1. Three rhesus monkeys were trained to perform a rapid (greater than 100 degrees/s) and a slow (less than 100 degrees/s) wrist movement guided by a visual cue. While the monkey performed wrist flexion or extension from a neutral position, Purkinje cell (P-cell) discharges were recorded from intermediate and lateral parts of lobules IV--VI of the cerebellum. 2. By the visually guided movement, we could control the direction of the wrist movement; the holding position at three different angles of the wrist joint: neutral, about 30 degrees flexed, and extended; and the velocity in four ranges: a) 10--30, b) 30--100, c) 100--300, and d) 300-650 degrees/s. 3. From 92 P-cells that significantly increased or decreased the discharge rate of simple spikes with task performance, we selected 45 P-cells ("response-locked" cells) as related to the wrist movement by statistical analyses of temporal correlation of P-cell activities to wrist movement. The direction of the frequency modulation (increase or decrease) was in a nonreciprocal fashion with oppositely directed wrist movements (flexion or extension) in 90% of the response-locked P-cells. The maintained frequencies at three holding positions did not significantly differ. 4. Nineteen P-cells changed their spike frequencies temporally locked to both rapid and slow wrist movements. By the discharge pattern in relation to the rapid and slow movements, these cells were classified into two groups. Discharge pattern in group I P-cells (n = 5) conformed very well to that of velocity, and a linear correlation between the instantaneous increase of the discharge rate and velocity was observed in analyses of individual trials. Group II cells showed increase (n = 9) or decrease (n = 5) of firing rate (20--50 spikes/s) larger than group I cells (less than 10 spikes/s) as long as the wrist was moving, even with very slow velocity (less than 30 degrees/s. The correlations between the increase of the discharge rate and the velocity in individual trials were less clear in group II than in group I cells. 5. The present study suggests the importance of the cerebellar cortex in controlling the slow limb movement as well as the rapid movement. The selected P-cells in this study also suggested that the velocity or some dynamic aspect related to the velocity of limb movement is the major information among the dissociated motion parameters coded by the simple-spike frequencies of the P-cells in the cerebellar hemisphere. Whether the latter suggestion represents an essential characteristic of all limb movement-related P-cells or reflects only a feature of a special subgroup among the movement-related cells should be clarified in future experiments.

Action potentials of cerebellar Purkinje cells were observed in intact monkeys during sleep and waking. Purkinje cells exhibit two sorts of action potentials, called simple and complex spikes, and these two sorts of spikes were differently affected by sleep. Simple-spike activity (generated by the parallel fiber inputs to the Purkinje cell) was highest during sleep with rapid eye movements as compared with both waking and sleep with electroencephalographic slow waves. In contrast, complex-spike activity (generated by the climbing fiber inputs to the Purkinje cell) was lowest during sleep with rapid eye movements. The complex action potential of the Purkinje cell consists of an initial large spike followed by one or more smaller secondary spikes, and the number of these secondary spikes was found to be independent of the background discharge frequency of the simple spike. This independence suggests a possible role of presynaptic factors rather than the excitability level of the Purkinje cell itself in determining the number of secondary discharges occurring in the complex spike.

Four rhesus monkeys were trained to perform visually guided wrist tracking movements (50). While they performed tasks by wrist flexion or extension from a neutral position, simple-spike (SS) and complex-spike (CS) discharges of a single Purkinje cell (P-cell) were recorded from intermediate and lateral parts of cerebellar hemispheres (lobules IV to VI) ipsilateral to the task-performing wrist. Of approximately 400 P-cells observed, 215 (54%) significantly increased or decreased their SS discharge rate during task performance (task-related P-cells). Of these, 161 were selected for analysis of CS activity; in these P-cells, we could reliably discriminate between CS and background SS by a spike discriminator. The 161 P-cells were further classified into response locked (n = 65) and poorly locked (n = 96) cells according to temporal coupling of the SS frequency modulation to the onset of wrist movements. About 60% of the response-locked P-cells showed a phasic increase (statistical significance level: P less than 0.01) of CS firing rate at the onset of wrist tracking movement. In a few P-cells, a phasic decrease (statistically insignificant) of CS firing rate was observed with the wrist movement. In most P-cells, an increase of CS firing rate was observed with both rapid- and slow-tracking wrist movements. The increase was larger with faster step-tracking movement than with slower ramp-tracking movement. In most P-cells, the CS activity increased with both wrist flexion and extension; in some cells, however, it increased only with either flexion or extension. In most of the response-locked P-cells, the increase of CS firing rate occurred during motor time, i.e., after the onset of the EMG change in prime movers and before the beginning of wrist tracking movement. The increase occurred phasically at the onset and/or at the recovery phase of SS frequency modulation. At neutral wrist position, the maintained frequency of the CS was 0.72 +/- 0.29 CS/s (mean and SD for 161 task-related P-cells). Compared with the frequency at neutral position, the CS frequency did not change tonically during maintained flexed or extended wrist position in any response-locked P-cells. There was no increase of CS firing rate when the monkey returned the handle to center position after completing the tracking task, even in P-cells that had shown a significant increase of CS activity during tracking.(ABSTRACT TRUNCATED AT 400 WORDS)

1. The activity of globus pallidus (GP) neurons (n = 1,117) was studied in two monkeys to reexamine the relation of neuronal activity to movement type (slow vs. fast) while they performed both a visually guided step and ramp wrist tracking task. To select neurons specifically related to wrist movements, we employed both a somatosensory examination of individual body parts and a statistical analysis of the strength of temporal coupling of neuronal discharges to active wrist movement. 2. Neuronal responses to somatosensory stimulation were studied in 1,000 high-frequency GP neurons, of which 686 exhibited clear responses to manipulation of body parts. Of the latter, 336 responded to passive manipulation of forelimb joints and 58 selectively to passive flexion or extension of the wrist. 3. In the external segment of GP (GPe), most neurons responding to passive wrist movement were found to be clustered in four to five adjacent, closely positioned (separated by 200 microns) tracks in single coronal planes. The clusters were irregular in shape with a maximal width of 800-1,000 microns. Separate clusters of neurons responsive to passive wrist movement were identified in planes 3 mm apart in one monkey and in planes 500 microns apart in the other. Multiple clusters of neurons were also found for neurons responsive to joints other than the wrist. These findings suggest a more discrete and complex representation of individual joints in the primate GP than previously conceived. 4. During the performance of the wrist flexion and extension task, 92 neurons showed clear and consistent changes in activity. For these neurons we measured, with a statistical method on a trial-by-trial basis, the strength of temporal coupling between the onset of active wrist movement and the onset of change in neuronal discharge rate. Fifteen neurons showed changes in activity time-locked to the onset of active wrist movement. 5. Twelve pallidal neurons were classified as "wrist-related" based on their movement-locked changes in discharge during task performance and their clear responses to passive wrist joint rotation on examination. All of these neurons exhibited statistically significant modulation of their discharge rate during both fast (peak velocity 97-205 degrees/s) and slow (peak velocity 20-62 degrees/s) wrist movements in the task. The amplitudes of modulation were larger during fast wrist movement than slow movement. These results suggest that the basal ganglia motor circuit plays a similar, rather than an exclusive, role in the control of slow and fast limb movements.

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