The vestibulo-ocular and the vestibulospinal reflexes: noradrenergic influences on the plastic changes which affect the cerebellar cortex during vestibular adaptation.
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The sources and pathways of the climbing fibers to the cerebellar posterior vermis were studied with comibined electrophysiological and anatomical methods in cats. Recording from identified cerebellar Purkinje cells, monosynaptic climbing fiber (CF) responses have been obtained both for stimulation of the inferior olive (IO) and various parts of the brain stem (BS). CF responses were found to of three types, IO only, BS only or both IO and BS. However the responses to BS stimulation were very few in number in comparison with IO or IO and BS types of responses. The latencies of the responses were shorter for the BS cases consistent with their distance from the cerebellum. A comparison of latencies and the relative responsiveness of the different area of the brain stem which were studied, indicate that part of the CF ascend through the pontine region and enter the cerebellum by way of the medium and superior penduncles. This finding is confirmed by the results of anatomical studies in which degenerating fibers were found in the molecular layer (using the Nauta technique) after lesion of the brachium pontis but not after lesions of the medial portion of the pons. Similarly, injection of radioactive leucine into the pontine nuclei failed to show any labeled fibers in the molecular layer. Horseradish peroxidase (HRP) was injected into localized regions of the posterior vermis after total bilateral destruction of the inferior peduncles. Large numbers of positive, marked cells were still found in the inferior olive. It is concluded that nearly all, if not all, the climbing fibers originate in the inferior olive and that they ascend to the cerebellum by way of all the peduncles.
We recorded from Purkinje cells in the anterior and posterior lobes of the cerebellar vermis in anesthetized cats during passive flexion and extension of the ipsilateral hind foot. Nearly 60% of the 302 cells recorded in lobules I-IV anterior and VII-IX posterior responded to this stimulus. In lobule V, generally considered to be the forelimb projection area, about a third of the 38 cells tested responded to hind foot movement. Cells in the anterior lobe typically exhibited different poststimulus patterns in response to foot flexion and extension while posterior cells usually had the same response to these stimuli. We also noted that cells recorded in the medial vermis (within 2.7 mm of the midline) generally had longer response latencies than cells recorded more laterally (2.7 mm of the midline) generally had longer response latencies than cells recorded more laterally (2.7-4.5 mm) in both lobes. The results illustrate a diffuse proprioceptive input to the cerebellar vermis which has different features in various cerebellar areas.
The cerebellum is subdivided into hundreds of discrete modules defined by their connectivity and molecular signatures. Cerebellar compartmentation arises very early in development through the formation of multiple populations of chemically distinct Purkinje cells that migrate in a coordinated fashion to form parasagittal bands of cells. Different Purkinje cell bands are then innervated by discrete subpopulations of cerebellar afferents. Because of its stereotyped and strikingly beautiful organization the cerebellum is an excellent model in which to explore genetic/epigenetic aspects of pattern formation in the central nervous system.
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