Cell death and the song control system: a model for how sex steroid hormones regulate naturally-occurring neurodegeneration.

The production, learning, and perception of song in songbirds are regulated by a series of discrete brain nuclei known as the song control system. In most songbird species, the song control system is sexually dimorphic, and these dimorphisms become more robust after birds have hatched. In seasonally breeding songbirds, the song control system grows and regresses depending upon breeding context. The development and seasonal plasticity of the song control system are dependent upon neurodegenerative processes, which can be ameliorated, at least in part, by circulating sex steroid hormones. I will describe two areas of song control system research that have provided important information about how hormonal control of cell death contributes to the shaping of behaviorally-relevant brain circuits. First, sexual dimorphism in the zebra finch song control system is robust and emerges partially due to substantial regression of female song control system nuclei during development. Second, in seasonally-breeding songbirds, the song control system regresses as birds transition from breeding to non-breeding conditions. In a controlled laboratory setting where hormones can be acutely withdrawn, these brain areas regress in only a matter of hours to days. Taken together, these results demonstrate that the study of cell death in the song control system provides an excellent opportunity for understanding how changes in circulating levels of sex steroids affect the degeneration of hormone-sensitive brain circuits.