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PubMed1989

Acquisition of classical conditioning without cerebellar cortex.

The left cerebellar cortex was surgically aspirated in rabbits who were then subsequently trained for classical conditioning of the nictitating membrane. All rabbits were trained sequentially on both eyes. Rabbits with the lesion confined to the cerebellar cortex were able to learn with the eye ipsilateral to the lesion although it took many times longer than reported for either naive rabbits or for rabbits first trained on the unlesioned, contralateral side. Rabbits with lesions that included the cerebellar cortex and the cerebellar interpositus nucleus did not learn with the eye ipsilateral to the lesion. Learning with the eye contralateral to either type of lesion was always very rapid. It is now clear on the basis of this and previous studies that cerebellar cortex, unlike the cerebellar interpositus nucleus, is not essential for acquisition or relearning/retention of classical conditioning. However, cerebellar cortex normally plays an important role since acquisition of classical eyeblink conditioning is prolonged and of poor quality in its absence.

Animals↗
PubMed1983

Scanning electron microscope recognition of intracortical climbing fiber pathways in the cerebellar cortex.

The cerebellar cortex of adult mouse, rat, fish and human has been explored by SEM in order to identify and trace the climbing fiber intracortical pathways and their synaptic relationships. Samples were processed by the freeze-fracture method for SEM using medium and rapid freezing rates. Climbing fibers appeared at the white matter as fine, wavy axonal processes with a characteristic cross-over bifurcation pattern. These features allowed us to differentiate them from the mossy fibers, which are distinguished as thick fibers with a straight course and a dichotomous arborization. In their course through the granular layer the climbing fibers exhibited glomerular and tendril collaterals. The glomerular collaterals appeared covered by the dendritic tips of several granule cells forming thin triangular climbing glomeruli. The tendril collaterals are seen as highly contoured axonal ramifications establishing axo-dendritic connections with the granule cell dendritic digits. Climbing fibers were also observed forming a pericellular plexus around the Golgi cell body. They cross the granular layer and reach the Purkinje cell perikaryon contributing to the formation of infraganglionic plexus and pericellular nest. The climbing fibers showed a typical sagittal compartmentalization and a contoured spiral or zigzag pathway in the course ascending along the Purkinje dendritic branchlets. Fine terminal climbing fiber tendrils synapsing on the tips of Purkinje dendritic spines were observed. The freeze-fracture method for SEM permits us to estimate with a more reliable degree of certainty the amount of branching or lateral collateralization of climbing fibers in the granular and molecular layers.

Animals↗
PubMed2001

The entire trajectories of single olivocerebellar axons in the cerebellar cortex and their contribution to Cerebellar compartmentalization.

The functional partitioning of the cerebellar cortex depends on the projection patterns of its afferent and efferent neurons. However, the entire morphology of individual projection neurons has been demonstrated in only a few classes of neurons in the vertebrate CNS. To investigate the contribution of the projection pattern of individual olivocerebellar axons to the cerebellar functional compartmentalization, we labeled individual olivocerebellar axons, which terminate in the cerebellar cortex as climbing fibers, with biotinylated dextran amine injected into the inferior olive in the rat, and completely reconstructed the entire trajectories of 34 olivocerebellar axons from serial sections of the cerebellum and medulla. Single axons had seven climbing fibers on average, which terminated at similar distances from the midline in a single or in multiple lobules. Cortical projection areas of adjacent olivary neurons were clustered as narrow but separate longitudinal segments and often innervated by collaterals of single neurons. Comparison of the cerebellar distribution of olivocerebellar axons arising from different sites within a single olivary subnucleus indicated that slightly distant neurons projected to complementary sets of such segments in a single longitudinal band. Several of these longitudinal bands formed a so-called parasagittal zone innervated by a subnucleus of the inferior olive. Single olivocerebellar axons projected rostrocaudally to segments within a single band but did not project mediolaterally to multiple bands. These results suggest fine substructural organization in the cerebellar compartmentalization that may represent functional units.

Animals↗
PubMedDate not supplied

Stimulus generalization of conditioned eyelid responses produced without cerebellar cortex: implications for plasticity in the cerebellar nuclei.

In Pavlovian eyelid conditioning and adaptation of the vestibulo-ocular reflex, cerebellar cortex lesions fail to completely abolish previously acquired learning, indicating an additional site of plasticity in the deep cerebellar or vestibular nucleus. Three forms of plasticity are known to occur in the deep cerebellar nuclei: formation of new synapses, plasticity at existing synapses, and changes in intrinsic excitability. Only a cell-wide increase in excitability predicts that learning should generalize broadly from a training stimulus to other stimuli capable of supporting learning, whereas the alternatives predict that learning should be relatively specific to the training stimulus. Here we show that deep nucleus plasticity, as assessed by conditioned eyelid responses produced without input from the cerebellar cortex, is relatively specific to the training conditioned stimulus (CS). We trained rabbits to a tone or light CS with periorbital stimulation as the unconditioned stimulus (US), and pharmacologically disconnected the cerebellar cortex during a posttraining generalization test. The short-latency conditioned responses unmasked by this treatment showed strong decrement along the dimension of auditory frequency and did not generalize across stimulus modalities. These results cannot be explained solely by a cell-wide increase in the excitability of deep nucleus neurons, and imply that an input-specific mechanism in the deep cerebellar nucleus operates as well.

Acoustic Stimulation↗
PubMed1986

Choline and ethanolamine phosphotransferase activities in glomerular particles isolated from bovine cerebellar cortex.

Isolated cerebellar glomeruli provide a relatively homogeneous subcellular fraction, which can be used to study the biochemical events related to chemical transmission within a well-characterized central synapse. Choline and ethanolamine phosphotransferase activities were identified and partially characterized in this nerve ending preparation. Choline phosphotransferase associated with the glomerular particles required Mg2+, while ethanolamine phosphotransferase required Mn2+ for optimal activities. Both enzymes were inhibited by exogenous Ca2+. The apparent Vmax values were 35.9 and 10.0 nmol/hr per mg protein for the choline and ethanolamine phosphotransferases, respectively. The apparent Km value for the CDPcholine substrate was 28.6 microM, and the Km for CDPethanolamine was 8.3 microM. Neither enzyme responded to the various adenine nucleotides, neurotransmitters or neurotransmitter agonists tested. However, exposure of the glomerular particles to cytidine nucleotides inhibited ethanolamine phosphotransferase activity and stimulated choline phosphotransferase activity.

Animals↗
PubMed1993

Neuronal-glial exchange of taurine during hypo-osmotic stress: a combined immunocytochemical and biochemical analysis in rat cerebellar cortex.

Rat cerebellar Purkinje cells show a high level of taurine-like immunoreactivity. Light-microscopic immunocytochemistry indicated that the level of taurine in these cells was substantially decreased in animals that had survived for 4 h after an intraperitoneal injection of distilled water. This treatment resulted in a 15-20% reduction in plasma osmolality. The changes in the Purkinje cells were accompanied by an increased immunolabeling of neighboring glial cells (Golgi epithelial cells). The changes in both cell types were reversed in animals whose plasma osmolality had been normalized by injections of hypertonic saline 4 h after the water loading. Adjacent sections incubated with a GABA antiserum did not exhibit any overt changes in response to the hypo-osmotic stress. Quantitative electron-microscopic analysis of ultrathin sections subjected to postembedding immunogold cytochemistry indicated that the Purkinje cells had lost 50-60% of their taurine contents after water loading and that the loss affected all intracellular compartments, including mitochondria and cytoplasmic matrix. The loss of taurine immunoreactivity from Purkinje cells was accompanied by an estimated 70-80% increase in the contents of immunoreactive taurine in adjacent glial cells. Biochemical recordings of tissue amino acids in a parallel series of animals revealed a 12% reduction in cerebellar taurine contents 4 h after water loading (value corrected for changes in specific gravity). This reduction had progressed to 32% after 8 h and was only partly prevented by normalization of plasma osmolality. The tissue levels of GABA and several other amino acids showed a decrease similar to that of taurine, while glutamine displayed a considerable increase after water loading. Our findings indicate that acute reductions in plasma osmolality cause a flux of taurine from Purkinje cells to glia, and that this flux is reversed upon normalization of plasma osmolality. These changes are superimposed on a decrease in the biochemically recorded tissue level of taurine. Unlike the cellular redistribution, this decrease was not reversible within the time frame of the present study, and it was not specific for taurine. Cellular redistribution of taurine may represent a rapid adjustment to osmotic perturbations in vivo. In addition, it may reflect a higher priority for neuronal compared with glial volume regulation.

Amino Acids↗
PubMed1981

Zones in the cerebellar cortex. Their organization and potential relevance to cerebellar stimulation.

The organization of zones in the cerebellar cortex, as reflected by the arrangement of cortical efferent and afferent fibers, was reviewed with special emphasis on the anterior lobe. There is conclusive evidence for the existence of at least six, and possibly as many as nine, ipsilateral rostrocaudally oriented cortical zones, each having specific afferent and efferent connections. The topography afferent and efferent fibers of the cerebellar cortex and/or the spatial arrangement of cortical zones have not been given appropriate attention in both experimental and clinical studies dealing with focal electrical stimulation of the cerebellum. It is suggested that inattention to the relationships between electrode placement and zones in the cerebellar cortex may be partially responsible for the sometimes capricious results during and/or subsequent cerebellar cortical stimulation.

Animals↗
PubMed1979

The organization of afferents to the cerebellar cortex in the cat: projections from the deep cerebellar nuclei.

The topography of the cerebellar nucleo-cortical projection was investigated in the cat by experiments employing the horseradish peroxidase (HRP) technique or by combined HRP-autoradiographic methods. The results of the HRP studies extend previous findings showing that neurons in the deep nuclei project to the cerebellar cortex in an orderly way. Thus, it appears that the cortex of the vermis-proper receives projections from neurons located predominately in the fastigial nucleus. Intermediate and lateral zones of mid-vermal cerebellar cortex are projected on by neurons located in the interposed and dentate nuclei. Crus II receives input from neurons located predominately in the dentate nucleus, while the paramedian lobule is projected on by neurons located in a large postero-dorsal sector of the interposed nucleus and in a smaller medial strip of the dentate nucleus. Neurons in the ventral part of the dentate nucleus and the lateral part of the interposed nucleus send fibers to the paraflocculus. The nucleo-cortical pathway to the flocculus and nodulus arises largely from a population of neurons located in a ventral region stretching from the medial border of the dentate nucleus to the lateral border of the fastigial nucleus. The results of experiments using the combined HRP-autoradiographic method show that clusters of neurons in the deep cerebellar nuclei project back to the cerebellar cortical areas from which they receive input, establishing a fairly precise feedback loop between the cerebellar cortex and deep nuclei.

Afferent Pathways↗
PubMed1991

Scanning electron microscopy of the granular layer of rat cerebellar cortex.

The rat cerebellar cortex has been studied with respect to its cytoarchitectural arrangement using Scanning Electron Microscope. We have distinguished several types of cells in the granular layer and characterized the granules and the Golgi cells. We also characterized the mossy fiber glomeruli which appears round or ovoid. The mossy fibers revealed their "en passant" nature. The climbing fibers formed tendril collaterals and glomeruli. The Golgi cell axon appeared "beaded" and it takes part in the formation of both types of glomeruli.

Animals↗
PubMed1982

Long-lasting depression of parallel fiber-Purkinje cell transmission induced by conjunctive stimulation of parallel fibers and climbing fibers in the cerebellar cortex.

In rabbit cerebellar cortex, local stimulation of parallel fibers induced field potentials with two negative peaks, n1 representing conducting spikes of parallel fibers and n2 postsynaptic excitation in dendrites of Purkinje cells and other cortical cells. Conjunctive stimulation of parallel fibers and climbing fibers at 4 Hz for 30-120 sec caused a significant depression of n2 potential which lasted for at least 1 h. Such an effect could not be produced by stimulation of climbing fibers or parallel fibers alone. These observations support the plasticity assumption in the Marr-Albus model of the cerebellum.

Animals↗
PubMed2004

Quantitative analysis of granule cell axons and climbing fiber afferents in the turtle cerebellar cortex.

The turtle cerebellar cortex is a single flat sheet of gray matter that greatly facilitates quantitative analysis of biotylinated dextran amine labeled granule cell and olivocerebellar axons and Nissl-stained granule and Purkinje neurons. On average, ascending granule cell axons are relatively thicker than their parallel fiber branches (mean +/- SD: 0.84 +/- 0.17 vs 0.64 +/- 0.12 microm, respectively). Numerous en passant swellings, the site of presynaptic contact, were present on both ascending and parallel fiber granule cell axons. The swellings on ascending axons (1.82 +/- 0.34 microm, n = 52) were slightly larger than on parallel fibers (1.43 +/- 0.24 microm, n = 430). In addition, per unit length (100 microm) there were more swellings on ascending axons (11.2 +/- 4.2) than on parallel fibers (9.7 +/- 4.2). Each parallel fiber branch from an ascending axon is approximately 1.5 mm long. Olivocerebellar climbing fiber axons followed the highly tortuous dendrites of Purkinje cells in the inner most 15-20% of the molecular layer. Climbing fibers displayed relatively fewer en passant swellings. The spatial perimeter of climbing fiber arbors (area) increased 72% from anteriorly (1797 microm2) to posteriorly (3090 microm2) and 104% from medially (1690 microm2) to laterally (3450 microm2). Differences in the size and spacing of en passant swellings on granule cell axons suggest that ascending axons may have a functionally more significant impact on the excitability of a limited number of radially overlying Purkinje cells than the single contacts by parallel fiber with multiple orthogonally aligned Purkinje cell dendrites. The spatially restricted distribution of climbing fibers to the inner most molecular layer, the paucity of en passant swellings, and different terminal arbor areas are enigmatic. Nevertheless, these finding provide important anatomical information for future optical imaging and electrophysiological experiments.

Afferent Pathways↗
PubMed1999

Metabotropic glutamate receptor 2/3 immunoreactivity in the developing rat cerebellar cortex.

In adult rat cerebellar cortex, the metabotropic glutamate receptors (mGluRs) 2 and 3 (mGluR2/3) are present in somata, dendrites, and terminals of Golgi cells as well as in presumed glial processes (Ohishi et al. [1994], Neuron 13:55-66). In the present study, spatiotemporal changes in immunostaining for mGluR2/3 were examined in postnatal rat cerebellar cortex. mGluR2/3-immunoreactive Golgi cell somata appeared first in the internal granular layer at postnatal day 3 (P3) and were restricted to lobules IX and X; however, by P5, they were present in all lobules. Immunoreactive Golgi cell axons were adult-like, appearing as tortuous fibers with clusters of varicosities. They were observed first in the internal granular layer at P7 and increased in number and complexity with time. It was confirmed that mGluR2/3-immunoreactive Golgi cell axon terminals belong to the synaptic glomerulus by P10. Immunoreactive Golgi cell dendrites extending into the molecular layer became prominent after P15. By that time, the immunostaining pattern was characteristic of Golgi cells, as seen typically in adults. Many intensely immunoreactive radial processes existed at birth (P0). These traversed the molecular and external granular layers, reaching the pial surface in every cerebellar lobule. Because they showed coimmunoreactivity for glial fibrillary acidic protein, they were confirmed to be Bergmann glial fibers. After P9, they began to lose immunoreactivity at the portion corresponding to the molecular layer, while an immunostained granular pattern appeared in that layer. Immunoreactive radial processes, however, remained in the external granular layer, and finally, at P21, they disappeared together along with the external granular layer. Granular staining in the molecular layer reached background levels at this time. These spatiotemporal changes in mGluR2/3 distribution suggested that there may be distinct roles for mGluR2/3 in Golgi cells and Bergmann glial cells during the early postnatal period. mGluR2/3 in Golgi cells might be associated closely with systemic maturation, whereas mGluR2/3 in Bergmann glia might be needed for neuron-glia interactions related to granule cell development.

Animals↗
PubMed2000

Roles of GABAergic inhibition and NMDA receptor subunits in the spatio-temporal integration in the cerebellar cortex of mice.

The cerebellar cortex consists of relatively small numbers of identified neuronal types, which form simple and well-defined layers. However, a direct high-resolution demonstration of spatio-temporal pattern of information transmission there has been lacking. Using an optical recording technique with a membrane-potential sensitive dye, we studied the spatio-temporal pattern of excitation propagation induced by white matter stimulation in the slice preparations. We focused on physiological roles of inhibitory synapses and N-methyl-D-aspartate (NMDA) receptors. White matter stimulation induced postsynaptic long-lasting depolarization in the granular layer and transient depolarization in the molecular layer, respectively. Inhibitory synapses modestly suppressed the amplitude of slow depolarization in the granular layer, whereas they exerted powerful lateral inhibition in the molecular layer. Using mutant mice deficient in NMDA receptor subunits NR2A and/or NR2C, we also demonstrated that the NR2A and NR2C subunits expressed in granule neurons contribute to the early and late components of slow depolarization respectively, and that both subunits cooperatively support the temporal summation of depolarization. Taking into account the anatomical organization of the cerebellar cortex, these results might suggest that the granular layer is specialized more in the temporal integration of input signals and the molecular layer in the spatial integration.

Animals↗
PubMed1975

The differential effect of cooling on responses of cerebellar cortex.

1. Responses of the cerebellar cortex in anaesthetized cats were evoked by mossy fibre and/or climbing fibre inputs, and the effects of graded cooling of the cerebellar cortex were investigated. Cooling was applied either globally by flooding the exposed cortex with cooled Ringer Locke, or in later experiments locally be passing cooled fluid through a silver tube in contact with the cerebellar cortex. The cortical temperature was continuously monitored by a thermistor inserted to a depth of 0.5 mm close to the recording site. 2. In the granular layer the cooling caused a large increase in the diphasic P1N1 wave generated by the afferent mossy fibre volley. The waves generated by synaptic excitation and discharge of granule cells, N2P2, were not diminished until the temperature fell towards 20 degrees C. In contrast the N3 wave of the molecular layer was largest with cooling in the range of 35 to 25 degrees C, often several times larger than at 38 to 40 degrees C. Associated with the enhanced N3 wave there was an enhanced N4 wave, which indicates an increased discharge by Purkynĕ cells. 3. Climbing fibre inputs generate a negative field potential in the molecular layer due to the powerful excitation of Purkynĕ cells. In contrast to the N3 potential this climbing fibre wave was largest at the higher temperatures 35-40 degrees C and declined progressively with cooling, being usually suppressed at moderate coolings of 31-27 degrees C. Intracellular recording revealed that the diminution was due both to the elimination of all but the first impulses of the normal burst discharge of the climbing fibre impulses and to the diminution of the synaptic excitation of a single climbing fibre impulse. 4. It is shown that the negative potentials produced in the molecular layer by combinations of mossy fibre and climbing fibre inputs can be very effectively distinguished by this differential effect of cooling. 5. The effects of cooling even to a severe level are immediately recoverable on warming. Repeated cooling has no untoward effects and there is no sign of the hysteresis reported for the cuneate nucleus. 6. There is a discussion of the factors that could cause cooling to differentiate between the actions of the mossy fibre and climbing fibre impulses on Purkynĕ cells.

Action Potentials↗
PubMed1995

Extinction of conditioned eyelid responses requires the anterior lobe of cerebellar cortex.

We test the hypothesis that the cerebellar cortex is required for the extinction of conditioned eyelid responses in rabbits trained using standard Pavlovian delay procedures. Following 10 daily training sessions during which rabbits achieved asymptotic performance, lesions of the ipsilateral hemisphere of the cerebellar cortex were made by aspiration. The target of these lesions was the anterior lobe, as suggested by previous observations that this region is necessary for the learning-dependent timing of conditioned eyelid responses (Perrett et al., 1993). We report that anterior lobe damage, as indicated by disrupted response timing and confirmed by tissue analysis, produces severe deficits in conditioned response extinction. Postlesion responses show no significant decline over ten training sessions, whereas response timing and extinction are unaffected by lesions that do not include the anterior lobe. These conditioned responses that do not extinguish display stimulus specificity, excluding the possibility that they are unlearned responses unmasked by cerebellar cortex lesions. These observations suggest that Pavlovian eyelid conditioning is mediated by synaptic plasticity in at least two sites and the anterior lobe of the cerebellar cortex influences one of these sites during extinction. Based on these and previous data, we propose the hypothesis that eyelid conditioning can involve plasticity in both the cerebellar cortex and interpositus nucleus and that plasticity in the nucleus is controlled by input from Purkinje cell activity in the cortex. This hypothesis is consistent with observations that the cerebellar cortex may not always be required for the expression of conditioned responses, but it is necessary for response timing and for extinction.

Animals↗