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.     

Depth of interocular suppression associated with continuous flash suppression, flash suppression, and binocular rivalry

When conflicting images are presented to the corresponding regions of the two eyes, only one image may be consciously perceived. In binocular rivalry (BR), two images alternate in phenomenal visibility; even a salient image is eventually suppressed by an image of low saliency. Recently, N. Tsuchiya and C. Koch (2005) reported a technique called continuous flash suppression (CFS), extending the suppression duration more than 10-fold. Here, we investigated the depth of this prolonged form of interocular suppression as well as conventional BR and flash suppression (FS) using a probe detection task. Compared to monocular viewing condition, CFS elevated detection thresholds more than 20-fold, whereas BR did so by 3-fold. In subsequent experiments, we dissected CFS into several components. By manipulating the number and timing of flashes with respect to the probe, we found that the stronger suppression in CFS is not due to summation between BR and FS but is caused by the summation of the suppression due to multiple flashes. Our results support the view that CFS is not a stronger version of BR but is due to the accumulated suppressive effects of multiple flashes.     

Staying focused: a functional account of perceptual suppression during binocular rivalry

Presenting different images to either eye can induce perceptual switching, with alternating disappearances of each image--a phenomenon called binocular rivalry. We believe that disappearances during binocular rivalry can be driven by a process that facilitates visibility near the point of fixation. As the point of fixation is tied neither to a particular stimulus nor to a specific eye, indifference to both would be an essential characteristic for the process we envisage. Many factors that influence disappearances during binocular rivalry scale with distance in depth from fixation. Of these, here we use blur. We break the links between this cue and both eye of origin and stimulus type. We find that perceptual dominance can track a better focused image as it is swapped between the eyes and that perceptual switches can be driven by alternating the focus of images fixed in each eye. This implies that, as a determinant of suppression selectivity, blur is functionally independent from both eye of origin and stimulus type. Our data and theoretical account suggest that binocular rivalry is not an irrelevant laboratory curiosity but, rather, that it is a product of a functional adaptation that promotes visibility in cluttered environments.     

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.     

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.     

Contrast masking in strabismic amblyopia: attenuation, noise, interocular suppression and binocular summation

To investigate amblyopic contrast vision at threshold and above we performed pedestal-masking (contrast discrimination) experiments with a group of eight strabismic amblyopes using horizontal sinusoidal gratings (mainly 3c/deg) in monocular, binocular and dichoptic configurations balanced across eye (i.e. five conditions). With some exceptions in some observers, the four main results were as follows. (1) For the monocular and dichoptic conditions, sensitivity was less in the amblyopic eye than in the good eye at all mask contrasts. (2) Binocular and monocular dipper functions superimposed in the good eye. (3) Monocular masking functions had a normal dipper shape in the good eye, but facilitation was diminished in the amblyopic eye. (4) A less consistent result was normal facilitation in dichoptic masking when testing the good eye, but a loss of this when testing the amblyopic eye. This pattern of amblyopic results was replicated in a normal observer by placing a neutral density filter in front of one eye. The two-stage model of binocular contrast gain control [Meese, T.S., Georgeson, M.A. & Baker, D.H. (2006). Binocular contrast vision at and above threshold. Journal of Vision 6, 1224-1243.] was 'lesioned' in several ways to assess the form of the amblyopic deficit. The most successful model involves attenuation of signal and an increase in noise in the amblyopic eye, and intact stages of interocular suppression and binocular summation. This implies a behavioural influence from monocular noise in the amblyopic visual system as well as in normal observers with an ND filter over one eye.     

ERP evidence that surface-based attention biases interocular competition during rivalry

J. F. Mitchell, G. R. Stoner, and J. H. Reynolds (2004) observed that exogenously cuing one of two superimposed transparent surfaces resulted in an enhanced perceptual bias for the cued surface during binocular rivalry. We investigated the neural bases of this effect by recording event-related potentials (ERPs). Subjects viewed two superimposed rotating transparent surfaces and compared the directions of two successive translations, either both of the same surface or one of each surface. Following the first translation, which cued attention to the translating surface, two surface images were removed-one from each eye (dichoptic viewing) or both from one eye (monocular viewing). Subjects were impaired at comparing the first and second translations when they occurred on different surfaces, and the impairment was greater during dichoptic viewing (rivalry). The P1 component (110-160 ms) of the ERP elicited by the second translation of the same surface was larger than for the different surface during dichoptic but not monocular viewing. Larger cueing effects were also observed for the subsequent posterior N1 (160-220 ms) and P2 (250-300 ms) components during rivalry than during monocular viewing. These results are in line with a hybrid model of rivalry whereby cuing one surface initiates an earlier interocular selection process when the competing surfaces are presented to separate eyes.     

The time course of interocular suppression in normal and amblyopic subjects

The authors measured the time course of interocular suppression of five normal subjects and eleven patients with amblyopia (strabismic and/or anisometropic). Orthogonal gratings were presented dichoptically for durations that ranged from 10-6000 msec. All normal observers reported fusion or superimposition of the orthogonal gratings for short stimuli and reported binocular rivalry for stimuli longer than 150 msec. At long presentation times, all amblyopes constantly suppressed the pattern that was presented to their amblyopic eye. Six amblyopes showed superimposition of the two patterns at short presentation times. Of these, three had time courses similar to those of normal observers; the other three had a much shorter onset of suppression (about 80 msec). The remaining five amblyopes perceived only the pattern of the dominant eye at short stimulus durations; at intermediate durations, they reported partial superimposition of the stimuli, whereas at the longest stimulus durations, again only the stimulus of the dominant eye was perceived. The results suggest that binocular rivalry in normal observers and strabismic suppression in amblyopes are mediated by different mechanisms. The heterogeneity of the time courses of suppression in amblyopes might result from differences in the disturbances of early visual development (age at onset of strabismus and/or anisometropia, origin, and therapy).     

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.     

Intra- and interocular colour-specific activation during dichoptic suppression

A new psychophysical method, based on dichoptic suppression, is introduced to study intra- and interocular colour activations. An experiment is performed which shows that activation of a specific colour in one eye may cancel suppression of the same colour in the ipsi-lateral eye or in the contra-lateral eye at non-corresponding retinal locations. A control experiment shows that the effect is not due to an overall response-bias to the activated colour. The colour-specific intra- and interocular activation might play a role in various colour-based percepts known in the literature.     

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.     

The spatial extent of binocular suppression in normal and strabismic subjects

The authors investigated and specified the spatial extent of binocular suppression for normal and strabismic subjects at differing retinal eccentricities. Increment luminance thresholds for detecting a probe line which was presented to one eye were measured under the influence of a contralateral suppressing line. Probe sensitivities were obtained as a function of positional disparity of the contralateral suppressing line. Two normals and two strabismics served as subjects. The suppression patterns for the normal and strabismic subjects were similar except for the width of the suppression area. The interocular suppression functions were U-shaped, and there were significant effects of eccentricity. The spatial extent of suppression for normal subjects was between 2 and 20 min arc at the fovea or between 5 and 40 min arc at an eccentricity of 2 degrees. The maximum disparity which produced suppression effects in the strabismics was about half of that obtained for the normals, in either the foveal or near peripheral conditions. The psychophysically obtained U-shaped functions appear to be similar to the neuro-physiological data from cat striate cortex by Kato, Bishop & Orban (1981). Therefore, the authors believe that these results may reflect the operation of an interocular version of the Westheimer function.     

Visual evoked potentials during suppression in exotropic and esotropic strabismics: strabismic suppression objectified

Background We performed an electrophysiological study in order to objectify suppression in strabismus. The extent of cortical involvement in the process of interocular suppression was also explored. Possible differences in the suppressive process of esotropic and exotropic strabismics were also studied.Methods An electroencephalographic recorder with eight leads was applied to the posterior one-third of the skull; three occipital, three parietal, and two temporal leads. We measured the activity of these visual cortical areas during stimulation of each eye under monocular as well as binocular viewing conditions with hemisinusoidal light pulses in a nature-like complex visual background. Recordings were made from six primary esotropic strabismic subjects and four primary exotropic and one consecutive exotropic strabismic subject. Also, five normal controls were studied.Results A characteristic, triphasic response complex was found at approximately 80 ms following the start of each light pulse under monocular viewing conditions in the dominant and the nondominant eye. However, under dichoptic viewing conditions in the nondominant eye of all esotropic cases as well as in the nondominant eye of three of five exotropic cases, this response complex was completely absent. They showed approximately 100% reduction of their cortical response activity.Conclusions These results show the vast extent of the cortex that is involved in the suppressive process, giving a good insight in the power of suppression.     

Stimulus specificity in spatially-extended interocular suppression

In typical binocular rivalry demonstrations, disparate images presented in corresponding locations to the two eyes are found to alternate perceptually over time. Alternation in perception can occur even if the images presented to the two eyes do not overlap, if they are sufficiently close in space. This implies a spatial spread in the interocular interaction. The current set of experiments explores how the luminance pattern of a target, in relation to a rivalrous suppressor, affects its susceptibility to suppression. It was found that the susceptibility to suppression of a target pattern was nonlinearly related to the amount of luminance variation along the target in the direction perpendicular to the suppressing stimulus. For instance, there was a strong effect of the orientation of the grating pattern within the target on the total time of suppression, with much more suppression for horizontal gratings than vertical gratings when suppressor bars were oriented vertically, regardless of the luminance pattern within the suppressors. Furthermore, it was shown that the inclusion of a spatial gap between the vertical suppressors and the central portion of the target does more than simply change the spatial relationships, it adds new figural information, such as vertically orientated edges in the targets, that modify the susceptibility to suppression of the target, thereby interfering with measurements of spatial interaction functions. All of the results are consistent with selectively suppressing stimulus information that would interfere with stereoscopic matching to aid the binocular fusion of disparate retinal images.     

Unraveling adaptation and mutual inhibition in perceptual rivalry

When the visual system is confronted with incompatible images in the same part of the visual field, the conscious percept switches back and forth between the rivaling stimuli. Such spontaneous flips provide important clues to the neuronal basis for visual awareness. The general idea is that two representations compete for dominance in a process of mutual inhibition, in which adaptation shifts the balance to and fro. The inherent nonlinear nature of the rivalrous flip-flop and its stochastic behavior, however, made it impossible to disentangle inhibition and adaptation. Here we report a general method to measure the time course, and asymmetries, of mechanisms involved in perceptual rivalry. Supported by model simulations, we show the dynamics of opponent interactions between mutual inhibition and adaptation. The findings not only provide new insight into the mechanism underlying rivalry but also offer new opportunities to study and compare a wide range of bistable processes in the brain and their relation to visual awareness.     

Age-related deficits in attentional control of perceptual rivalry

Some aspects of attentional processing are known to decline with normal aging. To understand how age affects the attentional control of perceptual stability, we investigated age-related changes in voluntarily controlled perceptual rivalry. Durations of the dominant percept, produced by an ambiguous Rubin vase-faces figure, were measured in conditions that required passive viewing and attentional control: holding and switching the dominant percept. During passive viewing, mean dominance duration in the older group was significantly longer (63%) than the dominance duration found in the young group. This age-related deficit could be due to a decline in the apparent strength of the alternating percepts as a result of higher contrast gain of visual cortical activity and a reduction in the amount of attentional resources allocated to the ambiguous stimulus in older people compared to young adults. In comparison to passive viewing, holding the dominant percept did not significantly alter the dominance durations in the older group, while the dominance durations in the young group were increased (∼100%). The dominance durations for both age groups in switch conditions were reduced compared to their passive viewing durations (∼40%). The inability of older people to voluntarily prolong the duration of the dominant percept suggests that they may have abnormal attentional mechanisms, which are inefficient at enhancing the effective strength of the dominant percept. Results suggest that older adults have difficulty holding attended visual objects in focus, a problem that could affect their ability to carry out everyday tasks.     

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.