Evolutionary transitions between sex-determining mechanisms: a review of theory

The extraordinary diversity of sex-determining mechanisms found in nature is thought to have arisen by the addition, modification or replacement of regulators at the upstream end of the sex-determining pathway. The spread of a novel regulator of sex determination can manifest itself by an evolutionary transition between environmental and genetic sex determination, for example, or between male and female heterogamety. Both kinds of transition have occurred frequently in the course of evolution. In this paper, various evolutionary forces acting on sex-determining mutations that can bias transitions in one direction or the other are reviewed. Furthermore, the adaptive significance of the main modes of sex determination are discussed, and the common principle underlying ultimate explanations for environmental sex determination, genetic sex determination and maternal control over sex determination in the offspring are highlighted. Most of the current theory concentrates on the population-genetic aspects of sex determination transitions, using models that do not reflect the developmental mechanisms involved in sex determination. However, the increasing availability of molecular data creates opportunities for the future development of mechanistic models that will further clarify how selection and developmental architecture interact to direct the evolution of sex determination genes.