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.     

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.     

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.     

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.     

Preserved gain control for luminance contrast during binocular rivalry suppression

Binocular rivalry elevates contrast increment thresholds for the detection of a transient stimulus presented to the suppressed eye, while thresholds measured during dominance are identical to those during monocular viewing (e.g. [Wales, R., & Fox, R. (1970). Increment detection thresholds during binocular rivalry suppression. Perception and Psychophysics, 8, 90-94]). It is well established that contrast increment thresholds depend on reference (pedestal) contrast. With high contrasts, increment thresholds increase with pedestal contrast, reflecting a gain control with sigmoidal non-linearity. We examined how this gain control mechanism operates during binocular rivalry (i.e., with and without perception of a pedestal mask). Subjects viewed a horizontal sine-wave grating (steady pedestal) and a radial checkerboard dichoptically. When the grating achieved a pre-specified phenomenal state (dominance or suppressed), subjects initiated the transient presentation (500-ms Gaussian pulse) of a contrast increment of the same spatial frequency. The pulse appeared in either the upper or lower half of the pedestal. Subjects indicated which half of the pedestal contained the pulse. Contrast increment thresholds were measured using a staircase method with various pedestal contrasts, which yielded threshold versus contrast (TvC) functions during dominance and suppression. The measured thresholds were reliably higher during suppression, but the rising slopes of TvC functions did not differ significantly between dominance and suppression (i.e., constant upward shift of TvC function). A control experiment demonstrated that the TvC function during dominance was identical to that during non-rivalry, monocular viewing. Evidently, the contrast gain control for transient luminance increment does not require the perception of pedestal contrast.     

Apparent motion can survive binocular rivalry suppression

For short-range motion, observers dichoptically viewed a random-dot cinematogram and a rival target. Upon keypress, the first frame of the cinematogram was replaced by the second frame. Observers judged the direction of motion, which was governed by the initial position of the central region. Performance was well above chance during both dominance and suppression. For long-range motion, observers rated the motion produced by sequentially flashing two small spots, with the first spot contained within a rivalrous region. Suppression reduced but did not prevent perception of this motion. Presenting the second motion frame to both eyes weakened both forms of motion.     

Saliency in a suppressed image affects the spatial origin of perceptual alternations during binocular rivalry

During binocular rivalry, perception alternates between dichoptically presented incompatible images. With larger images, such perceptual alternations will typically start locally and then gradually spread across the image, known as traveling waves of perceptual dominance. Several image-features (such as local contrast) are known to determine where in the image a traveling wave originates. Here we investigate whether orientation contrast in the suppressed image affects these spatial origin(s) of perceptual alternations. The results show that the origins are increasingly biased towards locations of increasing orientation contrast in the suppressed image. This increase in bias is related to the efficiency of visual search for the orientation contrast, tested offline: we find large biases towards orientation contrast when visual search for it is efficient, and small biases when search for it is inefficient. Our results imply that rivalry suppression is not homogenous across the suppressed image, but is dependent on local image-features in the suppressed image. The relation between spatial bias and visual search performance suggests that spatial origins of perceptual alternations are biased to salient locations in the suppressed image. Moreover, the finding that saliency affects the spatial origin of a perceptual alternation is in agreement with the idea that saliency is represented at a monocular, unconscious level of visual processing.     

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.     

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.     

Flash suppression and flash facilitation in binocular rivalry

We show that previewing one half image of a binocular rivalry pair can cause it to gain initial dominance when the other half is added, a novel phenomenon we term flash facilitation. This is the converse of a known effect called flash suppression, where the previewed image becomes suppressed upon rivalrous presentation. The exact effect of previewing an image depends on both the duration and the contrast of the prior stimulus. Brief, low-contrast prior stimuli facilitate, whereas long, high-contrast ones suppress. These effects have both an eye-based component and a pattern-based component. Our results suggest that, instead of reflecting two unrelated mechanisms, both facilitation and suppression are manifestations of a single process that occurs progressively during presentation of the prior stimulus. The distinction between the two phenomena would then lie in the extent to which the process has developed during prior stimulation. This view is consistent with a neural model previously proposed to account for perceptual stabilization of ambiguous stimuli, suggesting a relation between perceptual stabilization and the present phenomena.     

Dynamics of perceptual filling-in of visual phantoms revealed by binocular rivalry

How do selective and constructive visual mechanisms interact to determine the outcome of conscious perception? Binocular rivalry involves selective perception of one of two competing monocular images, whereas visual phantoms involve perceptual filling-in between two low-contrast collinear gratings. Recently, we showed that visual phantoms lead to neural filling-in of activity in V1 and V2, which can be dynamically gated by rivalry suppression (M. Meng, D. A. Remus, & F. Tong, 2005). Here, we used psychophysical methods to study the temporal dynamics of filling-in, by applying rivalry or flash suppression to trigger the suppression or appearance of visual phantoms. Experiments revealed that phantom filling-in involves an active, time-dependent process that depends on the phenomenal visibility of the phantom-inducing gratings. Shortly after the inducing gratings became dominant during rivalry, the likelihood of perceiving phantoms in the intervening gap increased over time, with larger gaps requiring more time for filling-in. In contrast, suppression of the inducing gratings promptly led to the disappearance of visual phantoms, with response times independent of gap size. The fact that binocular rivalry can prevent the formation of visual phantoms rules out the possibility that rivalry suppression occurs after the site of phantom filling-in. This study provides novel evidence that visual phantoms result from a slow time-dependent filling-in mechanism; possible models to account for its time course are discussed.     

Grouping visual features during binocular rivalry

During binocular rivalry, portions of one eye's view may be perceptually dominant while other portions are suppressed; at any given moment, overall dominance often resembles a patchwork mixture of the two eyes' views. This study investigates the potency of two Gestalt grouping cues--good continuation and common fate--to promote synchronous fluctuations in dominance of two, spatially separated rival targets. Two grating patches were presented to the left eye paired dichoptically with random-dot patches presented to corresponding right eye locations. The orientations of the two gratings were either collinear, parallel or orthogonal. Gratings underwent contrast modulations that were either correlated (identical contrast changes) or uncorrelated (independent contrast changes). Over 60 s trials, observers pressed one key when the left grating predominated, another when the right grating predominated and both keys when both were concurrently visible. Correlated contrast modulation promoted joint grating predominance relative to the uncorrelated conditions, an effect strongest for collinear gratings. Joint predominance depended strongly on the angular separation between gratings and the temporal phase-lag in contrast modulations. These findings may reflect neural interactions subserved by lateral connections between cortical hypercolumns.     

Inhibitory effect of binocular rivalry suppression is independent of orientation

A series of experiments examined the relationship between the inhibitory effect of binocular rivalry suppression and the difference in orientation of dichoptically viewed gratings. Using forced-choice techniques, it was found that test-probe detection was impaired during periods of suppression by an amount which was invariant with orientation differences. These data are inconsistent with a recently proposed neural model of binocular suppression based on inhibition between cortical feature detectors.     

Clinical suppression and binocular rivalry suppression: the effects of stimulus strength on the depth of suppression

In observers with abnormal binocular vision (such as strabismics or anisometropes) one eye's view is often suppressed. This clinical suppression serves to eliminate binocular diplopia and confusion. Suppression may also occur in observers with normal binocular vision, when the two eyes view disparate retinal images, a phenomenon known as binocular rivalry. When the image in an eye is suppressed, it is possible to determine the amount by which that suppressed stimulus is below the visibility threshold, or the depth of suppression. In the experiments presented here, the depth of suppression in an eye was measured as the strength of the stimulus in the contralateral eye (the stimulus inducing suppression) was varied. This was done for both clinical suppressors and normal observers undergoing binocular rivalry suppression. Independent changes were made to the contrast, the luminance, and the spatial frequency of the inducing stimulus. For both clinical suppression and binocular rivalry suppression, the depth of suppression was constant, regardless of the changes to the inducing stimulus.     

Effect of binocular rivalry suppression on initial ocular following responses

To study the effect of binocular rivalry (BR) suppression on the ocular following response (OFR), we recorded the OFR in both the suppressed and the dominant phases of BR. The BR was established using stationary horizontal/vertical grating patterns presented on two PC monitors. Once a subjective image of a vertical or horizontal grating pattern was perceived, subjects pressed a button to trigger an onset of brief horizontal movement (750 ms) of the vertical grating pattern and an offset of the horizontal pattern. The OFRs were recorded using a scleral search coil system at 1 kHz. The OFRs from the suppressed phases were significantly reduced compared to those from the dominant phases. The OFRs were asymmetrical to temporalward and nasalward motion in most conditions. We suggest that asymmetry of OFRs under the incomplete BR conditions may be a reflection of imbalance binocular inputs and processing in the visual system similar to asymmetrical optokinetic nystagmus in strabismic subjects. The latency of the OFR in deeper suppressed conditions was prolonged, suggesting that the interaction of BR and OFR may occur at multiple stages including an early stage of the visual processing. The OFR may have the potential for objective measurement of BR suppression in clinical evaluation of binocular function.     

Effect of binocular rivalry suppression on the motion aftereffect.

The relative loci within the visual system of the site of the motion aftereffect (MAE) and the site of binocular rivalry suppression was inferred by measuring the magnitude of the MAE when the inducing motion was phenomenally suppressed for > 50 per cent of the inspection period. The MAE magnitude was a function of the duration of physical impingement of the inducing stimulus; the state of suppression exerted no effect, thereby implying that the site of suppression does not occur before the site of the MAE. This result, together with other data, is interpreted to mean that the site of suppression is cortical.     

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.     

Conditions of perceptual selection and suppression during interocular rivalry in strabismic and normal cats

Presenting the two eyes with incongruent stimuli leads to the phenomenon of interocular rivalry. At any given time, one of the stimuli is perceptually suppressed in order to avoid double vision. In squinting subjects, rivalry occurs permanently also for congruent stimuli because of developmental rearrangement of cortical circuitry. In this study, we have investigated the dynamics and stimulus dependence of rivalry in six esotropic, four exotropic and three non-strabismic cats. As an indicator for perception, we used optokinetic nystagmus that was induced by moving gratings. The esotropic cats were tested for their visual acuity by means of a jumping stand procedure. The results show that one eye can dominate perception even if both eyes have equal visual acuity and are presented with stimuli of equal contrast. Strong eye dominance asymmetry was found in all but one of the tested cats. Notably, all three of the normal cats showed a clear asymmetry in perceptual selection. Measurements with varying contrast and velocity of the stimuli revealed that the influence of these parameters on perceptual selection was independent of the presence of strabismus. In all cats, the time during which a given eye dominated perception increased with the contrast and decreases with the velocity of the stimulus presented to this eye.     

Conflict defined by global gestalt can modulate binocular rivalry suppression

Binocular rivalry suppression is thought to necessarily require local interocular conflict: the presence of incompatible image elements, such as orthogonal contours, in retinally corresponding regions of two monocular displays. Whether suppression can also be driven by conflict at the level of spatially nonlocal surface or object representations is unclear. Here, we kept local contour conflict constant while varying global conflict, defined by the gestalt formed by the two monocular displays. Specifically, each eye was presented with a grid of image elements (crosses or plusses), placed such that the two eyes’ individual grid elements did not directly overlap but the grids as a whole did. In a “shared motion” condition, all elements moved in unison, inviting a gestalt made up of all elements across both eyes; in a “different motions” condition, the elements’ trajectories differed between eyes, inviting a gestalt of two overlapping surfaces, each associated with one eye. Perceptual disappearances of image elements occurred more readily in the different motions condition, an observation that could not be explained by any between-condition differences in local contour conflict. In a second experiment, we furthermore established that, whereas perceptual disappearances in the shared motion condition tended to involve a single element at a time, in the different motions condition, multiple elements belonging to the same gestalt often disappeared together. These findings indicate that, even though binocular rivalry may critically rely on inhibition due to locally incompatible image elements, this inhibition also depends on the global gestalt to which these elements contribute.