Sperm Displacement in the Yellow Dung Fly, Scatophaga stercoraria: An Investigation of Male and Female Processes.

Despite the ubiquity with which patterns of sperm utilization have been studied, the mechanisms underlying fertilization in insects are far from clear. One well-studied system is the yellow dung fly, in which the last male's ejaculate is thought to displace rival sperm from the female's sperm stores. Here we follow the movement of the copulating male's ejaculate through the female's reproductive tract using males labeled with different radioisotopes. We find that males ejaculate into the bursa copulatrix and that male-1 sperm are displaced from the spermathecae during copulation. The increase in male-2 ejaculate in the spermathecae matches the pattern of male-2 fertilization gain, indicating that only spermathecal sperm are utilized at fertilization. Previously we have analyzed this system with a direct model of sperm displacement in which the male displaces rival sperm from the spermathecae. The data, and morphology of the female, clearly preclude such a mechanism. Here we contrast this model with a new indirect model, in which the female facilitates displacement by exchange of sperm from the bursa copulatrix to the spermathecae. The two models give equivalent fits to the observed sperm utilization patterns because the rate of sperm transfer into the bursa copulatrix greatly exceeds the rate of sperm exchange with the spermathecae so that the concentration of the first male's sperm in the bursa remains considerably lower than that of the second male. These analyses provide a quantitative attempt to incorporate female processes into the analysis of sperm utilization patterns in insects.