Gaze capture by eye-of-origin singletons: interdependence with awareness

Where we look in visual tasks is determined by both bottom-up and top-down factors. One theory (Li, 1999a, 2002) suggests that visual area V1 creates a bottom-up saliency map, guiding gaze through extensive projections to the superior colliculus. V1 is the only visual cortical area that represents the eye of origin of an input and is also least associated with awareness; I therefore predicted that an ocular singleton (i.e., an item only shown to one eye among other items shown to the other eye) that is perceptually indistinct might nevertheless attract gaze. In visual searches for an orientation singleton target bar among uniformly oriented background bars, an ocular singleton non-target bar, at the same eccentricity as the target from the center of the search display, often captured the first search saccade. The chance of this capture was above 50% (e.g., 75%) when the eccentricity of the singletons was large and luminance did not vary between the bars, and it was below 50% when the eccentricity was smaller and luminance varied. After each search trial, observers reported whether an ocular singleton non-target (which was actually presented in half of the trials) had been shown. When different bars had similar luminance, misses numbered less than 50% and were independent of whether the gaze was captured by the ocular singleton. However, when luminance varied sufficiently between the bars, 50% were missed overall, albeit significantly less for those that captured gaze. The experiments in this work followed the guidelines in the Declaration of Helsinki.     

Binocular interactions in normal and anomalous binocular vision

Binocular interactions for grating patterns were investigated in humans with normal binocular vision and in humans with abnormal binocular visual experience due to strabismus and/or amblyopia via 1) comparison of monocular and binocular contrast thresholds; 2) interocular transfer of the threshold elevation aftereffect; and 3) dichoptic masking. Whereas the normal observers showed improved binocular over monocular contrast sensitivity (i.e., binocular summation) and substantial interocular transfer of the threshold elevation aftereffect, the abnormal observers showed an absence of binocular summation and no significant interocular transfer. The dichoptic masking experiments showed that a suprathreshold masking grating presented to one eye elevated the contrast threshold for gratings presented to the fellow eye, within a narrow range of spatial frequencies (about 1 octave wide at half height) and orientations, centered about the spatial frequency and orientation of the mask. The magnitude and bandwidth of this masking effect was similar in subjects with normal and abnormal binocular vision, occurring even when the masking grating was presented to the amblyopic eye. These effects depend upon the contrast of the masking grating. In individuals with normal binocular vision, a grating with subthreshold contrast presented to one eye reduces the contrast threshold for detection of gratings of similar spatial frequency and orientation presented to the fellow eye. No such subthreshold summation is evident in the amblyopic observers. We conclude that while strabismus and/or amblyopia disrupted the normal excitatory interactions between the two eyes, cortical inhibitory binocular connections were not disrupted.     

Interocular Transfer: The Dichoptic Flash-Lag Effect in Controls and Amblyopes

PurposeThe mammalian brain can take into account the neural delays in visual information transmission from the retina to the cortex when accurately localizing the instantaneous position of moving objects by motion extrapolation. In this study, we wanted to investigate whether such extrapolation mechanism operates in a comparable fashion between the eyes in normally sighted and amblyopic observers.MethodsTo measure interocular extrapolation, we adapted a dichoptic version of the flash-lag effect (FLE) paradigm, in which a flashed bar is perceived to lag behind a moving bar when their two positions are physically aligned. Twelve adult subjects with amblyopia and 12 healthy controls participated in the experiment. We measured the FLE magnitude of the subjects under binocular, monocular, and dichoptic conditions.ResultsIn controls, the FLE magnitude of binocular condition was significantly smaller than that of monocular conditions (P ≤ 0.023), but there was no difference between monocular and dichoptic conditions. Subject with amblyopia exhibited a smaller FLE magnitude in the dichoptic condition when the moving bar was presented to the amblyopic eye and the flash to the fellow eye (DA condition) compared to the opposite way around (DF condition), consistent with a delay in the processing of the amblyopic eye (P = 0.041).ConclusionsOur observations confirm that trajectory extrapolation mechanisms transfer between the eyes of normal observers. However, such transfer may be impaired in amblyopia. The smaller FLE magnitude in DA compared to DF in patients with amblyopia could be due to an interocular delay in the amblyopic visual system. The observation that normal controls present a smaller FLE in binocular conditions raises the question whether a larger FLE is or is not an indicator of better motion processing and extrapolation.     

Global shape discrimination at reduced contrast in enucleated observers

Previous research has shown that observers with early unilateral enucleation have selectively better sensitivity to luminance contrast than monocular viewing controls [González et al., 2002; Vision Research 36 (1) (1996) 175; Vision Research 36 (1996) 3011; Vision Research 37 (17) (1997) 2465]. We asked whether unilateral enucleation specifically enhances all levels of luminance processing. Enucleated observers, as well as binocular and monocular viewing controls, detected global shape in radial frequency (RF) patterns [Vision Research 38 (1998) 2555] at low contrast. Control observers were tested in two monocular conditions in which the stimulus was presented to one eye, while the fellow eye: (1) viewed a luminance-matched grey field or (2) was covered by a dark eye patch. Sensitivity to low-contrast global shape was equivalent in enucleated observers and binocular controls. More importantly, enucleated observers showed superior performance to that of controls in either monocular condition. At low contrast, the dichoptic control group was more sensitive than controls wearing an eye patch, which suggests that dichoptic viewing is a superior method of testing when comparing monocular control performance to that of monocularly deprived populations. The previously reported enhanced sensitivity to luminance-defined form in early enucleated observers also occurs for low-contrast global shape discrimination.     

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.     

Eye dominance effects in feature search

We studied the role of eye dominance in non-rivalry conditions, testing dichoptic visual search and comparing performance with target presented to the dominant or non-dominant eye. Using red-green glasses, subjects viewed an array of green and red lines of uniform orientation, with a differently oriented target line present on half the trials. Performance was significantly better when the dominant eye saw the target, especially when the opposite eye saw the distractors. This effect was reduced when only nearest-neighbor surrounding distractors were homogeneous. We conclude that the dominant eye has priority in visual processing, perhaps including inhibition of non-dominant eye representations.     

Looking behind a pathological blind spot in human retina.

Recent work suggests that dichoptic lateral interactions occur in the region of the visual field of one eye that corresponds to the physiological blind spot in the other eye (Tripathy, S. P., & Levi, D. M. (1994). The two-dimensional shape of spatial interaction zones in the parafovea. Vision Research, 34, 1127-1138.) Here we ask whether dichoptic lateral interactions occur in the region of the visual field of one eye that corresponds to a pathological blind spot, a retinal coloboma in the other eye. To address this question we had the observer report the orientation of a letter 'T' presented within this region in the presence of flanking 'T's presented to the other eye around the coloboma. A large drop in performance was seen due to the flanks, showing the existence of dichoptic lateral interactions in this monocular region. The presence of these dichoptic interactions in a region lacking direct retinal afferents from one eye is consistent with the proposition that long-range horizontal connections of the primary visual cortex mediate these interactions.     

Rod-cone interaction in monocular but not binocular pathways

Photopic background stimulation elevates scotopic increment thresholds (rod-cone interaction) at moderate background levels when both test and concentric disk-background stimuli enter the same eye (monocular condition) but not when they enter different eyes (dichoptic condition). Only when background levels are made extremely high is there any measurable dichoptic interaction, and this interaction does not resemble that observed monocularly. Rod-cone interaction, as usually studied, is a property of monocular pathways in human vision.     

No control in orientation search: the effects of instruction on oculomotor selection in visual search

The present study aimed to investigate whether people can selectively use salience information in search for a target. Observers were presented with a display consisting of multiple homogeneously oriented background lines and two orientation singletons. The orientation singletons differed in salience, where salience was defined by their orientation contrast relative to the background lines. Observers had the task to make a speeded eye movement towards a target, which was either the most or the least salient element of the two orientation singletons. The specific orientation of the target was either constant or variable over a block of trials such that observers had varying knowledge concerning the target identity. The results demonstrated that instruction - whether people were instructed to move to the most or the least salient item - only minimally affected the results. Short-latency eye movements were completely salience driven; here it did not matter whether people were searching for the most or least salient element. Long-latency eye movements were marginally affected by instruction, in particular when observers knew the target identity. These results suggest that even though people use salience information in oculomotor selection, they cannot use this information in a goal-driven manner. The results are discussed in terms of current models on visual selection.     

Dichoptic Perceptual Training and Sensory Eye Dominance Plasticity in Normal Vision

PurposeWe introduce a set of dichoptic training tasks that differ in terms of (1) the presence of external noise and (2) the visual feature implicated (motion, orientation), examining the generality of training effects between the different training and test cues and their capacity for driving changes in sensory eye dominance and stereoscopic depth perception.MethodsWe randomly assigned 116 normal-sighted observers to five groups (four training groups and one no training group). All groups completed both pre- and posttests, during which they were tested on dichoptic motion and orientation tasks under noisy and noise-free conditions, as well as a binocular phase combination task and two depth tasks to index sensory eye dominance and binocular function. Training groups received visual training on one of the four dichoptic tasks over 3 consecutive days.ResultsTraining under noise-free conditions supported generalization of learning to noise-free tasks involving an untrained feature. By contrast, there was a symmetric learning transfer between the signal-noise and no-noise tasks within the same visual feature. Further, training on all tasks reduced sensory eye dominance but did not improve depth perception.ConclusionsTraining-driven changes in sensory eye balance do not depend on the stimulus feature or whether the training entails the presence of external noise. We conjecture that dichoptic visual training acts to balance interocular suppression before or at the site of binocular combination.     

Learning to adapt: Dynamics of readaptation to geometrical distortions

The visual system can adapt to optical blur, whereby the adapted image is perceived as sharp. Here we show that adaptation reduces blur-induced biases in shape perception, with repeated adaptations (perceptual learning), leading to unbiased perception upon re-exposure to blur. Observers wore a cylindrical lens of +1.00 D on one eye, thus simulating monocular astigmatism. The other eye was either masked with a translucent blurred lens (monocular) or unmasked (dichoptic). Adaptation was tested in several repeated sessions with a proximity-grouping task, using horizontally or vertically arranged dot-arrays, without feedback, before, after, and throughout the adaptation period. A robust bias in global-orientation judgment was observed with the lens, in accordance with the blur axes. After the observer wore the lens for 2 h, there was no significant change in the bias, but after 4 h, the monocular condition, but not the dichoptic, resulted in reduced bias. The adaptation effect of the monocular 4-h adaptation was preserved, and even improved, when the lens was re-applied the next day, indicating learning. After-effects were observed under all experimental conditions except for the 4-h monocular condition, where learning took place. We suggest that, with long experience, adaptation is transferred to a long-term memory that can be instantly engaged when blur is re-applied, or disengaged when blur is removed, thus leaving no after-effects. The comparison between the monocular and dichoptic conditions indicates a binocular cortical site of plasticity.     

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.     

Reversed tilt effect for dichoptic stimulation in vertical meridian

The orientation of a test line appears shifted away from that of contextual contours. This repulsion in the orientation domain manifests itself in, for example, the tilt aftereffect, the simultaneous orientation contrast, and the Zöllner illusion. Attractive rather than repulsive interaction has been reported for collinear inducing configurations at small inducing angles. Here it is shown that the tilt induced in a monocular vertical foveal test line by a single centered inducing line is repulsive when both are in the same eye, but becomes attractive for dichoptic stimulation. This occurs for only a narrow range of test-line/inducing-line separations and varies somewhat between and, by ocularity, within observers. The phenomenon might be related to stereoscopy which involves the processing of dichoptic orientation differences around the vertical.     

Attention and scintillation

The scintillating grid is a recently discovered visual illusion, created by superimposing white discs on the intersections of orthogonal gray bars on a black background. Even though the entire display is stationary, observers perceive dark spots appearing and disappearing rapidly within some of the white discs. This scintillation effect is correlated with eye position and eye movements. Here we investigate whether covert shifts of attention, as revealed by cueing and visual search paradigms, can also affect the illusion. We find that the chance of a particular intersection scintillating is directly correlated with distance from the attentional focus, regardless of the location of the fixation point. This suggests that the dynamics of this illusion might reflect the distribution of attention in space and time.     

Perceptual learning to reduce sensory eye dominance beyond the focus of top-down visual attention

Perceptual learning is an important means for the brain to maintain its agility in a dynamic environment. Top-down focal attention, which selects task-relevant stimuli against competing ones in the background, is known to control and select what is learned in adults. Still unknown, is whether the adult brain is able to learn highly visible information beyond the focus of top-down attention. If it is, we should be able to reveal a purely stimulus-driven perceptual learning occurring in functions that are largely determined by the early cortical level, where top-down attention modulation is weak. Such an automatic, stimulus-driven learning mechanism is commonly assumed to operate only in the juvenile brain. We performed perceptual training to reduce sensory eye dominance (SED), a function that taps on the eye-of-origin information represented in the early visual cortex. Two retinal locations were simultaneously stimulated with suprathreshold, dichoptic orthogonal gratings. At each location, monocular cueing triggered perception of the grating images of the weak eye and suppression of the strong eye. Observers attended only to one location and performed orientation discrimination of the gratings seen by the weak eye, while ignoring the highly visible gratings at the second, unattended, location. We found SED was not only reduced at the attended location, but also at the unattended location. Furthermore, other untrained visual functions mediated by higher cortical levels improved. An automatic, stimulus-driven learning mechanism causes synaptic alterations in the early cortical level, with a far-reaching impact on the later cortical levels.     

A binocular site for contrast-modulated masking

Contrast-modulated (CM) gratings, composed of two luminance-modulated sinusoids of similar spatial frequency, mask the detection of test sinusoids at the difference frequency. However, the mechanism underlying masking by CM gratings remains poorly understood. In this paper, we aimed to determine whether the masking of 1 cycle deg(-1) LM test gratings by a 1 cycle deg(-1) beat (formed from a pair of carriers at 8 and 9 cycles deg(-1)) occurs in monocular channels or after the site of binocular combination, or both. Threshold elevations for the detection of a 1 cycle deg(-1) test grating were obtained for a number of stimulus conditions, including: (1) dichoptic CM (both 8 and 9 cycles deg(-1) mask components presented to one eye, with the 1 cycle deg(-1) test grating to the other); (2) dichoptic variant (8 and 9 cycles deg(-1) mask gratings presented to separate eyes, with the 1 cycle deg(-1) test grating presented to one eye); (3) binocular CM (all mask and test gratings presented to both eyes). As a control, masking magnitude was also measured for LM mask gratings of similar frequency (1 cycle deg(-1)) and effective contrast (3%) to that of the beat. For both LM and CM masks, the dichoptic condition yielded threshold elevations that were similar or greater than the binocular condition. When 8 and 9 cycles deg(-1) mask components were presented to separate eyes (the dichoptic variant condition), no beat pattern was visible and no elevations in detection threshold occurred. The results demonstrate that, like LM masking, detection of a target in the presence of a CM mask does not involve purely monocular mechanisms. Further, that the site of CM masking must occur beyond the stage at which monocular matching for stereopsis takes place. This is consistent with other studies which suggest that dichoptic masking is contingent on stereo matching, and thus occurs relatively late in the hierarchy of binocular visual processing.     

Interocular Suppression as Revealed by Dichoptic Masking Is Orientation-Dependent and Imbalanced in Amblyopia

PurposeWe investigate the orientation tuning of interocular suppression using a dichoptic masking paradigm in adult controls and amblyopes.MethodsFourteen adults with anisometropic or mixed amblyopia and 10 control adults participated in our study. Contrast sensitivity was measured by presenting a target Gabor in the tested eye and mean luminance in the untested eye (monocular) and by presenting a target in the tested eye and a bandpass oriented filtered noise in the other eye (masked). Interocular suppression was defined as the thresholds difference between the monocular and masked conditions for each eye. Interocular suppression was measured under parallel and orthogonal suppression configurations. The peak spatial frequency of the target and mask was 0.25 c/d in experiment 1 (low), 1.31 c/d in experiment 2 (mid), and 6.87 c/d in experiment 3 (high).ResultsThe masking suppression induced by the amblyopic eye was less strong than that induced by the fellow eye. The suppression from the fellow eye was similar to that observed in the controls. Interocular suppression under parallel configuration was less strong than under orthogonal configuration in amblyopes at low and mid spatial frequency, but not at high spatial frequency.ConclusionsWe demonstrate that the abnormal interocular masking in amblyopia displays the expected characteristic of orientation selectivity expected of normal controls at low and mid spatial frequency, but not at high spatial frequency. The dichoptic masking imbalance between the eyes of amblyopes results in a net suppression of the amblyopic eye during binocular viewing, modeling clinical suppression.     

Interocular interactions during acuity measurement in children and adults, and in adults with amblyopia

The binocular interactions that occur during dichoptic and binocular viewing were investigated using a letter acuity task in normally sighted children (age range 6-14 years) and adults, and in adults with anisometropic amblyopia. Our aims were to investigate the nature of binocular interactions that occur in each group, and the extent to which the characteristics of binocular interactions differ across the groups. The non-tested eye was occluded during monocular (baseline) viewing, and was allowed to view a uniform stimulus with fusion lock in dichoptic viewing. In adults and children with normal vision, acuity under dichoptic viewing was unchanged relative to monocular baseline in the dominant eyes, while acuity of the non-dominant eye improved under dichoptic viewing relative to baseline. The magnitude of dichoptic change in the non-dominant eyes was similar in the two normally sighted groups, but the dichoptic advantage was found to decrease with increasing age within the children tested. Binocular acuity was better than monocular acuity in normal subjects, and a decrease in binocular summation with age was noted within the age range of the children tested. In contrast, the amblyopic observers showed no change in acuity with viewing conditions. The results demonstrate development of interocular interactions during childhood, and wide inter-individual variation in pattern of interocular interactions among anisometropic amblyopic adults.     

Two-dimensional spatial distortions in human strabismic amblyopia

Two-dimensional space perception was measured in the central visual field of amblyopic subjects under special consideration of spatial distortions and spatial uncertainty. The subjects had to construct circles with different radii around a given fixation point. Subjects with deep strabismic amblyopia were showed to exhibit considerable distortion in the amblyopic eye. Each subject exhibited an individual distortion pattern. The dominant eye showed distortions similar to those seen in normal observers. Subjects with mild strabismic amblyopia, strabismic alternators and anisometropic amblyopes did not exhibit any significant spatial distortions. Data from dichoptic localization experiments indicate that anomalous retinal correspondence might contribute to monocular spatial distortions.     

Simultaneous binocular integration of the visual tilt effect in normal and stereoblind observers

Apparent displacement of an edge from the subjective visual vertical is typically induced by the simultaneous inspection of a large rotating textured background. In the present series of experiments we have tested the binocular integration of such an effect in normal and stereoblind observers in whom inspection of the rotating background and vision of the vertical edge are presented to different eyes. In normal subjects there was no difference between the binocular and the dichoptic viewing condition while in stereoblind subjects there was a clearcut decrease in the strength of the tilt effect under dichoptic conditions. A residual tilt effect, however, was still present in the latter group under dichoptic conditions and this shows that binocular interactions related to visual tilt can be dissociated from those required for stereopsis.