Primary motor cortex influences on the descending and ascending systems

Topography of Purkinje cell compartments and mossy fiber terminal fields in lobules II and III of the rat cerebellar cortex: spinocerebellar and cuneocerebellar projections.

Many mossy fiber afferent projections to the rat cerebellar cortex terminate in parasagittal bands. In particular, the anterior lobe vermis of the cerebellum contains alternating bands of mossy fibers from the spinal cord and external cuneate nuclei. The cerebellar cortical efferents, the Purkinje cells, are also organized in parasagittal bands. These can be revealed by immunochemical staining for the antigen zebrin II, which is selectively expressed by bands of Purkinje cells. In some cases, the boundaries between mossy fiber terminal fields align with identified transitions between zebrin+/- sets of Purkinje cells, whereas others are located within apparently homogeneous Purkinje cell compartments. Two theories can explain the terminal-field topography: In one view, mossy fiber terminals segregate during development, because growth cones from different sources compete for common territory. Alternatively, mossy fiber growth cones directly recognize chemically distinct target territories, and activity-dependent mechanisms play only minor roles. To explore these issues, two sets of experiments were performed. First, the terminal-field map of the neonatal spinocerebellar projection was compared to the Purkinje cell compartmentation as revealed by anticalbindin immunocytochemistry. Second, subsets of spinocerebellar mossy fiber afferents were ablated early in postnatal development, and the consequences for the neighboring cuneocerebellar terminal fields were mapped in the adult with reference to the zebrin II+/- compartments. These experiments revealed no evidence that competitive interactions constrain the mossy fiber terminal-field distribution but, rather, suggest that the organization of the mossy fiber projections follows the compartmentation of the Purkinje cells.

Developing mossy fiber terminal fields in the rat cerebellar cortex may segregate because of purkinje cell compartmentation and not competition

1) Experiments were designed to detect how static parameters of natural, passive hand movements are encoded and integrated within the cerebellar cortex. For this purpose unit activity was recorded extracellularly from presumed mossy fibres (MF), presumed granule cells (GrC) and from Purkinje cells (PC) discharging with simple spikes (SS) and complex spikes (CS). With respect to the PC, our interest was focussed primarily on the SS activity. The recordings were performed in the intermediate part of the cerebellar anterior lobe of decerebrate cats. The animal's forepaw was passively moved around the wrist joint by an electronically controlled device. The movements were exactly reproducible so that peristimulus time histograms of the unit activity could be constructed. 2) At the input level (MF) and at the first level of integration within the cerebellar cortex (GrC), patterns with similar discharge characteristics were found. Such patterns could, to a limited extent, also be detected at the cerebellar output (SS of PC). However, in most cases of SS discharge, patterns were found with weak correlation between the tonic activity and static parameters of the movements. 3) Absolute paw position, amplitude, and duration of movements were found to be related over wide ranges to the activities of MF and GrC. Absolute position is directly encoded by tonic discharge during the low or high holding phases. Beside this, units were found without a correlation between the tonic discharge and the position of the nonmoving paw. However, in these units it was sometimes observed that the information about the momentary position or the information about the mean position was sometimes conveyed exclusively during the proceeding upward or downward movement. Thus, information about static parameters was transmitted only at times when a dynamic parameter (such as velocity) occurred. This type of position information encoding is termed “indirect mode of transmission”. 4) A specific relationship between SS unit activity of PC and the absolute position of the forepaw or amplitude of the movement could be found primarily by using multiple ramps instead of single ramp movements. This was observed even if both types of ramp movements had the same velocity, individual amplitude, and tested range. However, on multiple ramp movements the paw generally remained for a shorter period at a specific position level as compared to the single ramp movements. 5) Apart from this timing phenomenon, a late movement response was observed, which results in a specific type of position information encoding on multiple ramp functions. 6) These results indicate that static parameters of a passive limb movement are conveyed via the MF input to the cerebellar cortex. Patterns related to these parameters undergo a change within the MF-, GrC-, Parallel fibre-, PC-system. Different modes of encoding these parameters were observed depending primarily on the neuronal niveau within the cerebellar cortex. Tonic discharge related e.g. to limb position was found at MF and GrC level. Such patterns resemble, at least to a certain extent, those obtained from different peripheral receptors. The high tonic SS activity never showed such a strong relationship to static parameters as observed at the input level; static parameters could be resolved only within relatively short periods of time, especially during the dynamic phases of the movement or during short periods immediately following these phases. This implies that the function of this part of the cerebellum, which is to provide correction signals, should be considered as a more dynamic process characterized by evaluating predominantly information about the momentary ongoing movement.

Cerebellar unit responses of the mossy fibre system to passive movements in the decerebrate cat. I. Responses to static parameters.

In the cat, somatosensory nuclei send substantial projections to the inferior olive, where they terminate in a somatotopic fashion. Although the organization of the cat inferior olive has been used to interpret data from other species, published data suggest this organization may not occur universally. The present study investigated whether the inferior olive in albino rats and cynomolgus monkeys receives the same brainstem somatosensory inputs, whether these inputs are organized somatotopically and, if so, how the organization compares with that in the cat. Projections from the gracile, cuneate and spinal trigeminal nuclei were labeled with wheat germ agglutinin conjugated to horseradish peroxidase or with biotinylated dextran. The results were compared with data from cats (Berkley and Hand [1978] J. Comp. Neurol. 180:253-264).

In the rat and monkey, the gracile, cuneate and spinal trigeminal nuclei all project to the contralateral inferior olive, where each nucleus has a distinct preferred terminal field. As in the cat, projections to the medial accessory olive and caudal dorsal accessory olive did not terminate in a precisely organized fashion. Projections to the rostral dorsal accessory olive, however, formed a clear somatotopic map. These somatotopic maps differed from those in the cat in that input from the trigeminal nucleus was confined rostrally, so that the caudal end only received input from the gracile and cuneate nuclei.

These data indicate that similar organizational principles characterize the somatosensory projections to the inferior olives of the three species. Nevertheless, distinct species differences occur with regard to the details of this organization. © 1996 Wiley-Liss, Inc.

Gracile, cuneate, and spinal trigeminal projections to inferior olive in rat and monkey

Distribution of neurons in the main cuneate nucleus projecting to the inferior olive in the cat. Evidence that they differ from those directly projecting to the cerebellum.

A. Alonso

Papers

Distribution of neurons in the main cuneate nucleus projecting to the inferior olive in the cat. Evidence that they differ from those directly projecting to the cerebellum.