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.     

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.     

Contrast sensitivity of form and motion discrimination during binocular rivalry

Binocular rivalry, which is induced by presenting the two eyes with incompatible stimuli, results in periods where one eye's stimulus is seen and the other stimulus is suppressed. We measured the depth of suppression in two ways, with very different results. First, two similar forms were briefly presented to one eye: the difference in shapes required to discriminate the forms was substantially greater during suppression than during dominance. Second, the two forms were made sufficiently different in shape to be easily distinguishable at high contrast, and contrast was lowered to find the threshold for discrimination of the forms. Contrast sensitivity did not differ between the suppression and dominance states. These results were replicated with a motion discrimination task: suppression markedly worsened the ability to distinguish increases from decreases in speed but did not elevate the minimum contrast required for the same task. We interpret the results in terms of steep contrast-response functions in visual cortex beyond the primary area.     

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.     

Adaptive center-surround interactions in human vision revealed during binocular rivalry

We used binocular rivalry as a psychophysical probe to explore center-surround interactions in orientation, motion and color processing. Addition of the surround matching one of the rival targets dramatically altered rivalry dynamics. For all visual sub-modalities tested, predominance of the high-contrast rival target matched to the surround was greatly reduced-a result that disappeared at low contrast. At low contrast, addition of the surround boosted dominance of orientation and motion targets matched to the surround. This contrast-dependent modulation of center-surround interactions seems to be a general property of the visual system and may reflect an adaptive balance between surround suppression and spatial summation.     

Increasing depth of binocular rivalry suppression along two visual pathways

Binocular rivalry refers to the alternating perception that occurs when the two eyes are presented with incompatible stimuli: one monocular image is seen exclusively for several seconds before disappearing as the other image comes into view. The unseen stimulus is physically present but is not perceived because the sensory signals it elicits are suppressed. The neural site of this binocular rivalry suppression is a source of continuing controversy. We psychophysically tested human subjects, using test probes designed to selectively activate the visual system at a variety of processing stages. The results, which apply to both form and motion judgements, show that the sensitivity loss during suppression increases as the subject's task becomes more sophisticated. We conclude that binocular rivalry suppression is present at a number of stages along two visual cortical pathways, and that suppression deepens as the visual signal progresses along these pathways.     

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.     

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.     

When crowding meets binocular rivalry: Challenges for object perception

Both crowding and binocular rivalry impair object perception, but their influence on object perception has so far only been investigated in separate fields. Three experiments investigated the joint influences of crowding and rivalry on object perception (orientation discrimination). Experiment 1 investigated how crowding and rivalry influence orientation discrimination together. Experiment 2 tested whether rivalry between flankers affects crowding using an orientation discrimination task. Experiment 3 tested whether crowding affects the temporal dynamics of the rivalry between a target and a rival stimulus. In Experiment 1, judgments of target orientation were more impaired when crowding and rivalry were simultaneously induced than when they were separately induced and their effects were combined. In Experiment 2, judgments of target orientation were impaired even when flankers were undergoing rivalry, thus highlighting the importance of the presence of flankers. Experiment 3 showed that flankers presented in the neighborhood of a target undergoing rivalry shortened target dominance and prolonged target suppression. The augmented impairments of object perception found in Experiments 1 and 3 suggest that crowding and rivalry interact, presumably through signal suppression. The adverse effect of flankers shown in Experiment 2 suggests that inappropriate feature integration may have additionally contributed to this interaction.     

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].     

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.     

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.     

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.     

Strength and coherence of binocular rivalry depends on shared stimulus complexity

Presenting incompatible images to the eyes results in alternations of conscious perception, a phenomenon known as binocular rivalry. We examined rivalry using either simple stimuli (oriented gratings) or coherent visual objects (faces, houses etc). Two rivalry characteristics were measured: Depth of rivalry suppression and coherence of alternations. Rivalry between coherent visual objects exhibits deep suppression and coherent rivalry, whereas rivalry between gratings exhibits shallow suppression and piecemeal rivalry. Interestingly, rivalry between a simple and a complex stimulus displays the same characteristics (shallow and piecemeal) as rivalry between two simple stimuli. Thus, complex stimuli fail to rival globally unless the fellow stimulus is also global. We also conducted a face adaptation experiment. Adaptation to rivaling faces improved subsequent face discrimination (as expected), but adaptation to a rivaling face/grating pair did not. To explain this, we suggest rivalry must be an early and local process (at least initially), instigated by the failure of binocular fusion, which can then become globally organized by feedback from higher-level areas when both rivalry stimuli are global, so that rivalry tends to oscillate coherently. These globally assembled images then flow through object processing areas, with the dominant image gaining in relative strength in a form of 'biased competition', therefore accounting for the deeper suppression of global images. In contrast, when only one eye receives a global image, local piecemeal suppression from the fellow eye overrides the organizing effects of global feedback to prevent coherent image formation. This indicates the primacy of local over global processes in rivalry.     

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.     

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.     

Attending to auditory signals slows visual alternations in binocular rivalry

A previous study has shown that diverting attention from binocular rivalry to a visual distractor task results in a slowing of rivalry alternation rate between simple orthogonal orientations. Here, we investigate whether the slowing of visual perceptual alternations will occur when attention is diverted to an auditory distractor task, and we extend the investigation by testing this for two kinds of binocular rivalry stimuli and for the Necker cube. Our results show that doing the auditory attention task does indeed slow visual perceptual alternations, that the slowing effect is a graded function of attentional load, and that the attentional slowing effect is less pronounced for grating rivalry than for house/face rivalry and for the Necker cube. These results are explained in terms of supramodal attentional resources modulating a high-level interpretative process in perceptual ambiguity, together with a role for feedback to early visual processes in the case of binocular rivalry.     

Eye rivalry and object rivalry in the intact and split-brain

Both the eye of origin and the images themselves have been found to rival during binocular rivalry. We presented traditional binocular rivalry stimuli (face to one eye, house to the other) and Diaz-Caneja stimuli (half of each image to each eye) centrally to both a split-brain participant and a control group. With traditional rivalry stimuli both the split-brain participant and age-matched controls perceived more coherent percepts (synchronised across the hemifields) than non-synchrony, but our split-brain participant perceived more non-synchrony than our controls. For rival stimuli in the Diaz-Caneja presentation condition, object rivalry gave way to eye rivalry with all participants reporting more non-synchrony than coherent percepts. We have shown that splitting the stimuli across the hemifields between the eyes leads to greater eye than object rivalry, but that when traditional rival stimuli are split as the result of the severed corpus callosum, traditional rivalry persists but to a lesser extent than in the intact brain. These results suggest that communication between the early visual areas is not essential for synchrony in traditional rivalry stimuli, and that other routes for interhemispheric interactions such as subcortical connections may mediate rivalry in a traditional binocular rivalry condition.     

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.     

Perceptual dominance during binocular rivalry is prolonged by a dynamic surround

We examined whether dynamic stimulation that surrounds a rival target influences perceptual alternations during binocular rivalry. We presented a rival target surrounded by dynamic random-dot patterns to both eyes, and measured dominance durations for each eye’s rival target. We found that rival target dominance durations were longer when surrounds were dynamic than when they were static or absent. Additionally, prolonged dominance durations were more apparent when the dynamic surround was alternately presented between the two eyes than when it was presented simultaneously to both eyes. These results indicate that dynamic stimulation that surrounds a rival target plays a role in maintaining the current perceptual state, and causes less perceptual alternations during binocular rivalry. Our findings suggest that dynamic signals on the retina may suppress rivalry, and thus provide useful information for stabilizing perceptions in daily life.