The Orientation Selectivity of Dichoptic Masking Suppression is Contrast Dependent in Amblyopia

PurposeWe aimed to study the effect of stimulus contrast on the orientation selectivity of interocular interaction in amblyopia using a dichoptic masking paradigm.MethodsEight adults with anisometropic or mixed amblyopia and 10 control adults participated in our study. The contrast threshold in discriminating a target Gabor in the tested eye was measured with mean luminance in the untested eye, as well as with a bandpass oriented filtered noise in the other eye at low spatial frequency (0.25 c/d). Threshold elevation, which represents interocular suppression, was assessed using a the dichoptic masking paradigm (i.e. the contrast threshold difference between the target only and masked conditions), for each eye. Orientation selectivity of the interocular suppression as reflected by dichoptic masking was quantified by the difference between the parallel and orthogonal masking configurations. Two levels of mask''s contrast (3 times or 10 times that of an individual''s contrast threshold) were tested in this study.ResultsThe strength of dichoptic masking suppression was stronger at high, rather than low mask contrast in both amblyopic and control subjects. Normal controls showed orientation-dependent dichoptic masking suppression both under high and low contrast levels. However, amblyopes showed orientation-tuned dichoptic masking suppression only under the high contrast level, but untuned under the low contrast level.ConclusionsWe demonstrate that interocular suppression assessed by dichoptic masking is contrast-dependent in amblyopia, being orientation-tuned only at high suprathreshold contrast levels of the mask.     

Monoptic and dichoptic metacontrast across the vertical meridian

That type of visual masking known as metacontrast was investigated with a viewing arrangement that imaged the test stimulus and the masking stimulus on different sides of the vertical meridian for both monoptic and dichoptic conditions. If the human visual pathways are in fact organized such that there is a “sharp” nasotemporal division along the vertical meridian, then in this experiment the test stimulus and the masking stimulus ought to have been available to different primary visual cortices, and any masking observed must have occurred beyond area 17. Such across-the-meridian masking was present in both the monoptic and dichoptic conditions; its magnitude was slightly greater monoptically than dichoptically when the test stimulus and the masking stimulus were juxtaposed, but monoptic and dichoptic masking were approximately equal when the test and masking stimuli were separated by 9.5′ or 19′ visual angle. Some difficulties with the classical accounts of the vertical meridian are discussed in relation to these results.     

Interocular suppression is gated by interocular feature matching

We present a new form of contrast masking in which the target is a patch of low spatial frequency grating (0.46 c/deg) and the mask is a dark thin ring that surrounds the centre of the target patch. In matching and detection experiments we found little or no effect for binocular presentation of mask and test stimuli. But when mask and test were presented briefly (33 or 200 ms) to different eyes (dichoptic presentation), masking was substantial. In a 'half-binocular' condition the test stimulus was presented to one eye, but the mask stimulus was presented to both eyes with zero-disparity. This produced masking effects intermediate to those found in dichoptic and full-binocular conditions. We suggest that interocular feature matching can attenuate the potency of interocular suppression, but unlike in previous work (McKee, S. P., Bravo, M. J., Taylor, D. G., & Legge, G. E. (1994) Stereo matching precedes dichoptic masking. Vision Research, 34, 1047) we do not invoke a special role for depth perception.     

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.     

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.     

Importance of phase alignment for interocular suppression

We measured contrast thresholds for Gabor targets in the presence of maskers which had higher or lower spatial frequencies than the targets. A high-pass fractal masker elevated target contrast thresholds at low and intermediate pedestal contrasts in both monocular and dichoptic modes of presentation, suggesting that the masking occurs after a monocular processing stage. Moreover we found that a high-pass checkerboard masker elevated thresholds at the low and intermediate pedestal contrasts and that most of this threshold elevation disappeared when the phase of the masker’s spatial components were scrambled. This masking was effective only in the dichoptic presentation, not in the monocular presentation. These results indicate that phase alignment of the high spatial frequency components plays a crucial role for interocular suppression. We speculate that phase alignments signal the existence of a luminance contour in the monocular image and that this signal suppresses processing of information in the other eye when there is no corresponding signal in that eye.     

Brightness perception and filling-in

Three experiments were performed in which a stimulus with homogeneous color and luminance was masked by a second stimulus containing contours. In the first experiment the target was a large white disk and the mask was a white circle concentric with the disk but of smaller radius. We found that the mask had a large (up to 2 log unit) suppressive effect on the brightness of the target, but only inside the radius of the mask. With monoptic presentation of target and mask, the greatest suppression was observed with an SOA of 50-100 msec. With dichoptic presentation the strongest suppression was obtained with simultaneous stimuli. The second experiment demonstrated that the latest time at which masking was effective was correlated with the distance between the edge of the target stimulus and the contour in the mask. One possible explanation of the results from these two experiments is that the masking contour is interfering with the propagation of a brightness signal from the target's border. In the third experiment gaps were introduced into the masking circle. Surprisingly, even with rather large gaps there was significant suppression of brightness in the center of the target. We have encountered difficulties attempting to account for these findings with known physiological mechanisms such as lateral inhibition. A qualitative explanation of the results that looks promising is a two-component process involving brightness filling-in and smoothing to satisfy fixed boundary conditions at contours.     

Development of rivalry and dichoptic masking in human infants

PURPOSE: To examine the development of rivalry, dichoptic masking, and binocular interactions in infants more than 5 months of age using the visual evoked potential (VEP). METHODS: VEPs were recorded in 35 infants between 5 and 15 months of age and 23 adults between 13 and 59 years of age. Counterphasing, sinusoidal, 1 cycle/deg gratings were presented dichoptically. Responses from each eye were isolated by "tagging" each half-image with a different temporal frequency (5 or 7.5 Hz). Observers were presented with fixed 80% contrast gratings in each eye in experiment 1. Rivalry was detected on the basis of a negative correlation between the simultaneously measured response amplitudes at the second harmonics of the two eye-tagging frequencies. In a second analysis of the same data, response amplitudes recorded under dichoptic viewing conditions were compared to those obtained in a monocular control condition (dichoptic masking). In experiment 2, a 40% fixed-contrast grating was presented to one eye, whereas the other eye viewed a grating that was swept in contrast from 1% to 67%. Dichoptic masking was measured as the reduction in the fixed-grating response caused by the variable contrast grating. RESULTS: Experiment 1: although adults showed evidence of VEP amplitude alternations between the eyes for cross-oriented half-images (physiological rivalry), infants did not. This immature response to rivalrous stimuli occurred despite the presence of responses at nonlinear combination frequencies recorded with gratings of the same orientation in each eye, a definitive indication of binocular interaction. In addition, both iso- and cross-oriented half-images produced less dichoptic masking in infants than in adults in this experiment. Experiment 2: dichoptic masking in the infants was equivalent to that seen in adults with parallel gratings in the two eyes; however, masking with cross-oriented configurations was approximately five times weaker in the infants relative to the adults. CONCLUSIONS: The authors have identified a set of stimulus conditions under which infants between 5 and 15 months of age fail to demonstrate physiological rivalry despite the presence of binocular interactions. The observed lack of binocular rivalry may be the result of a specific immaturity in dichoptic, cross-orientation suppression.     

Metacontrast masking between cyclopean and luminance stimuli

This study investigated the functional equivalence between cyclopean (global stereoscopic) and luminance (local stereoscopic) stimuli. To do so, a metacontrast masking paradigm was employed to determine the level of perceptual interaction between the two stimulus domains. Four target and mask combinations were used: cyclopean target-cyclopean mask, luminance target-luminance mask, cyclopean target-luminance mask, and luminance target-cyclopean mask. Substantial intradomain masking occurred, with the form of masking being similar for both domains. Moreover, significant interdomain masking also occurred, in equal measure for the cyclopean and luminance stimuli, although the magnitude of masking was one-half that of intradomain masking. These results imply that there is a functional equivalence at some stage of the visual system between the mechanisms representing cyclopean and luminance stimuli.     

Simultaneous electroretinograms and visually evoked potentials from adult amblyopes in response to a pattern stimulus.

Electroretinograms (ERGs) and visually evoked potentials (VEPs) were recorded simultaneously from each eye of three adult amblyopes. A spatially alternating checkerboard pattern stimulus of constant mean luminance was used to eliminate the effect of stray light on the ERG. The VEP was affected in the amblyopic eye of all subjects. In two subjects the VEP amplitude was reduced; in the third subject the amplitude was not attenuated, but the waveform of the VEP was markedly altered. Photopic ERGs recorded from the normal and amblyopic eye of each subject with an unpatterned flashing light were equal in amplitude. However, ERGs elicited by a patterned stimulus were affected in the amblyopic eye of all three subjects; the after-potential showed larger reductions in amplitude than the b-wave. These results suggest there may be some retinal involvement in human amblyopia.     

Binocular integration of contrast information in amblyopia.

We asked whether suppression in amblyopia could be accounted for by dichoptic masking as described in normals, operating in the presence of a contrast threshold difference between the two eyes. A dichoptic masking paradigm was employed to investigate binocular interaction in a mixed group of amblyopic subjects. Normal dichoptic masking was not seen after threshold differences between the two eyes were accounted for in the majority of subjects studied. We found that the binocular dysfunction did not merely follow as a consequence of the known monocular loss and that it depended upon the aetiology of the amblyopia and the spatial frequency of the stimulus.     

Luminance processing in apparent motion, Vernier offset and stereoscopic depth

We obtained (apparently) linear responses to luminance from three special displays of apparent motion, Vernier offset and stereoscopic depth. In our motion stimulus a dark and a light bar exchanged luminances repetitively on a grey surround. Motion was attributed to the bar that differed more from the surround, that is, on a dark surround the light bar appeared to jump, and on a light surround the dark bar appeared to jump. The apparent motion disappeared when the luminance of the surround lay halfway between that of the bars--on a linear, not a logarithmic scale. Similar results were obtained for special Vernier offset and stereo stimuli. These results cannot be explained if all luminances are processed within the same luminance pathway and that pathway transforms input luminance using non-linear compression. However, the apparent linearity of our results could arise from opposite and equal non-linearities cancelling out within separate ON- and OFF-spatial luminance pathways. A second set of experiments presented one bar separately into each eye on different surrounds (dichoptic presentation of competing apparent motion signals) or manipulated the display spatially so that different surrounds were associated with different bars (binocular presentation of competing Vernier targets). Results showed that apparent motion and Vernier signals of equal Weber contrast (normalisation of linear difference to surround luminance) evoked equal-motion and equal Vernier offset strengths. Given that motion and Vernier strength followed Weber's law, we infer that the ON- and OFF-pathways transform luminance non-linearly. Our third experiment presents an example of a brightness bisection task in which we were able to influence the bisection steps, to follow either a linear or non-linear series. The benefits of parsing the visual scene so that visual information is processed within two opposite luminance pathways is discussed.     

Metacontrast masking is specific to luminance polarity

A 1 degrees -spot was flashed up on a screen, followed by a snugly fitting annular mask. We measured the amount of masking as a function of stimulus luminance. The surround was always mid-gray, the masking ring was either black or white, and the luminance of the spot target ranged from 0% to 100% of white in 4% steps. Observers reported the apparent lightness of the masked spot by adjusting a matching spot. Results: A black annular mask made all spots that were darker than the gray surround appear to be transparent, that is, of the same luminance as the surround (complete masking). The black ring had virtually no masking effect on spots that were lighter than the surround. Conversely, a white ring made all spots that were lighter than the gray surround look apparently the same luminance as the surround (complete masking), but had virtually no masking effect on spots that were darker than the surround. In summary, a black ring masked spatial decrements but not increments, whilst a white ring masked spatial increments but not decrements. Thus masking occurred only when the spot and the ring had the same luminance polarity. This same-polarity masking still occurred when the target spot was larger than the 'donut hole' of the masking ring, so that the target and ring partly overlapped. This ruled out simple edge-cancellation theories. Instead, masking disrupts the filling-in process that normally propagates inward from the edges of a spot [Vision Res. 31 (7-8) (1991) 1221]. We conclude that metacontrast masking occurs within, but not between, separate visual ON and OFF pathways.     

Post-retinal processing of background luminance

It is generally thought that mean luminance and low spatial frequency information in a visual image are sharply attenuated at the retina, due to processes of light adaptation and the spatial filtering effects of lateral inhibition. Our results from interocular luminance masking suggest, however, that cortical masking effects play a primary role in the attenuation of low frequency sensitivity. Results also revealed that interocular luminance masking saturates and that semisaturation occurs where left and right eye luminances are equal, implying that the test luminance limits the effectiveness of the mask through interocular gating.     

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.     

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 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.     

Utrocular discrimination is not sufficient for utrocular identification

The distinction between reliability and validity is critical in examining utrocular identification. Four experiments demonstrated that two cues that lead to reliable discrimination do not lead to valid identification. Experiment 1 showed that, in the condition in which a stimulation of the right eye produced a visual direction toward the right and a stimulation of the left eye toward the left, there was a preponderance of correct responses. In the condition in which a stimulation of the right eye produced a visual direction toward the left and the left eye toward the right, there was a preponderance of incorrect responses. Experiment 2 showed that covariation of responses with visual direction decreased when feedback was provided because subjects sought other cues. Experiment 3, which included binocularly deficient subjects, showed that a feeling-in-the-eye is associated with the eye stimulated by a greater change in luminance rather than the eye stimulated by the target stimulus. When the luminance change was greater in the target eye, the feeling led to reliably correct responses, but when the luminance change was greater in the nontarget eye, it led to reliably incorrect responses. Experiment 4 indicated that the proportion of correct responses covaries with the degree of change in the luminance of the nontarget eye. The responses varied from reliably incorrect identifications, through unreliable identifications, to reliably correct identifications. These findings are consistent with the idea that stimulation of either eye is "projected" to the cyclopean eye.     

The function of contour in metacontrast.

The stimulus characteristics which constitute edge information for the visual system were studied by investigating the manner in which edge luminance is related to reduction in apparent brightness in metacontrast masking. Both targets and masks were briefly-flashed, nonoverlapping, rectangular apertures in which the luminance was modulated spatially in the horizontal dimension. The edge gradient was either gradual, a step function, or a step function with an additional spatial transient. The results showed that reduction in apparent brightness of the target is strongly influenced by the edge luminance of both the target and the mask. The data are discussed in terms of possible underlying mechanisms and with respect to other apparent brightness phenomena.     

Stereopsis and binocular rivalry from dichoptic stereograms

We employed stereograms having disparity and identical luminance and contrast, or disparate stereograms having binocular differences in luminance (luminance dichopticity) combined with either intermediate or extreme binocular differences in target-background contrast (contrast dichopticity). Stereoscopic depth was reported for the identical and dichoptic half-images with intermediate contrast dichopticity although attenuated depth estimates arose from the latter patterns. Binocular rivalry rather than depth was reported for the disparate patterns that carried luminance and extreme contrast dichopticities. The present results indicate that disparity information is. modified by accentuated dichoptic patterns.