Sexual differentiation of the vocal control system of birds

Birds evolved neural circuits of various complexities in relation to their capacity to produce learned or unlearned vocalizations. These vocalizations, in particular those that function in the realm of reproduction, are frequently sexually dimorphic, both in vocal learners (songbirds, parrots, some hummingbirds) and vocal nonlearners (all other birds). In many cases, the development and/or the adult differentiation of vocalizations of sociosexual function is sensitive to sex hormones, androgens and estrogens. The underlying mechanisms have been studied in detail in songbirds, a bird group that comprises about half of all bird species. Next to unlearned calls, songbirds produce learned songs that require forebrain vocal control areas that express receptors for androgens and estrogens. These forebrain vocal areas are sexually dimorphic in many species, but a clear relation between the degree of "brain sex" and sex differences in vocal pattern is lacking, except that a minimum number of vocal neurons is necessary to sing learned songs. Genetic brain-intrinsic mechanisms are likely to determine the neuron pools that develop into forebrain song control areas. Subsequently, gonadal steroid hormones, androgens and estrogens, modulate the fate of these neurons and thus the functionality of the vocal control systems. Further action of gonadal hormones, and may be other factors signaling the sociosexual and physical environment, affect the phenotype of vocal control areas in adulthood. Despite the clear evidence of hormone dependency of both adult vocalizations and phenotypes of vocal neuron pools, their causal relation is little understood.