When every sperm counts: factors affecting male fertility in the honeybee Apis mellifera

The seminal fluid of male Drosophila contains a cocktail of proteins that have striking effects on male and female fitness. In D. melanogaster, seminal fluid proteins affect female receptivity, ovulation, oogenesis, sperm storage, sperm competition and mating plug formation. In addition, the seminal fluid contains antibacterial peptides and protease inhibitors. Some seminal fluid-encoding genes also show high rates of evolutionary change, exhibiting both significant between-species divergence and within-species polymorphism. Seminal fluid protein genes are expressed only in males, begging the question of how and why the reproductive processes of females are influenced by males. In this review I address these issues by bringing together evidence for the function, evolution, diversification, and maintenance of variation in, seminal fluid-mediated traits.