Center-surround inhibition deepens binocular rivalry suppression

When dissimilar stimuli are presented to each eye, perception alternates between both images--a phenomenon known as binocular rivalry. It has been shown that stimuli presented in proximity of rival targets modulate the time each target is perceptually dominant. For example, presenting motion to the region surrounding the rival targets decreases the predominance of the same-direction target. Here, using a stationary concentric grating rivaling with a drifting grating, we show that a drifting surround grating also increases the depth of binocular rivalry suppression, as measured by sensitivity to a speed discrimination probe on the rival grating. This was especially so when the surround moved in the same direction as the grating, and was slightly weaker for opposed directions. Suppression in both cases was deeper than a no-surround control condition. We hypothesize that surround suppression often observed in area MT (V5)-a visual area implicated in visual motion perception-is responsible for this increase in suppression. In support of this hypothesis, monocular and binocular surrounds were both effective in increasing suppression depth, as were surrounds contralateral to the probed eye. Static and orthogonal motion surrounds failed to add to the depth of rivalry suppression. These results implicate a higher-level, fully binocular area whose surround inhibition provides an additional source of suppression which sums with rivalry suppression to effectively deepen suppression of an unseen rival target.     

Stimulus flicker alters interocular grouping during binocular rivalry

When the two eyes are presented with sufficiently different stimuli, the stimuli will engage in binocular rivalry. During binocular rivalry, a subject's perceptual state alternates between awareness of the stimulus presented to the right eye and that presented to the left eye. There are instances in which competition is not eye-based, but instead takes place between stimulus features, as is the case in flicker and switch rivalry (F&S). Here we investigate another such instance, interocular grouping, using a Diaz-Caneja type stimulus in conjunction with synchronous stimulus flicker. Our results indicate that stimulus flicker increases the total duration of interocularly bound percepts, and that this effect occurs for a range of temporal flicker frequencies. Furthermore, the use of contrast-inversion flicker causes a decrease of total dominance duration of the interocularly bound percepts. We argue that different flickering regimes can be used to differentially stimulate lower and higher levels of visual processing involved in binocular rivalry. We propose that the amount of interocularly combined pattern-completed percept can be regarded as a measure of the level at which binocular rivalry is resolved.     

Local binocular fusion is involved in global binocular rivalry

We examined whether interocular inhibition in binocular rivalry could occur at the interocular intersection of horizontal and vertical rectangular patches which are locally fusible but globally rivalrous between the two eyes. We measured contrast increment (and decrement) thresholds of a monocularly presented probe which was presented on the horizontal patch corresponding to the intersection. We found that the threshold was higher when the horizontal patch was perceptually suppressed than when it was dominant. In addition, threshold elevation did not occur when both patches were dominant, or when the horizontal patch was viewed in isolation. These results indicate that interocular inhibition occurs at the potentially fusible region, and the determination of binocular fusion or binocular rivalry does not depend on physical stimulus but rather perceptual state at the time.     

On the relation between dichoptic masking and binocular rivalry

When our two eyes view incompatible images, the brain invokes suppressive processes to inhibit one image, and favor the other. Two phenomena are typically observed: dichoptic masking (reduced sensitivity to one image) for brief presentations, and binocular rivalry (alternation between the two images), over longer exposures. However, it is not clear if these two phenomena arise from a common suppressive process. We investigated this by measuring both threshold elevation in simultaneous dichoptic masking and mean percept durations in rivalry, whilst varying relative stimulus orientation. Masking and rivalry showed significant correlations, such that strong masking was associated with long dominance durations. A second experiment suggested that individual differences across both measures are also correlated. These findings are consistent with varying the magnitude of interocular suppression in computational models of both rivalry and masking, and imply the existence of a common suppressive process. Since dichoptic masking has been localised to the monocular neurons of V1, this is a plausible first stage of binocular rivalry.     

Both monocular and binocular signals contribute to motion rivalry

There is an ongoing debate on whether binocular rivalry involves competition among monocular cells or binocular cells. We investigated this issue psychophysically with two specially designed test stimuli. One test stimulus contained monocular motion signals but greatly reduced binocular motion signals, while the other contained binocular motion signals but no monocular motion signals. For comparison, we also employed a normal rivalrous control containing both monocular and binocular motion signals, and a non-rivalrous flicker-noise control with neither monocular nor binocular motion signals. We found that binocular rivalry for the two test stimuli was significantly reduced compared with the normal rivalrous control, but not completely eliminated compared with the non-rivalrous control. Therefore, both monocular and binocular motion signals appear to contribute to motion rivalry, suggesting that motion rivalry must involve competition among both monocular and binocular cells.     

A fresh look at interocular grouping during binocular rivalry

During binocular rivalry, observers sometimes perceive one complete visual object even though component features of that perceptually dominant object are distributed between the two eyes and are in rivalry against other, dissimilar features. This interocular grouping cannot be explained by models of rivalry in which one eye or the other is completely dominant at any given moment. But perhaps global interocular grouping is achieved by simultaneous local eye dominance, wherein portions of one eye's view and complementary portions of the other eye's view become dominant simultaneously. To test this possibility, we performed two experiments using relatively large, complex figures as rival targets. In one experiment we used an "eye-swap" technique to confirm that within given, local spatial regions of rivalry it was the region of an eye--not a given stimulus feature--that was usually dominant. In a second experiment, we measured dominance durations for multiple, local zones of rivalry and then created 1-min animations of a global "montage" in which dominance within local regions was governed by the distributions of dominance measured empirically. These animations included significant periods of time during which global interocular grouping was evident; observers viewed these animations intermixed with actual rivalry displays, and the resulting tracking data confirmed the similarity in global dominance of the two display types. Thus interocular grouping during rivalry does not rule out local, eye-based rivalry, although synergistic and top-down influences almost certainly provide additional force in the promotion of interocular grouping.     

Monocular rivalry exhibits three hallmarks of binocular rivalry: Evidence for common processes

Binocular rivalry occurs when different images are presented one to each eye: the images are visible only alternately. Monocular rivalry occurs when different images are presented both to the same eye: the clarity of the images fluctuates alternately. Could both sorts of rivalry reflect the operation of a general visual mechanism for dealing with perceptual ambiguity? We report four experiments showing similarities between the two phenomena. First, we show that monocular rivalry can occur with complex images, as with binocular rivalry, and that the two phenomena are affected similarly by the size (Experiment 1) and colour (Experiment 2) of the images. Second, we show that the distribution of dominance periods during monocular rivalry has a gamma shape and is stochastic (Experiment 3). Third, we show that during periods of monocular-rivalry suppression, the threshold to detect a probe (a contrast pulse to the suppressed stimulus) is raised compared with during periods of dominance (Experiment 4). The threshold elevation is much weaker than during binocular rivalry, consistent with monocular rivalry’s weak appearance. We discuss other similarities between monocular and binocular rivalry, and also some differences, concluding that part of the processing underlying both phenomena is a general visual mechanism for dealing with perceptual ambiguity.     

A bias for looming stimuli to predominate in binocular rivalry

Concentric gratings that expand outwards are seen for a greater period of time relative to contracting gratings when engaged in binocular rivalry. During binocular rivalry (BR), which is a fluctuation in visual awareness between different images presented separately to each eye, equivalent images tend to be seen in equal proportion over the observation period. When one eye's image is particularly salient, brighter, or moving, this equality is curtailed, and the stronger image predominates. Here a specific direction of motion is found to predominate over another of equal speed. This tendency is consistent with the ability of looming objects to orient attention, coupled with previous accounts of the role of stimulus-driven attention in BR.     

Inter-ocular transfer of stimulus cueing in dominance selection at the onset of binocular rivalry

Recent work investigated the influence of exogenous attention on initial percept dominance at the onset of binocular rivalry. It was reported that cueing attention to one of two binocularly presented transparent stimuli immediately prior to rivalrous viewing provided the cued stimulus with a competitive advantage in subsequent binocular rivalry. This effect was independent of the eye containing the cued stimulus during the rivalry phase. In this recent work, the attention cue was always presented to both eyes. This leaves unclear the extent to which cueing affects binocular and/or monocular stimulus representations. To disambiguate this issue, we compared the cueing strength when the cue was presented ipsi-, contra- or bi-laterally with respect to the eye containing the cued stimulus during subsequent binocular rivalry. Besides replicating previous findings, we found that stimulus cueing readily transfers across eyes, suggesting that binocular mechanisms mediate exogenous attention effects on dominance selection at the onset of binocular rivalry.     

Feature-based activation and suppression during binocular rivalry

In the past decade, effects of pattern coherence have indicated that perception during binocular rivalry does not result solely from reciprocal inhibitory competition between monocular channels. In this study we were interested in feature selectivity both during dominance and during suppression. The first experiment shows that a suppressed stimulus perceptually appears earlier when it shares features with a visible stimulus than when it does not. Subsequently, our second experiment suggests a reversal of this effect when similarity is exhibited with a suppressed stimulus. These findings hint at a role for both selective enhancing (Experiment 1) and selective inhibitory cortical mechanisms (Experiment 2) in causing image rivalry. From a phenomenological perspective these results suggest that we are not only selectively aware but also selectively unaware of specific features in the visual scene.     

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 JW's hemifields, rivalry synchronized for similarly oriented stimuli, the same as happened for intact-brain observers. When the patches were in JW's opposite hemifields, there was no synchronizing of rivalry, unlike what happened for intact-brain observers. This suggests that rivalry processed in JW's two hemispheres is independent. We conclude that rivalry is processed fully within each hemisphere.     

Interactions between binocular rivalry and Gestalt formation

A question raised a long time ago in binocular rivalry research is whether the phenomenon of binocular rivalry is purely determined by local stimulus properties or that global stimulus properties also play a role. More specifically: do coherent features in a stimulus influence rivalrous behavior? After decades of underexposure of the subject, recently this question seemed to be answered in the affirmative. This paper presents additional evidence for an influence of coherent features. In an experiment in which eye movements cannot bias conclusions it is demonstrated that Gestalt formation influences binocular rivalry positively, i.e., stronger Gestalts have longer total dominance times. Gestalt formation appears to intervene in the states of dominance ("what"), not directly in the dominance durations ("how long"). This generates questions about the nature of interactions between binocular rivalry and Gestalt formation. Gestalt formation seems to be fed by signals that are generated after binocular convergence and only leaves its mark on binocular rivalry by feedback to monocular channels, a conclusion which has been drawn before by Alais and Blake [Alais, D., & Blake, R. (1998). Interaction between global motion and local binocular rivalry. Vision research 38, 637-644].     

An integrative model of binocular vision: a stereo model utilizing interocularly unpaired points produces both depth and binocular rivalry

Half-occluded points (visible only in one eye) are perceived at a certain depth behind the occluding surface without binocular rivalry, even though no disparity is defined at such points. Here we propose a stereo model that reconstructs 3D structures not only from disparity information of interocularly paired points but also from unpaired points. Starting with an array of depth detection cells, we introduce cells that detect unpaired points visible only in the left eye or the right eye (left and right unpaired point detection cells). They interact cooperatively with each other based on optogeometrical constraints (such as uniqueness, cohesiveness, occlusion) to recover the depth and the border of 3D objects. Since it is contradictory for monocularly visible regions to be visible in both eyes, we introduce mutual inhibition between left and right unpaired point detection cells. When input images satisfy occlusion geometry, the model outputs the depth of unpaired points properly. An interesting finding is that when we input two unmatched images, the model shows an unstable output that alternates between interpretations of monocularly visible regions for the left and the right eyes, thereby reproducing binocular rivalry. The results suggest that binocular rivalry arises from the erroneous output of a stereo mechanism that estimates the depth of half-occluded unpaired points. In this sense, our model integrates stereopsis and binocular rivalry, which are usually treated separately, into a single framework of binocular vision. There are two general theories for what the "rivals" are during binocular rivalry: the two eyes, or representations of two stimulus patterns. We propose a new hypothesis that bridges these two conflicting hypotheses: interocular inhibition between representations of monocularly visible regions causes binocular rivalry. Unlike the traditional eye theory, the level of the interocular inhibition introduced here is after binocular convergence at the stage solving the correspondence problem, and thus open to pattern-specific mechanisms.     

Center-surround interactions in visual motion processing during binocular rivalry

When each eye is confronted with a dissimilar stimulus, the percept will generally alternate between the two. This phenomenon is known as binocular rivalry. Although binocular rivalry occurs at locations where targets overlap spatially, the area surrounding rivalrous targets can modulate their dominance. Here we show that during binocular rivalry of oppositely moving gratings, a surrounding grating moving in the same direction as one of the two leads to increased dominance of the opposite direction of motion in the center. This increased dominance of the opposite direction in the center was observed irrespective of the eye to which the surround was presented. Inspection of the results for different conditions reveals that the preference for the opposite direction of motion cannot be explained by a single mechanism operating beyond binocular fusion. We therefore suggest that this phenomenon is the outcome of center-surround interactions at multiple levels along the pathway of visual motion processing.     

A binocular rivalry study of motion perception in the human brain

The relationship between brain activity and conscious visual experience is central to our understanding of the neural mechanisms underlying perception. Binocular rivalry, where monocular stimuli compete for perceptual dominance, has been previously used to dissociate the constant stimulus from the varying percept. We report here fMRI results from humans experiencing binocular rivalry under a dichoptic stimulation paradigm that consisted of two drifting random dot patterns with different motion coherence. Each pattern had also a different color, which both enhanced rivalry and was used for reporting which of the two patterns was visible at each time. As the perception of the subjects alternated between coherent motion and motion noise, we examined the effect that these alternations had on the strength of the MR signal throughout the brain. Our results demonstrate that motion perception is able to modulate the activity of several of the visual areas which are known to be involved in motion processing. More specifically, in addition to area V5 which showed the strongest modulation, a higher activity during the perception of motion than during the perception of noise was also clearly observed in areas V3A and LOC, and less so in area V3. In previous studies, these areas had been selectively activated by motion stimuli but whether their activity reflects motion perception or not remained unclear; here we show that they are involved in motion perception as well. The present findings therefore suggest a lack of a clear distinction between 'processing' versus 'perceptual' areas in the brain, but rather that the areas involved in the processing of a specific visual attribute are also part of the neuronal network that is collectively responsible for its perceptual representation.     

Caloric vestibular stimulation reveals discrete neural mechanisms for coherence rivalry and eye rivalry: a meta-rivalry model

Binocular rivalry is an extraordinary visual phenomenon that has engaged investigators for centuries. Since its first report, there has been vigorous debate over how the brain achieves the perceptual alternations that occur when conflicting images are presented simultaneously, one to each eye. Opposing high-level/stimulus-representation models and low-level/eye-based models have been proposed to explain the phenomenon, recently merging into an amalgam view. Here, we provide evidence that during viewing of Díaz-Caneja stimuli, coherence rivalry -- in which aspects of each eye's presented image are perceptually regrouped into rivalling coherent images -- and eye rivalry operate via discrete neural mechanisms. We demonstrate that high-level brain activation by unilateral caloric vestibular stimulation shifts the predominance of perceived coherent images (coherence rivalry) but not half-field images (eye rivalry). This finding suggests that coherence rivalry (like conventional rivalry according to our previous studies) is mediated by interhemispheric switching at a high level, while eye rivalry is mediated by intrahemispheric mechanisms, most likely at a low level. Based on the present data, we further propose that Díaz-Caneja stimuli induce 'meta-rivalry' whereby the discrete high- and low-level competitive processes themselves rival for visual consciousness.     

Subjective contours and binocular rivalry suppression

Binocular rivalry probably involves distributed neural processes, some responsible for dominance, others for suppression and still others for fluctuations in perception. Focusing on the suppression process, the present study asks whether neural events underlying rivalry suppression take place prior to, or subsequent to those underlying the synthesis of subjective contours. Specifically, we examined whether (i) a subjective contour could prematurely return a suppressed target to dominance and (ii) whether suppression of a Kanizsa-type inducer precludes the formation of a subjective contour. Suppression durations were not abbreviated by the subjective contour, but suppression did prevent the formation of a subjective contour. Evidently suppression precedes the synthesis of subjective contours in the visual processing hierarchy.     

Temporal frequency and contrast tagging bias the type of competition in interocular switch rivalry

The nature of competition underlying perceptual alternations in binocular rivalry remains controversial. Interocular swapping of rivalrous stimuli can result in either slow irregular perceptual alternations that bridge multiple interocular switches or fast regular alternations that are time locked to the stimulus exchanges. We labeled either the inputs to the eyes or the individual rivalrous stimuli using temporal frequency and contrast tagging. Tagging of eye-of-origin signals enhanced the fast regular perceptual alternations associated with eye rivalry, while stimulus tagging shifted perception towards slow irregular alternations characteristic of stimulus rivalry. Thus, the type of competition in binocular rivalry can be biased based on additional cues in the visual inputs. The results are consistent with a model in which the brain combines information across multiple visual features to resolve ambiguities in visual inputs.     

Exogenous attention and endogenous attention influence initial dominance in binocular rivalry

We investigated the influence of exogenous and endogenous attention on initial selection in binocular rivalry. Experiment 1 used superimposed +/-45 degrees gratings viewed dioptically for 3s, followed by a brief contrast increment in one of the gratings to direct exogenous attention to that grating. After a brief blank period, dichoptic stimuli were presented for various durations (100-700 ms). Exogenous attention strongly influenced which stimulus was initially dominant in binocular rivalry, replicating an earlier report (Mitchell, Stoner, & Reynolds. (2004). Object-based attention determines dominance in binocular rivalry. Nature, 429, 410-413). In Experiment 2, endogenous attention was manipulated by having participants track one of two oblique gratings both of which independently and continuously changed their orientations and spatial frequencies during a 5s period. The initially dominant grating was most often the one whose orientation matched the grating correctly tracked using endogenous attention. In Experiment 3, we measured the strength of both exogenous and endogenous attention by varying the contrast of one of two rival gratings when attention was previously directed to that grating. The contrast of the attended grating had to be reduced by an amount in the neighborhood of 0.3 log-units, to counteract attention's boost to initial dominance. Evidently both exogenous and endogenous attention can influence initial dominance of binocular rivalry, effectively boosting the stimulus strength of the attended rival stimulus.     

Minimal physiological conditions for binocular rivalry and rivalry memory

Binocular rivalry entails a perceptual alternation between incompatible stimuli presented to the two eyes. A minimal explanation for binocular rivalry involves strong competitive inhibition between neurons responding to different monocular stimuli to preclude simultaneous activity in the two groups. In addition, strong self-adaptation of dominant neurons is necessary to enable suppressed neurons to become dominant in turn. Here a minimal nonlinear neural model is developed incorporating inhibition, self-adaptation, and recurrent excitation. The model permits derivation of an equation for mean dominance duration as a function of the underlying physiological variables. The dominance duration equation incorporates an explicit representation of Levelt's second law. The same equation also shows that introduction of a simple compressive response nonlinearity can explain Levelt's fourth law. Finally, addition of brief, recurrent synaptic facilitation to the model generates properties of rivalry memory.