Illusory, motion, and luminance-defined contours interact in the human visual system.

Psychophysical studies of interactions between contours defined by different image attributes report that luminance-defined and illusory contours show little if any interaction. Because the contours defined by these attributes may vary in perceptual saliency, we employed the tilt aftereffect (TAE) and a cross-adaptation procedure to evaluate interaction effects between luminance-defined and illusory contours under varying saliency conditions as well as to explore the interaction between illusory and motion-defined contours. When contour salience of the adaptation or test stimuli was modified by the addition of various amounts of static noise, we observed a TAE for all combinations of contour types including the novel motion-illusory and illusory-motion pairs. The interactions demonstrated between the contour classes in this as well as other studies suggests contour invariance in the orientation domain.

Binocular immobilization induced by paralysis of the extraocular muscles of one eye: evidence for an interocular proprioceptive mechanism.

1. Spontaneous changes in the positions of both eyes were measured in anesthetized cats before and after infusion of paralytic or anesthetic agents directly into the extraocular muscle capsule behind one eye. 2. Before retrobulbar injection, large changes in eye position were observed in both eyes. The position changes of each eye were independent of each other and often in opposite directions. 3. Application of a paralytic agent into the orbit of one eye reduced the spontaneous position changes in that eye to a very low level. Unexpectedly, the changes in the position of the other, untreated eye were also greatly reduced. 4. Control experiments demonstrated that the binocular immobilization observed after monocular paralysis was not due to 1) leakage of the paralytic agent from the orbit of the treated eye to the orbit of the untreated eye or 2) systemic vascular transport of the paralytic agent. 5. Passive movement of the paralyzed eye did not induce movement of the untreated eye. 6. Application of a local anesthetic agent rather than a paralytic agent to one eye resulted in reversible reduction of movement only in the treated eye, but not in the untreated eye. 7. The results suggest that the reduction of movement in one eye produced by application of a paralytic agent to the other eye is mediated by an afferent neural signal derived from extraocular muscle proprioceptors possibly gated by a central motor control signal.

Behavioral determination of the spatial selectivity of contrast adaptation in cats: some evidence for a common plan in the mammalian visual system.

An aftereffects paradigm was used to behaviorally measure contrast sensitivity of cats to gratings of three different test spatial frequencies after adaptation to gratings of various spatial frequencies, contrasts, and durations. Post-adaptation reductions in sensitivity occurred even after short periods of adaptation (less than 7 s) and could be as large as 1.0 log unit under some conditions. The magnitude of the adaptation effect varied monotonically with (1) adaptation grating contrast, (2) duration, and (3) the contrast sensitivity for the test grating. Average half-width (at half-height) of the spatial-frequency tuning curves constructed from the data was 1.4 octaves, and was not dependent upon the level of adaptation or the spatial frequency of the test grating. Post-adaptation psychometric functions of the cats showed reduced slopes and maxima suggesting that, unlike humans, in cats apparent contrast grows more slowly with increases in physical contrast after contrast adaptation. All of the characteristics observed are in excellent agreement with electrophysiologically measured properties of neurons in striate cortex of cats. In addition, there was a remarkable similarity of the cat tuning functions, both in shape and bandpass, to those measured in man with a similar paradigm suggesting that (1) the two visual systems are sufficiently similar to make the cat a useful spatial vision model and (2) there is a common functional plan to all mammalian visual systems despite significant anatomical differences between species.

Integration regions for vernier acuity in the cat: comparative aspects of cortical scaling.

The spatial region involved in vernier acuity (integration region) was behaviorally determined in cats and compared with published human data and cortical scaling parameters for the two species. Integration region was estimated from vernier thresholds obtained with: (1) gaps of various sizes between the contiguous ends of the offset contours (gap effects); and (2) irrelevant contours placed at various distances from the vernier offset (interference effects). The gap studies showed that thresholds decreased as the gap grew from 0 to 4-7 min but then increased progressively with separations greater than 7 min. In the interference studies, thresholds were increased 2-6 times when the irrelevant contours were located within 40 min of the offset. Comparing these data with available human measures showed large species differences. Converting the interference distances for both species into cortical distances using published estimates of cortical magnification factor did not resolve the discrepancy. However, a comparison of the cat data with published human data, taken at an eccentricity at which the estimated human cortical magnification factor equals that of the cat, i.e. 7 deg, showed the effects of gap size and interference distance on vernier thresholds to be remarkably similar between the two species. This similarity supports the hypothesis that positional acuity depends upon mechanisms in striate cortex, and also suggests that such acuity may be mediated by a generic neural mechanism common to mammals.

Evoked potential estimates of the time course of adaptation and recovery to counterphase gratings.

Scalp-recorded evoked potentials (VEP) were sequentially sampled in humans during adaptation to and recovery from prolonged viewing of counterphase sinusoidal grating targets. The sum of the power at the first and second harmonics of the Fourier-transformed VEP components was found to decrease during adaptation and increase during recovery. Time constants (T) for the adaptation and recovery processes as estimated from exponential functions ranged from 2.9 to 19 sec, varying non-monotonically with the spatial frequency and contrast of the stimulus. The observed T values are shorter than those reported in psychophysical studies of adaptation but overlap estimates derived from single cell studies. An unexpected finding was the occurrence of a 3-6 sec delay in the appearance of the maximum VEP response after the onset of the adaptation stimulus. The delay occurred in all subjects and at all spatial frequencies when moderate to high adapting contrasts (e.g. greater than 0.2) were used. The data support a feature-selective, multi-channel lateral inhibitory model of spatial vision and suggest the presence of tonic inhibition between the channels.

Evoked potentials elicited by brief vernier offsets: estimating vernier thresholds and properties of the neural substrate.

The characteristics of the neural generator producing an evoked potential in response to the brief presentation of a vernier offset was investigated in three experiments. In the first study, averaged evoked potentials (EPs) recorded in response to a single vernier offset stimulus consisting of a horizontal line which changed from colinearity to noncolinearity for 100 msec every 1.5 sec were compared to responses elicited by other vernier configurations consisting of: stimuli with multiple offsets; stimuli presented in different orientations; targets with different offset features; and with the simple displacement of a colinear line. The results showed that a single vernier offset elicited a robust response if the offset was located in the central zone (1 degree) of the target. Other features of the target configuration were unimportant. Displacement of a colinear line over the same range without an offset evoked little, if any, response. In the second study, EPs were recorded in response to a single offset target which varied in magnitude from 21 to 82 sec of visual angle on different trials. The latency and amplitude of the EP response varied systematically with the amplitude of the vernier offset. Plots of EP amplitude against log of the offset magnitude were linear over the range of offsets employed. Straight lines fitted to the data and extrapolated to zero amplitude provided estimates of vernier threshold. These estimates agreed closely with psychophysical measures taken with the same targets and confirm the initial observations by Levi et al. (1983). In the third experiment, irrelevant contours were added to the vernier target in various spatial and temporal configurations. The addition of stationary, contiguous contours to the vernier target reduced the amplitude of the EP response when the contours were within 4-8 min of the offset, producing progressively less EP attenuation with increasing distance from the offset. However, brief presentation of the irrelevant contour (e.g. a single line passing through the offset) with an onset asynchrony relative to the vernier offset stimulus appropriate to assure simultaneity of the line-elicited EP and the offset-elicited EP yielded an enhanced response, i.e. to two responses added algebraically. The long latency of the offset evoked response and the summation results of the EPs generated by an offset and by a briefly presented linear contour suggests independence of the neural generators producing the response to these two targets.

Orientation sensitivity of the cat assessed from evoked potentials: central and peripheral contributions.

To estimate contour-orientation sensitivity of the cat and the degree to which precortical processing contributes to such estimates, the amplitude of visually evoked potentials (VEP) recorded from the visual cortex of cats in response to a visual stimulus (S2) presented at various intervals after presentation of another visual stimulus (S1) was measured under several conditions. Recordings were made when both stimuli were presented through one eye (monoptic condition) or when S1 was presented to one eye and S2 to the other (dichoptic condition). In some experiments, simultaneous recordings were made from the optic tract and visual cortex. The stimuli were pairs of sinusoidal gratings with a spatial frequency of 0.5 cycles/deg and of various orientations. Each stimulus was presented by stepping the grating contrast from 0.0 (adapting field) to 0.5 for 50 ms. The intervals between the presentation of the two test stimuli (S1 and S2) was varied from 0 to 1,550 ms, and on different trials the orientation of the S2 grating relative to that of S1 was varied from 0 to 90 degrees. Results showed that under monoptic conditions, the VEP to the second stimulus (S2) was reduced by presentation of the first stimulus (S1) when the interstimulus interval was less than 200 ms, whereas under dichoptic conditions, the response to S2 was reduced with interstimulus intervals less than 75 ms. The response reduction was always in a forward direction (e.g., reduced S2 response), increased in magnitude with decreases in the interstimulus interval, and was larger under monoptic conditions than under dichoptic conditions. The response reduction produced monoptically was orientation selective in that it was greatest when the orientation of S1 and S2 was the same, and it recovered by half when the orientation differed by 6 to 15 degrees (orientation half-band pass). In some cortical recordings, the orientation-selective response reduction was superimposed on a response reduction that was not selective for S2 orientation. Stimultaneous recording in the optic tract also showed a response reduction of S2 response that was not orientation selective, suggesting that precortical neural elements contribute to the cortical VEP. With dichoptic stimulus presentation an orientation-nonspecific response reduction was obtained. We hypothesized that binocular inhibitory effects, resulting from disparate retinal input, produced this surprising finding. The results demonstrate that the VEP recorded at the cortex can be used to estimate orientation sensitivity, but that response interactions in peripheral (precortical) neural elements can contribute to such estimates.

Stimulus-dependent form discrimination deficits in cats with superior colliculus lesions.

Previous studies have shown that cats with damaged superior colliculi are slower than normal in learning new visual discriminations based upon shape but show no losses in retention of discriminations learned prior to the tectal lesion. To elucidate the possible mechanisms producing this deficit, the rate at which cats with superior colliculus lesions learned to discriminate between targets differing either in (1) global features, or (2) a single, uniquely localized, feature was measured. Cats were first tested with problems shown in earlier studies to produce acquisition deficits and those with superior colliculus lesions showed the expected deficits. The rate of acquisition of form discriminations based on global stimulus features in these cats was, however, comparable to normals and sham-operated controls. When tested with targets differing only in a single, localized feature, acquisition of the discrimination was severely impaired compared to controls. The observed deficits did not depend upon damage to pretectum or total ablation of the superior colliculus. The results support the view that the superior colliculus is a major contributor to visuomotor integration (e.g. foveation of salient stimulus features), and that such visuomotor behavior is an important determinant in form discrimination learning.

Contrast sensitivity for vertically and obliquely oriented gratings as a function of grating area.

Contrast sensitivity for both vertically and obliquely oriented gratings increased with increases in stimulus area to an asymptotic value. Sensitivity grew more slowly for oblique gratings and reached an asymptotic value at a larger area than for vertical gratings. For equal areas, oblique gratings always yielded poorer sensitivity. The results suggest a larger spatial summation area for obliquely oriented gratings.

Intracortical processing of visual contour information in cats.

Fine slices made through the gray matter perpendicular to the surface of cortical area 17-18 in the cat produced no change in behavioral measures of grating acuity, a modest reduction in vernier acuity and a proportionately larger effect on contour orientation acuity. The introduction of vertical gaps in the vernier acuity targets produced increases in the vernier offset threshold proportional to the gap size both before and after cortical damage, except at the largest vertical gap values employed (1-1.5 degrees) where no change in thresholds was observed after surgery. Neither the preoperative vernier thresholds nor the postoperative changes in vernier thresholds were predicted by the measured contour orientation thresholds and thus an orientation discrimination model of vernier acuity was not supported. The results of this study support the view that intracortical processing in area 17-18 is important in both vernier and orientation acuity, but the neural mechanisms mediating the two capacities are different.

Striate cortex and visual acuity functions in the cat.

Using a two-choice visual discrimination paradigm, thresholds for size (gratings), parallelness (parallel vs. non-parallel lines), contour alignment (vernier offset), and angularity (polygon figures) were behaviorally determined in cats before and after ablations of portions of the geniculo-cortical system. Animals with a total loss of cortical area 17, and with a loss, in some cases, of up to 90% of areas 18 (with and without infringement into area 19), showed about a 30% reduction in grating acuity, a three-fold increase in parallelness and angularity thresholds, and a total loss of contour alignment ability. Control animals with ablations sparing area 17 showed no significant threshold changes. All animals were able to learn classic form discriminations postoperatively, but those with area 17-18 lesions at a somewhat slower than normal rate. Control procedures indicated that all tested discrimination capabilities did not depend on luminance differences between targets, local flux cues within the targets, or on the animals' use of residual portions of area 17 representing the peripheral visual field. Since the cat has multiple thalamo-cortical visual pathways, the results of the present study are consistent with the hypothesis that pathways parallel to the geniculo-striate system are capable of processing spatial information of considerable detail. The results also suggest, however, that the geniculo-striate system is uniquely necessary for the processing of the finest attributes of spatial contours.

Visual acuity functions and pattern discrimination in the destriate cat.

The study provides evidence that the cat is capable of processing visual information of some detail after removal of area 17 and most of area 18. This postoperative discrimination of complex spatial stimuli is not based on the use of such cues as luminance differences or local flux cues. From the deficits which follow extensive lesions of areas 17-18, it appears as though these cortices participate in the detection of fine details. We found a modest increase in the threshold of grating acuity, moderate loss in orientation acuity and extensive deficit in a task requiring topographic alignment of contours (vernier offset). Further evidence for the substantial preservation of spatial vision in the destriate cat is provided by experiments which show that perceptual grouping of rectilinear arrays of figural elements (dots or line segments) into obliquely oriented rows is largely unaffected by the lesion, even when the grouping is initiated by near-threshold proximity cues. Since grouping effects are felt to be involved in the organization of the visual field into the figure-ground dichotomy, these results indicate that the neural mechanisms subserving the initial stage of form perception lie outside of areas 17 and 18. Consistent with the increased acuity thresholds of destriate cats, these animals have deficits in several pattern discriminations that require fine-grained spatial analysis The results suggest that areas 17-18 serve as a high-spatial frequency analyzer, but are not essential to pattern and form recognition.

Movement discrimination capacities in the cat.

Cats were trained to discriminate moving from nonmoving targets or one direction of movement from another. Various stimulus changes, e.g., size, direction, and rate, were then introduced as a test for generalization of the dimension of movement. Thresholds for detection for minimal movement were also determined. The results showed that (a) for cats, discrimination of movement is more difficult than discriminations based on brightness; (b) the dimension of movement is completely generalized across stimulus configuration but incompletely generalized for direction of movement; (c) the mean movement detection threshold was found to be 3.3 degrees/sec; (d) the thresholds for minimal movement and direction of movement were essentially identical; and (e) stimulus-viewing strategies were found to play an important role in the threshold determinations. The results of the generalization tests are consistent with the physiological properties of neurons found to be sensitive to movement in the cat visual system. The movement threshold values were found to lend support to the view that resolution and slow movement thresholds are correlated.

The role of superior colliculus in vision: visual form discrimination in cats with superior colliculus ablations.

The rate of acquisition of visual form discriminations in cats with superior colliculus damage was measured in a testing apparatus requiring a minimal amount of stimulus localization in space. In animals with bilateral superior colliculus ablations, acquisition was imparied relative to acquisition in normal animals. In these animals, the extent of the lesion was related to the severity of the deficit. In split-brain preparations with unilateral superior colliculus ablations, acquisition with the eye ipsilateral to the lesion was impaired relative to acquisition with eye contralateral to the lesion. The deficits observed did not depend upon damage to pretectum. The results support the conclusion that the superior colliculus participates in certain aspects of form vision.

Responses of single cells in cat visual cortex to prolonged stimulus movement: neural correlates of visual aftereffects.

1. The activity of single cortical cells in area 17 of anesthetized and unanesthetized cats was recorded in response to prolonged stimulation with moving stimuli. 2. Under the appropriate conditions, all cells observed showed a progressive response decrement during the stimulation period, regardless of cell classification, i.e., simple, complex, or hypercomplex. 3. The observed response decrement was shown to be largely cortical in origin and could be adequately described with an exponential function of the form R = Rf +(R1-Rf)e-t/T. Time constants derived from such calculations yielded values ranging from 1.92 to 12.45 s under conditions of optimal-stimulation. 4. Most cells showed poststimulation effects, usually a brief period of reduced responsiveness that recovered exponentially. Recovery was essentially complete in about 5-35 s. 5. The degree to which stimuli were effective at inducing response was shown to have significant effects on the magnitude of the response decrement. 6. Several cells showed neural patterns of response and recovery that suggested the operation of intracortical inhibitory mechanisms. 7. A simple two-process model that adequately describes the behavior of all the studied cells is presented. 8. Because the properties of the cells studied correlate well with human psychophysical measures of contour and movement adaptation and recovery, a causal relationship to similar neural mechanisms in humans is suggested.