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D B Bender

Publications and source records attributed to D B Bender.

At least 19 recordsLinked to original sources

Effect of corticotectal tract lesions on relative motion selectivity in the monkey superior colliculus.

Many cells in the superficial layers of the monkey superior colliculus are sensitive to the relative motion between a small target moving through the classic receptive field and a textured, moving background pattern that fills the visual field beyond the classic receptive field. The cells respond well when motion of the target differs from that of the background, but their responses are suppressed when the target moves in phase with the background. To determine whether this relative motion sensitivity depends on input to the colliculus from visual cortex, we studied colliculus cells in immobilized, anesthetized monkeys after unilateral thermocoagulation, or anesthetic blockade, of the corticotectal tract at the level of the pulvinar. In the colliculus ipsilateral to the corticotectal tract lesions, relative motion sensitivity was significantly reduced when compared either with the colliculus in intact animals or with the colliculus contralateral to the lesion. However, a moving-background stimulus still had a modest suppressive effect compared with a stationary background ("background motion sensitivity"), as is the case for intact animals. Anesthetic blockade of the corticotectal tract had similar effects; relative motion sensitivity, but not background motion sensitivity, was lost following injection of mepivacaine or bupivacaine. Pulvinar cell loss alone, induced by kainic acid injection, had no effect on relative motion sensitivity in the colliculus. The corticotectal tract lesions, but not the anesthetic injections, also had minor effects on flash-evoked responses and spontaneous discharge rates; these effects may reflect a retrograde response of some tectopulvinar cells to injury of their axons by the corticotectal tract lesions. In the colliculus opposite the corticotectal tract lesion, relative motion sensitivity was similar to that in normal animals. However, responses in the presence of a moving background were enhanced, suggesting that removal of cortical input to one colliculus may disinhibit the contralateral colliculus, a phenomenon reminiscent of the Sprague effect in the cat. We conclude that while cortical input to the colliculus may contribute little to the classic receptive field properties of superficial-layer cells, it clearly does contribute to relative motion sensitivity.

Anesthetics, Local

Bilateral projections from the parabigeminal nucleus to the superior colliculus in monkey.

We examined the distribution of labeled neurons in the parabigeminal nucleus of the monkey following injections of retrograde fluorescent tracers into the superior colliculus. The extent of the visual field representation included in the injection site was assessed from the location of labeled cells in striate cortex. The results suggest a rough topographic organization of the parabigeminal nucleus, with the lower quadrant represented anteriorly and the upper quadrant posteriorly. We also found bilateral projections from the parabigeminal nucleus to both superior colliculi, but the crossed projection appeared to terminate only in that part of the colliculus where the vertical meridian is represented. Parabigeminal cells with a crossed projection were larger than those projecting to the ipsilateral colliculus. The results suggest that the organization of the monkey's parabigemino-tectal system is fundamentally similar to that of many other vertebrates.

Amidines

Selectivity for relative motion in the monkey superior colliculus.

1. Cells in the superficial layers of the colliculus were studied in immobilized monkeys anesthetized with nitrous oxide. We examined sensitivity to the relative motion between two stimuli: a small target in a cell's receptive field and a large random-dot background pattern that filled most of the visual field outside the receptive field. 2. Most cells were nonselective for either target direction or speed when the background pattern was stationary but were selective for both direction and speed relative to a moving background. Selectivity for relative motion was independent of the absolute direction and speed of both target and background. When both moved at the same speed in the same direction, the response evoked by the target was strongly suppressed. Changing the background direction relative to the target reduced the suppression; suppression was minimal when the two moved in opposite directions. Selectivity for relative direction was broad: the average tuning width at half-amplitude was 136 degrees. When target and background moved in the same direction, increasing or decreasing background speed relative to the target likewise reduced suppression. Average tuning width for relative speed was 1.4 log units. 3. Selectivity for relative motion was a global phenomenon. Suppression was present even when the background pattern was excluded from a region 10 times the receptive-field diameter. However, suppression gradually diminished with increasing distance between the receptive field and the background pattern. 4. Relative motion selectivity was most common in the deeper part of the superficial layers. Ninety percent of the cells below the middle of the stratum griseum superficiale were selective for relative direction, whereas above this level only 45% of the cells were. 5. Cells in the magnocellular and parvocellular layers of the lateral geniculate nucleus did not show selectivity for relative direction. 6. We suggest that the lower one-half of the superficial grey layer and the stratum opticum together constitute a subdivision of the superior colliculus that is specialized to detect strong discontinuities in relative motion. Descending input by way of the corticotectal tract is probably essential for the detection process. the projections from this tectal motion zone to the pulvinar, and from there to prestriate cortex, may provide a feedback pathway through which motion discontinuities such as occur at dynamic occlusion boundaries can influence local feature detection by cortical neurons.

Animals

Saccadic eye movements following kainic acid lesions of the pulvinar in monkeys.

Behavioral and anatomical experiments have suggested that the pulvinar might play a role in the generation of saccadic eye movements to visual targets. To test this idea, we trained monkeys to make visually-guided saccades by requiring them to detect the dimming of a small target. We used three different saccade paradigms. On single-step trials, saccades were made from a central fixation point (FP) to a target at 12, 24 or 36 degrees to the left or right. On overlap trials, the FP remained lit during presentation of a target at 12 or 24 degrees. On double-step trials, the target stepped first to 24 degrees, and then back to 12 degrees on the same side. Animals were trained to criterion, received kainic acid lesions of the pulvinar, and were retested on all three tasks. The lesions were very large, destroying almost all of the visually responsive pulvinar. They also encroached on the lateral geniculate nucleus, thereby producing small foveal scotomas, and this resulted in some behavioral changes, including difficulty in maintaining fixation on the target and in detecting its dimming. Results on the saccade tests suggest that the pulvinar is not crucial for initiation of saccadic eye movements. Saccade latency and amplitude were unimpaired on both single-step and overlap trials. Saccadic performance was also normal on double-step trials. In a second experiment, we measured the average length of fixations during spontaneous viewing of a complex visual scene. Fixation lengths did not differ from those of unoperated control monkeys. We suggest that the neglect, increased saccadic latencies, and prolonged fixations attributed to pulvinar damage in previous studies were probably the result instead of inadvertent damage to tectal afferents. The present results, together with single unit data, point to a role for the pulvinar not in the generation of saccades, but rather in the integration of saccadic eye movements with visual processing.

Animals

Comparison of saccadic eye movements in humans and macaques to single-step and double-step target movements.

Human and monkey saccade amplitude and latency, in response to 12-36 degrees target steps, differed substantially despite nearly identical experimental conditions. On single-step trials, monkeys did not undershoot targets, and latencies were insensitive to stimulus and contextual factors. Human saccades did undershoot, their latency was longer, and both undershoot and latency were affected by stimulus variables and experimental context. On double-step trials, the second target step altered primary saccade amplitude when the step occurred as little as 40 msec prior to saccade onset for both humans and monkeys. However, humans and monkeys showed somewhat different amplitude transition functions, and monkeys showed little evidence of parallel programming.

Adult

Comparison of the effects of superior colliculus and pulvinar lesions on visual search and tachistoscopic pattern discrimination in monkeys.

In order to investigate whether pulvinar lesions produce behavioral impairments similar to those that follow superior colliculus lesions, monkeys were tested on a visual search task before and after receiving radiofrequency lesions of either the superior colliculus or pulvinar. The animals searched for a small target pattern within an array of varying numbers of irrelevant patterns. After receiving colliculus lesions, the animals showed marked post-operative increases in either search time, percent errors, or both. By contrast, pulvinar lesions had little or no effect on visual search performance. Similarly, in learning to search for a target they had not previously seen, animals with colliculus lesions were impaired relative to unoperated controls, whereas pulvinar-lesioned animals did not differ from controls. In an attempt to confirm the finding that pulvinar lesions impair tachistoscopic pattern discrimination, we determined exposure-duration thresholds of pulvinar- and colliculus-lesioned monkeys for performance of a pattern discrimination. The thresholds of the colliculus-lesioned monkeys were elevated 20-fold relative to controls. By contrast, thresholds of the pulvinar-lesioned monkeys were normal. We conclude that the pulvinar is not critical for the attentional processes in which the superior colliculus participates.

Animals

Global visual processing in the monkey superior colliculus.

Neurons were recorded in the superficial layers of the superior colliculus in anesthetized monkeys. As classically described, cells were non-selective for target direction and speed when the target moved through an empty visual field. However, these same cells were sensitive to target direction and speed relative to a textured moving background. The target's response was suppressed when its direction and speed were similar to that of the background, irrespective of the absolute direction of background movement.

Animals

Anterograde degeneration in the superior colliculus following kainic acid and radiofrequency lesions of the macaque pulvinar.

Several studies have reported behavioral deficits following thermocoagulation of the primate pulvinar. However, these deficits may have resulted from damage to corticotectal fibers as they pass through the pulvinar. To evaluate this possibility and to determine whether kainic acid can be used to destroy pulvinar cells without damaging corticotectal fibers, we compared anterograde degeneration in the superior colliculus following kainic acid and radiofrequency lesions of the pulvinar. Kainic acid injections into the pulvinar produced total loss of neuronal perikarya within the inferior and lateral pulvinar. Four to 7 days following the kainic acid lesions, terminal and fiber degeneration within the superior colliculus was no greater than that produced by control injections of saline. By contrast, thermocoagulation lesions of the inferior and lateral pulvinar produced dense fiber and terminal degeneration throughout the superficial and intermediate layers of the superior colliculus. We conclude that whereas thermocoagulation of the pulvinar severely damages the corticotectal tract, kainic acid lesions spare these fibers of passage. Thus kainic acid lesions should provide an effective tool for studying the functional significance of the pulvinar.

Animals

Effects of kainic acid and radiofrequency lesions of the pulvinar on visual discrimination in the monkey.

Monkeys with thermocoagulation or kainic acid lesions of the pulvinar and unoperated control monkeys were tested in two tasks: pattern discrimination retention and color discrimination learning in which the stimuli were located at the response sites or were separated spatially from them (S-R separation). The monkeys with kainic acid pulvinar lesions were mildly impaired in retention of the pattern discrimination, but were unimpaired in the color discrimination tasks with or without the S-R separation. The monkeys with thermocoagulation lesions, like monkeys with superior colliculus lesions in a prior study, were severely impaired in performing one of the color discrimination tasks with S-R separation. These findings suggest that: (a) the inferior pulvinar, unlike the superior colliculus, does not contribute to the performance of discriminations involving S-R separation; and (b) corticotectal projections traversing the pulvinar and destroyed by the thermocoagulation lesions are crucial to the performance of discriminations involving S-R separation. The results of an earlier experiment also suggested that interruption of corticotectal fibers passing through the pulvinar impairs performance in another task sensitive to superior colliculus lesions--spatial localization of light flashes. Thus, corticotectal projections may be crucial for the contribution of the colliculus to performance in a variety of visual tasks.

Animals

Visual activation of neurons in the primate pulvinar depends on cortex but not colliculus.

Superior colliculus lesions had little effect on the visual response of neurons in the monkey inferior pulvinar. By contrast, striate cortex lesions eliminated the visual response of all inferior pulvinar neurons for a period of 3 weeks after the lesion. At longer survival times, a few pulvinar neurons responded to small light spots, but sensitivity to orientation and direction of movement never returned. Thus striate cortex, rather than the colliculus, appears to be responsible for the visual properties of pulvinar cells.

Animals

Localization and detection of visual stimuli in monkeys with pulvinar lesions.

Since the pulvinar receives a major ascending projection of the superior colliculus, pulvinar lesions might produce behavioral impairments resembling those that follow colliculus lesions. To test this possibility, we examined the effect of pulvinar lesions in monkeys on the localization and detection of brief light flashes, a task in which monkeys with colliculus lesions are severely impaired. Some of the pulvinar-lesioned monkeys showed localization impairments similar to those in monkeys with colliculus lesions. However, histological analyses of the lesions suggested that these deficits were related not to the pulvinar damage per se, but rather to interruption of corticotectal fibers that pass through the pulvinar. We conclude that the pulvinar is not critical for the ability to locate and detect brief visual stimuli.

Animals

Localization and detection of visual stimuli following superior colliculus lesions in rhesus monkeys.

Rhesus monkeys were trained to fixate a central stimulus and to detect and localize a 50 msec light flash presented 6-80 degrees on either side of the central stimulus. Following large lesions of the superior colliculus, they showed persistent deficits in localizing flashes presented 43-80 degrees from the fixation stimulus. However, they were not consistently impaired when the flashes were presented more centrally, and their performance with peripheral stimuli improved when the stimulus duration was 1 sec. Thus, the superior colliculus appears to be necessary for the localization of brief visual stimuli in the far periphery.

Animals

Contributions of the corpus callosum and the anterior commissure to visual activation of inferior temporal neurons.

Most neurons in the inferior temporal cortex of the rhesus monkeys have visual receptive fields that extend across the vertical meridian well into both the contralateral and ipsilateral visual half-fields. We examined the role of different portions of the forebrain commissures in providing the ipsilateral input with the following results. (1) Combined section of the splenium and anterior commissure eliminated visual activation from the ipsilateral visual half-field. (2) Section of the splenium, with sparing of the anterior commissure, reduced the incidence of ipsilateral activation by about one-half. (3) Section of the anterior commissure, with sparing of the splenium, did not alter the incidence of ipsilateral activation. (4) Section of the non-splenial portions of the corpus callosum had no effect on the laterality of the receptive fields. Thus, both the splenium and the anterior commissure but not the non-splenial callosum can provide information from the ipsilateral visual field to neurons in inferior temporal cortex. These results are interpreted as suggesting that the converging input from the two visual half-fields onto single inferior temporal neurons provided by the forebrain commissures may mediate interhemispheric transfer of visual habits.

Animals

A visual deficit after superior colliculus lesions in monkeys.

The superior colliculus has been implicated in visual orientation and localization in several mammalian species. To examine this possibility in rhesus monkeys, we trained monkeys to respond to one of six locations signaled by a small light of 5 s duration that was turned off immediately before they were allowed to respond. The locations were directly behind the light. After tectal lesions they continued to perform this task normally, but showed a severe deficit when the duration of the signal was reduced to 1 s. However, when the stimulus remained on throughout the trial and the site of the stimulus and response were further separated, performance of the animals with tectal lesions was normal. We suggest that this deficit in the accuracy of reaching guided by a transient visual stimulus may be related to the role of the superior colliculus in the control of eye movements.

Animals