Robert P O'Shea

Contrast as a depth cue

One consequence of aerial perspective is that far objects have lower contrast than near objects. We tested the efficacy of contrast as a cue for depth perception by measuring the apparent, relative depth of two areas that differed in contrast with a background and in size. We tested monocularly and binocularly. Differences in contrast were achieved by making the areas different in luminance, than varying the luminance of the background. Subjects reported that the area having lower contrast with the background appeared farther than the area having higher contrast. Even when size opposed it, contrast continued to have a significant effect on depth perception. Monocular observation yielded larger apparent depth than binocular observation. Contrast is an effective depth cue in the absence of any other depth information. We suggest that contrast acts as a pictorial depth cue simulating the optical effects of aerial perspective.

Visual sensitivity underlying changes in visual consciousness

When viewing a different stimulus with each eye, we experience the remarkable phenomenon of binocular rivalry: alternations in consciousness between the stimuli [1, 2]. According to a popular theory first proposed in 1901, neurons encoding the two stimuli engage in reciprocal inhibition [3-8] so that those processing one stimulus inhibit those processing the other, yielding consciousness of one dominant stimulus at any moment and suppressing the other. Also according to the theory, neurons encoding the dominant stimulus adapt, weakening their activity and the inhibition they can exert, whereas neurons encoding the suppressed stimulus recover from adaptation until the balance of activity reverses, triggering an alternation in consciousness. Despite its popularity, this theory has one glaring inconsistency with data: during an episode of suppression, visual sensitivity to brief probe stimuli in the dominant eye should decrease over time and should increase in the suppressed eye, yet sensitivity appears to be constant [9, 10]. Using more appropriate probe stimuli (experiment 1) in conjunction with a new method (experiment 2), we found that sensitivities in dominance and suppression do show the predicted complementary changes.

The effect of spatial frequency and field size on the spread of exclusive visibility in binocular rivalry

We measured binocular rivalry between dichoptic, orthogonal, sinusoidal gratings both having spatial frequencies of 0.5, 1, 2, 4, 8 or 16 c deg-1 in fields ranging from 0.5 to 8 deg of visual angle in diameter. Total time that one or the other grating was exclusively visible had an inverted U-shaped relationship with spatial frequency, with the peak shifting to coarser spatial frequencies as the field size increased. We computed for each spatial frequency the maximum field size over which a criterion duration of exclusive visibility would spread. When expressed as areas, these sizes were inversely proportional to spatial frequency. This dependence of rivalry on spatial frequency is similar to those for stereopsis and fusion, consistent with the notion that all three binocular phenomena have a common mechanism.

On binocular alternation.

Diaz-Caneja (1928) made some prescient observations about binocular rivalry. Being in French, however, his paper remained largely unknown to the broader research community. His findings are similar to those reported very recently by contemporary researchers who had independently observed similar phenomena. Using concentric circles and parallel lines as stimuli, Diaz-Caneja presented half of each form to opposite eyes to provoke binocular rivalry. He observed periods in the ensuing binocular alternations in which rivalry occurred between the good Gestalt forms, despite the fact that they were distributed between the eyes. He proposed that each half of a good form generates synchronised oscillations in the visual system, and that this synchronisation enables the dichoptically viewed halves of the one form to be perceived as a whole.

Early correlates of visual awareness in the human brain: time and place from event-related brain potentials

When something appears, how soon is the first neural correlate of awareness of it, and where is that activity in the brain? To answer these questions, we measured the electroencephalogram under conditions in which visual stimuli changed identically but in which awareness differed. We manipulated awareness by using binocular rivalry between orthogonal gratings viewed one to each eye. Then we changed the orientation of the grating to one eye to be the same as that to the other eye. Because of the rivalry, sometimes this happened to the visible grating, producing a clear change in perceived orientation, and other times it happened to the invisible grating, producing no change in perceived orientation. This procedure allowed us to analyze time-locked topographic scalp and tomographic primary current densities of the event-related potentials to physically identical events differing in their perceptual consequences. When the change in orientation reached awareness, neural responses began at about 100 ms, spreading mainly from dorsal occipital areas. When the change in orientation did not reach awareness, neural responses also began at about 100 ms, but they were attenuated, particularly in the right fusiform gyrus. We place the earliest correlate of visual awareness following binocular rivalry in the ventrolateral occipitotemporal cortex.

Frontoparietal activity and its structural connectivity in binocular rivalry.

To understand the brain areas associated with visual awareness and their anatomical interconnections, we studied binocular rivalry with functional magnetic resonance imaging (fMRI) and diffusion tensor imaging (DTI). Binocular rivalry occurs when one image is viewed by one eye and a different image by the other; it is experienced as perceptual alternations between the two images. Our first experiment addressed problems with a popular comparison condition, namely permanent suppression, by comparing rivalry with binocular fusion instead. We found an increased fMRI signal in right frontal, parietal, and occipital regions during rivalry viewing. The pattern of neural activity differed from findings of permanent suppression comparisons, except for adjacent activity in the right superior parietal lobule. This location was near fMRI signal changes related to reported rivalry alternations in our second experiment, indicating that neighbouring areas in the right parietal cortex may be involved in different components of rivalry. In our second experiment, we used probabilistic tractography to detect white matter fibres between right-hemispheric areas that showed event-related fMRI signal changes time-locked to reported perceptual alternations during rivalry viewing. Most of these functionally defined areas were linked by probabilistic fibre tracts, some of which followed long-distance connections such as the inferior occipitofrontal fasciculus. Corresponding anatomical pathways might mediate communication within the functional network associated with changes in conscious perception during binocular rivalry.

Binocular rivalry in split-brain observers

During binocular rivalry, visual perception switches between a stimulus viewed by one eye and a different stimulus viewed by the other. We studied rivalry in split-brain observers to test two explanations. Rivalry could reflect switching of activity between the cerebral hemispheres, or switching by a structure in the right frontoparietal cortex. From these two theories, we predict no rivalry when stimuli are presented to a split-brain observers left hemisphere. Yet we found similar rivalry from the left and right hemispheres of the split-brain observers, consistent with switchings being mediated by low-level processes within each hemisphere.

Visual grouping on binocular rivalry in a split-brain observer

We studied the effects of visual grouping on binocular rivalry in the left and right hemispheres of a split-brain observer, JW. In Experiments 1 and 2, we compared responses to traditional rivalry stimuli (e.g., a red vertical grating presented to the left eye and a green horizontal grating presented to the right eye) with responses to Diaz-Caneja stimuli (i.e., half of each grating was presented to one eye and the other half to the other eye). As found for intact-brain observers, JW reported episodes of exclusive visibility of coherent stimuli (e.g., of a red vertical grating alternating with a green horizontal grating) with Diaz-Caneja stimuli that were fewer and briefer than with traditional stimuli. This occurred in both hemispheres, demonstrating that during binocular rivalry, contours from one eye can be grouped with those of the opposite eye to create a coherent percept, even in the isolated hemispheres of the split-brain observer. In Experiment 3, we studied the tendency of rivalry in adjacent patches to synchronize. When both patches were in one of JWs hemifields, rivalry synchronized for similarly oriented stimuli, the same as happened for intact-brain observers. When the patches were in JWs opposite hemifields, there was no synchronizing of rivalry, unlike what happened for intact-brain observers. This suggests that rivalry processed in JWs two hemispheres is independent. We conclude that rivalry is processed fully within each hemisphere.

Probing visual consciousness: Rivalry between eyes and images

During binocular rivalry, one stimulus is visible (dominant), while the other stimulus is invisible (suppressed); after a few seconds, perception reverses. To determine whether these alternations involve competition between the eyes or between the images, we measured suppression depth to monocular probes. We did so in conventional rival stimuli and in rival stimuli swapping between the eyes at 1.5 Hz (both sorts of rivalry were shown either with or without 18-Hz flicker). The conventional conditions cause rivalry that could involve either competition between the eyes or between the images or both. The eye-swapping conditions cause rivalry that could involve competition between the images. Probes were either a small spot or a contrast increment to one of the rival stimuli. Using both yes-no and forced-choice procedures, we found that conventional conditions yielded large suppression depth and that eye-swapping conditions yielded small suppression depth. Weak suppression during image rivalry is consistent with conventional rivalrys involving competition between eyes and between images and flicker-and-swap rivalrys involving little, if any competition between eyes and mainly competition between images.

On the Role of Attention in Binocular Rivalry: Electrophysiological Evidence

During binocular rivalry visual consciousness fluctuates between two dissimilar monocular images. We investigated the role of attention in this phenomenon by comparing event-related potentials (ERPs) when binocular-rivalry stimuli were attended with when they were unattended. Stimuli were dichoptic, orthogonal gratings that yielded binocular rivalry and dioptic, identically oriented gratings that yielded binocular fusion. Events were all possible orthogonal changes in orientation of one or both gratings. We had two attention conditions: In the attend-to-grating condition, participants had to report changes in perceived orientation, focussing their attention on the gratings. In the attend-to-fixation condition participants had to report changes in a central fixation target, taking attention away from the gratings. We found, surprisingly, that attending to rival gratings yielded a smaller ERP component (the N1, from 160–210 ms) than attending to the fixation target. To explain this paradoxical effect of attention, we propose that rivalry occurs in the attend-to-fixation condition (we found an ERP signature of rivalry in the form of a sustained negativity from 210–300 ms) but that the mechanism processing the stimulus changes is more adapted in the attend-to-grating condition than in the attend-to-fixation condition. This is consistent with the theory that adaptation gives rise to changes of visual consciousness during binocular rivalry.