Neuroendocrinology of song behavior and avian brain plasticity: Multiple sites of action of sex steroid hormones

Seasonal changes in the brain of songbirds are one of the most dramatic examples of naturally occuring neuroplasticity that have been described in any vertebrate species. In males of temperate-zone songbird species, the volumes of several telencephalic nuclei that control song behavior are significantly larger in the spring than in the fall. These increases in volume are correlated with high rates of singing and high concentrations of testosterone in the plasma. Several song nuclei express either androgen receptors or estrogen receptors, therefore it is possible that testosterone acting via estrogenic or androgenic metabolites regulates song behavior by seasonally modulating the morphology of these song control nuclei. However, the causal links among these variables have not been established. Dissociations among high concentrations of testosterone, enlarged song nuclei, and high rates of singing behavior have been observed. Singing behavior itself can promote cellular changes associated with increases in the volume of the song control nuclei. Also, testosterone may stimulate song behavior by acting in brain regions outside of the song control system such as in the preoptic area or in catecholamine cell groups in the brainstem. Thus testosterone effects on neuroplasticity in the song system may be indirect in that behavioral activity stimulated by testosterone acting in sites that promote male sexual behavior could in turn promote morphological changes in the song system.