Binocular Suppression Research Articles

Adaptation to invisible motion results in low-level but not high-level aftereffects

After prolonged exposure to moving stimuli, illusory motion is perceived in stimuli that do not contain consistent motion, a phenomenon termed the motion aftereffect (MAE). In this study, we tested MAEs under binocular suppression that renders the motion adaptor invisible for the entire adaptation period. We developed a variant of the continuous flash suppression method to reliably suppress target motion stimuli for durations longer than several tens of seconds. Here, we ask whether motion systems are functional in the absence of perception by measuring the MAE, a question difficult to address using binocular rivalry that accompanies a switch of percept between visible and invisible. Results show that both the MAEs with static and dynamic tests are attenuated with an invisible adaptor when the adaptor and the test stimulus are presented to the same eye. In contrast, when the test pattern was presented to the other eye, the dynamic MAE was observed in invisible adaptor conditions. These results indicate that low-level adaptation survives under total binocular suppression, a finding predicted by previous studies. In contrast, disappearance of interocular transfer in the dynamic MAE suggests that a high-level motion detector does not operate when the motion adaptor is rendered invisible.

Efficient stereo video coding based on frame skipping for real-time mobile applications

An efficient stereo video coding algorithm based on frame skipping is proposed for real-time mobile applications in this work. Based on the binocular suppression theory, the proposed algorithm skips a selected set of frames to save the encoding complexity and the transmission bit rate. At the decoder side, we reconstruct those skipped frames faithfully using the advanced bilateral interpolation method. Extensive simulation results demonstrate that the proposed algorithm provides better PSNR performances than the conventional stereo coding algorithm, while achieving realtime encoding and decoding.

Effects of binocular suppression on surround suppression

The responses of neurons in the primary visual cortex (V1) are generally inhibited by stimuli surrounding their classical receptive fields (CRF). This surround suppression can influence the visual perception of stimuli. For instance, the presence of a surround stimulus can decrease the apparent contrast of a central stimulus. A recent neurophysiological study in nonhuman primates suggests that two distinct mechanisms, early and late mechanisms, give rise to surround suppression. Here, we used binocular suppression to render the surround stimuli invisible and evaluated the effects of this masking on the two types of surround suppression. We found that the early mechanism was unsusceptible to, whereas the late mechanism was eliminated by, binocular suppression. The distinct effects of binocular suppression on the early and late mechanisms suggest that the two types of surround suppression arise from different neural substrates.

Feature-selective inhibition during binocular rivalry

It has been argued that binocular rivalry occurs as a result of competition between monocular channels (eg eye competition). However, some recent literature favours a pattern competition account of rivalry, in which rivalry occurs between incompatible (binocular) pattern representations. In this current study, we investigated feature-specific aspects of dichoptic suppression. Three coloured gratings were presented simultaneously next to each other to one eye on a dichoptically presented grid. One of the two flanking gratings had the same colour and orientation as the central grating. All three gratings were suppressed by presenting high-contrast elements to the other eye. After 600 ms, the flanking suppressors were removed and the flanking gratings became visible. Results show that the grating dissimilar to the suppressed central grating became visible first, both for the intraocular and interocular condition. This is evidence for feature-specific inhibition during binocular suppression. These findings support the idea that all presented visual information reaches higher visual areas and favour a pattern competition account of binocular rivalry.

Retinotopic maps and foveal suppression in the visual cortex of amblyopic adults

Amblyopia is a developmental visual disorder associated with loss of monocular acuity and sensitivity as well as profound alterations in binocular integration. Abnormal connections in visual cortex are known to underlie this loss, but the extent to which these abnormalities are regionally or retinotopically specific has not been fully determined. This functional magnetic resonance imaging (fMRI) study compared the retinotopic maps in visual cortex produced by each individual eye in 19 adults (7 esotropic strabismics, 6 anisometropes and 6 controls). In our standard viewing condition, the non-tested eye viewed a dichoptic homogeneous mid-level grey stimulus, thereby permitting some degree of binocular interaction. Regions-of-interest analysis was performed for extrafoveal V1, extrafoveal V2 and the foveal representation at the occipital pole. In general, the blood oxygenation level-dependent (BOLD) signal was reduced for the amblyopic eye. At the occipital pole, population receptive fields were shifted to represent more parafoveal locations for the amblyopic eye, compared with the fellow eye, in some subjects. Interestingly, occluding the fellow eye caused an expanded foveal representation for the amblyopic eye in one early-onset strabismic subject with binocular suppression, indicating real-time cortical remapping. In addition, a few subjects actually showed increased activity in parietal and temporal cortex when viewing with the amblyopic eye. We conclude that, even in a heterogeneous population, abnormal early visual experience commonly leads to regionally specific cortical adaptations.

Central factors contributing to para-contrast modulation of contour and brightness perception

Following up on a prior study of contour and brightness processing in visual masking (Breitmeyer et al., 2006), we investigated the effects of a binocular and dichoptic para-contrast masking on the visibility of the contour and brightness of a target presented to the other eye. Combined, the results support the contributions of several cortical processes to para-contrast: (1) two central sources of inhibition, one long-latency and prolonged and the other short-latency and brief; (2) binocular rivalry suppression; and (3) a facilitatory effect peaking at different SOAs for the contour and the brightness tasks, reflecting; (4) known properties of two separate cortical systems, one a fast contour-processing pathway and the other a slower brightness-processing pathway.

Cortical Effects of Brief Daily Periods of Unrestricted Vision During Early Monocular Form Deprivation

Experiencing daily brief periods of unrestricted vision during early monocular form deprivation prevents or reduces the degree of resulting amblyopia. To gain insight into the neural basis for these "protective" effects, we analyzed the monocular and binocular response properties of individual neurons in the primary visual cortex (V1) of macaque monkeys that received intermittent unrestricted vision. Microelectrode-recording experiments revealed significant decreases in the proportion of units that were dominated by the treated eyes, and the magnitude of this ocular dominance imbalance was correlated with the degree of amblyopia. The sensitivity of V1 neurons to interocular spatial phase disparity was significantly reduced in all treated monkeys compared with normal adults. With unrestricted vision, however, there was a small but significant increase in overall disparity sensitivity. Binocular suppression was prevalent in monkeys with constant form deprivation but significantly reduced by the daily periods of unrestricted vision. If neurons exhibited consistent responses to stimulation of the treated eye, monocular response properties obtained by stimulation of the two eyes were similar. These results suggest that the observed protective effects of brief periods of unrestricted vision are closely associated with the ability of V1 neurons to maintain their functional connections from the deprived eye and that interocular suppression in V1 may play an important role in regulating synaptic plasticity of these monkeys.

Rapid plasticity of binocular connections in developing monkey visual cortex (V1)

The basic sets of cortical connections are present at birth in the primate visual system. The maintenance and refinement of these innate connections are highly dependent on normal visual experience, and prolonged exposure to binocularly uncorrelated signals early in life severely disrupts the normal development of binocular functions. However, very little is known about how rapidly these changes in the functional organization of primate visual cortex emerge or what are the sequence and the nature of the abnormal neural events that occur immediately after experiencing binocular decorrelation. In this study, we investigated how brief periods of ocular misalignment (strabismus) at the height of the critical period alter the cortical circuits that support binocular vision. After only 3 days of optically imposed strabismus, there was a striking increase in the prevalence of V1 neurons that exhibited binocular suppression, i.e., binocular responses were weaker than monocular responses. However, the sensitivity of these neurons to interocular spatial phase disparity was not significantly altered. These contrasting results suggest that the first significant change in V1 caused by early binocular decorrelation is binocular suppression, and that this suppression originates at a site(s) beyond where binocular signals are initially combined.

Face Adaptation Depends on Seeing the Face

Retinal input that is suppressed from visual awareness can nevertheless produce measurable aftereffects, revealing neural processes that do not directly result in a conscious percept. We here report that the face identity-specific aftereffect requires a visible face; it is effectively cancelled by binocular suppression or by inattentional blindness of the inducing face. Conversely, the same suppression does not interfere with the orientation-specific aftereffect. Thus, the competition between incompatible or interfering visual inputs to reach awareness is resolved before those aspects of information that are exploited in face identification are processed. We also found that the face aftereffect remained intact when the visual distracters in the inattention experiment were replaced with auditory distracters. Thus, cross-modal or cognitive interference that does not affect the visibility of the face does not interfere with the face aftereffect. We conclude that adaptation to face identity depends on seeing the face.

Effect of temporal sparseness and dichoptic presentation on multifocal visual evoked potentials

Multifocal VEP (mfVEP) responses were obtained from 13 normal human subjects for nine test conditions, covering three viewing conditions (dichoptic and left and right monocular), and three different temporal stimulation forms (rapid contrast reversal, rapid pattern pulse presentation, and slow pattern pulse presentation). The rapid contrast reversal stimulus had pseudorandomized reversals of checkerboards in each visual field region at a mean rate of 25 reversals/s, similar to most mfVEP studies to date. The rapid pattern pulse presentation had pseudorandomized presentations of a checkerboard for one frame, interspersed with uniform grey frames, with a mean rate of 25 presentations/s per region per eye. The slow pattern pulse stimulus had six presentations/s per region per eye. Recording time was 5.3 min/condition. For dichoptic presentation slow pattern pulse responses were 4.6 times larger in amplitude than the contrast reversal responses. Binocular suppression was greatest for the contrast reversal stimulus. Consideration of the signal-to-noise ratios indicated that to achieve a given level of reliability, slow pattern pulse stimuli would require half the recording time of contrast reversal stimuli for monocular viewing, and 0.4 times the recording time for dichoptically presented stimuli. About half the responses to the slow pattern pulse stimuli had peak value exceeding five times their estimated standard error. Responses were about 20% smaller in the upper visual field locations. Space-time decomposition showed that responses to slow pattern pulse were more consistent across visual field locations. We conclude that the pattern pulse stimuli, which we term temporally sparse, maintain the visual system in a high contrast gain state. This more than compensates for the smaller number of presentations in the run, and provides signal-to-noise advantages that may be valuable in clinical application.

Interocular masking and summation indicate two stages of divisive contrast gain control

Our understanding of early spatial vision owes much to contrast masking and summation paradigms. In particular, the deep region of facilitation at low mask contrasts is thought to indicate a rapidly accelerating contrast transducer (eg a square-law or greater). In experiment 1, we tapped an early stage of this process by measuring monocular and binocular thresholds for patches of 1 cycle deg-1 sine-wave grating. Threshold ratios were around 1.7, implying a nearly linear transducer with an exponent around 1.3. With this form of transducer, two previous models (Legge, 1984 Vision Research 24 385 - 394; Meese et al, 2004 Perception 33 Supplement, 41) failed to fit the monocular, binocular, and dichoptic masking functions measured in experiment 2. However, a new model with two-stages of divisive gain control fits the data very well. Stage 1 incorporates nearly linear monocular transducers (to account for the high level of binocular summation and slight dichoptic facilitation), and monocular and interocular suppression (to fit the profound 42 Oral presentations: Spatial vision Thursday dichoptic masking). Stage 2 incorporates steeply accelerating transduction (to fit the deep regions of monocular and binocular facilitation), and binocular summation and suppression (to fit the monocular and binocular masking). With all model parameters fixed from the discrimination thresholds, we examined the slopes of the psychometric functions. The monocular and binocular slopes were steep (Weibull s˜3-4) at very low mask contrasts and shallow (s˜1.2) at all higher contrasts, as predicted by all three models. The dichoptic slopes were steep (s˜3-4) at very low contrasts, and very steep (s>5.5) at high contrasts (confirming Meese et al, loco cit.). A crucial new result was that intermediate dichoptic mask contrasts produced shallow slopes (s˜2). Only the two-stage model predicted the observed pattern of slope variation, so providing good empirical support for a two-stage process of binocular contrast transduction. [Supported by EPSRC GR/S74515/01]

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.

Binocular rivalry suppression does impede buildup of the motion aftereffect.

Binocular rivalry suppression does impede buildup of the motion aftereffect.

Amygdala responses to fearful and happy facial expressions under conditions of binocular suppression.

The human amygdala plays a crucial role in processing affective information conveyed by sensory stimuli. Facial expressions of fear and anger, which both signal potential threat to an observer, result in significant increases in amygdala activity, even when the faces are unattended or presented briefly and masked. It has been suggested that afferent signals from the retina travel to the amygdala via separate cortical and subcortical pathways, with the subcortical pathway underlying unconscious processing. Here we exploited the phenomenon of binocular rivalry to induce complete suppression of affective face stimuli presented to one eye. Twelve participants viewed brief, rivalrous visual displays in which a fearful, happy, or neutral face was presented to one eye while a house was presented simultaneously to the other. We used functional magnetic resonance imaging to study activation in the amygdala and extrastriate visual areas for consciously perceived versus suppressed face and house stimuli. Activation within the fusiform and parahippocampal gyri increased significantly for perceived versus suppressed faces and houses, respectively. Amygdala activation increased bilaterally in response to fearful versus neutral faces, regardless of whether the face was perceived consciously or suppressed because of binocular rivalry. Amygdala activity also increased significantly for happy versus neutral faces, but only when the face was suppressed. This activation pattern suggests that the amygdala has a limited capacity to differentiate between specific facial expressions when it must rely on information received via a subcortical route. We suggest that this limited capacity reflects a tradeoff between specificity and speed of processing.

Contrast Adaptation Effects under Interocular Suppression for Normal and Strabismic Observers

Our vision system receives two different images from our two eyes. Since these two images usually have high correlation they are fused into a single stable image. In binocular rivalry suppression a stimulus presented in one eye is suppressed and a different stimulus in the other eye is perceived for a certain time interval. For strabismic observers the suppression is continuous between different retinal images caused by misalignment of eye positions. Previous research investigated the effects of this binocular rivalry suppression on a number of visual processes. The present study looked at the effects of contrast adaptation for the adaptation stimulus presented to eyes under interocular suppression. We compared normal and strabismic observers and found that in both types, regardless of the existence of suppression, there was the effect of contrast adaptation. These findings provide evidence that the suppression occurs at a site in the visual system after the locus of contrast adaptation. The contrast adaptation strength differed in the two types of observers.

Binocular interactions in area 21a of the cat.

We investigated binocular suppression in area 21a cells of the anaesthetized cat using drifting sinusoidal gratings presented simultaneously to each eye. The grating presented to the dominant eye was always oriented optimally and the grating presented to the non-dominant eye was either at the same orientation, but at the least effective relative spatial phase, or orthogonal. The binocular response of approximately 80% of cells was less than the monocular dominant eye response when there was a mismatch in orientation or spatial location between stimuli. Response suppression in the two binocular stimulus conditions had a correlation coefficient (r) of 0.55. We propose a parsimonious model to account for the response facilitation and suppression by binocular stimulation of area 21a neurons.

Detection, prevention, and rehabilitation of amblyopia.

The necessity of visual preschool screening for reducing the prevalence of amblyopia is widely accepted. The beneficial results of large-scale screening programs conducted in Scandinavia are reported. Screening monocular visual acuity at 3.5 to 4 years of age appears to be an excellent basis for detecting and treating amblyopia and an acceptable compromise between the pitfalls encountered in screening younger children and the cost-to-benefit ratio. In this respect, several preschoolers' visual acuity charts have been evaluated. New recently developed small-target random stereotests and binocular suppression tests have also been developed with the aim of correcting the many false negatives (anisometropic amblyopia or bilateral high ametropia) induced by the usual stereotests. Longitudinal studies demonstrate that correction of high refractive errors decreases the risk of amblyopia and does not impede emmetropization. The validity of various photoscreening and videoscreening procedures for detecting refractive errors in infants prior to the onset of strabismus or amblyopia, as well as alternatives to conventional occlusion therapy, is discussed.

Residual binocular interactions in the striate cortex of monkeys reared with abnormal binocular vision.

We investigated the nature of residual binocular interactions in the striate cortex (V1) of monkey models for the two most common causes of visual dysfunction in young children, specifically anisometropia and strabismus. Infant rhesus monkeys were raised wearing either anisometropic spectacle lenses that optically defocused one eye or ophthalmic prisms that optically produced diplopia and binocular confusion. Earlier psychophysical investigations had demonstrated that all subjects exhibited permanent binocular vision deficits and, in some cases, amblyopia. When the monkeys were adults, the responses of individual V1 neurons were studied with the use of microelectrode recording techniques while the animals were anesthetized and paralyzed. The manner in which the signals from the two eyes were combined in individual cells was investigated by dichoptically stimulating both eyes simultaneously with drifting sine wave gratings. In both lens- and prism-reared monkeys, fewer neurons had balanced ocular dominances and greater numbers of neurons were excited by only one eye. However, many neurons that appeared to be monocular exhibited clear binocular interactions during dichoptic stimulation. For the surviving binocular neurons, the maximum binocular response amplitudes were lower than normal; fewer neurons, particularly complex cells, were sensitive to relative interocular spatial phase disparities; and the remaining disparity-sensitive neurons exhibited lower degrees of binocular interaction. In prism-reared monkeys, an unusually high proportion of complex cells exhibited binocular suppression during dichoptic stimulation. Binocular contrast summation experiments showed that for both cooperative and antagonistic binocular interactions, contrast signals from the two eyes were combined by individual neurons in a normal linear fashion in both lens- and prism-reared monkeys. The observed binocular deficits appear to reflect a reduction in functional inputs from one eye and/or spatial imprecision in the monocular receptive fields rather than an aberrant form of binocular interaction. In the prism-reared monkeys, the predominance of suppression suggests that inhibitory connections were, however, less susceptible to diplopia and confusion than excitatory connections. Overall, there were many parallels between V1 physiology in our monkey models and the residual vision of humans with anisometropia or strabismus.

Binocular spatial phase tuning characteristics of neurons in the macaque striate cortex.

We employed microelectrode recording techniques to study the sensitivity of individual neurons in the striate cortex of anesthetized and paralyzed monkeys to relative interocular image disparities and to determine the effects of basic stimulus parameters on these cortical binocular interactions. The visual stimuli were drifting sine wave gratings. After the optimal stimulus orientation, spatial frequency, and direction of stimulus movement were found, the cells' disparity tuning characteristics were determined by measuring responses as a function of the relative interocular spatial phase of dichoptic grating pairs. No attempts were made to assess absolute position disparities or horizontal disparities relative to the horopter. The majority (approximately 70%) of simple cells were highly sensitive to interocular spatial phase disparities, particularly neurons with balanced ocular dominances. Simple cells typically demonstrated binocular facilitation at the optimal phase disparity and binocular suppression for disparities 180 degrees away. Fewer complex cells were phase selective (approximately 40%); however, the range of disparity selectivity in phase-sensitive complex cells was comparable with that for simple cells. Binocular interactions in non-phase-sensitive complex cells were evidenced by binocular response amplitudes that differed from responses to monocular stimulation. The degree of disparity tuning was independent of a cell's optimal orientation or the degree of direction tuning. However, disparity-sensitive cells tended to have narrow orientation tuning functions and the degree of disparity tuning was greatest for the optimal stimulus orientations. Rotating the stimulus for one eye 90 degrees from the optimal orientation usually eliminated binocular interactions. The effects of phase disparities on the binocular response amplitude were also greatest at the optimal spatial frequency. Thus a cell's sensitivity to absolute position disparities reflects its spatial tuning characteristics, with cells sensitive to high spatial frequencies being capable of signaling very small changes in image disparity. On the other hand, stimulus contrast had relatively little effect on a cell's disparity tuning, because response saturation occurred at the same contrast level for all relative interocular phase disparities. Thus, as with orientation tuning, a cell's optimal disparity and the degree of disparity selectivity were invariant with contrast. Overall, the results show that sensitivity to interocular spatial phase disparities is a common property of striate neurons. A cell's disparity tuning characteristics appear to largely reflect its monocular receptive field properties and the interocular balance between excitatory and inhibitory inputs. However, distinct functional classes of cortical neurons could not be discriminated on the basis of disparity sensitivity alone.

Selective masking of the achromatic system can be cortically mediated

A luminance decrement selectively masks the achromatic system when presented in the same eye as the stimulus. The purpose of this study was to determine whether the masking effect can be elicited under dichoptic conditions. The achromatic and color-opponent systems were identified with monocularly presented 10- and 200-ms spectral increments. A luminance decrement mask was presented to the fellow eye. This mask was 50-ms in duration and either preceded the onset of the increment by 25 ms (forward masking) or followed the increment by 25 ms (backward masking). The spectral sensitivity functions obtained with 10-ms increments, either preceded or followed by the dichoptic mask, were profoundly different from those obtained with isolated 10-ms increments: the broadband achromatic function was replaced by a color-opponent function of lesser sensitivity. In comparison, the masks did not have a substantial effect on the form or sensitivity of the color-opponent functions obtained with 200-ms increments. These data reveal that a luminance decrement mask, presented under dichoptic viewing conditions, suppresses the achromatic system but not the color-opponent system. Therefore selective masking of the achromatic system by a luminance decrement can be mediated at a site after the signals from the two eyes are combined—the striate cortex or beyond. Implications for models of binocular, blink, and saccadic suppression are discussed.