Discharge Characteristics of Pursuit Neurons in MST During Vergence Eye Movements

2005 ◽  
Vol 93 (5) ◽  
pp. 2415-2434 ◽  
Author(s):  
Teppei Akao ◽  
Michael J. Mustari ◽  
Junko Fukushima ◽  
Sergei Kurkin ◽  
Kikuro Fukushima
Keyword(s):  
Eye Movements ◽  
Visual Motion ◽  
Significant Proportion ◽  
Visual Signals ◽  
Visual Responses ◽  
Discharge Characteristics ◽  
Temporal Area ◽  
Mst Neurons ◽  
Pursuit Neurons ◽  
Vergence Tracking

For small objects moving smoothly in space close to the observer, smooth pursuit and vergence eye movements maintain target images near the foveae to insure high-resolution processing of visual signals about moving objects. Signals for both systems must be synthesized for pursuit-in-three-dimensions (3D). Recent studies have shown that responses of the majority of pursuit neurons in the frontal eye fields (FEF) code pursuit-in-3D. This area is known to have reciprocal connections with the medial superior temporal area (MST) where frontal pursuit neurons are found. To examine whether pursuit-in-3D signals are already present in MST and how MST neurons discharge during vergence-tracking induced by a small spot, we examined discharge of MST pursuit neurons in 2 monkeys. Of a total of 219 pursuit neurons examined during both frontal pursuit and vergence-tracking, 61% discharged only for frontal pursuit, 18% only for vergence-tracking, and 21% for both. A majority of vergence-related MST neurons exhibited sensitivity to vergence eye velocity. Their discharge was maintained during brief blanking of a vergence target. About 1/3 of vergence-related MST neurons exhibited visual responses to spot motion in depth. The preferred directions for visual motion and vergence-tracking were similar in half of our population. Some of the remaining neurons showed opposite preferred directions. A significant proportion (29%) of vergence-related neurons discharged before onset of eye movements with lead times longer than 20 ms. The results in this and previous studies indicate differences in discharge characteristics of FEF and MST pursuit neurons, suggesting different roles for the two in pursuit-in-3D.

scholarly journals Modulation of Visual Signals in Macaque MT and MST Neurons During Pursuit Eye Movement

2009 ◽  
Vol 102 (6) ◽  
pp. 3225-3233 ◽  
Author(s):  
Leanne Chukoskie ◽  
J. Anthony Movshon
Keyword(s):  
Eye Movements ◽  
Eye Movement ◽  
Retinal Image ◽  
Visual Motion ◽  
Single Cells ◽  
Response Amplitude ◽  
Visual Signals ◽  
Image Motion ◽  
Visual Responses ◽  
Retinal Image Motion

Retinal image motion is produced with each eye movement, yet we usually do not perceive this self-produced “reafferent” motion, nor are motion judgments much impaired when the eyes move. To understand the neural mechanisms involved in processing reafferent motion and distinguishing it from the motion of objects in the world, we studied the visual responses of single cells in middle temporal (MT) and medial superior temporal (MST) areas during steady fixation and smooth-pursuit eye movements in awake, behaving macaques. We measured neuronal responses to random-dot patterns moving at different speeds in a stimulus window that moved with the pursuit target and the eyes. This allowed us to control retinal image motion at all eye velocities. We found the expected high proportion of cells selective for the direction of visual motion. Pursuit tracking changed both response amplitude and preferred retinal speed for some cells. The changes in preferred speed were on average weakly but systematically related to the speed of pursuit for area MST cells, as would be expected if the shifts in speed selectivity were compensating for reafferent input. In area MT, speed tuning did not change systematically during pursuit. Many cells in both areas also changed response amplitude during pursuit; the most common form of modulation was response suppression when pursuit was opposite in direction to the cell's preferred direction. These results suggest that some cells in area MST encode retinal image motion veridically during eye movements, whereas others in both MT and MST contribute to the suppression of visual responses to reafferent motion.

Pursuit and optokinetic deficits following chemical lesions of cortical areas MT and MST

1988 ◽  
Vol 60 (3) ◽  
pp. 940-965 ◽  
Author(s):  
M. R. Dursteler ◽  
R. H. Wurtz
Keyword(s):  
Eye Movements ◽  
Visual Field ◽  
Visual Motion ◽  
Ibotenic Acid ◽  
Motion Processing ◽  
Temporal Area ◽  
Target Speed ◽  
Pursuit Eye Movements ◽  
Medial Superior Temporal ◽  
Visual Motion Processing

1. Previous experiments have shown that punctate chemical lesions within the middle temporal area (MT) of the superior temporal sulcus (STS) produce deficits in the initiation and maintenance of pursuit eye movements (10, 34). The present experiments were designed to test the effect of such chemical lesions in an area within the STS to which MT projects, the medial superior temporal area (MST). 2. We injected ibotenic acid into localized regions of MST, and we observed two deficits in pursuit eye movements, a retinotopic deficit and a directional deficit. 3. The retinotopic deficit in pursuit initiation was characterized by the monkey's inability to match eye speed to target speed or to adjust the amplitude of the saccade made to acquire the target to compensate for target motion. This deficit was related to the initiation of pursuit to targets moving in any direction in the visual field contralateral to the side of the brain with the lesion. This deficit was similar to the deficit we found following damage to extrafoveal MT except that the affected area of the visual field frequently extended throughout the entire contralateral visual field tested. 4. The directional deficit in pursuit maintenance was characterized by a failure to match eye speed to target speed once the fovea had been brought near the moving target. This deficit occurred only when the target was moving toward the side of the lesion, regardless of whether the target began to move in the ipsilateral or contralateral visual field. There was no deficit in the amplitude of saccades made to acquire the target, or in the amplitude of the catch-up saccades made to compensate for the slowed pursuit. The directional deficit is similar to the one we described previously following chemical lesions of the foveal representation in the STS. 5. Retinotopic deficits resulted from any of our injections in MST. Directional deficits resulted from lesions limited to subregions within MST, particularly lesions that invaded the floor of the STS and the posterior bank of the STS just lateral to MT. Extensive damage to the densely myelinated area of the anterior bank or to the posterior parietal area on the dorsal lip of the anterior bank produced minimal directional deficits. 6. We conclude that damage to visual motion processing in MST underlies the retinotopic pursuit deficit just as it does in MT. MST appears to be a sequential step in visual motion processing that occurs before all of the visual motion information is transmitted to the brainstem areas related to pursuit.(ABSTRACT TRUNCATED AT 400 WORDS)

MST Neuronal Responses to Heading Direction During Pursuit Eye Movements

1999 ◽  
Vol 81 (2) ◽  
pp. 596-610 ◽  
Author(s):  
William K. Page ◽  
Charles J. Duffy
Keyword(s):  
Eye Movements ◽  
Smooth Pursuit ◽  
Visual Motion ◽  
Neuronal Population ◽  
Smooth Pursuit Eye Movements ◽  
Neuronal Responses ◽  
Temporal Area ◽  
Pursuit Eye Movements ◽  
Heading Direction ◽  
Monkey Visual Cortex

MST neuronal responses to heading direction during pursuit eye movements. As you move through the environment, you see a radial pattern of visual motion with a focus of expansion (FOE) that indicates your heading direction. When self-movement is combined with smooth pursuit eye movements, the turning of the eye distorts the retinal image of the FOE but somehow you still can perceive heading. We studied neurons in the medial superior temporal area (MST) of monkey visual cortex, recording responses to FOE stimuli presented during fixation and smooth pursuit eye movements. Almost all neurons showed significant changes in their FOE selective responses during pursuit eye movements. However, the vector average of all the neuronal responses indicated the direction of the FOE during both fixation and pursuit. Furthermore, the amplitude of the net vector increased with increasing FOE eccentricity. We conclude that neuronal population encoding in MST might contribute to pursuit-tolerant heading perception.

scholarly journals Functional Properties of Neurons in Macaque Area V3

1997 ◽  
Vol 77 (4) ◽  
pp. 1906-1923 ◽  
Author(s):  
Karl R. Gegenfurtner ◽  
Daniel C. Kiper ◽  
Jonathan B. Levitt
Keyword(s):  
Functional Properties ◽  
Visual Processing ◽  
Significant Proportion ◽  
Large Degree ◽  
Visual Signals ◽  
Motion Processing ◽  
Temporal Area ◽  
Area V2 ◽  
Layer 4 ◽  
Contrast Thresholds

Gegenfurtner, Karl R., Daniel C. Kiper, and Jonathan B. Levitt. Functional properties of neurons in macaque area V3. J. Neurophysiol. 77: 1906–1923, 1997. We investigated the functional properties of neurons in extrastriate area V3. V3 receives inputs from both magno- and parvocellular pathways and has prominent projections to both the middle temporal area (area MT) and V4. It may therefore represent an important site for integration and transformation of visual signals. We recorded the activity of single units representing the central 10° in anesthetized, paralyzed macaque monkeys. We measured each cell's spatial, temporal, chromatic, and motion properties with the use of a variety of stimuli. Results were compared with measurements made in V2 neurons at similar eccentricities. Similar to area V2, most of the neurons in our sample (80%) were orientation selective, and the distribution of orientation bandwidths was similar to that found in V2. Neurons in V3 preferred lower spatial and higher temporal frequencies than V2 neurons. Contrast thresholds of V3 neurons were extremely low. Achromatic contrast sensitivity was much higher than in V2, and similar to that found in MT. About 40% of all neurons showed strong directional selectivity. We did not find strongly directional cells in layer 4 of V3, the layer in which the bulk of V1 and V2 inputs terminate. This property seems to be developed within area V3. An analysis of the responses of directionally selective cells to plaid patterns showed that in area V3, as in MT and unlike in V1 and V2, there exist cells sensitive to the motion of the plaid pattern rather than to that of the components. The exact proportion of cells classified as being selective to color depended to a large degree on the experiment and on the criteria used for classification. With the use of the same conditions as in a previous study of V2 cells, we found as many (54%) color-selective cells as in V2 (50%). Furthermore, the responses of V3 cells to colored sinusoidal gratings were well described by a linear combination of cone inputs. The two subpopulations of cells responsive to color and to motion overlapped to a large extent, and we found a significant proportion of cells that gave reliable and directional responses to drifting isoluminant gratings. Our results show that there is a significant interaction between color and motion processing in area V3, and that V3 cells exhibit the more complex motion properties typically observed at later stages of visual processing.

Topographic and directional organization of visual motion inputs for the initiation of horizontal and vertical smooth-pursuit eye movements in monkeys

1989 ◽  
Vol 61 (1) ◽  
pp. 173-185 ◽  
Author(s):  
S. G. Lisberger ◽  
T. A. Pavelko
Keyword(s):  
Eye Movements ◽  
Visual Field ◽  
Visual Motion ◽  
Target Position ◽  
Target Motion ◽  
Visual Signals ◽  
Motion Processing ◽  
Pursuit Eye Movements ◽  
Visual Inputs ◽  
Initial Target

1. The goal of our study was to determine the properties of the visual inputs for pursuit eye movements. In a previous study we presented horizontal target motion along the horizontal meridian and showed that targets were more effective if they moved across the center of the visual field. We have now analyzed the topographic weighting of the inputs for pursuit in greater detail, using targets that moved in all directions and across a wide area of the visual field. 2. Monkeys were rewarded for tracking targets that started at 48 positions in the visual field. The initial positions were spaced equally around 4 circles that were centered at the position of fixation and had radii of 3, 6, 9, and 12 degrees. Targets moved horizontally or vertically at 30 degrees/s. We measured the smooth eye acceleration in the first 80 ms after the initiation of pursuit, before there had been time for visual feedback to affect the position or velocity of the retinal images from the target. 3. For both horizontal and vertical target motion, there were major differences between the early and late intervals in the first 80 ms of pursuit. In the first 20 ms eye acceleration was largely independent of initial target position. In later intervals eye acceleration decreased sharply as a function of initial target eccentricity. The later intervals also showed a pronounced toward/away asymmetry such that the initiation of pursuit was more vigorous for target motion toward than for motion away from the horizontal or vertical meridian. 4. Comparison of the topographic organization of the middle temporal visual area (MT) with our data on pursuit suggests that the topography of cortical maps is smoothed when the visual signals are transmitted to the pursuit system. For example, the superior visual hemifield is underrepresented in cortical motion processing areas, but target motion in the superior and inferior visual hemifields is equally effective for the initiation of pursuit. 5. We investigated the directional organization of the visual inputs for pursuit by presenting targets that started at 6 degrees eccentric and moved in 16 different directions. Horizontal target motion always evoked larger eye accelerations than did vertical target motion. Target motion in oblique directions evoked intermediate values of eye acceleration. 6. Our data show two classes of variation in pursuit performance. First, some subjects showed ideosyncratic variations that were restricted to one hemifield or one direction of target motion. We attribute these variations to differences among subjects in the physiology of visual pathways.(ABSTRACT TRUNCATED AT 400 WORDS)

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

1988 ◽  
Vol 60 (2) ◽  
pp. 580-603 ◽  
Author(s):  
H. Komatsu ◽  
R. H. Wurtz
Keyword(s):  
Eye Movements ◽  
Smooth Pursuit ◽  
Visual Motion ◽  
Receptive Fields ◽  
Motion Processing ◽  
Temporal Area ◽  
Pursuit Eye Movements ◽  
Visual Motion Processing ◽  
Visual Areas ◽  
Visual Properties

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.

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

1989 ◽  
Vol 62 (1) ◽  
pp. 31-47 ◽  
Author(s):  
H. Komatsu ◽  
R. H. Wurtz
Keyword(s):  
Eye Movements ◽  
Visual Motion ◽  
Directional Bias ◽  
Position Information ◽  
Temporal Area ◽  
Pursuit Eye Movements ◽  
Visual Motion Processing ◽  
Pursuit System ◽  
Pursuit Velocity ◽  
The Brain

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)

Distinct Nature of Directional Signals Among Parietal Cortical Areas During Visual Guidance

2002 ◽  
Vol 88 (4) ◽  
pp. 1777-1790 ◽  
Author(s):  
Emad N. Eskandar ◽  
John A. Assad
Keyword(s):  
Visual Motion ◽  
Hand Movement ◽  
Direction Selectivity ◽  
Visually Guided ◽  
Stimulus Motion ◽  
Directional Information ◽  
Temporal Area ◽  
Medial Superior Temporal ◽  
Direction Of Movement ◽  
Mst Neurons

We examined neuronal signals in the monkey medial superior temporal area (MST), the medial intraparietal area (MIP), and the lateral intraparietal area (LIP) during visually guided hand movements. Two animals were trained to use a joystick to guide a spot to a target. Many neurons responded in a direction-selective manner in this guidance task. We tested whether the direction selectivity depended on the direction of the stimulus spot or the direction of the hand movement. First, in some trials, the moving spot disappeared transiently. Second, the mapping between the hand direction and the spot direction was reversed on alternate blocks of trials. Third, we recorded the spot's movement while the animals moved the joystick and then played back that movement while the animals fixated without moving the joystick. Neurons in the three parietal areas conveyed distinct directional information. MST neurons were active and directional only on visible trials in both joystick-movement mode and playback mode and were not affected by the direction of hand movement. MIP neurons were mainly directional with respect to the hand movement, although some MIP neurons were also selective for stimulus direction. MIP neurons were much less active in playback mode. LIP neurons were active and directional in both joystick-movement mode and playback mode. Directional signals in LIP were unrelated to planning saccades. The selectivity of LIP neurons also became evident hundreds of milliseconds before the start of movement. Since the direction of movement was consistent throughout a block of trials, these signals could provide a prediction of the upcoming direction of motion. We tested this by alternating blocks of trials in which the direction was consistent or randomized. The direction selectivity developed earlier on trials in which the upcoming direction could be predicted. These results suggest that LIP neurons combine “bottom-up” visual motion signals with extraretinal, predictive signals about stimulus motion.

MST Neurons Code for Visual Motion in Space Independent of Pursuit Eye Movements

2007 ◽  
Vol 97 (5) ◽  
pp. 3473-3483 ◽  
Author(s):  
Naoko Inaba ◽  
Shigeru Shinomoto ◽  
Shigeru Yamane ◽  
Aya Takemura ◽  
Kenji Kawano
Keyword(s):  
Eye Movements ◽  
Eye Movement ◽  
Visual Motion ◽  
Neural Mechanism ◽  
Mt Neurons ◽  
Pursuit Eye Movements ◽  
Medial Superior Temporal ◽  
Counter Rotation ◽  
Induced Motion ◽  
Mst Neurons

When a person tracks a small moving object, the visual images in the background of the visual scene move across his/her retina. It, however, is possible to estimate the actual motion of the images despite the eye-movement-induced motion. To understand the neural mechanism that reconstructs a stable visual world independent of eye movements, we explored areas MT (middle temporal) and MST (medial superior temporal) in the monkey cortex, both of which are known to be essential for visual motion analysis. We recorded the responses of neurons to a moving textured image that appeared briefly on the screen while the monkeys were performing smooth pursuit or stationary fixation tasks. Although neurons in both areas exhibited significant responses to the motion of the textured image with directional selectivity, the responses of MST neurons were mostly correlated with the motion of the image on the screen independent of pursuit eye movement, whereas the responses of MT neurons were mostly correlated with the motion of the image on the retina. Thus these MST neurons were more likely than MT neurons to distinguish between external and self-induced motion. The results are consistent with the idea that MST neurons code for visual motion in the external world while compensating for the counter-rotation of retinal images due to pursuit eye movements.

scholarly journals Pursuit Speed Compensation in Cortical Area MSTd

2002 ◽  
Vol 88 (5) ◽  
pp. 2630-2647 ◽  
Author(s):  
Krishna V. Shenoy ◽  
James A. Crowell ◽  
Richard A. Andersen
Keyword(s):  
Eye Movements ◽  
Retinal Image ◽  
Visual Motion ◽  
Object Motion ◽  
Retinal Position ◽  
Temporal Area ◽  
Pursuit Eye Movements ◽  
Medial Superior Temporal ◽  
The Difference ◽  
Pursuit Velocity

When we move forward the visual images on our retinas expand. Humans rely on the focus, or center, of this expansion to estimate their direction of self-motion or heading and, as long as the eyes are still, the retinal focus corresponds to the heading. However, smooth pursuit eye movements add visual motion to the expanding retinal image and displace the focus of expansion. In spite of this, humans accurately judge their heading during pursuit eye movements even though the retinal focus no longer corresponds to the heading. Recent studies in macaque suggest that correction for pursuit may occur in the dorsal aspect of the medial superior temporal area (MSTd); neurons in this area are tuned to the retinal position of the focus and they modify their tuning to partially compensate for the focus shift caused by pursuit. However, the question remains whether these neurons shift focus tuning more at faster pursuit speeds, to compensate for the larger focus shifts created by faster pursuit. To investigate this question, we recorded from 40 MSTd neurons while monkeys made pursuit eye movements at a range of speeds across simulated self- or object motion displays. We found that most MSTd neurons modify their focus tuning more at faster pursuit speeds, consistent with the idea that they encode heading and other motion parameters regardless of pursuit speed. Across the population, the median rate of compensation increase with pursuit speed was 51% as great as required for perfect compensation. We recorded from the same neurons in a simulated pursuit condition, in which gaze was fixed but the entire display counter-rotated to produce the same retinal image as during real pursuit. This condition yielded the result that retinal cues contribute to pursuit compensation; the rate of compensation increase was 30% of that required for accurate encoding of heading. The difference between these two conditions was significant ( P < 0.05), indicating that extraretinal cues also contribute significantly. We found a systematic antialignment between preferred pursuit and preferred visual motion directions. Neurons may use this antialignment to combine retinal and extraretinal compensatory cues. These results indicate that many MSTd neurons compensate for pursuit velocity, pursuit direction as previously reported and pursuit speed, and further implicate MSTd as a critical stage in the computation of egomotion.

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