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One of the main conditions of the species splitting from a common precursor lineage is the prevention of a gene flow between diverging populations. The study of  Drosophila  interspecific hybrids allows to reconstruct the speciation mechanisms and to identify hybrid incompatibility factors that maintain post-zygotic reproductive isolation between closely related species. The regulation, evolution, and maintenance of the testis-specific  Ste-Su(Ste)  genetic system in  Drosophila melanogaster  is the subject of investigation worldwide. X-linked tandem testis-specific  Stellate  genes encode proteins homologous to the regulatory β-subunit of protein kinase CK2, but they are permanently repressed in wild-type flies by the piRNA pathway via piRNAs originating from the homologous Y-linked  Su(Ste)  locus. Derepression of  Stellate  genes caused by  Su(Ste)  piRNA biogenesis disruption leads to the accumulation of crystalline aggregates in spermatocytes, meiotic defects and male sterility. In this review we summarize current data about the origin, organization, evolution of the  Ste-Su(Ste)  system, and piRNA-dependent regulation of  Stellate  expression. The  Ste-Su(Ste)  system is fixed only in the  D. melanogaster  genome. According to our hypothesis, the acquisition of the  Ste-Su(Ste)  system by a part of the ancient fly population appears to be the causative factor of hybrid sterility in crosses of female flies with males that do not carry Y-linked  Su(Ste)  repeats. To support this scenario, we have directly demonstrated  Stellate  derepression and the corresponding meiotic disorders in the testes of interspecies hybrids between  D. melanogaster  and  D. mauritiana . This finding embraces our hypothesis about the contribution of the  Ste-Su(Ste)  system and the piRNA pathway to the emergence of reproductive isolation of  D. melanogaster  lineage from initial species.

Stellate Genes and the piRNA Pathway in Speciation and Reproductive Isolation of Drosophila melanogaster