Shifting spatial attention makes you flip: Exogenous visual attention triggers perceptual alternations during binocular rivalry

Although it has been argued that visual attention and the dynamics of binocular rivalry are closely linked, strong evidence for this proposition is still lacking. Here, we investigate how perceptual alternations during binocular rivalry are affected by spatial attention by employing a cuing paradigm. We show a tight link between the occurrence of perceptual alternations and the spatiotemporal properties of visual attention: Alternations occurred earlier and more frequently at locations where visual attention was summoned by an exogenous cue. We argue that cuing a location where rival images are presented leads to a transient increase in the effective contrast of these rival images. This transient increase in effective contrast increases the probability of an alternation at that location. Furthermore, we suggest that an occipito-fronto-parietal network known to be involved in selective attention and binocular rivalry mediates perceptual alternations by boosting the neural response at attended locations.     

Binocular rivalry alternation rate declines with age

The effect of aging on the time of spontaneous perceptual alternation in binocular rivalry was examined in 59 subjects. An earlier study reported the change of alternation time by comparing middle-age and elderly subjects. We also observed age-related prolongation in alternation time by comparing subjects in a lower age group (20-34 years) with those in both a middle-age group (35-49 years) and a higher age group (50-64 years). Aging of visual optical functions such as presbyopia or the reduction of contrast sensitivity has an accelerating effect over age and may not be related to the age-associated monotonic prolongation of alternation time in binocular rivalry. The origin of aging in binocular rivalry is still unclear but a neural interval generator for perceptual switching is suggested.     

Afterimages, binocular rivalry, and the temporal properties of dominance and suppression

When different contours are presented to the two eyes, an unstable percept, binocular rivalry, is the result. Parts of each set of contours may be seen but the two sets are not seen in the same place at the same time. The contours need not be physically present. Afterimages will produce binocular rivalry. Normal rivalry can be prevented if intermittent stimulation is used. Previous work has shown that orthogonal gratings, flashed for less than 150 ms and separated by more than 150 ms, will appear to fuse into a plaid or checkerboard pattern. In the present experiment this phenomenon is examined with afterimages used to produce rivalry. This abnormal 'fusion' is seen when negative afterimages are stroboscopically illuminated at less than 5 Hz. The results obtained with afterimages are predictable from the previous results obtained with stimuli external to the eye.     

Attention speeds binocular rivalry

During binocular rivalry, incompatible images presented dichoptically compete for perceptual dominance. It has long been debated whether binocular rivalry can be controlled by attention. Most studies have shown that voluntary control over binocular rivalry is limited. We sought to remove attention from binocular rivalry by presenting a concurrent task. Diverting attention slowed the rivalry alternation rate, and did so in proportion to the difficulty of the concurrent task. Even a very demanding distractor task, however, did not arrest rivalry alternations completely. Given that diverting attention was equivalent to lowering the contrast of the rival stimuli, the ability of attention to speed binocular rivalry is most likely due to an increase in the effective contrast of the stimuli through boosting the gain of the cortical response. This increase in effective contrast will ultimately lead to a perceptual switch, thereby limiting voluntary control. Thus, attention speeds rivalry alternations, but has no inherent control over the rivalry process.     

Monocular and binocular rivalry between contours.

The temporal characteristics of binocular and monocular rivalry between orthogonal gratings of the same or complementary colours were investigated. Rivalry was measured in terms of the dominance of either grating or the visibility of composites comprised of parts of both gratings. The total duration for which either grating was dominant was significantly longer in binocular rivalry between gratings of complementary colours. A comparison of binocular and monocular rivalry indicated considerable phenomenal differences between them. Dominance in binocular rivalry corresponds to the visibility of one grating alone; this occurs rarely in monocular rivalry, which is characterized by fluctuations in the distinctiveness of the gratings. The changes in distinctiveness are influenced by colour in a similar manner to that in binocular rivalry, and the frequencies of fluctuations are higher for gratings of complementary colours.     

Determinants of fusion of dichoptically presented orthogonal gratings

Liu, Tyler, and Schor (1992 Vision Research 32 1471-1479) reported the surprising finding that dichoptically presented orthogonal sine-wave gratings do not always produce binocular rivalry. Gratings of high spatial frequency, and especially of low contrast, fuse to produce a stable percept of a dichoptic plaid. Using a somewhat different perceptual task, we replicated those findings and extended them. The probability of a plaid percept is higher for square-wave gratings than for sine-wave gratings, and higher still for rectangular-wave gratings with high duty cycles (with very thin light or dark bars). Experiments were conducted to test whether this duty-cycle effect was due to changes in overall luminance, or in the size of the regions of luminance congruity (which may reduce the probability of rivalry), but no such effects could account for the results. The presence of locally conflicting contour information in the two eyes was shown to be an important determinant of rivalry onset, but, since removing such regions did not eliminate rivalry, other factors also have a role to play. The spatial frequency composition of the gratings is one such factor which is consistent with all of the findings we report.     

Predictive coding explains binocular rivalry: an epistemological review

Binocular rivalry occurs when the eyes are presented with different stimuli and subjective perception alternates between them. Though recent years have seen a number of models of this phenomenon, the mechanisms behind binocular rivalry are still debated and we still lack a principled understanding of why a cognitive system such as the brain should exhibit this striking kind of behaviour. Furthermore, psychophysical and neurophysiological (single cell and imaging) studies of rivalry are not unequivocal and have proven difficult to reconcile within one framework. This review takes an epistemological approach to rivalry that considers the brain as engaged in probabilistic unconscious perceptual inference about the causes of its sensory input. We describe a simple empirical Bayesian framework, implemented with predictive coding, which seems capable of explaining binocular rivalry and reconciling many findings. The core of the explanation is that selection of one stimulus, and subsequent alternation between stimuli in rivalry occur when: (i) there is no single model or hypothesis about the causes in the environment that enjoys both high likelihood and high prior probability and (ii) when one stimulus dominates, the bottom-up, driving signal for that stimulus is explained away while, crucially, the bottom-up signal for the suppressed stimulus is not, and remains as an unexplained but explainable prediction error signal. This induces instability in perceptual dynamics that can give rise to perceptual transitions or alternations during rivalry.     

Stochastic properties of binocular rivalry alternations

Previous researches have demonstrated that the successive phase durations in binocular rivalry are independent. These findings are confirmed and extended to chromatic stimuli. The nature of the function that is shown to describe the distribution of the dominance phase durations is consistent with the independence of successive phases and suggests that a parallel may exist between binocular rivalry and the perceptual reversal of ambiguous figures.

     

The role of voluntary and involuntary attention in selecting perceptual dominance during binocular rivalry

When incompatible images are presented to corresponding regions of each eye, perception alternates between the two monocular views (binocular rivalry). In this study, we have investigated how involuntary (exogenous) and voluntary (endogenous) attention can influence the perceptual dominance of one rival image or the other during contour rivalry. Subjects viewed two orthogonal grating stimuli that were presented to both eyes. Involuntary attention was directed to one of the grating stimuli with a brief change in orientation. After a short period, the cued grating was removed from the image in one eye and the uncued grating was removed from the image in the other eye, generating binocular rivalry. Subjects usually reported dominance of the cued grating during the rivalry period. We found that the influence of the cue declined with the interval between its onset and the onset of binocular rivalry in a manner consistent with the effect of involuntary attention. Finally, we demonstrated that voluntary attention to a grating stimulus could also influence the ongoing changes in perceptual dominance that accompany longer periods of binocular rivalry Voluntary attention did not increase the mean dominance period of the attended grating, but rather decreased the mean dominance period of the non-attended grating. This pattern is analogous to increasing the perceived contrast of the attended grating. These results suggest that the competition during binocular rivalry might be an example of a more general attentional mechanism within the visual system.     

Influence of spatial frequency, luminance, and duration on binocular rivalry and abnormal fusion of briefly presented dichoptic stimuli

Orthogonal gratings, presented dichoptically, do not fuse into a single percept. Parts of each are seen while other parts are suppressed in an unstable perception (binocular rivalry). However, it has been previously noted that, if the gratings are briefly flashed, they will appear to fuse into a plaid or checkerboard pattern. Three experiments are reported which have been designed to define more clearly the spatial and temporal parameters of this effect in the hope that this would lead to better understanding of the normal mechanisms of dominance and suppression. Stimuli appear fused if flashed for less than 150 ms. The effect is independent of substantial changes in spatial frequency and luminance. Single flashes that appear fused when presented in isolation produce rivalry if separated by intervals less than about 150 ms. Intervals greater than 150 ms produce continued abnormal fusion. Possible mechanisms are discussed.     

Selective attention and the control of binocular rivalry

The control of binocular rivalry (BR) was reintroduced as an example of selective attention after several decades. Ss passively observed BR in a stereoscopic tachistoscope and, at an undetermined phase of rivalry, received an instruction signal to attend to one of the rivalry patterns. Letter test stimuli were presented to either the instructed (I) eye or noninstructed (NI) eye 1, 3, 5, or 7 sec following the instruction signal. The difference between the eyes in recognition performance (I-NI) provided an objective measure of control, which was correlated with previously used measures based on subjective reports of rivalry alternation rates. Subjective and objective measures showed correlated increases as a result of practice of control. The fact that the objective measure reached a peak value at the 3-sec delay of test stimuli suggested a minimum time to shift attention and maximum time to maintain attention in BR. In addition, it was found that the control of BR results in a change of signal strength with no associated change in the use of confidence ratings, a result similar to that found with selective attention in dichotic listening.     

Binocular rivalry and surface-boundary processing

Theoretical and empirical studies show that the visual system relies on boundary contours and surface features (e.g. textures) to represent 3-D surfaces. When the surface to be represented has little texture information, or has a periodic texture pattern (grating), the boundary contour information assumes a larger weight in representing the surface. Adopting the premise that the mechanisms of 3-D surface representation also determine binocular rivalry perception, the current paper focuses on whether boundary contours have a similar role in binocular rivalry. In experiment 1, we tested the prediction that the visual system prefers selecting an image/figure defined by boundary contours for rivalry dominance. We designed a binocular rivalry stimulus wherein one half-image has a boundary contour defined by a grating disk on a background with an orthogonal grating orientation. The other half-image consists solely of the (same orientation) grating background without the grating disk, ie no boundary contour. Confirming our prediction, the predominance for the half-image with the grating disk is approximately 90%, despite the fact that the grating disk corresponds to an area with orthogonal grating in the fellow eye. The advantage of the grating disk is dramatically reduced to about 50% predominance when a boundary contour is added to the background-only half-image at the location corresponding to the grating disk. We attribute this reduced advantage to the formation of a corresponding binocular boundary contour. In experiment 2 the grating background was substituted by a random-dot background in a similar stimulus design. We found that the perceptual salience of the corresponding binocular boundary contours extracted by the interocular matching process is an important factor in determining the dynamics of binocular rivalry. Experiment 3 showed that vertical lines with uneven thickness and spacing as the background reduce the contribution of the monocular boundary contour of the grating disk in binocular rivalry, possibly through the formation of binocular boundary contours between the local edges (vertical components) of the vertical lines and the corresponding grating disk.     

The effect of orientation in binocular contour rivalry of real images and afterimages

Binocular rivalry was investigated using gratings of different orientations in three experiments. No consistent effects of orientation were found for predominance measures of rivalry between real images. Rivalrous afterimages, on the other hand, did exhibit orientation selectivity: vertical gratings were visible for longer than were 45-deg gratings. This effect was compared to the similar orientation selectivity found for monocular observation of grating afterimages. Comparisons of binocular rivalry between real images and afterimages were made in terms of the frequency distributions of the dominance periods.     

Seeing it both ways: Openness to experience and binocular rivalry suppression

Openness to experience is characterised by flexible and inclusive cognition. Here we investigated whether this extends to basic visual perception, such that open people combine information more flexibly, even at low-levels of perceptual processing. We used binocular rivalry, where the brain alternates between perceptual solutions and times where neither solution is fully suppressed, mixed percept. Study 1 showed that openness is positively associated with duration of mixed percept and ruled out the possibility of response bias. Study 2 showed that mixed percept increased following a positive mood induction particularly for open people. Overall, the results showed that openness is linked to differences in low-level visual perceptual experience. Further studies should investigate whether this may be driven by common neural processes.     

The temporal course of suppression during binocular rivalry

Orthogonally oriented sinusoidal luminance gratings were dichoptically presented to the observers' left and right eyes. During the subsequent binocular rivalry, a small target was briefly presented (4AFC) to probe the strength of interocular suppression at various temporal latencies. Both stationary and moving rivalrous patterns were investigated. The purpose of experiment 1 was to compare the temporal characteristics of stationary and motion rivalry (0 and 1.2 deg s-1), while that of experiment 2 was to examine rivalry suppression for higher speeds (2 and 4 deg s-1). In all cases, it was found that the strength of suppression remained essentially constant throughout a single phase of binocular rivalry. The results of the investigation also revealed that moving rivalrous patterns lead to greater magnitudes of interocular suppression than static patterns. Despite these differences in the strength of suppression, the results of both experiments show that the temporal characteristics of motion and static rivalry are essentially identical.     

Binocular rivalry and visual awareness: the role of attention

When the right eye and the left eye view dissimilar scenes, the observer does not experience a stable superimposed percept of the images presented to the two eyes, but instead perceives an alternation between the images seen by each eye. A critical question confronting this robust and intriguing phenomenon of binocular rivalry is how the visual system selects the image to be perceived (dominant). The current main-stream literature emphasizes a bottom-up explanation in which the rivalry stimulus with the higher contour strength has the advantage, and becomes dominant in rivalry. Nevertheless, some workers in the past have favored an attention-selection explanation for binocular rivalry. We investigated the role of attention in binocular rivalry by employing novel psychophysical paradigms which capitalized on several established phenomena (e.g. the Cheshire Cat effect, attention cueing, pop-out effect). Our results revealed two major aspects of attention modulation in binocular rivalry. We found that a dominant image is less likely to be suppressed when voluntary attention is directed to it. This suggests the role of voluntary attention in retaining the dominant image in visual awareness. Second, a rivalry stimulus is more likely to become dominant if accompanied by a pop-out cue (in the same eye and proximity). Since a pop-out cue attracts involuntary attention to its location/eye, this result suggests that cue-mediated involuntary attention can promote the ability of a rivalry stimulus to reach visual awareness.     

Emotional pictures predominate in binocular rivalry

This study used binocular rivalry to examine whether emotionally relevant pictures are processed preferentially in the visual system. Binocular rivalry occurs when two incompatible pictures are presented to both eyes. If the pictures cannot be merged to a single unambiguous percept, binocular rivalry results in an alternation of the two percepts. However, it has been controversial whether emotional picture content can influence a picture's relative predominance. Twenty pairs of emotional and neutral pictures from the International Affective Picture System (IAPS) were presented to 45 healthy participants. As hypothesised, emotional pictures significantly predominated over neutral pictures, both with respect to the first percept and the total viewing time across each trial. We conclude that the relative predominance of emotional pictures in binocular rivalry provides further support for the preferential processing of emotional stimuli in the visual system.     

Emotion space as a predictor of binocular rivalry

In our 1999 report, we examined robustness of a two-dimensional structure of facial expressions of emotion under the condition of some perceptual ambiguity, using a stereo-scope. The current study aimed to replicate and extend the previous work by adding facial photographs of different persons and by measuring participants' perception of stereoscopically presented faces. Multidimensional scaling provided a two-dimensional configuration of facial expressions comparable with the previous studies. Although binocular rivalry was a less frequent phenomenon, it was suggested that the distances between facial expressions in the derived space were a contributing factor in eliciting binocular rivalry.     

Conscious perception of flickering stimuli in binocular rivalry and continuous flash suppression is not affected by tACS-induced SSR modulation

The content of conscious perception is known to correlate with steady-state responses (SSRs), yet their causal relationship remains unclear. Can we manipulate conscious perception by directly interfering with SSRs through transcranial alternating current stimulation (tACS)? Here, we directly addressed this question in three experiments involving binocular rivalry and continuous flash suppression (CFS). Specifically, while participants (N = 24) viewed either binocular rivalry or tried to detect stimuli masked by CFS, we applied sham or real tACS across parieto-occipital cortex at either the same or a different frequency and phase as an SSR eliciting flicker stimulus. We found that tACS did not differentially affect conscious perception in the forms of predominance, CFS detection accuracy, reaction time, or metacognitive sensitivity, confirmed by Bayesian statistics. We conclude that tACS application at frequencies of stimulus-induced SSRs does not have perceptual effects and that SSRs may be epiphenomenal to conscious perception.     

Neural bases of binocular rivalry

During binocular rivalry, conflicting monocular images compete for access to consciousness in a stochastic, dynamical fashion. Recent human neuroimaging and psychophysical studies suggest that rivalry entails competitive interactions at multiple neural sites, including sites that retain eye-selective information. Rivalry greatly suppresses activity in the ventral pathway and attenuates visual adaptation to form and motion; nonetheless, some information about the suppressed stimulus reaches higher brain areas. Although rivalry depends on low-level inhibitory interactions, high-level excitatory influences promoting perceptual grouping and selective attention can extend the local dominance of a stimulus over space and time. Inhibitory and excitatory circuits considered within a hybrid model might account for the paradoxical properties of binocular rivalry and provide insights into the neural bases of visual awareness itself.