Subcortical contributions to head movements in macaques. II. Connections of a medial pontomedullary head-movement region.

1. In the companion article, a variety of head movements were elicited by stimulation in, and adjacent to, the gigantocellular reticular nucleus (Cowie and Robinson 1994). We refer to this area, caudal to the abducens nucleus, as the gigantocellular head movement region. In the present paper, the anatomical connections of this region, as determined by injections of wheat-germ agglutinin conjugated horseradish peroxidase (WGA-HRP), are reported. The majority of efferent and afferent connections were with areas related to head movements. 2. Efferent fibers from the region projected via two paths to the caudal medulla and upper cervical spinal cord. Labeled fibers descended in the anterolateral funiculus of the ipsilateral spinal cord to terminate in lateral parts of the ventral horn. A second pathway descended bilaterally in the medial longitudinal fasciculus to the anterior funiculi and medial portions of the ventral gray. These efferents paralleled the head-movement topography demonstrated physiologically. Other projections included efferents to the interstitial nucleus of Cajal, caudal field H of Forel, paramedian pontine reticular formation, and caudal vestibular nuclei. Other efferent fibers projected to the trigeminal, facial, and hypoglossal nuclei, as well as to the parvocellular reticular field, which contains interneurons for these motor groups. However, no efferent or afferent labeling involved the ocular motor nuclei. 3. Afferents to the gigantocellular head movement region arose mainly from head-movement areas. In all animals, labeled cells were found in the intermediate and deep layers of the caudal superior colliculus. Labeled neurons also were found in the caudal field H of Forel, interstitial nucleus of Cajal, pontine medial tegmentum including the pontine paramedian reticular formation, nucleus subcoeruleus, and vestibular nuclear complex. Caudally, filled cells were located in the parvocellular, magnocellular, dorsal, and ventral reticular nuclei, the supraspinal nucleus, and the upper cervical ventral horn. 4. In one animal, the ipsilateral frontal cortex contained retrogradely labeled neurons. These cells were found in layer V of cortical areas 4 and 6. Other afferent cells were found consistently in the periventricular and periaqueductal gray matter. 5. A control injection into the caudal vestibular nuclear complex showed projections to the gigantocellular reticular formation and labeled cells in the vestibular and parvocellular reticular nuclei. These observations show that the connections of the gigantocellular region are not typical of all head movement sites. 6. These data indicate that the gigantocellular head-movement region has the requisite efferent and afferent connections to function in the subcortical control of head, but not eye, movements.(ABSTRACT TRUNCATED AT 400 WORDS)