Corticofugal connections between the cerebral cortex and brainstem vestibular nuclei in the macaque monkey

The distribution of cortical efferent connections to brainstem vestibular nuclei was quantitatively analysed by means of retrograde tracer substances injected into different electrophysiologically identified parts of the brainstem vestibular nuclear complex of five Java monkeys (Macaca fascicularis). Three polysensory vestibular areas were found to have a substantial projection to the vestibular nuclei: area 2v located at the tip of the intraparietal sulcus, the parietoinsular vestibular cortex (PIVC) covering the most occipital part of the granular insula (Ig) and the retroinsular area (Ri or reipt), and the dorsolateral part of the somatosensory area 3a (“area 3aV” neck/trunk region). From physiological recording experiments, these three cortical fields were known to contain many neurons responding to stimulation of semicircular canals as well as to optokinetic (area 2v, PIVC) and somatosensory stimuli (PIVC, area 3a). These three regions form the inner cortical vestibular circuit. Besides these polysensory vestibular cortical fields, three other circumscribed cortical regions of the macaque brain were also found to project directly to the brainstem vestibular nuclei: a circumscribed part of the postarcuate premotor cortex (area 6pa), part of the agranular and the adjacent dysgranular cortex located around the cingulate sulcus (area 6c/23c), and a predominantly visual (optokinetic) association field located at the fundus of the lateral sulcus (area T3). These areas are known to have connections with the structures of the inner cortical vestibular circuit. Only a few efferent connections to the brainstem vestibular nuclei were found for the different parts of cytoarchitectonic area 7. Significant differences were found between the efferent innervation patterns of the axons originating in the six cortical areas mentioned and ending in the various compartments of the vestibular nuclear complex. Vestibular nuclei with a dominant output to the gaze motor system of the brainstem receive efferent connections preferably from the parietoinsular vestibular cortex. Vestibular structures with their primary output to skeletomotor centers, however, receive stronger efferent connections from areas 6pa and 3a. The ventrolateral nucleus, which sends efferent axons to both the oculomotor and skeletomotor systems of the brainstem and the spinal cord, also receives its main cortical efferents from the somatomotor area 6 and from area 3aV. Through these connections the corti