Interocular Grouping in Perceptual Rivalry Localized with fMRI.

Bistable perception refers to a broad class of dynamically alternating visual illusions that result from ambiguous images. These illusions provide a powerful method to study the mechanisms that determine how visual input is integrated over space and time. Binocular rivalry occurs when subjects view different images in each eye, and a similar experience called stimulus rivalry occurs even when the left and right images are exchanged at a fast rate. Many previous studies have identified with fMRI a network of cortical regions that are recruited during binocular rivalry, relative to non-rivalrous control conditions (termed replay) that use physically changing stimuli to mimic rivalry. However, we show here for the first time that additional cortical areas are activated when subjects experience rivalry with interocular grouping. When interocular grouping occurs, activation levels broadly increase, with a slight shift towards right hemisphere lateralization. Moreover, direct comparison of binocular rivalry with and without grouping highlights strong focused activity in the intraparietal sulcus and lateral occipital areas, such as right-sided retinotopic visual areas LO1 and IP2, as well as activity in left-sided visual areas LO1, and IP0-IP2. The equivalent analyses for comparable stimulus (eye-swap) rivalry showed very similar results; the main difference is greater recruitment of the right superior parietal cortex for binocular rivalry, as previously reported. Thus, we found minimal interaction between the novel networks isolated here for interocular grouping, and those previously attributed to stimulus and binocular rivalry. We conclude that spatial integration (i.e,. image grouping/segmentation) is a key function of lateral occipital/intraparietal cortex that acts similarly on competing binocular stimulus representations, regardless of fast monocular changes.

Dynamic perspective cues enhance depth perception from motion parallax.

Motion parallax, the perception of depth resulting from an observer's self-movement, has almost always been studied with random dot textures in simplified orthographic rendering. Here we examine depth from motion parallax in more naturalistic conditions using textures with an overall 1/f spectrum and dynamic perspective rendering. We compared depth perception for orthographic and perspective rendering, using textures composed of two types of elements: random dots and Gabor micropatterns. Relative texture motion (shearing) with square wave corrugation patterns was synchronized to horizontal head movement. Four observers performed a two-alternative forced choice depth ordering task with monocular viewing, in which they reported which part of the texture appeared in front of the other. For both textures, depth perception was better with dynamic perspective than with orthographic rendering, particularly at larger depths. Depth ordering performance with naturalistic 1/f textures was slightly lower than with the random dots; however, with depth-related size scaling of the micropatterns, performance was comparable to that with random dots. We also examined the effects of removing each of the three cues that distinguish dynamic perspective from orthographic rendering: (a) small vertical displacements, (b) lateral gradients of speed across the corrugations, and (c) speed differences in rendered near versus far surfaces. Removal of any of the three cues impaired performance. In conclusion, depth ordering performance is enhanced by all of the dynamic perspective cues but not by using more naturalistic 1/f textures.

Comparison of stimulus rivalry to binocular rivalry with functional magnetic resonance imaging.

When incompatible images are presented to each eye, a phenomenon known as binocular rivalry occurs in which the viewer's conscious visual perception alternates between the two images. In stimulus rivalry, similar perceptual alternations between rival images can occur even in the midst of fast image swapping between the eyes. Here, we used functional magnetic resonance imaging to directly compare brain activity underlying the two types of perceptual rivalry. Overall, we found that activity for binocular rivalry was always stronger and more widespread than that for stimulus rivalry-even more so during passive viewing conditions. In particular, the right superior parietal cortex and the right temporoparietal junction were prominently engaged for passive binocular rivalry. While both types of rivalry engaged higher tier visual regions such as the ventral temporal cortex during an active task, activity for stimulus rivalry was comparatively weak in early visual areas V1 to V3, presumably due to a weaker feed-forward signal due to both intraocular and interocular inhibition that may reduce effective contrast. In sum, only binocular rivalry produced perceptually vivid alternations, increased activation of the early visual cortex, and the coordinated engagement of dorsal stream regions, even when a task was not performed. These findings help characterize how stimulus rivalry fits within hierarchical models of binocular rivalry.

fMRI investigation of monocular pattern rivalry.

In monocular pattern rivalry, a composite image is shown to both eyes. The patient experiences perceptual alternations in which the two stimulus components alternate in clarity or salience. We used fMRI at 3T to image brain activity while participants perceived monocular rivalry passively or indicated their percepts with a task. The stimulus patterns were left/right oblique gratings, face/house composites, or a nonrivalrous control stimulus that did not support the perception of transparency or image segmentation. All stimuli were matched for luminance, contrast, and color. Compared with the control stimulus, the cortical activation for passive viewing of grating rivalry included dorsal and ventral extrastriate cortex, superior and inferior parietal regions, and multiple sites in frontal cortex. When the BOLD signal for the object rivalry task was compared with the grating rivalry task, a similar whole-brain network was engaged, but with significantly greater activity in extrastriate regions, including V3, V3A, fusiform face area (FFA), and parahippocampal place area (PPA). In addition, for the object rivalry task, FFA activity was significantly greater during face-dominant periods whereas parahippocampal place area activity was greater during house-dominant periods. Our results demonstrate that slight stimulus changes that trigger monocular rivalry recruit a large whole-brain network, as previously identified for other forms of bistability. Moreover, the results indicate that rivalry for complex object stimuli preferentially engages extrastriate cortex. We also establish that even with natural viewing conditions, endogenous attentional fluctuations in monocular pattern rivalry will differentially drive object-category-specific cortex, similar to binocular rivalry, but without complete suppression of the nondominant image.

How simultaneous is the perception of binocular depth and rivalry in plaid stimuli?

Psychophysical experiments have demonstrated that it is possible to perceive both binocular depth and rivalry in plaids (Buckthought and Wilson 2007, Vision Research47 2543-2556). In a recent study, we investigated the neural substrates for depth and rivalry processing with these plaid patterns, when either a depth or rivalry task was performed (Buckthought and Mendola 2011, Journal of Vision11 1-15). However, the extent to which perception of the two stimulus aspects was truly simultaneous remained somewhat unclear. In the present study, we introduced a new task in which subjects were instructed to perform both depth and rivalry tasks concurrently. Subjects were clearly able to perform both tasks at the same time, but with a modest, symmetric drop in performance when compared to either task carried out alone. Subjects were also able to raise performance levels for either task by performing it with a higher priority, with a decline in performance for the other task. The symmetric declines in performance are consistent with the interpretation that the two tasks are equally demanding of attention (Braun and Julesz 1998, Perception & Psychophysics60 1-23). The results demonstrate the impressive combination of binocular features that supports coincident depth and rivalry in surface perception, within the constraints of presumed orientation and spatial frequency channels.

Bistable percepts in the brain: FMRI contrasts monocular pattern rivalry and binocular rivalry.

The neural correlates of binocular rivalry have been actively debated in recent years, and are of considerable interest as they may shed light on mechanisms of conscious awareness. In a related phenomenon, monocular rivalry, a composite image is shown to both eyes. The subject experiences perceptual alternations in which the two stimulus components alternate in clarity or salience. The experience is similar to perceptual alternations in binocular rivalry, although the reduction in visibility of the suppressed component is greater for binocular rivalry, especially at higher stimulus contrasts. We used fMRI at 3T to image activity in visual cortex while subjects perceived either monocular or binocular rivalry, or a matched non-rivalrous control condition. The stimulus patterns were left/right oblique gratings with the luminance contrast set at 9%, 18% or 36%. Compared to a blank screen, both binocular and monocular rivalry showed a U-shaped function of activation as a function of stimulus contrast, i.e. higher activity for most areas at 9% and 36%. The sites of cortical activation for monocular rivalry included occipital pole (V1, V2, V3), ventral temporal, and superior parietal cortex. The additional areas for binocular rivalry included lateral occipital regions, as well as inferior parietal cortex close to the temporoparietal junction (TPJ). In particular, higher-tier areas MT+ and V3A were more active for binocular than monocular rivalry for all contrasts. In comparison, activation in V2 and V3 was reduced for binocular compared to monocular rivalry at the higher contrasts that evoked stronger binocular perceptual suppression, indicating that the effects of suppression are not limited to interocular suppression in V1.

A matched comparison of binocular rivalry and depth perception with fMRI.

Psychophysical experiments have demonstrated that it is possible to simultaneously perceive binocular depth and rivalry in plaids (A. Buckthought & H. R. Wilson, 2007). Here, we used fMRI at 3T to image activity in the visual cortex while human subjects perceived depth and rivalry from plaids. Six subjects performed either a rivalry or depth task. The spatial frequencies of the near-vertical and diagonal components were, respectively: 2.5, 6.4 cpd; 6.4, 2.5 cpd; or 6.4, 6.4 cpd. The network of activated cortical areas was very similar for the depth compared to the rivalry task. Nevertheless, regions of superior and inferior parietal cortices (including intraparietal sulcus) were activated more during the depth than the rivalry task, independent of spatial frequency, whereas a bias toward rivalry was seen in a lateral occipital region, superior temporal sulcus, and retrosplenial and ventral temporal cortices. Several retinotopic areas in the visual cortex showed a preference for the task with the higher (V1, V2, V3) or lower spatial frequency component (MT+), regardless of the depth or rivalry condition. Our results indicate that depth and rivalry are processed in a similar network of cortical areas and are perceived simultaneously by coexisting in different spatial channels. These results place constraints on binocular vision models.

Hysteresis effects in stereopsis and binocular rivalry.

Neural hysteresis plays a fundamental role in stereopsis and reveals the existence of positive feedback at the cortical level [Wilson, H. R., & Cowan, J. D. (1973). A mathematical theory of the functional dynamics of cortical and thalamic nervous tissue. Kybernetik 13(2), 55-80]. We measured hysteresis as a function of orientation disparity in tilted gratings in which a transition is perceived between stereopsis and binocular rivalry. The patterns consisted of sinusoidal gratings with orientation disparities (0 degrees, 1 degrees, 2 degrees,..., 40 degrees) resulting in various degrees of tilt. A movie of these 41 pattern pairs was shown at a rate of 0.5, 1 or 2 pattern pairs per second, in forward or reverse order. Two transition points were measured: the point at which the single tilted grating appeared to split into two rivalrous gratings (T1), and the point at which two rivalrous gratings appeared to merge into a single tilted grating (T2). The transitions occurred at different orientation disparities (T1=25.4 degrees, T2=17.0 degrees) which was consistent with hysteresis and far exceeded the difference which could be attributed to reaction time. The results are consistent with a cortical model which includes positive feedback and recurrent inhibition between neural units representing different eyes and orientations.

Interaction between binocular rivalry and depth in plaid patterns.

Binocular rivalry was studied using plaids which were the sum of orthogonal diagonal gratings plus identical vertical gratings in the two eyes. The rivalry alternations sped up as the spatial frequency difference between the vertical and diagonal gratings was increased above about one octave, but slowed down for smaller differences. The interaction between depth and rivalry was studied using similar plaids but with depth introduced in the vertical components. Depth and rivalry coexisted when the spatial frequency difference between the vertical and diagonal gratings was greater than about one octave, but rivalry slowed down and depth perception was reduced for smaller differences. Plaids consisting of square wave gratings were used to compare: (1) added gratings; (2) vertical gratings superimposed on (i.e. occluding) diagonal gratings; (3) diagonal gratings superimposed on vertical gratings. Rivalry alternations were fastest in condition (3), indicating that grouping effects played a role. The final experiment indicated that depth and rivalry coexisted within a spatial frequency band if the orientation difference between the vertical and diagonal components was 60-70 degrees . These results place constraints on models of stereopsis and rivalry, indicating that depth and rivalry can coexist in different spatial frequency and orientation bands but that each interferes with the other in the same band.

Stereoscopic depth discrimination with contrast windowed stimuli.

Depth discrimination with a shifted contrast window was compared to that with a fixed contrast window. Stereoscopic performance with the fixed window was limited to small disparities and varied with spatial frequency. Performance with the shifted window extended to larger disparities and was more similar for low and high spatial frequencies. The results depended upon window shape, indicating that edge blur is an important factor. Stereoscopic performance with shifted patterns was supported at disparities larger than a phase disparity model might predict, suggesting that a combination of position and phase disparity computations are used for the perception of stereoscopic depth.