Avian nucleus retroambigualis: cell types and projections to other respiratory-vocal nuclei in the brain of the zebra finch (Taeniopygia guttata).

In songbirds song production requires the intricate coordination of vocal and respiratory muscles under the executive influence of the telencephalon, as for speech in humans. In songbirds the site of this coordination is suspected to be the nucleus retroambigualis (RAm), because it contains premotor neurons projecting upon both vocal motoneurons and spinal motoneurons innervating expiratory muscles, and because it receives descending inputs from the telencephalic vocal control nucleus robustus archopallialis (RA). Here we used tract-tracing techniques to provide a more comprehensive account of the projections of RAm and to identify the different populations of RAm neurons. We found that RAm comprises diverse projection neuron types, including: 1) bulbospinal neurons that project, primarily contralaterally, upon expiratory motoneurons; 2) a separate group of neurons that project, primarily ipsilaterally, upon vocal motoneurons in the tracheosyringeal part of the hypoglossal nucleus (XIIts); 3) neurons that project throughout the ipsilateral and contralateral RAm; 4) another group that sends reciprocal, ascending projections to all the brainstem sources of afferents to RAm, namely, nucleus parambigualis, the ventrolateral nucleus of the rostral medulla, nucleus infra-olivarus superior, ventrolateral parabrachial nucleus, and dorsomedial nucleus of the intercollicular complex; and 5) a group of relatively large neurons that project their axons into the vagus nerve. Three morphological classes of RAm cells were identified by intracellular labeling, the dendritic arbors of which were confined to RAm, as defined by the terminal field of RA axons. Together the ascending and descending projections of RAm confirm its pivotal role in the mediation of respiratory-vocal control.