The distribution and frequency of vestibulospinal axons and boutons in the upper cervical spinal cord of the cat were investigated. The anterograde tracer Phaseolus vulgaris leucoagglutinin (PHA-L) was injected into discrete regions of the vestibular nuclei, including the medial and descending nuclei, as well as small regions of the lateral vestibular nucleus along its medial border with the medial vestibular nucleus. In contrast to previous reports, labelled vestibulospinal axons were not found to be restricted to the ventromedial and ventrolateral funiculi, but were also observed bilaterally in the lateral funiculi, the dorsolateral funiculi and the dorsal columns. The diameter of these axons ranged from 0.5 to 7.4 microns. Labelled boutons were found bilaterally from lamina IV to IX as well as in lamina X. Contralateral to the injection site, boutons were frequently found as far dorsal as lamina II. Ipsilaterally, boutons were found this far dorsal in only one experiment. There was a dense projection to the contralateral central cervical nucleus, while very few, if any, boutons were observed in the ipsilateral central cervical nucleus. In each experiment, the density of boutons was greater in the rostral cervical segments than in more caudal segments. The "new" vestibulospinal projections to the dorsal horn and central cervical nucleus were confirmed in separate experiments using retrograde transport of horseradish peroxidase. These results show that vestibulospinal axons project to the upper cervical spinal cord via multiple funicular paths. The rich terminations of these axons outside of the ventral horn, as well as in the neck motoneuron nuclei, indicate that vestibulospinal projections must play a wide variety of functions in addition to their well-documented role in the direct control of head movement.
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