Competition in bistable vision is attribute-specific

We employ ambiguous figures and rivalrous stimuli that have multiple ambiguous properties to show that the different attributes of an ambiguous stimulus can undergo independent switching dynamics. This suggests that competition is distributed and attribute-specific, consistent with the known functional segregation of visual processing. Conflicting evidence that binocular rivalry is an early or late visual process may be better understood as evidence for attribute-specific competition occurring at multiple stages of visual processing. Specifically, we show that whether perceptual selection during binocular rivalry is early and eye-based or late and percept-based depends on the particular ambiguous attributes of the rivalrous stimulus.     

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.     

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.     

Recovery from perceptual filling-in is gated by interocular matching

We examined how the binocular visual system behaves during perceptual filling-in. In these experiments an initial filled-in target was replaced with an interocularly matched (fusible) or unmatched (rivalrous) target immediately after the disappearance of the initial target induced by perceptual filling-in. We measured the times for the target to recover from the filling-in. We found that recovery times were faster when the target was replaced with an interocularly matched target than with an unmatched target: The matched visual input was immediately released from perceptual suppression by filling-in but the unmatched one was not. These results indicate that even during perceptual filling-in our visual system can use the information whether the visual inputs from the two eyes are interocularly matched or not, and the interocular matching stage (the initial stage of binocular fusion or binocular rivalry) is not inhibited by the perceptual filling-in processing. Our findings suggest that the interocular matching processing may serve to gate visual inputs accessing visual awareness.     

Determinants of visual awareness following interruptions during rivalry

The inability of the human visual system to fuse dissimilar patterns in corresponding regions of the two eyes results in stochastic alternation of perceptual dominance between the two patterns: rivalry. When rivalrous stimuli are presented intermittently their perception is stabilized (Leopold, Wilke, Maier, & Logothetis, 2002). This stability indicates the operation of some kind of perceptual memory across interruptions in stimulation. Here we examined the contents of this perceptual memory to quantify the relative contributions of different sources of information: eye-of-origin, orientation, and color. Stimuli were intermittently presented and, during each blank interruption, we swapped either the color, orientation, or eye of presentation of the gratings. Comparing the percepts reported before and after each interruption allowed us to establish what aspects of perception remained stable. During conventional binocular rivalry, the eye in which the stimulus was presented remained stable across 74% of interruptions. Stimulus color and orientation also had weaker significant effects. When eye-of-origin information was eliminated by alternating the patterns rapidly between the two eyes, stimulus color remained stable across 86% of interruptions. Stimulus orientation again had a weaker but significant effect. These results demonstrate that the mechanisms mediating perceptual stability across interruptions in rivalry can operate at both monocular and binocular levels, much like the mechanisms operating during continuous viewing of rivalrous stimuli. On the basis of this similarity, we speculate that perceptual memory across interruptions in rivalry may involve the same neural representations as visual competition during rivalry. If this is the case, the use of intermittent stimulation in rivalry might permit the investigation of aspects of the mechanisms underlying visual competition that remain hidden during continuous presentation.     

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.     

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.     

Distance in feature space determines exclusivity in visual rivalry

Visual rivalry is thought to be a distributed process that simultaneously takes place at multiple levels in the visual processing hierarchy. Also, the different types of rivalry, such as binocular and monocular rivalry, are thought to engage shared underlying mechanisms. We hypothesized that the amount of perceptual suppression during rivalry as measured by the total duration of fully exclusive perceptual dominance is determined by a distance in a neurally represented feature space. This hypothesis can be contrasted with the possibility that the brain constructs an internal model of the outside world using full-fledged object representations, and that perceptual suppression is due to an appraisal of the likelihood of the particular stimulus configuration at a high, object-based level. We applied color and stereo-depth differences between monocular rivalry stimulus gratings, and manipulated color and eye-of-origin information in binocular rivalry using the flicker & switch presentation paradigm. Our data show that exclusivity in visual rivalry increases with increased difference in feature space without regard for real-world constraints, and that eye-of-origin information may be regarded as a segregating feature that functions in a manner similar to color and stereo-depth information. Moreover, distances defined in multiple feature dimensions additively and independently increase the amount of perceptual exclusivity and coherence in both monocular and binocular rivalry. We conclude that exclusivity in visual rivalry is determined by a distance in feature space that is subtended by multiple stimulus features.     

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.     

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.     

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.     

Functional hierarchies of nonconscious visual processing

A number of psychophysical techniques can be used to eliminate the registration of stimuli in visual awareness and to study the dynamics of conscious and nonconscious information processing in the visual system. However, little is known about how these techniques relate to each other. We chose to compare binocular rivalry, induced by orthogonal gratings presented separately to the two eyes, and metacontrast suppression, produced when a target stimulus is followed by a spatially surrounding mask stimulus, to investigate relative levels and correlates of nonconscious processing. Combined with prior results, our findings indicate that binocular rivalry expresses its suppressive effects prior to the level at which the mechanism of metacontrast does. Implications for theories of masking and interpretations of the loss or perceptual effects when stimulus visibility is suppressed by different psychophysical methods are discussed.     

Effect of transient versus sustained activation on interocular suppression

Switches in perceptual dominance resulting from either binocular rivalry or flash suppression likely involve some mechanism of interocular suppression, although it is unclear from past research whether different mechanisms are involved in the two cases. Using monocular, centrally fixated sinusoidal gratings surrounded by contiguous annuli of rivalrous gratings, suppression of the entire central grating was possible using either technique. However, the magnitude of the suppression was unaffected by the presence of an ipsilateral surround for flash suppression, yet, for binocular rivalry, suppression no longer occurred when the surrounds were fusible. Nevertheless, computational modeling demonstrates that the differences between the techniques may be attributable to the sustained versus transient stimulation of the contralateral surround, with the magnitude of the suppression proportional to the activation of the contralateral surround. Consistent with this, suppression extends over a greater distance at the onset of the contralateral surround than during sustained rivalry. Therefore, it is likely that perceptual dominance in both binocular rivalry and flash suppression is based on the same mechanism of interocular suppression.     

The effects of transcranial magnetic stimulation on visual rivalry

One extensively investigated form of perceptual bistability is binocular rivalry--When dissimilar patterns are presented one to each eye, these patterns compete for perceptual dominance. Here, we report that transcranial magnetic stimulation (TMS) over early visual areas induces alternations during binocular rivalry. The effect of TMS on binocular rivalry was retinotopic, suggesting that rivalry mechanisms are localized in the cortical representation of visual space. The timing of perturbations was highly dependent on individual differences in rivalry alternation frequencies, with more delayed effects found in slower alternators. This finding suggests that both binocular rivalry and TMS dynamics might be contingent on individual differences among observers. We performed an analogous set of experiments by replacing TMS with transient visual stimulation. The results, however, qualitatively and quantitatively differed from those reported with TMS. Finally, we found that TMS over early visual areas does not produce any time-locked effects on another dynamical process--eye-swapping stimulus rivalry. These findings constitute the first causative evidence that binocular rivalry is contingent on neural activity in early visual areas and suggest that binocular rivalry and stimulus rivalry have different neural correlates, supporting multilevel theories of visual rivalry.     

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.     

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 rivalry's involving competition between eyes and between images and flicker-and-swap rivalry's involving little, if any competition between eyes and mainly competition between images.     

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].     

How does binocular rivalry emerge from cortical mechanisms of 3-D vision?

Under natural viewing conditions, a single depthful percept of the world is consciously seen. When dissimilar images are presented to corresponding regions of the two eyes, binocular rivalry may occur, during which the brain consciously perceives alternating percepts through time. How do the same brain mechanisms that generate a single depthful percept of the world also cause perceptual bistability, notably binocular rivalry? What properties of brain representations correspond to consciously seen percepts? A laminar cortical model of how cortical areas V1, V2, and V4 generate depthful percepts is developed to explain and quantitatively simulate binocular rivalry data. The model proposes how mechanisms of cortical development, perceptual grouping, and figure-ground perception lead to single and rivalrous percepts. Quantitative model simulations of perceptual grouping circuits demonstrate influences of contrast changes that are synchronized with switches in the dominant eye percept, gamma distribution of dominant phase durations, piecemeal percepts, and coexistence of eye-based and stimulus-based rivalry. The model as a whole also qualitatively explains data about the involvement of multiple brain regions in rivalry, the effects of object attention on switching between superimposed transparent surfaces, monocular rivalry, Marroquin patterns, the spread of suppression during binocular rivalry, binocular summation, fusion of dichoptically presented orthogonal gratings, general suppression during binocular rivalry, and pattern rivalry. These data explanations follow from model brain mechanisms that assure non-rivalrous conscious percepts.     

Mechanisms selectively engaged in rivalry: normal vision habituates, rivalrous vision primes

When rivalrous stimuli are presented intermittently, perception stabilises. This indicates the operation of perceptual memory across interruptions in stimulation. Here we show that a percept under non-rivalrous and rivalrous conditions has qualitatively different effects on subsequent rivalrous vision. When an image is perceived under rivalrous viewing, that image is more likely to be perceived in later rivalrous viewing: an effect of stabilisation or priming. When the same image is perceived under non-rivalrous viewing conditions, it is less likely to be perceived again during subsequent rivalrous viewing: an effect of adaptation or habituation. When these stimuli possess different attributes to those in subsequent vision their effect declines. This suggests that visual rivalry might recruit mechanisms that are not engaged in 'normal' non-rivalrous vision but perhaps dedicated to the resolution of competing sensory information.     

Early correlates of visual awareness following orientation and colour rivalry

Binocular rivalry occurs when dissimilar images are presented to corresponding retinal regions of the two eyes: visibility alternates irregularly between the two images, interspersed by brief transitions when parts of both may be visible. We measured event-related potentials (ERPs) following binocular rivalry by changing the stimulus viewed by one eye to be identical to that in the other eye, eliciting binocular fusion. Because of the rivalry, observers either saw the change, when it happened to the visible stimulus, or did not see the change, when it happened to the invisible stimulus. The earliest ERP differences between visible and invisible changes occurred after about 100 ms (P1) when the rivalry was between stimuli differing in orientation, and after about 200 ms (N1) when the rivalry was between stimuli differing in colour. These differences originated from ventro-lateral temporal and prefrontal areas. We conclude that the rivalling stimulus property influences the timing of modulation of correlates of visual awareness in a property-independent cortical network.