Organization of monkey superior colliculus: enhanced visual response of superficial layer cells

1976 ◽  
Vol 39 (4) ◽  
pp. 745-765 ◽  
Author(s):  
R. H. Wurtz ◽  
C. W. Mohler

1. Cells in the superficial layers of monkey superior colliculus respond more vigorously to a spot of light falling in their receptive fields when the monkey uses that spot of light as the target for a saccadic eye movement. Our purpose in these experiments was to investigate the characteristics of this enhancement effect. While monkeys fixated, we determined the response of a cell to a stimulus falling in its receptive field. Then we determined the response of the cell to the same stimulus when the monkey made a saccade to the stimulus or near to it. 2. The enhancement of the visual response is spatially limited. The receptive field of a cell always shows enhancement throughout its extent and frequently shows a slight expansion. Saccades made near to a stimulus in the visual receptive field, but not to it, also lead to an enhancement of that visual stimulus; an area around the excitatory center of the receptive field where such enhancement occurs was referred to as the enhancement field of the cell. An enhanced response in one part of the visual field was not accompanied by depressed responses associated with saccades to other parts of the visual field. 3. The enhancement effect is temporally limited; it begins 200-300 ms before the eye movement, as determined by the increasing response to 50-ms light pulses presented at varying intervals before the eye movement. The degree of enhancement intensifies when the visual stimulus is turned on closer in time to the onset of the saccade. A buildup of the enhancement also occurs on successive trials as does the response of eye movement-related cells in the intermediate layers. 4. The enhancement response is not present in the upper quarter-millimeter of the superficial layers, suggesting that the effect is not present in retinal afferents which terminate primarily in this area of the superficial layers. The enhancement effect is seen throughout the visual field; the foveal area was not tested. 5. In order to determine the relation of the enhancement effect to the monkey's behavioral response, we required the monkey to make a hand response rather than an eye movement-response to the visual stimuli. Cells did not show a clear enhancement with such a hand response. Results of these experiments indicate that the enhancement effect is dependent on the type of response the monkey makes to the stimulus and is probably specifically related to eye movements. Since the enhancement of visual response seems likely to be related specifically to eye movements both on physiological and behavioral grounds, the response-free term "attention" is probably inappropriate for the phenomenon. 6. The hypothesis advanced in the preceding paper that eye movement-related activity from intermediate and deep colliculus layers is directed upward to converge with visually related activity in the superficial layers is extended to include an input from cells in these deeper layers (or their afferents) to the superficial layer cells...

1995 ◽  
Vol 73 (5) ◽  
pp. 1988-2003 ◽  
Author(s):  
M. F. Walker ◽  
E. J. Fitzgibbon ◽  
M. E. Goldberg

1. Previous experiments have shown that visual neurons in the lateral intraparietal area (LIP) respond predictively to stimuli outside their classical receptive fields when an impending saccade will bring those stimuli into their receptive fields. Because LIP projects strongly to the intermediate layers of the superior colliculus, we sought to demonstrate similar predictive responses in the monkey colliculus. 2. We studied the behavior of 90 visually responsive neurons in the superficial and intermediate layers of the superior colliculus of two rhesus monkeys (Macaca mulatta) when visual stimuli or the locations of remembered stimuli were brought into their receptive fields by a saccade. 3. Thirty percent (18/60) of intermediate layer visuomovement cells responded predictively before a saccade outside the movement field of the neuron when that saccade would bring the location of a stimulus into the receptive field. Each of these neurons did not respond to the stimulus unless an eye movement brought it into its receptive field, nor did it discharge in association with the eye movement unless it brought a stimulus into its receptive field. 4. These neurons were located in the deeper parts of the intermediate layers and had relatively larger receptive fields and movement fields than the cells at the top of the intermediate layers. 5. The predictive responses of most of these neurons (16/18, 89%) did not require that the stimulus be relevant to the monkey's rewarded behavior. However, for some neurons the predictive response was enhanced when the stimulus was the target of a subsequent saccade into the neuron's movement field. 6. Most neurons with predictive responses responded with a similar magnitude and latency to a continuous stimulus that remained on after the saccade, and to the same stimulus when it was only flashed for 50 ms coincident with the onset of the saccade target and thus never appeared within the cell's classical receptive field. 7. The visual response of neurons in the intermediate layers of the colliculus is suppressed during the saccade itself. Neurons that showed predictive responses began to discharge before the saccade, were suppressed during the saccade, and usually resumed discharging after the saccade. 8. Three neurons in the intermediate layers responded tonically from stimulus appearance to saccade without a presaccadic burst. These neurons responded predictively to a stimulus that was going to be the target for a second saccade, but not to an irrelevant flashed stimulus. 9. No superficial layer neuron (0/27) responded predictively when a stimulus would not be brought into their receptive fields by a saccade.(ABSTRACT TRUNCATED AT 400 WORDS)


1989 ◽  
Vol 61 (5) ◽  
pp. 1064-1084 ◽  
Author(s):  
R. A. Boch ◽  
M. E. Goldberg

1. We recorded from 257 neurons in the banks of the posterior third of the principal sulcus of two rhesus monkeys trained to look at a fixation point and make saccades to stimuli in the visual periphery. Sixty-six percent (220/257) discharged or were suppressed in association with one or more aspects of the tasks we used. 2. Fifty-eight percent (151/257) of the neurons responded to the appearance of a spot of light in some part of the contralateral visual field. Cells did not seem to have absolute requirements for stimulus shape, size, or direction of motion. 3. Thirty-six percent (29/79) of visually responsive neurons tested quantitatively gave an enhanced response to the stimulus in the receptive field when the monkey had to make a saccade to the stimulus when its appearance was synchronous with the disappearance of the fixation point (synchron task). Twenty-nine percent (19/57) of the neurons gave an enhanced response to the stimulus when the monkey had to make a saccade to the stimulus some time after it appeared (delayed-saccade task). In general, enhancement in the synchron task correlated well with enhancement in the delayed-saccade task. 4. Enhancement was spatially specific. It did not occur when the monkey made a saccade to a stimulus outside the receptive field even though there was a stimulus within the receptive field. 5. Twenty-three percent (27/117) of neurons studied in the delayed-saccade task gave two bursts, one at the appearance of the stimulus and a second one around the saccade. This second burst generally did not occur when the monkey made the same saccade to a remembered target, but instead required the presence of the visual stimulus, and so we describe it as a reactivation of the visual response. Reactivation was also spatially specific. 6. The latency from reactivation to the beginning of the saccade ranged from 160 ms before the saccade to the beginning of the saccade. Reactivation usually continued for several hundred milliseconds after the saccade, sometimes for the duration of the trial. 7. Reactivation and enhancement are not the same mechanism. Although some cells showed both phenomena there was no correlation between enhancement and reactivation. 8. Cells that showed reactivation in the saccade task also showed reactivation at a weaker level in a suppressed-saccade task. In this task the monkeys had to hold fixation despite the disappearance of the fixation point and the continued presence of the peripheral stimulus.(ABSTRACT TRUNCATED AT 400 WORDS)


1980 ◽  
Vol 50 (2) ◽  
pp. 631-636
Author(s):  
Evans Mandes

Post-exposural eye movements were studied in 32 adults and 24 7-yr.-old children. Stimuli were binary figures exposed tachistoscopically in both visual fields simultaneously. The data showed significant correlations between direction of eye movement and locus of recognition for both children and adults. No significant differences were found in frequencies of eye movements of children and adults. The data are interpreted in terms of the facilitative effects of post-exposural eye movements upon perception for both groups.


1976 ◽  
Vol 39 (4) ◽  
pp. 722-744 ◽  
Author(s):  
C. W. Mohler ◽  
R. H. Wurtz

1. We investigated the characteristics of cells in the intermediate layers of the superior colliculus that increase their rate of discharge before saccadic eye movements. Eye movements were repeatedly elicited by training rhesus monkeys to fixate on a spot of light and to make saccades to other spots of light when the fixation spot was turned off. 2. The eye movement cells showed consistent variations with their depth within the colliculus. The onset of the cell discharge led the eye movement by less time and the duration of the discharge was shorter as the cell was located closer to the dorsal edge of the intermediate layers. The movements fields (that area of the visual field where a saccade into the area is preceded by a burst of cell discharges) of each successive cell also became smaller as the cells were located more dorsally. The profile of peak discharge frequency remained fairly flat throughout the movement field of the cells regardless of depth of the cell within the colliculus. 3. A new type of eye movement-related cell has been found which usually lies at the border between the superficial and intermediate layers. This cell type, the visually triggered movement cell, increased its rate of discharge before saccades made to a visual stimulus but not before spontaneous saccades of equal amplitude made in the light or the dark. A vigorous discharge of these cells before an eye movement was dependent on the presence of a visual target; the cells seemed to combine the visual input of superficial layer cells and the movement-related input of the intermediate layer cells. The size of the movement fields of these cells were about the same size as the visual fields of superficial layer cells just above them...


2005 ◽  
Vol 94 (4) ◽  
pp. 2491-2503 ◽  
Author(s):  
Keith A. Schneider ◽  
Sabine Kastner

The superior colliculus (SC) is a multimodal laminar structure located on the roof of the brain stem. The SC is a key structure in a distributed network of areas that mediate saccadic eye movements and shifts of attention across the visual field and has been extensively studied in nonhuman primates. In humans, it has proven difficult to study the SC with functional MRI (fMRI) because of its small size, deep location, and proximity to pulsating vascular structures. Here, we performed a series of high-resolution fMRI studies at 3 T to investigate basic visual response properties of the SC. The retinotopic organization of the SC was determined using the traveling wave method with flickering checkerboard stimuli presented at different polar angles and eccentricities. SC activations were confined to stimulation of the contralateral hemifield. Although a detailed retinotopic map was not observed, across subjects, the upper and lower visual fields were represented medially and laterally, respectively. Responses were dominantly evoked by stimuli presented along the horizontal meridian of the visual field. We also measured the sensitivity of the SC to luminance contrast, which has not been previously reported in primates. SC responses were nearly saturated by low contrast stimuli and showed only small response modulation with higher contrast stimuli, indicating high sensitivity to stimulus contrast. Responsiveness to stimulus motion in the SC was shown by robust activations evoked by moving versus static dot stimuli that could not be attributed to eye movements. The responses to contrast and motion stimuli were compared with those in the human lateral geniculate nucleus. Our results provide first insights into basic visual responses of the human SC and show the feasibility of studying subcortical structures using high-resolution fMRI.


1996 ◽  
Vol 76 (5) ◽  
pp. 2841-2852 ◽  
Author(s):  
C. L. Colby ◽  
J. R. Duhamel ◽  
M. E. Goldberg

1. Posterior parietal cortex contains neurons that are visually responsive and active in relation to saccadic eye movements. We recorded from single neurons in a subregion of parietal cortex, the lateral intraparietal area (LIP), in alert rhesus monkeys. To characterize more completely the circumstances under which LIP neurons are responsive, we used five tasks designed to test the impact of sensory, motor, and cognitive factors. We obtained quantitative data in multiple tasks in 91 neurons. We measured neural activity during central fixation and in relation to stimulus onset and saccade onset. 2. LIP neurons have visual responses to the onset of a stationary stimulus in the receptive field. These visual responses occurred both in tasks that require a subsequent eye movement toward the stimulus and in tasks in which eye movements are not permitted, indicating that this activity is sensory rather than presaccadic. 3. Visual responses were enhanced when the monkey had to use information provided by the stimulus to guide its behavior. The amplitude of the sensory response to a given stimulus was increased in a task in which the monkey would subsequently make a saccade to the location signaled by the stimulus, as compared with the amplitude of the visual response in a simple fixation task. 4. The visual response was also enhanced when the monkey attended to the stimulus without looking at it. This result shows that enhancement does not reflect saccade preparation because the response is enhanced even when the monkey is not permitted to make a saccade. Instead, enhancement reflects the allocation of attention to the spatial locus of the receptive field. 5. Many LIP neurons had saccade-related activity in addition to their visual responses. The visual response for most neurons was stronger than the saccade-related activation. 6. Saccade-related activity was independent of visual activity. Similar presaccadic activity was observed in trials that included a recent visual stimulus (memory-guided saccade task) and in trials with no visual stimulus (learned saccade task). 7. We observed increases in activity during fixation in tasks in which the monkey could anticipate the onset of a behaviorally significant stimulus. LIP neurons usually showed low levels of background firing in the fixation task during the period before stimulus onset. This background activity was increased in the peripheral attention and memory-guided saccade tasks during the period when the monkey was waiting for a behaviorally relevant stimulus to appear. 8. The results from these several tasks indicate that LIP neurons are activated in a variety of circumstances and are not involved exclusively in sensory processing or motor planning. The modulation of sensory responses by attention and anticipation suggests that cognitive factors play a major role in parietal function.


2020 ◽  
Vol 10 (1) ◽  
Author(s):  
Lukasz Grzeczkowski ◽  
Heiner Deubel ◽  
Martin Szinte

Abstract Across saccadic eye movements, the visual system receives two successive static images corresponding to the pre- and the postsaccadic projections of the visual field on the retina. The existence of a mechanism integrating the content of these images is today still a matter of debate. Here, we studied the transfer of a visual feature across saccades using a blanking paradigm. Participants moved their eyes to a peripheral grating and discriminated a change in its orientation occurring during the eye movement. The grating was either constantly on the screen or briefly blanked during and after the saccade. Moreover, it either was of the same luminance as the background (i.e., isoluminant) or anisoluminant with respect to it. We found that for anisoluminant gratings, the orientation discrimination across saccades was improved when a blank followed the onset of the eye movement. Such effect was however abolished with isoluminant gratings. Additionally, performance was also improved when an anisoluminant grating presented before the saccade was followed by an isoluminant one. These results demonstrate that a detailed representation of the presaccadic image was transferred across saccades allowing participants to perform better on the transsaccadic orientation task. While such a transfer of visual orientation across saccade is masked in real-life anisoluminant conditions, the use of a blank and of an isoluminant postsaccadic grating allowed to reveal its existence.


1977 ◽  
Vol 40 (1) ◽  
pp. 156-173 ◽  
Author(s):  
M. Schlag-Rey ◽  
J. Schlag

1. Visual responses and eye movement-related activities were studied in single neurons of the thalamic internal medullary lamina (IML) of alert cats. The animals faced a tangent screen on which stationary or moving spots of light were presented. Of 95 units, 26% discharged in relation to photic stimuli but not eye movement, 6% in relation to eye movement but not photic stimuli, and 68% in relation to both. These units were intermixed in the same region. 2. Visual responses varied from transient to sustained. IML units were not found particularly sensitive to stimulus movement when the eyes were fixed. Strong and consistent responses could be elicited by extremely dim and weakly contrasted stationary stimuli (e.g.) 3.4 mcd/m2, 2.6% of illumination background) binocularly viewed. Receptive fields (from 250 to 800 deg2) were determined, in absence of eye movements, by computing the position of effective stimuli relative to the point of fixation of the gaze. An area of greatest responsiveness in the receptive field of most units could be detected on the basis of either higher probability of response, minimum latency, greater number of spikes in initial transient burst, or stronger sustained activity. Whole fields or their areas of greatest responsiveness were located on the side toward which saccades were accompanied by increased firing of the unit. 3. On trials in which a delay occurred between stimulus presentation and the cat's targeting saccade, the majority of the units studied changed their activity twice: after the stimulus and before the eye movement. In 16 units, the presaccadic activation occurred only with targeting, not with spontaneous saccades. 4. These results suggest that cells in the IML region of the cat play a significant role in the control of visually elicited eye movements. The resemblance of these cells to the monkey's tectual cells is discussed and hypotheses are proposed a) to relate the receptive field characteristics to the targeting operation, and b) to account for the double activation--sensory and motor--of many IML cells.


1988 ◽  
Vol 60 (2) ◽  
pp. 604-620 ◽  
Author(s):  
W. T. Newsome ◽  
R. H. Wurtz ◽  
H. Komatsu

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)


1991 ◽  
Vol 65 (6) ◽  
pp. 1464-1483 ◽  
Author(s):  
S. Funahashi ◽  
C. J. Bruce ◽  
P. S. Goldman-Rakic

1. Single-neuron activity was recorded from the prefrontal cortex of monkeys performing saccadic eye movements in oculomotor delayed-response (ODR) and visually guided saccade (VGS) tasks. In the ODR task the monkey was required to maintain fixation of a central spot throughout the 0.5-s cue and 3.0-s delay before making a saccadic eye movement in the dark to one of four or eight locations where the visual cue had been presented. The same locations were used for targets in the VGS tasks; however, unlike the ODR task, saccades in the VGS tasks were visually guided. 2. Among 434 neurons recorded from prefrontal cortex within and surrounding the principal sulcus (PS), 147 changed their discharge rates in relation to saccadic eye movements in the ODR task. Their response latencies relative to saccade initiation were distributed between -192 and 460-ms, with 22% exhibiting presaccadic activity and 78% exhibiting only postsaccadic activity. Among PS neurons with presaccadic activity, 53% also had postsaccadic activity when the monkey made saccadic eye movements opposite to the directions for which the presaccadic activity was observed. 3. Almost all (97%) PS neurons with presaccadic activity were directionally selective. The best direction and tuning specificity of each neuron were estimated from parameters used to fit a Gaussian tuning curve function. The best direction for 62% of the neurons with presaccadic activity was toward the contralateral visual field, with the remaining neurons having best directions toward the ipsilateral field (23%) or along the vertical meridian (15%). 4. Most postsaccadic activity of PS neurons (92%) was also directionally selective. The best direction for 48% of these neurons was toward the contralateral visual field, with the remaining neurons having best directions toward the ipsilateral field (36%) or along the vertical meridian (16%). Eighteen percent of the neurons with postsaccadic activity showed a reciprocal response pattern: excitatory responses occurred for one set of saccade directions, whereas inhibitory responses occurred for roughly the opposite set of directions. 5. Sixty PS neurons with saccade-related activity in the ODR task were also examined in a VGS task. Forty of these neurons showed highly similar profiles of directional specificity and response magnitude in both tasks, 13 showed saccade-related activity only in the ODR task, and 7 changed their response characteristics between the ODR and VGS tasks.(ABSTRACT TRUNCATED AT 400 WORDS)


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