Brainstem reticular nuclei that project to the cerebellum in rats: a retrograde tracer study

Abstract

The nuclear origins of projections from the brainstem reticular formation to the cerebellum were examined using four retrograde tracer substances: horseradish peroxidase, wheat germ agglutinin-horseradish peroxidase conjugate, Fluoro-Gold, and rhodamine beads. Tracer injections were made into each of the three major longitudinal zones of the cerebellar cortex (vermis, paravermal hemisphere, and lateral hemisphere) as well as into the various deep cerebellar nuclei. Counts of retrogradely labeled cells were done on a large sample of select cases. The data generated by these cell counts indicate that the strongest reticulocerebellar projections arise from the three specialized pre-cerebellar reticular nuclei: the lateral reticular nucleus, the medullary paramedian reticular nucleus, and the reticulotegmental nucleus. The presumed noradrenergic locus coeruleus (A6 cell group) was also densely packed with retrogradely labeled neurons. However, strong reticulocerebellar projections also arose from other presumed catecholamine cell groups such as those in the ventrolateral medulla (the A1/C1 complex) and the caudal pons (A5). Substantial cerebellar projections originated from most of the various presumed serotonergic brainstem raphe cell groups (particularly raphe obscurus in the medulla), as well as from the presumed cholinergic Ch5 cell group (the pedunculopontine pars compactus nucleus). Labeled cells were also seen in several nonaminergic isodendritic reticular nuclei thought to be involved in visuomotor activity (e.g. paragigantocellularis dorsalis, raphe interpositus, and the pontine dorsomedial tegmental area), as well as in the lateral reticular zone of the medulla and lower pons (reticularis dorsalis and parvocellularis). Tracer injections into the deep nuclei produced relatively greater numbers of labeled neurons in large-celled medial reticular nuclei associated with skeletomotor activity, such as gigantocellularis, magnocellularis, and pontis caudalis. Reticular nuclei conspicuous in their lack of projections to the cerebellum included reticularis ventralis in the medulla, pontis oralis, and both subdivisions of the midbrain reticular formation (cuneiformis and subcuneiformis). As a whole, the various isodendritic reticular nuclei project most strongly to midline cerebellar structures (vermal cortex or fastigial nuclei), less strongly to the paravermal cortex or interposed nuclei, and least strongly to the lateral cortex or dentate nucleus. Within individual reticular nuclei, the morphology of labeled neurons is identical to that reported previously by this laboratory subsequent to spinal or cortical HRP injections, thus strengthening this laboratory's hypothesis that the various brainstem reticular nuclei can be distinguished on the basis of neuronal morphology.