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.     

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.     

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.     

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.     

Binocular rivalry: a time dependence of eye and stimulus contributions

In binocular rivalry, the visual percept alternates stochastically between two dichoptically presented stimuli. It is established that both processes related to the eye of origin and binocular, stimulus-related processes account for these fluctuations in conscious perception. Here we studied how their relative contributions vary over time. We applied brief disruptions to rivalry displays, concurrent with an optional eye swap, at varying time intervals after one stimulus became visible (dominant). We found that early in a dominance phase the dominant eye determined the percept by stabilizing its own contribution (regardless of the stimulus), with an additional yet weaker stabilizing contribution of the stimulus (regardless of the eye). Their stabilizing contributions declined in parallel with time so that late in a dominance phase the stimulus (and in some cases also the eye-based) contribution turned negative, favoring a perceptual (or ocular) switch. Our findings show that depending on the time, first processes related to the eye of origin and then those related to the stimulus can have a greater net influence on the stability of the conscious percept. Their co-varying change may be due to feedback from image- to eye-of-origin representations.     

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.     

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.     

The influence of chromatic context on binocular color rivalry: perception and neural representation

The predominance of rivalrous targets is affected by surrounding context when stimuli rival in orientation, motion or color. This study investigated the influence of chromatic context on binocular color rivalry. The predominance of rivalrous chromatic targets was measured in various surrounding contexts. The first experiment showed that a chromatic surround's influence was stronger when the surround was uniform or a grating with luminance contrast (chromatic/black grating) compared to an equiluminant grating (chromatic/white). The second experiment revealed virtually no effect of the orientation of the surrounding chromatic context, using chromatically rivalrous vertical gratings. These results are consistent with a chromatic representation of the context by a non-oriented, chromatically selective and spatially antagonistic receptive field. Neither a double-opponent receptive field nor a receptive field without spatial antagonism accounts for the influence of context on binocular color rivalry.     

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.     

The initial interactions underlying binocular rivalry require visual awareness

Current theories of binocular vision suggest that the neural processes that resolve interocular conflict do not involve a single brain region but occur at multiple stages of visual processing. Here, using an adaptation paradigm, we explore the initial mechanisms involved in selecting a stimulus for perceptual dominance during binocular rivalry. When one or both eyes briefly viewed an adapting grating stimulus prior to the presentation of the adapting grating to one eye and an orthogonal, non-adapted grating to the other eye, participants more often reported perceptual dominance of the non-adapted grating. Crowding reduced awareness of the adapting grating. On trials in which subjects were unaware of the orientation of the adaptor grating, there was no effect of the adaptor on perceived dominance during rivalry; participants were just as likely to report dominance of the adapted or non-adapted grating. This implies that the initial events in binocular rivalry involve later stages of visual processing.     

Attention-based perceptual learning increases binocular rivalry suppression of irrelevant visual features

Perceptual learning refers to an improvement on a perceptual task after repeated exposure to a stimulus. It has been shown that attention can play an important role in perceptual learning. Recently, it has been suggested that training can lead to increased suppression of information that is continuously irrelevant, and that this attention-based suppression plays an important role in more efficient noise exclusion. Here we investigate this claim. Observers performed a visual speed-discrimination task for 5 consecutive days. After training, sensitivity to motion directions that were relevant, irrelevant, or neutral toward the training task was assessed by measuring motion coherence thresholds. In addition, perceptual dominance during binocular rivalry was assessed for combinations of the three motion directions. The results showed that sensitivity to the task-relevant feature increased due to training. That is, motion coherence thresholds were selectively lowered for the task-relevant feature. Interestingly, the feature that was task-irrelevant during training was more strongly suppressed during binocular rivalry: The mean perceptual dominance of this feature was selectively decreased. Our results show that task-irrelevant information that potentially interferes with the primary task during learning gets more strongly suppressed. Furthermore, our results add new evidence in support of the claim that mechanisms involved in visual attention and binocular rivalry overlap.     

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.     

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.     

Binocular rivalry between identical retinal stimuli with an induced color difference

An open question in color rivalry is whether alternation between two colors is caused by a difference in receptoral stimulation or a difference in the neural representation of color appearance. This question was examined with binocular rivalry between physically identical lights that differed in appearance due to chromatic induction. Perceptual alternation was measured between gratings of the same chromaticity; each one was presented within a different patterned surround that caused the gratings, one to each eye, to appear unequal in hue because of chromatic induction. The gratings were presented dichoptically with binocular disparity so the rivalrous gratings appeared in front of the surround. Perceptual alternation in hue was found for the two physically identical chromaticities. Stereoscopic depth also was perceived, corroborating binocular neural combination despite color rivalry (Treisman, 1962). The results show that color rivalry is resolved after color-appearance shifts caused by chromatic context, and that color rivalry does not require competing unequal cone excitations from the rivalrous stimuli.     

The relationship between binocular rivalry and strabismic suppression

Increment-threshold spectral sensitivity functions were determined for normal observers during binocular rivalry and for esotropic observers during strabismic suppression and under viewing conditions that normally induce binocular rivalry. Depending on the spatial and temporal properties of the test stimulus, the normal observers exhibited a wavelength-specific loss in sensitivity during the suppression phase of rivalry, which suggests that binocular rivalry differentially attenuates the sensitivity of the chromatic mechanisms relative to the luminance mechanisms. In contrast, regardless of the test stimulus dimensions, the esotropic observers did not manifest a wavelength-specific loss in sensitivity either during strabismic suppression or under conditions that normally induce binocular rivalry. The different patterns of suppression shown by the normal and esotropic subjects suggest that strabismic observers do not demonstrate normal binocular rivalry, and that strabismic suppression and normal binocular rivalry suppression are mediated by different neural mechanisms.     

Stochastic resonance in binocular rivalry

When a different image is presented to each eye, visual awareness spontaneously alternates between the two images--a phenomenon called binocular rivalry. Because binocular rivalry is characterized by two marginally stable perceptual states and spontaneous, apparently stochastic, switching between them, it has been speculated that switches in perceptual awareness reflect a double-well-potential type computational architecture coupled with noise. To characterize this noise-mediated mechanism, we investigated whether stimulus input, neural adaptation, and inhibitory modulations (thought to underlie perceptual switches) interacted with noise in such a way that the system produced stochastic resonance. By subjecting binocular rivalry to weak periodic contrast modulations spanning a range of frequencies, we demonstrated quantitative evidence of stochastic resonance in binocular rivalry. Our behavioral results combined with computational simulations provided insights into the nature of the internal noise (its magnitude, locus, and calibration) that is relevant to perceptual switching, as well as provided novel dynamic constraints on computational models designed to capture the neural mechanisms underlying perceptual switching.     

Eye- and feature-based modulation of onset rivalry caused by the preceding stimulus

Pre-exposure to a stimulus can modulate initial perceptual dominance experienced in binocular rivalry with brief test stimuli (onset rivalry). This study investigated this modulating effect using both color and pattern stimuli. We confirmed separate contributions of eye- and feature-based suppressions and showed that their relative strength varied with temporal parameters. Eye-based suppression was stronger with a short test duration (10 ms) and shorter ISIs between the preceding and test stimuli. On the other hand, feature-based suppression grew with ISI and was more pronounced with a longer test duration (200 ms). We also investigated the nature of the modulating effect associated with feature-based suppression using chromatic gratings of high luminance contrast. Results revealed that different features of the preceding stimulus (i.e., color and orientation) exerted nearly independent effects on onset rivalry. However, different features shared their fate in competitive interactions for perceptual dominance; when one feature became dominant, the other also dominated. These findings suggest that competitive interactions for perceptual dominance and the modulation of these interactions are mediated at least partially by different mechanisms. Overall, the present findings are consistent with a theoretical view that initial dominance is established through competitive interactions at multiple levels of 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.     

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.     

Dichoptic masking and binocular rivalry share common perceptual dynamics

Two of the strongest tools to manipulate visual awareness of potentially salient stimuli are binocular rivalry and dichoptic masking. Binocular rivalry is induced by presenting incompatible images to the two eyes over prolonged periods of time, leading to an alternating perception of the two images. Dichoptic masking is induced when two images are presented once in rapid succession, leading to the perception of just one of the images. Although these phenomena share some key characteristics, most notably the ability to erase from awareness potentially very salient stimuli, their relationship is poorly understood. We investigated the perceptual dynamics during long-lasting dynamic stimulation leading to binocular rivalry or dichoptic masking. We show that the perceptual dynamics during dichoptic masking conditions meet the classifiers used to classify a process as binocular rivalry; that is, (1) Levelt's 2nd proposition is obeyed; (2) perceptual dominance durations follow a gamma distribution; and (3) dominance durations are sequentially independent. We suggest that binocular rivalry and dichoptic masking may be mediated by the same inhibitory mechanisms.