Sex differences in the volume of avian song control nuclei: comparative studies and the issue of brain nucleus delineation.

Two goals of research on neural sex differences are to establish the behavioral function of such sex differences and to identify precisely what features differ between males and females. Comparative studies of sex differences in the volume of brain nuclei within the songbird vocal control circuit provide one way to address these goals. Informative comparisons can be either inter-specific or intra-specific. Inter-specific comparisons of species within the songbird suborder allow one to establish how species variation in the degree to which there is a sex difference in nuclear volume relates to species variation in the degree to which there is a sex difference in vocal behavior. Intraspecific comparisons of sex differences in nuclear volume involve the comparison of a variety of histochemical methods to define nuclei and describe a nucleus within a species. Sex differences in nuclear volume have now been measured for at least some song control nuclei in 10 different passerine species. In species with more complex male than female song, the volume of key song control nuclei is on average larger in males than in females. However, future studies will require more refined measures of vocal behavior and perceptual abilities to make more precise correlations between brain and behavior. In European starlings (Sturnus vulgaris), the volume of the vocal control nucleus, area X was found to be on average 1.95 times bigger in males than in females based on Nissl stained sections. Variation in neurotransmitter receptor density as determined by quantitative receptor autoradiography can also be used to define clearly the boundaries of a nucleus. When the boundaries of area X in male and female starlings were defined based on variation in muscarinic cholinergic and alpha 2-adrenergic receptor densities, volumetric estimates were obtained that are nearly identical to those obtained with the use of Nissl stains. Intra-specific comparisons of this sort extend our knowledge concerning the neurochemical basis of sex differences in nuclear volume. The wide application of this method would greatly increase our understanding of neural sex differences.