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Insects are the only known animals in which sexual differentiation is controlled by sex-specific splicing. The  doublesex  transcription factor produces distinct male and female isoforms, which are both essential for sex-specific development.  dsx  splicing depends on  transformer , which is also alternatively spliced such that functional Tra is only present in females. This pathway has evolved from an ancestral mechanism where  dsx  was independent of  tra  and expressed and required only in males. To reconstruct this transition, we examined three basal, hemimetabolous insect orders: Hemiptera, Phthiraptera, and Blattodea. We show that  tra  and  dsx  have distinct functions in these insects, reflecting different stages in the changeover from a transcription-based to a splicing-based mode of sexual differentiation. We propose that the canonical insect  tra-dsx  pathway evolved via merger between expanding  dsx  function (from males to both sexes) and narrowing  tra  function (from a general splicing factor to dedicated regulator of  dsx ).

Hemimetabolous insects elucidate the origin of sexual development via alternative splicing