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N J Gandhi

Publications and source records attributed to N J Gandhi.

10 recordsLinked to original sources
PubMed2001

Experimental control of eye and head positions prior to head-unrestrained gaze shifts in monkey.

A coordinated movement of the eyes and head in the head-unrestrained condition is often used to change orientation between targets. Under natural conditions, these gaze shifts are typically generated with the eyes roughly centered in the orbits. To achieve experimental control of eye and head positions, a miniature laser was mounted on the head implants of monkeys that were trained to point the head to one target and direct gaze to another before generating a head-unrestrained gaze shift to a third target (dissociation paradigm). For comparison, monkeys were also required to make gaze shifts between stimuli, without any constraints on eye and head positions (standard paradigm). Analyses indicated that movement parameters, limited to horizontal gaze shifts, were similar for both behavioral conditions. Thus, the proposed technique and behavioral paradigm, when used in conjunction with electrophysiological and pharmacological experiments, may facilitate the study of neural control of gaze.

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PubMed2000

Activity in deep intermediate layer collicular neurons during interrupted saccades.

The activity of neurons located in the deep intermediate and adjacent deep layers (hereafter called just deep intermediate layer neurons) of the superior colliculus (SC) in monkeys was recorded during saccades interrupted by electrical stimulation of the brainstem omnipause neuron (OPN) region. The goal of the experiment was to determine if these neurons maintained their discharge during the saccadic interruption, and thus, could potentially provide a memory trace for the intended movement which ends accurately on target in spite of the perturbation. The collicular neurons recorded in the present study were located in the rostral three-fifths of the colliculus. Most of these cells tended to show considerable presaccadic activity during a delayed saccade paradigm, and, therefore, probably overlap with the population of SC cells called buildup neurons or prelude bursters in previous studies. The effect of electrical stimulation in the OPN region (which interrupted ongoing saccades) on the discharge of these neurons was measured by computing the percentage reduction in a cell's activity compared to that present during non-interrupted saccades. During saccade interruption about 70% of deep intermediate layer neurons experienced a major reduction (30% or greater) in their activity, but discharge recovered quickly after the termination of the stimulation as the eyes resumed their movement to finish the saccade on the target. Therefore, the pattern of activity recorded in most of the deep intermediate layer neurons during interrupted saccades qualitatively resembled that previously reported for the saccade-related burst neurons which tend to be located more dorsally in the intermediate layer. In contrast, some of our cells (30%) showed little or no perturbation in their activity caused by the saccade interrupting stimulation. Because all the more dorsally located burst neurons and the majority of our deep intermediate layer neurons show a total or major suppression in their discharge during interrupted saccades, it seems unlikely that the colliculus by itself could maintain an accurate memory of the desired saccadic goal or the remaining dynamic motor error required to account for the accuracy of the resumed movement which occurs following the interruption. However, it remains possible that the smaller proportion of our neurons whose activity was not perturbed during interrupted movements could play a role in the mechanisms underlying saccade accuracy in the interrupted saccade paradigm. Interrupted saccades have longer durations than normal saccades to the same target. Therefore, we investigated whether the discharge of our deeper collicular cells was also necessarily prolonged during interrupted saccades, and, if so, how the prolongation compared to the prolongation of the saccade. Sixty percent of our sample neurons showed a prolongation in discharge that was approximately the same as the prolongation in saccade duration (difference < 15 ms in magnitude). The, observation that temporal discharge in our neurons was perturbed to roughly match saccadic temporal perturbation suggests that dynamic feedback about ongoing saccadic motion is provided to the colliculus, but does not necessarily imply that this structure is the site responsible for the computation of dynamic motor error.

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PubMed1999

Comparison of saccades perturbed by stimulation of the rostral superior colliculus, the caudal superior colliculus, and the omnipause neuron region.

Over the past decade, considerable research efforts have been focused on the role of the rostral superior colliculus (SC) in control of saccades. The most recent theory separates the deeper intermediate layers of the SC into two functional regions: the rostral pole of these layers constitutes a fixation zone and the caudal region comprises the saccade zone. Sustained activity of fixation neurons in the fixation zone is argued to maintain fixation and help prevent saccade generation by exciting the omnipause neurons (OPNs) in the brain stem. This hypothesis is in contrast to the traditional view that the SC contains a topographic representation of the saccade motor map on which the rostral pole of the SC encodes signals for generating small saccades (<2 degrees ) instead of preventing them. There is therefore an unresolved controversy about the specific role on the most rostral region of the SC, and we reexamined its functional contribution by quantifying and comparing spatial and temporal trajectories of 30 degrees saccades perturbed by electrical stimulation of the rostral pole and more caudal regions in the SC and of the OPN region. If the rostral pole serves to preserve fixation, then saccades perturbed by stimulation should closely resemble interrupted saccades produced by stimulation of the OPN region. If it also contributes to saccade generation, then the disrupted movements would better compare with redirected saccades observed after stimulation of the caudal SC. Our experiments revealed two significant findings: 1) the locus of stimulation was the primary factor determining the perturbation effect. If the directions of the target-directed saccade and stimulation-evoked saccade were aligned and if the stimulation was delivered within approximately the rostral 2 mm (<10 degrees amplitude) of SC, the ongoing saccade stopped in midflight but then resumed after stimulation end to reach the original visually specified goal with close to normal accuracy. When stimulation was applied at more caudal sites, the ongoing saccade directly reached the target location without stopping at an intermediate position. If the directions differed considerably, both initial and resumed components were typically observed for all stimulation sites. 2) A quantitative analysis of the saccades perturbed from the fixation zone showed significant deviations from their control spatial trajectories. Thus they resembled redirected saccades induced by caudal SC stimulation and differed significantly from interrupted saccades produced by OPN stimulation. The amplitude of the initial saccade, latency of perturbation, and spatial redirection were greatest for the most caudal sites and decreased gradually for rostral sites. For stimulation sites within the rostral pole of SC, the measures formed a smooth continuation of the trends observed in the saccade zone. As these results argue for the saccade zone concept, we offer reinterpretations of the data used to support the fixation zone model. However, we also discuss scenarios that do not allow an outright rejection of the fixation zone hypothesis.

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PubMed1999

Activity of the brain stem omnipause neurons during saccades perturbed by stimulation of the primate superior colliculus.

Stimulation of the rostral approximately 2 mm of the superior colliculus (SC) during a large, visual target-initiated saccade produces a spatial deviation of the ongoing saccade and then stops it in midflight. After the termination of the stimulation, the saccade resumes and ends near the location of the flashed target. The density of collicular projections to the omnipause neuron (OPN) region is greatest from the rostral SC and decreases gradually for the more caudal regions. It has been hypothesized that the microstimulation excites the OPNs through these direct connections, and the reactivation of OPNs, which are normally silent during saccades, stops the initial component in midflight by gating off the saccadic burst generator. Two predictions emerge from this hypothesis: 1) for microstimulation triggered on the onset of large saccades, the time from stimulation onset to resumption of OPN discharge should decrease as the stimulation site is moved rostral and 2) the lead time from reactivation of OPNs to the end of the initial saccade on stimulation trials should be equal to the lead time of pause end with respect to the end of control saccades. We tested this hypothesis by recording OPN activity during saccades perturbed by stimulation of the rostral approximately 2 mm of the SC. The distance of the stimulation site from the most rostral extent of the SC and the time of reactivation with respect to stimulation onset were not significantly correlated. The mean lead of reactivation of OPNs relative to the end of the initial component of perturbed saccades (6.5 ms) was significantly less than the mean lead with respect to the end of control (9.6 ms) and resumed saccades (10.4 ms). These results do not support the notion that the excitatory input from SC neurons-in particular, the fixation neurons in the rostral SC-provide the major signal to reactivate OPNs and end saccades. An alternative, conceptual model to explain the temporal sequence of events induced by stimulation of the SC during large saccades is presented. Other OPN activity parameters also were measured and compared for control and stimulation conditions. The onset of pause with respect to resumed saccade onset was larger and more variable than the onset of pause with respect to control saccades, whereas pause end with respect to the end of resumed and control saccades was similar. The reactivated discharge of OPNs during the period between the end of the initial and the onset of the resumed saccades was at least as strong as that following control movements. This latter observation is interpreted in terms of the resettable neural integrator hypothesis.

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PubMed1998

Two-dimensional saccade-related population activity in superior colliculus in monkey.

The two-dimensional distribution of population activity in the superior colliculus (SC) during saccadic eye movements in the monkey was estimated using radial basis functions. To make these ensemble activity estimates, cells in the deeper layers of the SC were recorded over much of the rostrocaudal (caudal to 3.8 mm from the rostral tip), mediolateral extent of this structure. The dynamic movement field of each cell was determined at 2-ms intervals around the time of saccades for a wide variety of horizontal and oblique movements. Collicular neurons were divided into partially overlapping dorsal and ventral cell layers on the basis of recorded depth in SC. The pattern of presaccadic activity was used as an additional discriminant to sort the cells in the two layers into separate burst (dorsal) and buildup (ventral) cell classes. Rostrocaudal and medioventral cell location on the colliculus was estimated from the optimal target vector for a cell's visual response rather than from the optimal motor vector. The former technique was more reliable for locating some buildup neurons because it produced locations that compared better with the locations suggested by electrical stimulation. From the movement field data and from the estimates of each cell's anatomic location, a similar algorithm was used to compute the two-dimensional population activity in the two layers of the SC during horizontal and oblique saccades. A subset of the sample of neurons, located near the horizontal meridian of the SC, first was used to compute one-dimensional dynamic population activity estimates for horizontal saccades to allow partial comparison to previous studies. Statistical analyses on the one-dimensional data were limited to saccades of </=20 degrees. The analyses indicated that while there was a small rostrally directed shift in the center of gravity of the distributed activity in the buildup cell layer, there was little support for the theory of a systematic rostrally directed spread of the leading edge of the activity. The two-dimensional results extend the previous one-dimensional estimates of collicular activity during saccades. Discharge in the burst layer was invariant in size for all saccade vectors and symmetrically arranged about a center of gravity that did not move during saccades. The size of the active area in the buildup layer grew modestly with saccade amplitude, whereas the distribution of activity was skewed toward the rostral end of the SC for saccades larger than 10 degrees. There was a small, but consistent shift in the center of gravity of the two-dimensional activity that was directed along the horizontal meridian (for horizontal movements) or an oblique meridian (for oblique movements) of the SC. However, the spread of activity during a saccade was as large or larger in the mediolateral direction as it was in the rostral direction. The results indicate that changes in activity occur in an extended zone on the SC, and in all directions but caudal, in the buildup layer during saccades and do not support the idea of a rostrally directed spread of activity as a dynamic control mechanism for saccades. Our results and those of previous investigators of collicular population activity may be limited by stationarity concerns in that the cells used to estimate population activity were recorded in several monkeys over an extended period of time to obtain a sufficient spatial sample.

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PubMed1997

Spatial distribution and discharge characteristics of superior colliculus neurons antidromically activated from the omnipause region in monkey.

One proposed role of the superior colliculus (SC) in oculomotor control is to suppress or excite the activity of brain stem omnipause neurons (OPNs) to initiate or terminate saccades, respectively. Although connections from the SC to the OPNs have been demonstrated, the spatial distribution and discharge characteristics of the projecting neurons from the SC remain unknown. We mapped the spatial distribution of the deeper-layer neurons of the SC by stimulating the region of the OPNs to identify antidromic projections and found that the density of direct projections from the SC to the OPNs was greatest in the most rostral region and decreased gradually for more caudal sites. On the basis of saccade-related discharge characteristics, the antidromically driven neurons were predominantly fixation and buildup neurons. The spatially distributed SC projections to the OPNs and the discharge characteristics of the SC neurons suggest that the direct projections from SC to OPNs are excitatory. Finally, we propose how excitation and disfacilitation from SC activity can contribute to modulation of OPN response and control saccades.

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PubMed1996

Discharge of superior collicular neurons during saccades made to moving targets.

1. The discharge of neurons in the deeper layers of the monkey superior colliculus was recorded during saccades made to stationary and to smoothly moving visual targets. 2. All neurons that discharged for saccades made to stationary targets also discharged during saccades made to moving targets, but there was a systematic shift in the saccade vector yielding maximal activity (i.e. center of the movement field) of collicular neurons for the latter class of movements. The shift moved the center of the movement fields toward larger-amplitude pursuit saccades for target motion away from the fovea, in comparison with saccades made to stationary targets. However, the discharge at the center of the movement field for pursuit saccades was 14% lower when averaged over the sample of recorded cells. 3. The saccades made during pursuit tracking of moving visual stimuli have different dynamics than saccades made to stationary targets. At similar amplitudes pursuit saccades are slower, and their velocity profiles often show secondary velocity peaks or inflection points and have longer-duration decelerating phases. 4. The combined experimental observations of a change in saccade dynamics and the shift in movement fields in collicular neurons for pursuit saccades are compatible with the hypothesis that saccades made to moving targets are controlled by neural processing in two partially separate pathways. In this theory, one path is concerned with correction of a presaccadic retinal position error (a path that includes the colliculus) and another path is concerned with position extrapolations based on the velocity of the moving target (a path that does not include the colliculus).

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PubMed1995

Open-loop simulations of the primate saccadic system using burst cell discharge from the superior colliculus.

Saccade-related burst neurons (SRBNs) in the monkey superior colliculus (SC) have been hypothesized to provide the brainstem saccadic burst generator with the dynamic error signal and the movement initiating trigger signal. To test this claim, we performed two sets of open-loop simulations on a burst generator model with the local feedback disconnected using experimentally obtained SRBN activity as both the driving and trigger signal inputs to the model. First, using neural data obtained from cells located near the middle of the rostral to caudal extent of the SC, the internal parameters of the model were optimized by means of a stochastic hill-climbing algorithm to produce an intermediate-sized saccade. The parameter values obtained from the optimization were then fixed and additional simulations were done using the experimental data from rostral collicular neurons (small saccades) and from more caudal neurons (large saccades); the model generated realistic saccades, matching both position and velocity profiles of real saccades to the centers of the movement fields of all these cells. Second, the model was driven by SRBN activity affiliated with interrupted saccades, the resumed eye movements observed following electrical stimulation of the omnipause region. Once again, the model produced eye movements that closely resembled the interrupted saccades produced by such simulations, but minor readjustment of parameters reflecting the weight of the projection of the trigger signal was required. Our study demonstrates that a model of the burst generator produces reasonably realistic saccades when driven with actual samples of SRBN discharges.

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PubMedDate not supplied

Endpoint accuracy in saccades interrupted by stimulation in the omnipause region in monkey.

Electrical stimulation of the omnipause neuron region (OPN) at saccade onset results in interrupted saccades (IS)- eye movements which pause in midflight, resume after a brief period, and end near the target location. Details on the endpoint accuracy of IS do not exist, except for a brief report by Becker et al. (1981). Their analysis emphasized the accuracy of IS relative to the visual target which remained on during the interrupted period. We instead quantified the metric properties of IS relative to nonstimulated saccades during a target flash paradigm. Our results show that IS tend to be slightly hypermetric relative to the nonstimulated saccades to the same target location. The amount of overshoot is not correlated with target eccentricity. Detailed analyses also indicate that the standard deviations of the endpoint in IS are not significantly larger than those for nonstimulated saccades, although there was a much larger variability produced in eye position during the interruption. Both these latter observations support the notion that saccades are controlled by an internal negative feedback system. Also, the size of the remaining motor error during the interrupted period is one factor influencing when an IS resumes, but the variability in this measure is large particularly for smaller motor errors. Recent results have suggested that the resettable neural integrator involved in the feedback loop may be reset after each saccade through an exponential decay process. To probe the properties of the neural integrator, we varied the duration of interruption between the initial and resumed saccades and sought a systematic overshoot in the final eye position with increasing interruption period and variable initial saccade size. Our results showed the neural integrator does not decay during the pause period of interrupted saccades.

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