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R H Wurtz

Publications and source records attributed to R H Wurtz.

At least 19 recordsLinked to original sources
PubMed1992

Neuronal correlates of optic flow stimulation.

Neurons in a region of monkey extrastriate cortex, MSTd, respond to the components of optic flow stimulation. Some of these neurons (single-component neurons) are selective for a single type of motion such as inward- or outward-radial motion. Other neurons respond to multiple types of rotation, for example, rightward planar, clockwise circular, and inward radial. Rather than forming discrete groups, we think these neurons represent a continuum covering the range from single-component sensitivity to multiple-component sensitivity. By combining the optic flow stimuli, we have also been able to recognize that such combinations alter the response of cells in the continuum to varying degrees. At this point, while our evidence is consistent with the hypothesis that cells in area MSTd contribute to the processing of optic flow stimuli, we do not know whether these neurons do in fact serve this function. As in all single-cell recording experiments, even those in awake animals performing tasks closer to real-world tasks than we have succeeded in emulating here, the activity of the cell in relationship to the visual stimulation is simply a correlate of the optic flow stimulation and may or may not contribute to the processing of optic flow stimulation upon which behavior depends. Further information on a number of characteristics of these cells might clarify their role. Information on such factors as whether heading in the environment is conveyed by individual neurons, or whether this property is more likely to be conveyed over a population of neurons, and the role of changes in the point of fixation of the eyes are critical points. Generation of behavior on the basis of the optic flow stimulation and determination that this behavior is modified by selective lesion of MSTd would also strengthen the argument that visual motion processing in this area is related to analyzing optic flow information.

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PubMed1992

Role of the rostral superior colliculus in active visual fixation and execution of express saccades.

1. In the rostral pole of the monkey superior colliculus (SC) a subset of neurons (fixation cells) discharge tonically when a monkey actively fixates a target spot and pause during the execution of saccadic eye movements. 2. To test whether these fixation cells are necessary for the control of visual fixation and saccade suppression, we artificially inhibited them with a local injection of muscimol, an agonist of the inhibitory neurotransmitter gamma-aminobutyric acid (GABA). After injection of muscimol into the rostral pole of one SC, the monkey was less able to suppress the initiation of saccades. Many unwanted visually guided saccades were initiated less than 100 ms after onset of a peripheral visual stimulus and therefore fell into the range of express saccades. 3. We propose that fixation cells in the rostral SC form part of a fixation system that facilitates active visual fixation and suppresses the initiation of unwanted saccadic eye movements. Express saccades can only occur when activity in this fixation system is reduced.

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PubMed1992

Disparity sensitivity of neurons in monkey extrastriate area MST.

We tested the disparity sensitivity of neurons from the medial superior temporal area (MST) in awake behaving monkeys. While the monkey looked at a fixation spot on a screen in front of it, random dot stimuli moved in the preferred direction of the cell under study, and the disparity of the dots made the stimuli appear to move in a frontoparallel plane in front of, on, or behind the screen. Over 90% of the 272 neurons studied were sensitive to the disparity of the visual stimulus. Of those disparity-sensitive cells, 95% were most responsive either to near stimuli (stimuli with crossed disparities appearing to move in front of the screen) or to far stimuli (stimuli with uncrossed disparities appearing to move behind the screen). In a smaller sample of the disparity-sensitive cells, we found cells whose preferred direction of stimulus motion reversed as the disparity of the stimulus reversed. For example, a cell that responded best to rightward motion for near stimuli responded best to leftward motion for far stimuli. We found that 40% of the disparity-sensitive cells had this disparity-dependent direction selectivity. This disparity-dependent direction selectivity was maintained over the entire range of speeds tested (6-56 degrees/sec). We tested whether the disparity sensitivity of the neurons indicated the distance of the stimulus from the screen where the monkey was fixating (relative depth) or the distance of the stimulus from the monkey (absolute depth) by having the monkey fixate at different depths in front of or behind the screen. For most MST neurons, the changes in vergence did not alter the disparity response, indicating that the disparity sensitivity of these neurons conveyed information on depth relative to the plane of fixation. We conclude that the disparity characteristics of cells in the dorsomedial MST are those expected of a system serving primarily coarse rather than fine stereopsis. The correlation between disparity selectivity and direction selectivity in these neurons, as well as their other properties, suggests a role in signaling the direction of self-motion of the observer through the environment.

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PubMed1991

Location of saccade-related neurons in the macaque superior colliculus.

The locations of saccade-related neurons were studied in the superior colliculi of two adult rhesus monkeys (Macaca mulatta) by placing marking lesions at the sites of physiologically characterized cells and comparing these histologically identified sites with the collicular laminae and acetylcholinesterase (AChE)-rich patches. Three major conclusions were drawn on the basis of 39 histologically identified sites at which saccade-related neurons were recorded. First, saccade-related neurons were distributed from the ventral half of the optic layer through the deep gray layer, and were most concentrated in the intermediate gray and white layers. Second, there was a clear relationship between the discharge characteristics of these saccade-related neurons and the depths at which they were found. Neurons having presaccadic bursts, defined as clipped and partially-clipped, tended to be encountered more dorsally, and neurons that did not have bursts (unclipped) were encountered more ventrally. Although cells having different discharge characteristics seemed to be organized along a dorsoventral axis, there was no compelling evidence that these properties were specified by their laminar locations. Third, there was no clear correlation between the locations of saccade-related neurons and the distribution of individual AChE-rich patches. Saccade-related cells were found both in the caudal superior colliculus where patches were located and in the rostral superior colliculus where patches were not found; both within and between the two tiers of AChE-rich patches in the caudal superior colliculus; and both within and between individual AChE-rich patches. However, the depth-level at which saccade-related neurons occurred generally matched the region bounded by the two tiers of AChE-rich patches in the intermediate and deep layers, and the dorsal and ventral extent of saccade-related neurons was the same as that of the AChE-rich patches.

Acetylcholinesterase↗
PubMed1991

Sensitivity of MST neurons to optic flow stimuli. I. A continuum of response selectivity to large-field stimuli.

1. Neurons in the dorsomedial region of the medial superior temporal area (MSTd) have large receptive fields that include the fovea, are directionally selective for moving visual stimuli, prefer the motion of large fields to small spots, and respond to rotating and expanding patterns of motion as well as frontal parallel planar motion. These characteristics suggested that these neurons might contribute to the analysis of the large-field optic flow stimulation generated as an observer moves through the visual environment. 2. We tested the response of MSTd neurons in two awake monkeys by systematically presenting a set of translational and rotational stimuli to each neuron. These 100 X 100 degrees stimuli were the motion components from which all optic flow fields are derived. 3. In 220 single neurons we found 23% that responded primarily to one component of motion (planar, circular, or radial), 34% that responded to two components (planocircular or planoradial, but never circuloradial), and 29% that responded to all three components. 4. The number of stimulus components to which a neuron responded was unrelated to the size or eccentricity of its receptive field. 5. Triple-, double-, and single-component neurons varied widely in the strength of their responses to the preferred components. Grouping these neurons together revealed that they did not form discrete classes but rather a continuum of response selectivity. 6. This continuum was apparent in other response characteristics. Direction selectivity was weakest in triple-component neurons, strongest in single-component neurons. Significant inhibitory responses were less frequent in triple-component neurons than in single-component neurons. 7. There was some indication that the neurons of similar component classes occupied adjacent regions within MSTd, but all combinations of component and direction selectivity were occasionally found in immediate juxtaposition. 8. Experiments on a subset of neurons showed that the speed of motion, the dot density, and the number of different speed planes in the display had little influence on these responses. 9. We conclude that the selective responses of many MSTd neurons to the rotational and translational components of optic flow make these neurons reasonable candidates for contributing to the analysis of optic flow fields.

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PubMed1991

Sensitivity of MST neurons to optic flow stimuli. II. Mechanisms of response selectivity revealed by small-field stimuli.

1. In these experiments we examined the receptive field mechanisms that support the optic flow field selective responses of neurons in the dorsomedial region of the medial superior temporal area (MSTd). Our experiments tested the predictions of two hypotheses of optic flow field selectivity. The direction mosaic hypothesis states that these receptive fields contain a set of planar direction-selective subfields that match the local directions of motion within optic flow fields. The vector field hypothesis states that these receptive fields are uniquely sensitive to distributed properties of planar, circular, or radial optic flow fields. 2. Experiments using large-field stimuli revealed that some neurons showed changes in optic flow field selectivity depending on the position of the stimulus in the receptive field; these are position-dependent responses. However, other neurons maintained the same optic flow field selectivities in spite of changes in stimulus position; these are position-invariant responses. We have used the position dependence or invariance of optic flow field selectivity as a way of testing the direction mosaic and vector field hypotheses. Position dependence is more consistent with the direction mosaic hypothesis, whereas position invariance is more consistent with the vector field hypothesis. 3. To test for position effects, we examined the optic flow field selectivity of small subfields within the large receptive fields of 160 MSTd neurons. First, we centered small-field optic flow stimuli of various sizes over the same position in the receptive field. Most MSTd neurons showed decreasing response amplitude with decreasing stimulus size but maintained optic flow field selectivity. 4. We then placed small-field stimuli at various positions within the large receptive field of these MSTd neurons. Position-invariant response selectivity was most prominent in single-component neurons, suggesting that they were more consistent with the vector field hypothesis. Position-dependent response selectivity was most prominent in triple-component neurons, suggesting that they were more consistent with the direction mosaic hypothesis. However, the variations in planar direction preference throughout the receptive field of these triple-component neurons were not consistent with a direction mosaic explanation of the large-field circular or radial selectivity observed. 5. Small-field position studies also demonstrated the existence of zones within the receptive field in which either direction-selective inhibitory or direction-selective excitatory responses predominated. The degree of overlap between these zones increased from nonselective to triple- to double- and finally to single-component neurons.(ABSTRACT TRUNCATED AT 400 WORDS)

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PubMed1991

Recovery of function after lesions in the superior temporal sulcus in the monkey.

1. Ibotenic acid lesions in the monkey's middle temporal area (MT) and the medial superior temporal area (MST) in the superior temporal sulcus (STS) have previously been shown to produce a deficit in initiation of smooth-pursuit eye movements to moving visual targets. The deficits, however, recovery within a few days. In the present experiments we investigated the factors that influence that recovery. 2. We tested two aspects of the monkey's ability to use motion information to acquire moving targets. We used eye-position error as a measure of the monkey's ability to make accurate initial saccades to the moving target. We measured eye speed within the first 100 ms after the saccade to evaluate the monkey's initial smooth pursuit. 3. We determined that pursuit recovery was not dependent specifically on the use of neurotoxic lesions. Although the rate of recovery was slightly altered by replacing the usual neurotoxin (ibotenic acid) with another neurotoxin [alpha-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid (AMPA)] or with an electrolytic lesion, pursuit recovery still occurred within a period of days to weeks. 4. There was a relationship between the size and location of the lesion and the recovery time. The time to recovery for eye-position error and initial eye speed increased with the fraction of MT removed. Whether the rate of recovery and size of lesions within regions on the anterior bank were related was unresolved. 5. We found that a large AMPA lesion within the STS that removed all of MT and nearly all of MST drastically altered the rate of recovery. Recovery was incomplete more than 7 mo after the lesion. Even with this lesion, however, the monkey's ability to use motion information for pursuit was not completely eliminated. 6. The large lesion also included parts of areas V1, V2, V3, and V4, but analysis of the visual fields associated with this lesion indicated that these areas probably did not have a substantial effect on recovery. 7. We tested whether visual motion experience of the monkey after a lesion was necessary for recovery by limiting the monkey's experience either by using a mask or by using 4-Hz stroboscopic illumination. In one monkey the eye-position error component of pursuit was prolonged to greater than 2 wk, but recovery of eye speed was not. Reduced motion experience had little effect on recovery in the other two monkeys. These results suggest that such visual motion experience is not necessary for the recovery of pursuit.(ABSTRACT TRUNCATED AT 400 WORDS)

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PubMed1991

Superior colliculus neurons mediate the dynamic characteristics of saccades.

1. The locus of activity within the superior colliculus (SC) is related to the desired displacement of the eye. Current hypotheses suggest that the location of this locus of activity determines the amplitude of the saccade and that the level of activity at this locus determines eye velocity. We present evidence that suggests that, although the locus determines the amplitude of the saccade, the level of activity in the colliculus encodes dynamic motor error (the difference between desired and current eye displacement). 2. We categorized 86 neurons in the intermediate and deep layers of the superior colliculus of two rhesus monkeys by their activity in relation to the end of saccadic eye movements. In 36% of the cells (n = 31), activity was completely cut off by the end of the saccade (clipped cells). For 53% of cells (n = 46), the major burst of activity ceased by the end of the saccade, but activity continued for 30-100 ms after the end of the movement (partially clipped cells). The remaining 10% of the cells (n = 9) had no clear burst of activity (unclipped cells) but rather had activity that increased gradually before the saccade and then slowly decreased for up to 100 ms after the saccade. These categories were part of a continuum of cell types rather than discrete classes of cells. 3. We first determined whether this new categorization of cells revealed a special relation between the discharge of clipped and partially clipped cells and saccadic amplitude and peak velocity. As expected, we found a steady increase in spike count as saccadic amplitude increased up to the center of the movement field, and an increase in peak spike discharge as peak velocity increased up to a maximum radial eye velocity. Variability in the cell discharge was substantially greater than the variability of saccadic amplitude or peak velocity. We concluded that these single point or averaged measures did not reveal any new functional relationship of these cells. 4. We then examined the relationship of the temporal pattern of discharge of clipped and partially clipped cells to instantaneous changes in radial error and radial velocity. There was a monotonic decay in spike discharge with declining radial error. In contrast, there was a complex, multivalued relationship between spike discharge and radial velocity; collicular cells produced two different values of spike discharge for the same velocity, one during acceleration and the other during deceleration of the eye during a saccade.(ABSTRACT TRUNCATED AT 400 WORDS)

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PubMed1990

The role of disparity-sensitive cortical neurons in signalling the direction of self-motion.

Movement of an observer through the environment generates motion on the retina. This optic flow provides information about the direction of self-motion, but only if it contains differential motion of elements at different depths. If the observer tracks a stationary object while moving in a direction different from his line of sight, the images of objects in the foreground and in the background move in opposite directions. We have found neurons in the cerebral cortex of monkeys that prefer one direction of motion when the disparity of a stimulus corresponds to foreground motion and prefer the opposite direction when the disparity corresponds to background motion. We propose that these neurons contribute a signal about the direction of self-motion.

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PubMed1989

Modulation of pursuit eye movements by stimulation of cortical areas MT and MST.

1. Many cells in the superior temporal sulcus (STS) of the monkey that represent the foveal region of the visual field discharge during pursuit eye movements. Damage to these areas produces a deficit in the maintenance of pursuit eye movements when the target towards the side of the brain with the lesion. In the present experiments, we electrically stimulated these areas to better localize and understand the mechanisms underlying this directional pursuit deficit. 2. Monkeys were trained to pursue a moving target using a step-ramp task in which the target first stepped to an eccentric position and then moved smoothly across the screen. Trains of stimulation were applied after the monkey had begun to pursue the target to study stimulation effects of maintenance of pursuit. 3. Stimulation during pursuit frequently produced eye acceleration toward the side of the brain stimulated. Eye speed increased during pursuit toward the side stimulated and decreased during pursuit away from the side stimulated. This increase in velocity toward the side of the brain where stimulation presumably activated cells is consistent with the decrease in pursuit velocity toward the side of the brain after cells were removed by chemical lesions. 4. The increase or decrease in pursuit speed following stimulation produced a slip of the target on the retina. The pursuit system seemed to be insensitive to this slip during the period of stimulation, however, since the effect of stimulation during pursuit of a stabilized image (open-loop condition) was similar to that resulting from stimulation under normal pursuit conditions (closed-loop). This insensitivity to visual motion during stimulation suggests that the stimulation substitutes for that visual input. 5. The separation of eye and target position that resulted from stimulation did produce catch-up saccades. This provides added evidence that alteration of middle temporal area (MT) and medial superior temporal area (MST) modifies visual-motion but not visual-position information. 6. Stimulation that produced eye acceleration during pursuit produced only a slight effect during fixation of a stationary target. The effectiveness of the stimulation also increased as the speed of the pursuit increased between 5 and 25 degrees/s. These observations, which show that pursuit velocity altered the effect of stimulation, suggest that the stimulation acted on visual motion processing before information about the pursuit movement itself is incorporated. Since this stimulation produces directional pursuit effects, we hypothesize that the directional bias for pursuit originates in the visual signal conveyed to the pursuit system.(ABSTRACT TRUNCATED AT 400 WORDS)

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PubMed1988

Superior colliculus neurons provide the saccadic motor error signal.

Studies of the intermediate layers of the superior colliculus have suggested that it provides a desired change in eye position signal (delta E) for the generation of saccadic eye movements. Recent evidence, however, has shown that some neurons in these layers may be related to the velocity of saccades. We present single cell recordings from the intermediate layers of monkey superior colliculus that are consistent with the hypothesis that many superior colliculus neurons provide instead a motor error signal, em. Our hypothesis about the function of these cells places them inside the local feedback loop controlling the waveform of the saccade.

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PubMed1988

A grid system and a microsyringe for single cell recording.

The designs of two instruments are presented which have proven to be useful in single cell and chemical injection studies performed in awake monkeys. The first is a plastic grid that acts as a guide to produce parallel penetrations with either a microelectrode or microsyringe. The second is a syringe for injecting microliter quantities of a solution that also allows recording of neuronal activity.

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PubMed1988

Relation of cortical areas MT and MST to pursuit eye movements. I. Localization and visual properties of neurons.

1. Among the multiple extrastriate visual areas in monkey cerebral cortex, several areas within the superior temporal sulcus (STS) are selectively related to visual motion processing. In this series of experiments we have attempted to relate this visual motion processing at a neuronal level to a behavior that is dependent on such processing, the generation of smooth-pursuit eye movements. 2. We studied two visual areas within the STS, the middle temporal area (MT) and the medial superior temporal area (MST). For the purposes of this study, MT and MST were defined functionally as those areas within the STS having a high proportion of directionally selective neurons. MST was distinguished from MT by using the established relationship of receptive-field size to eccentricity, with MST having larger receptive fields than MT. 3. A subset of these visually responsive cells within the STS were identified as pursuit cells--those cells that discharge during smooth pursuit of a small target in an otherwise dark room. Pursuit cells were found only in localized regions--in the foveal region of MT (MTf), in a dorsal-medial area of MST on the anterior bank of the STS (MSTd), and in a lateral-anterior area of MST on the floor and the posterior bank of the STS (MST1). 4. Pursuit cells showed two characteristics in common when their visual properties were studied while the monkey was fixating. Almost all cells showed direction selectivity for moving stimuli and included the fovea within their receptive fields. 5. The visual response of pursuit cells in the several areas differed in two ways. Cells in MTf preferred small moving spots of light, whereas cells in MSTd preferred large moving stimuli, such as a pattern of random dots. Cells in MTf had small receptive fields; those in MSTd usually had large receptive fields. Visual responses of pursuit neurons in MST1 were heterogeneous; some resembled those in MTf, whereas others were similar to those in MSTd. This suggests that the pursuit cells in MSTd and MST1 belong to different subregions of MST.

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PubMed1988

Relation of cortical areas MT and MST to pursuit eye movements. II. Differentiation of retinal from extraretinal inputs.

1. We investigated cells in the middle temporal visual area (MT) and the medial superior temporal area (MST) that discharged during smooth pursuit of a dim target in an otherwise dark room. For each of these pursuit cells we determined whether the response during pursuit originated from visual stimulation of the retina by the pursuit target or from an extraretinal input related to the pursuit movement itself. We distinguished between these alternatives by removing the visual motion stimulus during pursuit either by blinking off the visual target briefly or by stabilizing the target on the retina. 2. In the foveal representation of MT (MTf), we found that pursuit cells usually decreased their rate of discharge during a blink or during stabilization of the visual target. The pursuit response of these cells depends on visual stimulation of the retina by the pursuit target. 3. In a dorsal-medial region of MST (MSTd), cells continued to respond during pursuit despite a blink or stabilization of the pursuit target. The pursuit response of these cells is dependent on an extraretinal input. 4. In a lateral-anterior region of MST (MST1), we found both types of pursuit cells; some, like those in MTf, were dependent on visual inputs whereas others, like those in MSTd, received an extraretinal input. 5. We observed a relationship between pursuit responses and passive visual responses. MST cells whose pursuit responses were attributable to extraretinal inputs tended to respond preferentially to large-field random-dot patterns. Some cells that preferred small spots also had an extraretinal input. 6. For 92% of the pursuit cells we studied, the pursuit response began after onset of the pursuit eye movement. A visual response preceding onset of the eye movement could be observed in many of these cells if the initial motion of the target occurred within the visual receptive field of the cell and in its preferred direction. In contrast to the pursuit response, however, this visual response was not dependent on execution of the pursuit movement. 7. For the remaining 8% of the pursuit cells, the pursuit discharge began before initiation of the pursuit eye movement. This occurred even though the initial motion of the target was outside the receptive field as mapped during fixation trials. Our data suggest, however, that such responses may be attributable to an expansion of the receptive field that accompanies enhanced visual responses.(ABSTRACT TRUNCATED AT 400 WORDS)

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