Evolution of cytoplasmic sex ratio distorters: Effect of paternal transmission

Eukaryotic organisms carry various genetic factors the so-called cytoplasmic genetic elements (CGEs), in their cytoplasm. Numerous examples are known in which CGEs possess the ability to control sex determination of their host organisms and cause sex ratio distortion (SRD). In general, CGEs are inherited maternally from female hosts, via egg cytoplasm to offspring. Thus, the elements tend to evolve abilities to avoid entrance into "dead-end" males. Previous theoretical studies have revealed that, as long as maternal transmission is perfect, CGEs evolve the highest levels of ability to cause SRD. However, it is recently reported that some CGEs transmit from male to offspring through infection to female in mating. This raises the question of how such a paternal contribution alters selective forces and SRD evolution. In the present study, the evolutionary process of SRD ability of CGEs was analyzed theoretically. The main finding is that paternal transmission results in evolution towards intermediate levels of SRD. Further, coexistence was observed of different CGEs inducing different levels of SRD. These results point to the importance of paternal transmission in the evolution of CGEs.