Topography of cerebellar nuclear projections to the brain stem in the rat.

The organization of the cerebellum is characterized by a number of parallel and parasagittally ordered olivocorticonuclear modules; as such, the cerebellar nuclei basically function as output system of these modules. The present study provides a comprehensive and detailed description of the organization of the connections from the cerebellar nuclei to the brain stem in the rat. Thirteen small injections with the anterograde tracer Phaseolus vulgaris leucoagglutinin or biotinylated dextran amine which were centered on various aspects of the cerebellar nuclear complex are described and are illustrated with serial plots detailing the distribution of labeled varicosities throughout the brain stem. In every case at least 1,000 an up to 36,000 varicosities were plotted. All injections resulted in some or heavy labeling concentrated within specific regions of the contralateral inferior olivary complex and, usually, in some labeling of the contralateral ventrolateral thalamus. However, apart from these two areas it is shown that the cerebellar projections are generally very widespread and may be found throughout the entire brain stem. Below, only a survey of main projection areas will be given. Terminal arborizations originating from the rostral part of the medial cerebellar nucleus are mostly found in the caudal half of the brain stem with emphasis on the vestibular nuclear complex, whereas its caudal part rather connects to midbrain areas. Terminals that originate from the dorsolateral protuberance of the medial cerebellar nucleus are distributed more evenly throughout the brain stem and are mostly confined to reticular areas. The interstitial cell groups, interspersed between the medial and both interposed cerebellar nuclei, provide major projections to the ipsilateral vestibular nuclear complex and contralateral mesodiencephalic regions. However, reticular areas are also targeted over a large rostrocaudal range. The medial part of the posterior interposed nucleus sends most projections to the caudomedial red nucleus, prerubral regions and parvicellular reticular formation, all contralateral to the injection site. Projections that originate from more laterally placed injections are directed, apart from the inferior olivary complex, to the rostral half of the contralateral brain stem, where most labeled varicosities are found in the superior colliculus and zona incerta. The anterior interposed nucleus specifically targets the inferior olive, the red nucleus, the pontine reticulotegmental nucleus, the prectectum and the ventrolateral thalamic nucleus. More laterally placed injections also project to the ipsilateral parvicellular reticular formation and deep layers of the spinal trigeminal complex. The latter areas are more specifically targeted by the dorsolateral hump. In addition, its projections are found in the red nucleus and pretectum but do not seem to reach the ventrolateral thalamus. Projections from the lateral cerebellar nucleus are all characterized by a widespread distribution of terminals. Especially, the caudal aspect of the nucleus sends, apart from projections to the deep mesencephalic nucleus, red nucleus, periaquaductal gray, pretectum, prerubral area, and several thalamic regions, prominent projections to the caudal brain stem which terminate in the inferior olive and gigantocellular reticular formation. Projections from the ventral, parvicellular part of the nucleus are mostly, but not exclusively, directed to the rostral half of the brain stem and mainly terminate in the pararubral area, accessory oculomotor nuclei, pretectal areas, zona incerta, and in the parafascicular and ventrolateral thalamic nuclei. We conclude that the impact of the cerebellar nuclei on the brain stem is widespread; projections from different regions of the same cerebellar nucleus may show important differences in distribution of labeled terminals. On the other hand, injections placed in different cerebellar nuclei may result in a simila