Axonal connections of the medial magnocellular nucleus of the anterior neostriatum in zebra finches.

The medial magnocellular nucleus of the anterior neostriatum (mMAN) is a small cortical nucleus which was previously identified as a component of the neural circuitry controlling vocal behavior in songbirds based on its efferent connection to the High Vocal Center (HVC), a major song control nucleus (Nottebohm et al. [1982] J. Comp. Neurol. 207:344-357; Bottjer et al. [1989] J. Comp. Neurol. 279:312-326). We have conducted tract tracing experiments (using wheat-germ agglutinin-horseradish peroxidase (WGA-HRP), the carbocyanine dye DiI, and biocytin) to determine the complete pattern of afferent and efferent connections of mMAN in adult male zebra finches. We confirmed the existence of an efferent projection from mMAN to HVC and discovered a novel projection to the region medial to caudal HVC called paraHVC (pHVC). Injections of retrograde tracers into mMAN showed that afferent input to mMAN originates from the dorsomedial nucleus of the posterior thalamus (DMP). Injections of DiI into DMP produced anterograde label over mMAN, thus confirming the DMP-to-mMAN projection. Interestingly, this anterograde label extended beyond the region of mMAN defined by HVC-projecting neurons into the immediately surrounding cortex. This extended terminal field of DMP efferents indicates that mMAN encompasses a core population of projection neurons surrounded by a shell of non-HVC-projecting neurons, both of which receive input from the dorsal thalamus. Analysis of retrograde DiI label resulting from DMP injections revealed two major sources of afferent input to DMP originating in regions of the archistriatum and hypothalamus. Inputs to DMP were distributed throughout the dorsal archistriatum and included the area that receives a projection from the parvicellular shell region of the lateral magnocellular nucleus of the anterior neostriatum, a song control nucleus, as well as the dorsal portion of the robust nucleus of the archistriatum, the motor-cortical output of the song control system. The projections from song control regions of the archistriatum to DMP may feed information back into telencephalic song control circuitry via the DMP-->mMAN-->HVC/pHVC pathway. The other source of afferent input to DMP is located in the external cellular stratum of the lateral hypothalamus (SCE). This newly delineated SCE-->DMP-->mMAN-->HVC/pHVC pathway is the first report of a hypothalamic brain region neuroanatomically integrated with song control circuitry. Because hypothalamic brain regions are important for homeostasis and regulating behavior, the trans-synaptic circuitry of mMAN may help to integrate information about the bird's internal state, such as sexual maturation, with song learning and production.