Cerebellar Purkinje cell simple spike discharge encodes movement velocity in primates during visuomotor arm tracking. (1/3218)

Pathophysiological, lesion, and electrophysiological studies suggest that the cerebellar cortex is important for controlling the direction and speed of movement. The relationship of cerebellar Purkinje cell discharge to the control of arm movement parameters, however, remains unclear. The goal of this study was to examine how movement direction and speed and their interaction-velocity-modulate Purkinje cell simple spike discharge in an arm movement task in which direction and speed were independently controlled. The simple spike discharge of 154 Purkinje cells was recorded in two monkeys during the performance of two visuomotor tasks that required the animals to track targets that moved in one of eight directions and at one of four speeds. Single-parameter regression analyses revealed that a large proportion of cells had discharge modulation related to movement direction and speed. Most cells with significant directional tuning, however, were modulated at one speed, and most cells with speed-related discharge were modulated along one direction; this suggested that the patterns of simple spike discharge were not adequately described by single-parameter models. Therefore, a regression surface was fitted to the data, which showed that the discharge could be tuned to specific direction-speed combinations (preferred velocities). The overall variability in simple spike discharge was well described by the surface model, and the velocities corresponding to maximal and minimal discharge rates were distributed uniformly throughout the workspace. Simple spike discharge therefore appears to integrate information about both the direction and speed of arm movements, thereby encoding movement velocity.  (+info)

Eye movement deficits following ibotenic acid lesions of the nucleus prepositus hypoglossi in monkeys II. Pursuit, vestibular, and optokinetic responses. (2/3218)

The eyes are moved by a combination of neural commands that code eye velocity and eye position. The eye position signal is supposed to be derived from velocity-coded command signals by mathematical integration via a single oculomotor neural integrator. For horizontal eye movements, the neural integrator is thought to reside in the rostral nucleus prepositus hypoglossi (nph) and project directly to the abducens nuclei. In a previous study, permanent, serial ibotenic acid lesions of the nph in three rhesus macaques compromised the neural integrator for fixation but saccades were not affected. In the present study, to determine further whether the nph is the neural substrate for a single oculomotor neural integrator, the effects of those lesions on smooth pursuit, the vestibulo-ocular reflex (VOR), vestibular nystagmus (VN), and optokinetic nystagmus (OKN) are documented. The lesions were correlated with long-lasting deficits in eye movements, indicated most clearly by the animals' inability to maintain steady gaze in the dark. However, smooth pursuit and sinusoidal VOR in the dark, like the saccades in the previous study, were affected minimally. The gain of horizontal smooth pursuit (eye movement/target movement) decreased slightly (<25%) and phase lead increased slightly for all frequencies (0.3-1.0 Hz, +/-10 degrees target tracking), most noticeably for higher frequencies (0.8-0.7 and approximately 20 degrees for 1.0-Hz tracking). Vertical smooth pursuit was not affected significantly. Surprisingly, horizontal sinusoidal VOR gain and phase also were not affected significantly. Lesions had complex effects on both VN and OKN. The plateau of per- and postrotatory VN was shortened substantially ( approximately 50%), whereas the initial response and the time constant of decay decreased slightly. The initial OKN response also decreased slightly, and the charging phase was prolonged transiently then recovered to below normal levels like the VN time constant. Maximum steady-state, slow eye velocity of OKN decreased progressively by approximately 30% over the course of the lesions. These results support the previous conclusion that the oculomotor neural integrator is not a single neural entity and that the mathematical integrative function for different oculomotor subsystems is most likely distributed among a number of nuclei. They also show that the nph apparently is not involved in integrating smooth pursuit signals and that lesions of the nph can fractionate the VOR and nystagmic responses to adequate stimuli.  (+info)

Optimality of position commands to horizontal eye muscles: A test of the minimum-norm rule. (3/3218)

Six muscles control the position of the eye, which has three degrees of freedom. Daunicht proposed an optimization rule for solving this redundancy problem, whereby small changes in eye position are maintained by the minimum possible change in motor commands to the eye (the minimum-norm rule). The present study sought to test this proposal for the simplified one-dimensional case of small changes in conjugate eye position in the horizontal plane. Assuming such changes involve only the horizontal recti, Daunicht's hypothesis predicts reciprocal innervation with the size of the change in command matched to the strength of the recipient muscle at every starting position of the eye. If the motor command to a muscle is interpreted as the summed firing rate of its oculomotor neuron (OMN) pool, the minimum-norm prediction can be tested by comparing OMN firing rates with forces in the horizontal recti. The comparison showed 1) for the OMN firing rates given by Van Gisbergen and Van Opstal and the muscle forces given by Robinson, there was good agreement between the minimum-norm prediction and experimental observation over about a +/-30 degrees range of eye positions. This fit was robust with respect to variations in muscle stiffness and in methods of calculating muscle innervation. 2) Other data sets gave different estimates for the range of eye-positions within which the minimum-norm prediction held. The main sources of variation appeared to be disagreement about the proportion of OMNs with very low firing-rate thresholds (i.e., less than approximately 35 degrees in the OFF direction) and uncertainty about eye-muscle behavior for extreme (>30 degrees ) positions of the eye. 3) For all data sets, the range of eye positions over which the minimum-norm rule applied was determined by the pattern of motor-unit recruitment inferred for those data. It corresponded to the range of eye positions over which the size principle of recruitment was obeyed by both agonist and antagonist muscles. It is argued that the current best estimate of the oculomotor range over which minimum-norm control could be used for conjugate horizontal eye position is approximately +/-30 degrees. The uncertainty associated with this estimate would be reduced by obtaining unbiased samples of OMN firing rates. Minimum-norm control may result from reduction of the image movement produced by noise in OMN firing rates.  (+info)

Visuomotor processing as reflected in the directional discharge of premotor and primary motor cortex neurons. (4/3218)

Premotor and primary motor cortical neuronal firing was studied in two monkeys during an instructed delay, pursuit tracking task. The task included a premovement "cue period," during which the target was presented at the periphery of the workspace and moved to the center of the workspace along one of eight directions at one of four constant speeds. The "track period" consisted of a visually guided, error-constrained arm movement during which the animal tracked the target as it moved from the central start box along a line to the opposite periphery of the workspace. Behaviorally, the animals tracked the required directions and speeds with highly constrained trajectories. The eye movements consisted of saccades to the target at the onset of the cue period, followed by smooth pursuit intermingled with saccades throughout the cue and track periods. Initially, an analysis of variance (ANOVA) was used to test for direction and period effects in the firing. Subsequently, a linear regression analysis was used to fit the average firing from the cue and track periods to a cosine model. Directional tuning as determined by a significant fit to the cosine model was a prominent feature of the discharge during both the cue and track periods. However, the directional tuning of the firing of a single cell was not always constant across the cue and track periods. Approximately one-half of the neurons had differences in their preferred directions (PDs) of >45 degrees between cue and track periods. The PD in the cue or track period was not dependent on the target speed. A second linear regression analysis based on calculation of the preferred direction in 20-ms bins (i.e., the PD trajectory) was used to examine on a finer time scale the temporal evolution of this change in directional tuning. The PD trajectories in the cue period were not straight but instead rotated over the workspace to align with the track period PD. Both clockwise and counterclockwise rotations occurred. The PD trajectories were relatively straight during most of the track period. The rotation and eventual convergence of the PD trajectories in the cue period to the preferred direction of the track period may reflect the transformation of visual information into motor commands. The widely dispersed PD trajectories in the cue period would allow targets to be detected over a wide spatial aperture. The convergence of the PD trajectories occurring at the cue-track transition may serve as a "Go" signal to move that was not explicitly supplied by the paradigm. Furthermore, the rotation and convergence of the PD trajectories may provide a mechanism for nonstandard mapping. Standard mapping refers to a sensorimotor transformation in which the stimulus is the object of the reach. Nonstandard mapping is the mapping of an arbitrary stimulus into an arbitrary movement. The shifts in the PD may allow relevant visual information from any direction to be transformed into an appropriate movement direction, providing a neural substrate for nonstandard stimulus-response mappings.  (+info)

Short-latency vergence eye movements induced by radial optic flow in humans: dependence on ambient vergence level. (5/3218)

Radial patterns of optic flow, such as those experienced by moving observers who look in the direction of heading, evoke vergence eye movements at short latency. We have investigated the dependence of these responses on the ambient vergence level. Human subjects faced a large tangent screen onto which two identical random-dot patterns were back-projected. A system of crossed polarizers ensured that each eye saw only one of the patterns, with mirror galvanometers to control the horizontal positions of the images and hence the vergence angle between the two eyes. After converging the subject's eyes at one of several distances ranging from 16.7 cm to infinity, both patterns were replaced with new ones (using a system of shutters and two additional projectors) so as to simulate the radial flow associated with a sudden 4% change in viewing distance with the focus of expansion/contraction imaged in or very near both foveas. Radial-flow steps induced transient vergence at latencies of 80-100 ms, expansions causing increases in convergence and contractions the converse. Based on the change in vergence 90-140 ms after the onset of the steps, responses were proportional to the preexisting vergence angle (and hence would be expected to be inversely proportional to viewing distance under normal conditions). We suggest that this property assists the observer who wants to fixate ahead while passing through a visually cluttered area (e.g., a forest) and so wants to avoid making vergence responses to the optic flow created by the nearby objects in the periphery.  (+info)

Common 3 and 10 Hz oscillations modulate human eye and finger movements while they simultaneously track a visual target. (6/3218)

1. A 10 Hz range centrally originating oscillation has been found to modulate slow finger movements and anticipatory smooth eye movements. To determine if an interaction or linkage occurs between these two central oscillations during combined visuo-manual tracking, frequency and coherence analysis were performed on finger and eye movements while they simultaneously tracked a visual target moving in intermittently visible sinusoidal patterns. 2. Two different frequencies of common or linked oscillation were found. The first, at 2-3 Hz, was dependent on visual feedback of target and finger tracking positions. The second, at around 10 Hz, still occurred when both target and finger positions were largely obscured, indicating that this common oscillation was generated internally by the motor system independent of visual feedback. Both 3 and 10 Hz oscillation frequencies were also shared by the right and left fingers if subjects used these together to track a visual target. 3. The linking of the 10 Hz range oscillations between the eyes and finger was task specific; it never occurred when eye and finger movements were made simultaneously and independently, but only when they moved simultaneously and followed the target together. However, although specific for tracking by the eyes and fingers together, the linking behaviour did not appear to be a prerequisite for such tracking, since significant coherence in the 10 Hz range was only present in a proportion of trials where these combined movements were made. 4. The experiments show that common oscillations may modulate anatomically very distinct structures, indicating that single central oscillations may have a widespread distribution in the central nervous system. The task-specific manifestation of the common oscillation in the eye and finger suggests that such mechanisms may have a functional role in hand-eye co-ordination.  (+info)

Projections and firing properties of down eye-movement neurons in the interstitial nucleus of Cajal in the cat. (7/3218)

To clarify the role of the interstitial nucleus of Cajal (INC) in the control of vertical eye movements, projections of burst-tonic and tonic neurons in and around the INC were studied. This paper describes neurons with downward ON directions. We examined, by antidromic activation, whether these down INC (d-INC) neurons contribute to two pathways: a commissural pathway to the contralateral (c-) INC and a descending pathway to the ipsilateral vestibular nucleus (i-VN). Stimulation of the two pathways showed that as many as 74% of neurons were activated antidromically from one of the pathways. Of 113 d-INC neurons tested, 44 were activated from the commissural pathway and 40 from the descending pathway. No neurons were activated from both pathways. We concluded that commissural and descending pathways from the INC originate from two separate groups of neurons. Tracking of antidromic microstimulation in the two nuclei revealed multiple low-threshold sites and varied latencies; this was interpreted as a sign of existence of axonal arborization. Neurons with commissural projections tended to be located more dorsally than those with descending projections. Neurons with descending projections had significantly greater eye-position sensitivity and smaller saccadic sensitivity than neurons with commissural projections. The two groups of INC neurons increased their firing rate in nose-up head rotations and responded best to the rotation in the plane of contralateral posterior/ipsilateral anterior canal pair. Neurons with commissural projections showed a larger phase lag of response to sinusoidal rotation (54.6 +/- 7.6 degrees ) than neurons with descending projections (45.0 +/- 5.5 degrees ). Most neurons with descending projections received disynaptic excitation from the contralateral vestibular nerve. Neurons with commissural projections rarely received such disynaptic input. We suggest that downward-position-vestibular (DPV) neurons in the VN and VN-projecting d-INC neurons form a loop, together with possible commissural loops linking the bilateral VNs and the bilateral INCs. By comparing the quantitative measures of d-INC neurons with those of DPV neurons, we further suggest that integration of head velocity signals proceeds from DPV neurons to d-INC neurons with descending projections and then to d-INC neurons with commissural projections, whereas saccadic velocity signals are processed in the reverse order.  (+info)

Optic flow selectivity in the anterior superior temporal polysensory area, STPa, of the behaving monkey. (8/3218)

Earlier studies of neurons in the anterior region of the superior temporal polysensory area (STPa) have demonstrated selectivity for visual motion using stimuli contaminated by nonmotion cues, including texture, luminance, and form. The present experiments investigated the motion selectivity of neurons in STPa in the absence of form cues using random dot optic flow displays. The responses of neurons were tested with translation, rotation, radial, and spiral optic flow displays designed to mimic the types of motion that occur during locomotion. Over half of the neurons tested responded significantly to at least one of these displays. On a cell by cell basis, 60% of the neurons tested responded selectively to rotation, radial, and spiral motion, whereas 20% responded selectively to translation motion. The majority of neurons responded maximally to single-component optic flow displays but was also significantly activated by the spiral displays that contained their preferred component. Moreover, there was a bias in the selectivity of the neurons for radial expansion motion. These results suggest that neurons within STPa are contributing to the analysis of optic flow. Furthermore, the preponderance of cells selective for radial expansion provides evidence that this area may be specifically involved in the processing of forward locomotion and/or looming stimuli. Finally, these results provide carefully controlled physiological evidence for an extension and specialization of the motion-processing pathway into the anterior temporal lobe.  (+info)