Seasonal and hormonal modulation of neurotransmitter systems in the song control circuit.

In the years following the discovery of the song system, it was realized that this specialized circuit controlling learned vocalizations in songbirds (a) constitutes a specific target for sex steroid hormone action and expresses androgen and (for some nuclei) estrogen receptors, (b) exhibits a chemical neuroanatomical pattern consisting in a differential expression of various neuropeptides and neurotransmitters receptors as compared to surrounding structures and (c) shows pronounced seasonal variations in volume and physiology based, at least in the case of HVC, on a seasonal change in neuron recruitment and survival. During the past 30 years numerous studies have investigated how seasonal changes, transduced largely but not exclusively through changes in sex steroid concentrations, affect singing frequency and quality by modulating the structure and activity of the song control circuit. These studies showed that testosterone or its metabolite estradiol, control seasonal variation in singing quality by a direct action on song control nuclei. These studies also gave rise to the hypothesis that the probability of song production in response to a given stimulus (i.e. its motivation) is controlled through effects on the medial preoptic area and on catecholaminergic cell groups that project to song control nuclei. Selective pharmacological manipulations confirmed that the noradrenergic system indeed plays a role in the control of singing behavior. More experimental work is, however, needed to identify specific genes related to neurotransmission that are regulated by steroids in functionally defined brain areas to enhance different aspects of song behavior.