validate the Two-by-One model in  w Mel-infected  D .  melanogaster , establish a robust system for transgenic studies of CI in a model system, and represent the first case of completely engineering male and female animal reproduction to depend upon bacteriophage gene products.

Two-By-One model of cytoplasmic incompatibility: Synthetic recapitulation by transgenic expression of cifA and cifB in Drosophila