Barto, A. G.; Fagg, A. H.; Sitkoff, N., and Houk, J. C. A cerebellar model of timing and prediction in the control of reaching. Neural Computation. 1999; 11:565-94.
Abstract:  A simplified model of the cerebellum was developed to explore its potential for adaptive, predictive control based on delayed feedback information. An abstract representation of a single Purkinje cell with multistable properties was interfaced, using a formalized premotor network, with a simulated single degree-of-freedom limb. The limb actuator was a nonlinear spring-mass system based on the nonlinear velocity dependence of the stretch reflex. By including realistic mossy fiber signals, as well as realistic conduction delays in afferent and efferent pathways, the model allowed the investigation of timing and predictive processes relevant to cerebellar involvement in the control of movement. The model regulates movement by learning to react in an anticipatory fashion to sensory feedback. Learning depends on training information generated from corrective movements and uses a temporally asymmetric form of plasticity for the parallel fiber synapses on Purkinje cells.

Braitenberg, V. The cerebellum and the physics of movement: Some speculations. Glickstein, M.; Yeo, C., and Stein, J. Cerebellum and Neuronal Plasticity. New York: Plenum Press; 1987; pp. 193-207

Eccles, J. C.; Sabah, N. H.; Schmidt, R. F., and Taborikova, H. Mode of operation of the cerebellum in the dynamic loop control of movement. Brain Res. 1972; 4073-80.

Fortier, P. A.; Kalaska, J. F., and Smith, A. M. Cerebellar neuronal activity related to whole-arm reaching movements in the monkey. J.Neurophysiol. 1989; 62:198-211.
Abstract:  1. Three monkeys were trained to make whole-arm reaching movements from a common central starting position toward eight radially arranged targets disposed at 45 degrees intervals. A sample of 312 cerebellar neurons with proximal-arm receptive fields or discharge related to shoulder or elbow movements was studied in the task. The sample included 69 Purkinje cells, 115 unidentified cortical cells, 65 interpositus neurons, and 63 dentate units.  2. The reaching task was divided into three movement-related epochs: a reaction time, a movement time, and holding over the target. All neurons demonstrated significant changes in discharge during one or more of these three epochs. Almost all of the cells (95%) showed a significant change in activity during the movement, whereas 68-69% of the cells showed significant changes from premovement activity during the reaction time and holding periods. 3. During the combined reaction time-movement period, 231/312 cells were strongly active in the task. Of these, 151 cells (65.4%) demonstrated unimodal directional responses. Sixty-three had a reciprocal relation to movement direction, whereas 88 showed only graded increases or decreases in activity. A further 37 cells (16.0%) were nondirectional, with statistically uniform changes in discharge in all eight directions. The remaining 43 cells (18.6%) showed significant differences in activity for different directions of movement, but their response patterns were not readily classifiable. 4.  The proportion of directional versus nondirectional cells was consistent across the four cell populations. However, graded response patterns were more common and reciprocal responses less common among Purkinje and dentate neurons than among unidentified cortical cells and interpositus neurons. 5.  The distribution of preferred directions of the population of cerebellar neurons covered all possible movement directions away from the common central starting position in the horizontal plane. When the preferred direction of each cell in the sample population was aligned, the mean direction-related activity of the cerebellar population formed a bell-shaped tuning curve for the activity recorded during both the reaction time and the movement, as well as during the time the arm maintained a fixed posture over the targets. A vector representation also showed that the overall activity of the cerebellar population during normal reaching arm movements generated a signal that varied with movement direction. 6. These results demonstrate that the cerebellum generates a signal that varies with the direction of movement of the proximal arm during normal aimed reaching movements and is consistent with a role in the control of the activity of muscles or muscle groups generating these movements.

Fu, Q. G.; Flament, D.; Coltz, J. D., and Ebner, T. J. Relationship of cerebellar purkinje cell simple spike discharge to movement kinematics in the monkey. Journal of Neurophysiology. 1997 Jul; 78(1):478-91.
Abstract:  The simple spike discharge of 231 cerebellar Purkinje cells in ipsilateral lobules V and VI was recorded in three monkeys trained to perform a visually guided reaching task requiring movements of different directions and distances. The discharge of 179 cells was significantly modulated during movement to one or more targets. Mean simple spike rate was fitted to a cosine function for direction tuning, a simple linear function for distance modulation, and a multiple linear regression model that included terms for direction, distance, and target position. On the basis of the fit to the direction and distance models, there were more distance-related than direction-related Purkinje cells. The simple spike discharge of most direction-related cells modulated at only one target distance. The preferred directions for the simple spike tuning were not uniformly distributed across the workspace. The discharge of most distance-related cells modulated along only one movement direction. On the basis of the multiple linear regression model, simple spike discharge was also correlated with target position, in addition to direction and distance. Approximately half of the Purkinje cells had simple spike activity associated with only a single parameter, and only a small fraction of the cells with all three. The multiple regression model was extended to evaluate the correlations as a function of time. Considerable overlap occurred in the timing of the simple spike correlations with the parameters. The latency for correlation with movement direction occurred mainly in a 500-ms interval centered on movement onset. The correlations with target position also occurred around movement onset, in the range of -200-500 ms. Distance correlations were more variable, with onset latencies from -500 to 1,000 ms. These results demonstrate that the simple spike discharge of cerebellar Purkinje cells is correlated with movement direction, distance, and target position. Comparing these results to motor cortical discharge shows that the correlations with these parameters were weaker in Purkinje cell simple spike discharge, and that, for the majority of Purkinje cells, the simple spike discharge was significantly related to only a single movement parameter. Other differences between simple spike responses and those of motor cortical cells include the nonuniform distribution of preferred directions and the extensive overlap in the timing of the correlations. These differences suggest that Purkinje cells process, encode, and use kinematic information differently than motor cortical neurons.

Georgopoulos, A. P. New concepts in generation of movement. Neuron. 1994; 13:257-268.
 

Higgins, D. C. The cerebellum and initiation of movement: The stretch reflex. Yale J.Biol.Med. 1987; 60:123-131.
Abstract:  Studies of the stretch reflex in decerebrate cats indicate a phase advance of peak sinusoidal tension in steady-state cycles between 0.1 and 10 Hz. This phase advance is reduced in acute and chronic cerebellectomy, as shown in previous investigations. Also, the augmentation of muscle peak tension in initial sinusoidal stretch cycles at 0.5-5 Hz has been found to be reduced during the time of reflex and motor instability in the several months following cerebellar ablation. This report shows the increased amplitude and phase lead of integrated electromyographic activity in initiating sinusoidal stretch cycles in the decerebrate cat.  These reflex aspects are demonstrated in relation to the discharge of neurons in the dorsal spinocerebellar tract and of cerebellar cortical Purkinje cells in initial sinusoidal cycles. The intensity and phase advance of the discharge in dorsal spinocerebellar tract neurons is altered little, but these features are usually increased in Purkinje cells during initial stretches compared to continuous cycling. In terms of overall motor control, these findings are compatible with concepts of movement control, modulated by the cerebellum, in which the discharge of antagonist motor neurons is regulated in concert with that of agonist muscles upon initiation and termination of movement.

Ivry, R. B.; Keele, S. W., and Diener, H. C. Dissociation of the lateral and medial cerebellum in movement timing and movement execution. Exp.Brain Res. 1988; 73:167-180.
Abstract:  In a previous study (Ivry and Keele, in press), cerebellar patients were found to be impaired on both a motor and a perceptual task which required accurate timing. This report presents case study analyses of seven patients with focal lesions in the cerebellum. The lesions were predominantly in the lateral, hemispheric regions for four of the patients.  For the remaining three patients, the lesions were centered near the medial zone of the cerebellum. The clinical evaluation of the patients also was in agreement with the different lesion foci: lateral lesions primarily impaired fine motor coordination, especially apparent in movements with the distal extremities and medial lesions primarily disturbed balance and gait. All of the patients were found to have increased variability in performing rhythmic tapping when tapping with an effector (finger or foot) ipsilateral to the lesion in comparison to their performance with a contralateral effector.  Separable estimates of a central timekeeper component and an implementation component were derived from the total variability scores following a model developed by Wing and Kristofferson (1973). This analysis indicated that the poor performance of patients with lateral lesions can be attributed to a deficit in the central timing process. In contrast, patients with medial lesions are able to accurately determine when to make a response, but are unable to implement the response at the desired time. A similar dissociation between the lateral and medial regions has been observed on a time perception task in patients with cerebellar atrophy. It is concluded that the lateral regions of the cerebellum are critical for the accurate functioning of an internal timing system.

Kitazawa, S.; Kimura, T., and Yin, P. B. Cerebellar complex spikes encode both destinations and errors in arm movements. Nature. 1998 Apr 2; 392(6675):494-497.
Abstract:  Purkinje cells of the cerebellum discharge complex spikes, named after the complexity of their waveforms, with a frequency of approximately 1 Hz during arm movements. Despite the low frequency of firing, complex spikes have been proposed to contribute to the initiation of arm movements or to the gradual improvement of motor skills. Here we recorded the activity of Purkinje cells from the hemisphere of cerebellar lobules IV-VI while trained monkeys made short-lasting reaching movements (of approximately 200 milliseconds in duration) to touch a visual target that appeared at a random location on a tangent screen. We examined the relationship between complex-spike discharges and the absolute touch position, and between complex-spike discharges and relative errors in touching the screen. We used information theory to show that the complex spikes occurring at the beginning of the reach movement encode the absolute destination of the reach, and the complex spikes occurring at the end of the short-lasting movements encode the relative errors. Thus, complex spikes convey multiple types of information, consistent with the idea that they contribute both to the generation of movements and to the gradual, long-term improvement of these movements.

Lou, J.-S. and Bloedel, J. R. A study of cerebellar cortical involvement in motor learning using a new avoidance conditioning paradigm involving limb movement. Brain Res. 1988; 445:171-174.
Abstract:  These experiments were performed to examine the relationship between the simple and complex spike responses of 3-5 simultaneously recorded Purkinje cells during the acquisition, performance and extinction of a conditioned forelimb movement in decerebrate, unanesthetized ferrets. The data demonstrate parallel, correlated changes in simple and complex spike responses throughout the experimental period. Since the evaluated Purkinje cells were examined in the cerebellar cortical region that contains neurons highly modulated by the intermittent application of the conditioning stimulus, these findings argue against an induction of a long-lasting modification in simple spike responses by the climbing fiber input as the basis for this type of motor learning.

Lu, X. F.; Hikosaka, O., and Miyachi, S. Role of monkey cerebellar nuclei in skill for sequential movement. J.Neurophysiol. 1998 May; 79(5):2245-2254.
Abstract:  To examine whether the cerebellum is involved in learning and memory of visuomotor sequences, we trained two monkeys on a sequential button press task and inactivated different portions of the cerebellar nuclei by injecting a small amount of muscimol (-aminobutyric acid agonist). Before the injection experiments started, the monkeys had learned a set of sequences (n = 21 and 12) extensively. After each injection, we had the monkeys perform the learned sequences and, in addition, learn new sequences. We found deficits in learning/memory by the injections into the dorsal and central part of the dentate nucleus. The number of errors increased significantly for the learned sequences but not for the new sequences. This effect was present only when the hand ipsilateral to the muscimol injection was used.  Consistent with this result, anticipatory saccades, the occurrence of which is correlated closely with motor skill, also became less frequent particularly when the ipsilateral hand was used. No effect on learning/memory was observed after injections into the ventral or lateral parts of the dentate nucleus, interpositus nucleus, or fastigial nucleus. In contrast, hand movements became slower after ipsilateral injections at all of the injection sites. These results suggest that, among the cerebellar nuclei, the dentate nucleus, especially its dorsal and central regions, is related to the storage and/or retrieval of long-term memory for motor skill.

MacKay, W. A. Cerebellar nuclear activity in relation to simple movements. Exp.Brain Res. 1988; 71:47-58.
Abstract:  Single unit activity in the fastigial, interpositus and dentate cerebellar nuclei was recorded in relation to simple elbow flexion and extension movements in two macaque monkeys. In common with proximal muscle activity, 94% of the task-related neurons had qualitatively similar discharge patterns for both directions of forearm movement. In many cases the flexion and extension discharge was virtually identical, but some cells had a distinct directional bias. The very few neurons which were directionally specific were located in the dentate and interpositus. Two had tonic activity well correlated to elbow angle. Task-related changes in discharge rate occurred earliest in dentate and latest in fastigial, but almost always during the period of concomitant proximal and elbow EMG changes. Correlations of phasic activity with movement velocity were uniformly weak. Many eye movement-related neurons were encountered in the fastigial, dentate and y-group nuclei. Fastigial eye cells, both bursting and tonic, tended to be highly direction specific, whereas dentate eye cells were usually omnidirectional and variable. For both arm and eye cerebellar cells, the directional preferences of phasic and tonic discharge, in the same neuron, could be opposed to one another.

MacKay, W. A. Unit activity in the cerebellar nuclei related to arm reaching movements. Brain Res. 1988; 442:240-254.
Abstract:  Single units in the fastigial, interpositus and dentate nuclei of two stump-tail macaque monkeys were studied in relation to a right arm, visually guided reaching task. Of 638 recorded cells, 149 showed activity changes correlated to the task, including 24 in the contralateral fastigial and interpositus. Reach-related discharge patterns fell into two broad categories, tonic and phasic. Tonic responses were maintained throughout the reach with no observable relation to kinematic parameters. Most of the task-related activity occurred during the upward lift of the arm toward the target button, with a drop-off as the arm was lowered toward the rest plate.  Phasic response cells fired bursts (or suppressed discharge) at specific points in the arm trajectory, most commonly during the lift phase. Many had a sharp drop in discharge when the shoulder flexion torque was transiently reversed to decelerate the arm. For either type, restricted directional specificity was rarely seen in any nucleus, and correlations with recorded EMGs were weak. Visual responses to target button illumination were observed in both the fastigial and dentate nuclei, but did not necessarily correspond with the button giving the best movement-related response. Task-related activity changes started earliest in the fastigial nuclei and latest in the interpositus nuclei. The data suggested that cerebellar output facilitates motor centers in a rather general manner, but at precisely determined times.

Mason, C. R.; Miller, L. E.; Baker, J. F., and Houk, J. C. Organization of reaching and grasping movements in the primate cerebellar nuclei as revealed by focal muscimol inactivations. J.Neurophysiol. 1998 Feb; 79(2):537-554.
Abstract:  Two monkeys were trained to point to targets and to retrieve fruit bits from a Kluver board, bottles, and tubes. Once proficient in the tasks, the macaques underwent aseptic surgical implantation of a recording chamber over the cerebellar nuclei on the side of their preferred hand. After recovery from surgery, a series of mapping penetrations were completed to identify task-related areas within the cerebellar nuclei. Muscimol (4- 16 microgram; 1-2 microgram/microliter) was pressure injected at different sites within the forelimb zone, and the resultant deficits were observed as the monkeys performed the behavioral tasks. Quantitative measures of task performance were supplemented by direct observation of live and videotaped performance.  The locations of nuclear inactivation sites were reconstructed from marking lesions and tracks visible in histological sections. Injections placed in the cerebellar interpositus nucleus and adjacent regions of dentate caused a variety of deficits in forelimb function. A prominent anteroposterior specialization was apparent within the forelimb zone of this intermediate nuclear region. Injections into the anterior interpositus nucleus and adjacent dentate impaired preshaping of the hand and the manipulation of objects, whereas injections placed more posteriorly in posterior interpositus nucleus and adjacent dentate produced deficits in the aiming of reach and the stability of the arm. During anterior injections, the monkeys failed to adequately extend their fingers in preparation for target contact, as documented for >85% of the reaches in the pointing task of monkey J. Up to 38% of the fruit bits it attempted to retrieve from the Kluver board were dropped. In comparison, during posterior inactivations, 15% were dropped and during control conditions 3% were dropped. The monkeys made significantly greater pointing errors during posterior inactivations (11 times for monkey J and 4 times for monkey C) than during anterior inactivations (8 times for monkey J and 2 times for monkey C). We refer to the region producing hand deficits as the anterior hand zone and the region producing reaching deficits as the posterior reach zone. These results are discussed in relation to the problem of achieving spatiotemporal coordination in the large population of nuclear cells that participate in any given movement. The results do not favor the hypothesis that coordination is achieved through a selection of Purkinje cells along beams of parallel fibers. Instead, it is proposed that distal and proximal musculature is coordinated by the adaptive influences of climbing fiber input to Purkinje cells. We envision a relatively nonspecific recruitment of anterior and posterior nuclear cells due to positive feedback in the limb premotor network, which then is shaped into an appropriate spatiotemporal pattern of discharge through the inhibitory input from Purkinje cells.

Milak, M. S.; Shimansky, Y.; Bracha, V., and Bloedel, J. R. Effects of inactivating individual cerebellar nuclei on the performance and retention of an operantly conditioned forelimb movement. Journal of Neurophysiology. 1997 Aug; 78(2):939-59.
Abstract:  These experiments were designed to examine the effects of inactivating separately each of the major cerebellar nuclear regions in cats on the execution and retention of a previously learned, operantly conditioned volitional forelimb movement. The experiments test the postulates that the cerebellar nuclei, and particularly the interposed nuclei, contribute substantially to the spatial and temporal features of the interjoint coordination required to execute the task and that the engram necessary for the retention of this task is not located in any one of the cerebellar nuclei. All cats were trained to perform a task in which they were required to reach for and grasp a vertical bar at the sound of a tone and move the bar to a reward zone through a template consisting of two straight grooves in the shape of an inverted "L." After the task was learned, the effects of inactivating separately each nuclear region (the fastigial, interposed, and dentate nuclei) using muscimol microinjections were determined. Data were analyzed by quantifying several features of the movement's kinematics and by determining changes in the organization of the reaching component of the movement using an application of dimensionality analysis, an analysis that examines the correlation among the changes in joint angles and limb segment positions during the task. The retention of the previously learned task also was assessed after each injection. Injections of each nuclear region affected temporal and spatial features of the learned movement. However, the largest effects resulted from inactivating the interposed nuclei. These effects included an increased length of the reach trajectory, an accentuated deviation of the wrist trajectory from a straight line, cyclic movement of the distal extremity as the target was approached, a difficulty in grasping the bar, altered temporal features of the movement, and a highly characteristic change in the dimensionality measurements. The changes in dimensionality reflected a decreased correlation (linear interdependence) of the joint angular velocities coupled with an increased correlation among the linear velocities of markers located on the joints themselves. Related but less consistent changes in dimensionality resulted from fastigial injections.  The motor sequence required to negotiate the template could be executed after the nuclear microinjections, indicating that retention of the motor sequence was not affected by the inactivation of any of the cerebellar nuclei.  However, in two of the five animals, some decreases in performance were observed after dentate injection that were not characteristic of changes related to an effect on retention. These data suggest that the cerebellum plays an important role in regulating the consistent, stereotypic organization of complex goal-directed movements, including the temporal correlation among joint angle velocities. The data also indicate that the retention of the task is not dependent on any of the individual cerebellar nuclear regions. Consequently, these structures are unlikely to be critical storage sites for the engram established during the learning of this task.

Rao, S. M.; Harrington, D. L.; Haaland, K. Y.; Bobholz, J. A.; Cox, R. W., and Binder, J. R. Distributed neural systems underlying the timing of movements. Journal of Neuroscience. 1997 Jul 15; 17(14):5528-35.
Abstract:  Timing is essential to the execution of skilled movements, yet our knowledge of the neural systems underlying timekeeping operations is limited. Using whole-brain functional magnetic resonance imaging, subjects were imaged while tapping with their right index finger in synchrony with tones that were separated by constant intervals [Synchronization (S)], followed by tapping without the benefit of an auditory cue [Continuation (C)]. Two control conditions followed in which subjects listened to tones and then made pitch discriminations (D). Both the S and the C conditions produced equivalent activation within the left sensorimotor cortex, the right cerebellum (dorsal dentate nucleus), and the right superior temporal gyrus (STG). Only the C condition produced activation of a medial premotor system, including the caudal supplementary motor area (SMA), the left putamen, and the left ventrolateral thalamus. The C condition also activated a region within the right inferior frontal gyrus (IFG), which is functionally interconnected with auditory cortex. Both control conditions produced bilateral activation of the STG, and the D condition also activated the rostral SMA. These results suggest that the internal generation of precisely timed movements is dependent on three interrelated neural systems, one that is involved in explicit timing (putamen, ventrolateral thalamus, SMA), one that mediates auditory sensory memory (IFG, STG), and another that is involved in sensorimotor processing (dorsal dentate nucleus, sensorimotor cortex).

Robinson, F. R. Role of the cerebellum in movement control and adaptation. Curr. Opinion Neurobiol. 1995; 5:755-762.
Abstract:  Three recent discoveries have substantially improved our knowledge of cerebellar function. First, the forelimb regions of the interpositus nuclei specialize in control of one particular limb movement, reach to grasp.  Second, a new model indicates that vestibulo-ocular reflex adaptation requires neural changes in both the cerebellum and the brainstem. Finally, the caudal fastigial nucleus uses both short- and long-term influences to maintain saccade accuracy.

Schweighofer, N.; Arbib, M. A., and Kawato, M. Role of the cerebellum in reaching movements in humans. I. Distributed inverse dynamics control. Eur.J.Neurosci. 1998 Jan; 10(1):86-94.
Abstract:  This study focuses on the role of the motor cortex, the spinal cord and the cerebellum in the dynamics stage of the control of arm movement. Currently, two classes of models have been proposed for the neural control of movements, namely the virtual trajectory control hypothesis and the acquisition of internal models of the motor apparatus hypothesis. In the present study, we expand the virtual trajectory model to whole arm reaching movements. This expanded model accurately reproduced slow movements, but faster reaching movements deviated significantly from the planned trajectories, indicating that for fast movements, this model was not sufficient. These results led us to propose a new distributed functional model consistent with behavioural, anatomical and neurophysiological data, which takes into account arm muscles, spinal cord, motor cortex and cerebellum and is consistent with the view that the central nervous system acquires a distributed inverse dynamics model of the arm. Previous studies indicated that the cerebellum compensates for the interaction forces that arise during reaching movements. We show here how the cerebellum may increase the accuracy of reaching movements by compensating for the interaction torques by learning a portion of an inverse dynamics model that refines a basic inverse model in the motor cortex and spinal cord.

Van Kan, P. L. E.; Gibson, A. R., and Houk, J. C. Movement-related inputs to intermediate cerebellum of the monkey. J.Neurophysiol. 1993; 69:74-94.
Abstract:  1. The primary goal of this study was to characterize the information about single-joint forelimb movements supplied to intermediate cerebellar cortex by mossy fibers. Discharge of mossy fibers and Golgi cells was studied while monkeys operated six devices that required movements about specific joints.  Additional control experiments in anesthetized cats and monkeys established criteria for identification of mossy fibers and Golgi cells. 2. The control experiments demonstrate that mossy fibers can be distinguished from Purkinje and Golgi cells by the waveshapes of their action potentials. Asynaptic activation from the inferior cerebellar peduncle, in combination with histological localization of recording sites in granular layer or subcortical white matter, verified that mossy fibers produce a variety of waveshapes that are characterized by brief initial phases and relatively small amplitudes.  The same waveshapes were observed for the mossy fiber recordings from awake monkeys, and many identified mossy fibers had sensory properties similar to those found in the awake animals. From these combined criteria, we conclude that the recordings in the awake animals were from mossy fibers. Golgi cells, recorded exclusively in the granular layer of cerebellar cortex, were characterized by action potentials of longer duration and larger amplitude as compared with mossy fibers, and none were asynaptically activated from the inferior cerebellar peduncle. 3. Units were isolated while the monkeys made free-form and tracking movements. We studied movement-related discharge of 80 mossy fibers and 12 Golgi cells. Mossy fibers showed high modulations during use of at least one of the six manipulanda and had clear preferences for movement about a specific joint, although they often showed consistent but weaker firing during movement about a neighboring joint. Separation of movements by more than one joint produced a large reduction in discharge:  shoulder units never fired well to movements of the finger, and finger units never fired well to movement of the shoulder. 4. The tracking task required maintenance of fixed limb positions (a static phase) as well as movements between these positions (a dynamic phase). Of 80 mossy fibers, 18% had purely tonic discharge patterns, 63% were phasic-tonic, and 20% were purely phasic.  Discharge patterns were reciprocal (45%), bidirectional (42%), or unidirectional (13%). 5. Eighty percent of the mossy fibers exhibited tonic discharge that was significantly (P < 0.01) correlated with joint angle (r = 0.65 +/- 0.19, mean +/- SD), and about one third had phasic components that were significantly correlated with movement velocity.  Eleven mossy fibers were tested for correlations between the duration of the phasic discharge component and movement duration, and all revealed significant positive correlations.  The onset time of mossy fiber discharge was distributed approximately equally about the onset time of movement.  Thus discharge of about one fourth of 80 units significantly (P < 0.05) led movement onset and one third significantly lagged.  6.  Twenty-nine movement-related mossy fibers were tested for sensory responsiveness by manipulation of joints and / or by mechanical disturbances of device position during static phases of the tracking task.  In most (69%) cases, passive responses were of the same polarity as the modulations in discharge during comparable active movements; in 17% of the cases, they were of opposite polarity.  Four units (14%) failed to respond to passive movement.  7.  Recordings from 12 Golgi cells revealed properties strikingly different from mossy fibers.  All showed phasic discharge without tonic components, and most cells showed bidirectional discharge patterns during both active and passive movements.  Four of six cells showed modulations in discharge of equal magnitude during use of proximal and distal devices.  These properties are consistent with extensive convergence of mossy fibers on individual Golgi cells.  8.  It is clear from our results that mossy fibers provide intermediate cerebellum with position, velocity, and direction information about movement of individual forelimb joints.  Several characteristics of the signals indicate that they may contain information derived from efference as well as afference.  9.  A comparison of input information supplied by mossy fibers with output signals in the nucleus interpositus suggests that the intermediate cerebellum incorporates position and velocity information from individual joints, together with other inputs, into phasic signals related to coordinated movements of the entire limb.

Wang, J.-J.; Kim, J. H., and Ebner, T. J. Climbing fiber afferent modulation during a visually guided, multi-joint arm movement in the monkey. Brain Res. 1987; 410:323-329.
Abstract:  During a visually guided, multi-joint voluntary arm movement Purkinje cell simple and complex spike activity was recorded from the ipsilateral hemisphere and intermediate zone of the cerebellum in the rhesus monkey. The task consisted of moving a manipulandum over a horizontal video screen.  Manipulandum (hand position) was represented by a cursor on the screen, the animal required to place the manipulandum within displayed start and target boxes. Purkinje cell complex spike discharge was examined using two paradigms. In the first the animal moved the manipulandum from a start box to a target box. In the second the animal was required to modify an ongoing movement and place the cursor within a repositioned target box. A majority of the cells (44/74) exhibited a statistically significant increase in the probability of complex spike discharge at various times during the movement.  The increase was observed when the movement trajectory was redirected (36/44) and/or during the initial portion of the movement (27/44). These results suggest the climbing fiber afferent system is routinely involved in the execution of multi-joint movements especially when the movement is redirected. Possibilities include that climbing fiber afferent input is required when the motor state changes and/or during errors in motor performance.

Wang, J. J.; Shimansky, Y.; Bracha, V., and Bloedel, J. R. Effects of cerebellar nuclear inactivation on the learning of a complex forelimb movement in cats. J.Neurophysiol. 1998 May; 79(5):2447-2459.
Abstract:  The purpose of this study was  to determine the effects of inactivating concurrently the  cerebellar interposed and dentate nuclei on the capacity of cats to acquire and retain a complex, goal-directed forelimb movement. To assess the effects on acquisition, cats were required to learn to move a vertical manipulandum bar through a two-segment template with a shape approximating an inverted "L" after the injection of muscimol (saline for the control group) in the interposed and dentate cerebellar nuclei. During training periods, they were exposed progressively to more difficult templates, which were created by decreasing the angle between the two segments of the template. After determining the most difficult template the injected animals could learn within the specified time and performance constraints, the retraining phase of the experiment was initiated in which the cats were required to execute the same sequence of templates in the absence of any injection. This stage of the experiment assessed retention and determined the extent of any relearning required to execute the task at criterion levels. Next, the animals were overtrained without any injection on the most difficult template they could perform. Finally, to determine the effects of nuclear inactivation on retention after extensive retraining, their capacity to perform the same template was determined after muscimol injection in the interposed and dentate nuclei. The findings show that during the inactivation of the dentate and interposed nuclei the animals could learn to execute the more difficult templates. However, when required to execute the most difficult template learned under muscimol on the day after injections were discontinued, the cats had to "relearn" (reacquire) the movement. Finally, when the cerebellar nuclei were inactivated after the animals learned the task in the absence of any injections during the retraining phase, retention was not blocked. The data indicate that the intermediate and lateral cerebellum are not required either for learning this type of complex voluntary movement or for retaining the capacity to perform the task once it is learned. Nevertheless, when the cerebellum becomes available for executing a task learned in the absence of this structure, reacquisition of the behavior usually is necessary. It is hypothesized that the relearning observed after acquisition during muscimol inactivation reflects the tendency of the system to incorporate the cerebellum into the interactions responsible for the learning and performance of a motor sequence that is optimal for executing the task.

Waterhouse, B. D. and McElligott, J. G. Simple spike activity of Purkinje cells in the posterior vermis of awake cats during spontaneous saccadic eye movements. Brain Res.Bull. 1980; 5:159-168.
Abstract:  Extracellular recordings were made from 151 cerebellar cortical cells in the posterior vermis of 12 awake cats. Thirty-two percent (n = 48) of these cells modulated their activity with respect to the onset of spontaneous saccadic eye movements. Thirty-five cells in this group were positively identified as Purkinje cells and manifested changes in simple spike activity that were related to saccade onset. These included short excitatory, inhibitory, or biphasic changes that were superimposed on background tonic firing rates (avg. = 54 spikes/sec). Such changes were recorded before as well as after the onset of a saccade. Sixty-five percent (n = 22) of these cells were related to horizontal and vertical saccades in more than one direction of motion. These cells were randomly distributed throughout the posterior vermis and manifested no anatomical topographic organization with respect to the direction of saccadic eye movement. The results of this study suggest that lobules VI and VII of the cerebellar vermis participate in both the initiation and execution of spontaneous saccades in preferred directions.