The effect of pupil size on binocular summation at suprathreshold conditions

PURPOSE: To determine the influence of retinal illuminance on monocular or binocular visual reaction time (VRT). METHODS: On two normal subjects, uniform circular stimuli were presented with respect to a reference stimulus at the fovea under suprathreshold conditions, and the detection of positive and negative luminance variations was recorded. Binocular and monocular reaction times were measured (by the index finger pressing on a mouse key) for viewing with both natural and artificial pupils. RESULTS: Binocular reaction times were shorter than monocular reaction times; nevertheless, this binocular-summation effect was less marked in trials with the artificial pupil. Analyses of binocular-summation ratios for contrast changes for both pupil types indicated maximum and minimum binocular-saturation values depending on contrast variations in both positive and negative luminance changes. CONCLUSIONS: Binocular summation can be influenced by pupil size under suprathreshold conditions. Results are discussed in terms of retinal illuminance and cortical pupil response mechanisms.     

Induced aniseikonia diminishes binocular contrast sensitivity and binocular summation.

PURPOSE: To study the effect on binocular contrast sensitivity and binocular summation of aniseikonia induced by size (afocal) lenses. METHODS: In 18 young emmetropic observers, the monocular and binocular contrast sensitivity function was measured under normal conditions and after inducing different degrees of aniseikonia (2%, 3%, and 5% and for 6 observers, 8%) in the right eye. The spatial frequencies tested were 2.4, 3.7, 6.0, 9.2, 12, 15, 20, and 24 cpd. RESULTS: The results reveal a significant decline in binocular contrast sensitivity and binocular summation for 5% aniseikonia, this decline being more pronounced for intermediate and high spatial frequencies. CONCLUSIONS: These results, together with others on different binocular functions, show the importance of aniseikonia in binocular vision. On this basis, we recommend that aniseikonia be considered in surgical processes such as cataract and refractive surgery, whereby aniseikonia could be induced and binocular performance subsequently diminished.     

Reaction time as a measure of temporal summation at suprathreshold levels

Two subjects performed simple reaction-time responses to green-blue stimuli of varying luminance and duration under binocular and monocular observation. Reaction time varied as a power function of luminance with simple fractional exponents: an exponent of 1/3 for binocular vision and an exponent of 2/5 for monocular vision. The luminance required to produce ten criterion reaction times was calculated, and the luminance-duration relationship was examined. The critical duration was a function of the criterion reaction time: as the criterion reaction time increases, the critical duration increases monotonically. The binocular critical duration was found to be longer than the monocular critical duration.     

Integration of motion information during binocular rivalry.

When two moving gratings are superimposed in normal viewing they often combine to form a pattern that moves with a single direction of motion. Here, we investigated whether the same mechanism underlies pattern motion when drifting gratings are presented independently to the two eyes. We report that, with relatively large circular grating patches (4 deg), there are periods of monocular dominance in which one eye's orientation alone is perceived, usually moving orthogonal to the contours (component motion). But, during the transitions from one monocular view to the other, a fluid mosaic is perceived, consisting of contiguous patches, each containing contours of only one of the gratings. This entire mosaic often appears to move in a single direction (pattern motion), just as when two gratings are literally superimposed. Although this implies that motion signals from the perceptually suppressed grating continue to influence the perception of motion, an alternative possibility is that it reflects a strategy that involves integrating directional information from the contiguous single-grating patches. To test between these possibilities, we performed a second experiment with very small grating stimuli that were about the same size as the contiguous single-grating patches in the mosaic (1-deg diameter). Despite the fact that the form of only one grating was perceived, we report that pattern motion was still perceived on about one third of trials. Moreover, a decrease in the occurrence of pattern motion was apparent when the contrast and spatial frequency of the gratings were made more different from each other. This phenomenon clearly demonstrates an independent binocular interaction for form and motion.     

Sine-wave gratings: scale invariance and spatial integration at suprathreshold contrast

Thresholds for the detection of contrast differences and spatial frequency differences present in a sine-wave grating with suprathreshold contrast were measured as a function of the size of the grating and the number of cycles in the grating. For gratings with the same number of cycles, these thresholds are found to be independent of the size of the grating over the entire range of sizes tested (1.5 log unit): the thresholds do not change when the stimulus is enlarged by some scale factor. We call this phenomenon "scale invariance". The thresholds decrease with increasing number of grating cycles, at least up to the point where 16 cycles are present. This support the view that "spatial pooling" or "spatial integration" mechanisms act over a larger area than the "functional summation area" found in threshold contrast measurements (about 10 cycles high and wide).     

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.     

Resolution of binocular rivalry: Perceptual misbinding of color

Is neural binding of color and form required for perception of a unified colored object? Individual cells selectively tuned to both color and orientation are proposed to moot the binding problem. This study reveals perceptual misbinding of color, thereby revealing separate neural representations of color and form followed by a subsequent binding process. Low luminance-contrast, rivalrous chromatic gratings were presented dichoptically. Each grating had alternating chromatic and gray stripes (e.g., red/gray in the left eye, green/gray in the right eye). Observers viewed the two rivalrous, 2 cpd gratings for 1 min. The duration of exclusive visibility was measured for four percepts: left-eye stimulus, right-eye stimulus, fusion of the two colors, or a two-color grating (e.g. a red/green grating). The percept of a two-color grating (misbinding) was observed with Michelson luminance contrast in the grating up to 20%. In general, for a given level of luminance contrast either misbinding (low luminance contrast) or color mixture (high luminance contrast) was observed, but not both of them. The perceived two-color gratings show that two rivalrous chromaticities are both represented neurally when color and form are combined to give a unified percept. "Resolution" of competing chromatic signals from the two eyes is not restricted to color dominance and color mixture. The transition from misbinding to color mixture by increasing luminance contrast shows that luminance edges have an important role in correct localization of color.     

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.     

Temporal characteristics of binocular rivalry: visual field asymmetries

Very little is known about the mechanisms that drive the alternation between the two views during binocular rivalry. A key property of the rivalry process is the rate at which the two views alternate. Understanding the factors that affect the rate of the alternation is critical to the final understanding of the underlying process. Using a circular and a radial grating as the rivalry stimuli, we observed a significantly faster binocular rivalry when stimuli were presented in the right visual field than that in the left visual field for the right-handed observers, and a reversed asymmetry for the left-handed observers. In both groups, rivalry was faster for stimuli presented in the lower visual field than that in the upper visual field. This pattern of results suggests that (1) rivalry is likely a locally driven process and (2) the visual brain in the left hemisphere may be the faster one of the two hemispheres in right-handed people.     

Effects of dominant and nondominant eyes in binocular rivalry.

PURPOSE: To investigate the relation between sighting and sensory eye dominance and attempt to quantitatively examine eye dominance using a balance technique based on binocular rivalry. METHODS: The durations of exclusive visibility of the dominant and nondominant eye target in binocular rivalry were measured in 14 subjects. The dominant eye was determined by using the hole-in-card test (sighting dominance). In study 1, contrast of the target in one eye was fixed at 100% and contrast of the target in the other eye was varied from 100% to 80% to 60% to 40% to 20%, when using rectangular gratings of 1, 2, and 4 cycles per degree (cpd) at 2 degrees, 4 degrees , and 8 degrees in size. In study 2, contrast of the target in the nondominant eye was fixed at 100% and contrast of the target in the dominant eye was varied from 100% to 80% to 60% to 40% to 20%, when using a rectangular grating of 2 cpd at 4 degrees in size. RESULTS: In study 1, the total duration of exclusive visibilities of the dominant eye target; that is, the target seen by the eye that had sighting dominance was longer compared with that of the nondominant eye target. When using rectangular gratings of 4 cpd, mean total duration of exclusive visibility of the dominant eye target was statistically longer than that of the nondominant eye target (p < 0.05). In study 2, reversals (in which duration of exclusive visibility of the nondominant eye becomes longer than the dominant eye when the contrast of the dominant eye target is decreased) were observed for all contrasts except for 100%. CONCLUSIONS: The dominant sighting eye identified by the hole-in-card test coincided with the dominant eye as determined by binocular rivalry. The contrast at which reversal occurs indicates the balance point of dominance and seems to be a useful quantitative indicator of eye dominance to clinical applications.     

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.     

Troxler effect with dichoptic stimulus presentations: evidence for binocular inhibitory summation and interocular suppression

Whether the Troxler effect (TE) has to do with interocular suppression and/or summation was studied with dichoptically matched (binocular or dioptic) and unmatched (monocular) stimulus presentations. Perceptual disappearance was found to occur more slowly under the binocular condition (mean=14.2s) than the monocular condition (mean=8.4s), but much faster than predicted by probability summation of the experimentally obtained latencies and durations of the TE in the monocular conditions (>27 s), suggesting a binocular inhibitory summation, the opposite of the binocular summation found with detection and contrast matching tasks [(Blake, R., & Fox, R. (1973). The psychological inquiry into binocular summation. Perception and Psychophysics, 14, 161-185; Blake, R., Sloane, M., & Fox, R. (1981). Further developments in binocular summation. Perception &Psychophysics 30, 266-276.)]. In addition, Ss with poorer stereoacuity took longer to see the disappearance in the monocular condition, and showed a larger disparity between the TEs from the two monocular conditions, suggesting a contribution of interocular suppression to the TE.     

Optokinetic nystagmus as an objective indicator of binocular rivalry.

When each eye is separately stimulated by moving contours that generate antagonistic optokinetic nystagmus, vigorous phenomenal rivalry results, yet the nystagmus eye movements remain yoked and shift direction in accord with the eye that is phenomenally dominant. The feasibility of using this nystagmus rivalry as an indicator of phenomenal dominance was examined. The high correlation between phenomenal report and nystagmus rivalry, together with stochastic and stimulus similarities between nystagmus rivalry and conventional rivalry, support using nystagmus as an objective indicator.     

Endogenous attention selection during binocular rivalry at early stages of visual processing

Directing attention to one of two superimposing surfaces composed of dot fields rotating in opposing directions facilitates processing of brief translations of the attended surface [Valdes-Sosa, M., Bobes, M. A., Rodriguez, V., & Pinilla, T. (1998). Switching attention without shifting the spotlight object-based attentional modulation of brain potentials. Journal of Cognition and Neuroscience, 10(1), 137-151]. Here we used ERP recordings to investigate the mechanisms of endogenous attentional selection of such competing dot surfaces under conditions of dichoptic viewing (one surface to each eye) and monocular viewing (both surfaces to one eye). Under dichoptic conditions, which induced binocular rivalry, translations of the attended surface presented to one eye elicited enhanced visual P1 and N1 ERP components relative to translations of the unattended surface presented to the other eye. In comparison, during monocular viewing the attended surface translations elicited a significantly larger N1 component in the absence of any P1 modulation. These results indicate that processing of the attended surface is biased at an earlier level in extrastriate visual cortex under conditions of inter-ocular versus intra-ocular competition.     

Spatial zones of binocular rivalry in central and peripheral vision.

This paper presents results from psychophysical experiments on human binocular rivalry in central and peripheral vision. Results show that the incidence of periods of exclusive visibility of a given eye's rival target increased with decreasing target size, and for a given sized target exclusive visibility increased with retinal eccentricity. Control measures confirmed that these results were not attributable solely to reduced peripheral acuity, to Troxler's effect, or to spatial frequency. We computed the minimum-sized stimulus that would lead to a criterion level of exclusive visibility of one or the other eye; this we term the spatial zone of binocular rivalry. The change in estimated size of spatial zones of rivalry with eccentricity compares favorably with estimates of human cortical magnification. We propose a model that assumes concentrically organized zones of rivalry. These zones do not function independently, but instead exhibit a high degree of mutual excitatory cooperativity. The model has multiple solutions for the foveal zone size, but the best fits predict a diameter of 5.3 or 7.3 min of visual angle; these values dovetail nicely with our empirical estimates of the foveal zone size.     

The absence of a measurable “critical band” at low suprathreshold contrasts

Subjects adjusted the contrast of a computer-generated test grating until it appeared to have the same overall contrast as a 6.25-c/deg grating of fixed contrast. Results show that the relative phase of the components of a compound test grating does not affect the perceived contrast of the grating. This suggests that independent mechanisms may process the components, as is the case at contrast threshold. The component contrasts, however, appear to be combined by a square-law summation mechanism below 20% contrast, which results in the absence of a measurable “critical band” in grating perception at such contrast levels. Thus, these results are also consistent with a simple single-channel model in which perceived contrast is directly related to root-mean-square contrast. It is not possible to reject either the single- or multiple-channel hypothesis on the basis of these experiments.     

No evidence for widespread synchronized networks in binocular rivalry: MEG frequency tagging entrains primarily early visual cortex

We investigated the spatio-temporal dynamics of the steady-state-visual-evoked field during perceptual switches in binocular rivalry using MEG. Several authors have previously used frequency tagging in MEG studies on binocular rivalry and have claimed to have found a widespread network of synchronized areas that are entrained by the stimulus, reaching up to frontal regions. Moreover, it has been claimed that the results prove that becoming aware of a stimulus is established by increased intra- and interhemispheric synchronization of brain areas, separated by large distances. Our results dovetailed nicely with previous findings such as power and coherence modulations as a function of perceptual state. However, while we also found stimulus-entrained activity across the entire scalp, a phase analysis revealed that the spatially extended nature of the frequency tag was produced by a limited set of occipital sources. Furthermore, we provide evidence that the coherence results from earlier studies are ambiguous in that they likely measured coherence between different sensors whose signals were dominated by the same sources. We conclude that the claims about widespread synchronized networks to consciously perceive flicker stimuli are currently unconvincing.     

Functional asymmetries of the human visual system as revealed by binocular rivalry and binocular brightness matching.

Two techniques, a binocular rivalry task and a binocular brightness matching task, were designed to yield indices of asymmetry (relative dominance weights) for the two eyes, the crossed and uncrossed visual pathways, and the two cerebral hemispheres. Twenty subjects with normal vision were run on all conditions. Intercorrelations of the dominance weights obtained by the two methods showed no relationship between the two methods, but produced three hypotheses about visual functioning: (1) the left hemisphere appears more dominant for rivalry; (2) the right hemisphere appears more dominant for brightness matching; (3) the uncrossed visual pathways are dominant over the crossed pathways in the binocular rivalry task.