Convergence of somatosensory and auditory projections in the avian torus semicircularis, including the central auditory nucleus

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Abstract
Projections of dorsal column, spinal, and cochlear nuclei upon the central nucleus of the torus semicircularis (otherwise known as nucleus mesencephalicus lateralis, pars dorsalis, or MLd) and upon other toral nuclei were investigated in pigeon by anterograde and retrograde tracing and electrophysiological methods. The anatomical results showed that caudal regions of the dorsal column nuclei and medial lamina V of the upper four cervical spinal segments have extensive projections upon the contralateral central auditory nucleus and upon other nuclei of the torus, in particular the core portion of the preisthmic superficial area of Puelles et al. (L. Puelles, C. Rrobles, M. Martiez‐de‐la‐Torre, and S. Martinez, 1994, J. Comp. Neurol. 340:98–125). The projections of nucleus angularis were found to terminate throughout most of the contralateral central nucleus except the dorsomedial portion at rostral levels, where the majority of the projections of nucleus laminaris were concentrated. Nucleus angularis (and to a lesser extent nucleus laminaris) was also found to have substantial projections to certain noncentral toral nuclei, in particular to the caudomedial shell nucleus of Puelles et al. (1994). As shown positively with both Nissl and cytochrome oxidase staining and negatively with substance P labeling, this nucleus is a medial extension of more caudal regions of the central nucleus, and it is suggested that it should be included as part of the auditory midbrain. The electrophysiological results confirmed the anatomical findings by showing that evoked potentials and multiunit activity can be recorded throughout the central and noncentral toral nuclei by using electrical stimulation of the radial nerve and auditory click stimuli. The core portion of the preisthmic superficial area, however, can be regarded as a distinct somatosensory nucleus of the midbrain. It is concluded that there is substantial convergence of somatosensory and auditory inputs within both central auditory and noncentral nuclei of the torus semicircularis in pigeon.