Crowding under scotopic and photopic vision in albino and normal-sighted participants.

Crowding is a phenomenon in which the ability to recognize an object in a clutter deteriorates. It is, therefore, a fundamental aspect of object recognition and crucial in deciphering resolution. For visually impaired individuals, deficiency in crowding has a tremendous effect on vision and may reflect and predict the amount of deterioration in vision. It is well established that albinos suffer much more from crowding than normally sighted individuals under daylight luminance conditions. However, to our knowledge, this study is the first to investigate crowding in albino participants under low light conditions. In this study, we explored the crowding effect in a group of albino participants (n = 9) and a control group of normally sighted participants (n = 9). Crowding was conducted under daylight (photopic vision) and low light (scotopic vision). We measured the visual acuity threshold under crowding in three-letter spacing (0.5, 1, and 1.5) and compared it to a single target. Results indicate that albino participants experienced stronger crowding than the control under the photopic condition, while crowding under the scotopic condition was apparent in the albino but abolished for the control group. These findings highlight the importance of considering luminance when discussing the visually impaired population in general. In particular, it suggests that crowding in albinism is based on a peripheral-like mechanism and may indicate a cessation in visual development.

Probing the Bottleneck of Awareness Formed by Foveal Crowding: A Neurophysiological Study.

Crowding occurs when an easily identified isolated stimulus is surrounded by stimuli with similar properties, making it very difficult to identify. Crowding is suggested as a mechanism that creates a bottleneck in object recognition and awareness. Recently, we showed that brief presentation times at the fovea resulted in a significant crowding effect on target identification, impaired the target's color awareness, and resulted in a slower reaction time. However, when tagging the target with a red letter, the crowding effect is abolished. Crowding is widely considered a grouping; hence, it is pre-attentive. An event-related potential (ERP) study that investigated the spatial-temporal properties of crowding suggested the involvement of higher-level visual processing. Here, we investigated whether ERP's components may be affected by crowding and tagging, and whether the temporal advantage of ERP can be utilized to gain further information about the crowding mechanism. The participants reported target identification using our standard foveal crowing paradigm. It is assumed that crowding occurs due to a suppressive effect; thus, it can be probed by changes in perceptual (N1, ~160 ms) and attentive (P3 ~300-400 ms) components. We found a suppression effect (less negative ERP magnitude) in N1 under foveal crowding, which was recovered under tagging conditions. ERP's amplitude components (N1 and P3) and the behavioral proportion correct are highly correlated. These findings suggest that crowding is an early grouping mechanism that may be combined with later processing involving the segmentation mechanism.

Space, time, and dynamics of binocular interactions.

Binocular summation (BS), defined as the superiority of binocular over monocular visual performance, shows that thresholds are about 40% (a factor of 1.4) better in binocular than in monocular viewing. However, it was reported that different amounts of BS exist in a range from 1.4 to 2 values because BS is affected by the spatiotemporal parameters of the stimulus. Lateral interactions can be defined as the neuron's ability to affect the neighboring neurons by either inhibiting or exciting their activity. We investigated the effect of the spatial and temporal domains on binocular interactions and BS under the lateral masking paradigm and how BS would be affected by lateral interactions via a lateral masking experiment. The two temporal alternative forced-choice (2TAFC) method was used. The stimuli consisted of a central vertically oriented Gabor target and high-contrast Gabor flankers positioned in two configurations (orthogonal or collinear) with target-flanker separations of either 2 or 3 wavelengths (λ), presented at 4 different presentation times (40, 80, 120, and 200 ms) using a different order of measurements across the different experiments. Opaque lenses were used to control the monocular and binocular vision. BS is absent at close distances (2λ), depending on the presentation time's order, for the collinear but not for the orthogonal configuration. However, BS exists at more distant flankers (collinear and orthogonal, 3λ). BS is not uniform (1.4); it depends on the stimulus condition, the presentation times, the order, and the method that was used to control the monocular and binocular vision.

Abnormal basic visual processing functions in binocular fusion disorders.

Heterophoria is a common type of binocular fusion disorder that consists of a latent eye misalignment with potential consequences on daily activities such as reading or working on a computer (with CVS). Crowding, a type of contextual modulation, can also impair reading. Our recent studies found an abnormal pattern of low-level visual processing with larger perceptive fields (PF) in heterophoria. The PF is the fundamental processing unit of human vision and both masking and crowding depend on its size. We investigated how heterophoria would impact the PF's size via a lateral masking experiment and consequently affect the foveal crowding at different letter-spacings (the crowding zone). More specifically, we explored the relationship between crowding, lateral masking, the PF's size, and the amount of heterophoria. The binocular horizontal PF's size was larger with heterophoric subjects, in agreement with our previous study. We found a stronger crowding and an extended crowding zone associated with slower response times; this shows that the processing of letter identification under both crowded and uncrowded conditions requires more processing effort in heterophoric individuals. In agreement with previous studies, we found a correlation between the crowding zone and the PF's size; each was strongly correlated with the amount of phoria. These findings resemble those involving the PF size and the extended crowding found at the fovea in amblyopia and young children. We suggest that these findings could help explain the inter-observers' variability found in the masking literature, and the reading difficulties often encountered in subjects with high heterophoria.

Traces of early developmental bias in the adult brain.

During the first 2 years of life, there is a high prevalence of optical distortions in the human eye, causing vertical blur on the retina (astigmatism), which is naturally resolved by the age of 5; thus, it is not treated. Here we determined the possible long-term effects on visual grouping resulting from optical distortions during the development of visual perception. Our results show a clear directional bias in shape perception for optically corrected astigmatic adults, compared with non-astigmatic ones, with remarkably slow decision times. These effects can be explained by a mismatch between the developmental timescales of different components in the visual system.

Meridional binocular rivalry reveals a trace of uncorrected oblique input during development in the adult brain.

Binocular rivalry (BR) is a visual perception phenomenon that occurs when each eye perceives different images and stimuli, causing alternating monocular dominance. To measure BR, many studies have used two monocular conflicting images to induce monocular alternations. Here we chose a group of participants with oblique astigmatism (OA) and who produced blur on the orthogonal oblique meridian in each eye, resulting in two conflicting images, which may enhance the stimulation of monocular alternations. Our results show that OA participants tend to have a high rate of BR when viewing natural images, whereas the control group does not have BR for the same images. We suggest that this low ability to fuse could indicate the presence of a trace due to uncorrected vision during the critical period, which could be retained in the adult brain.

Extended perceptive field revealed in humans with binocular fusion disorders.

Binocular vision disorders or dysfunctions have considerable impact on daily visual activities such as reading. Heterophoria (phoria) is a latent eye misalignment (with a prevalence of up to 35%) that appears in conditions that disrupt binocular vision and it may affect the quality of binocular fusion. Our recent study, which used lateral masking (LM), suggests that subjects with binocular fusion disorders (horizontal phoria) exhibit an asymmetry and an abnormal pattern of both binocular and monocular lateral interactions, but only for the horizontal meridian (HM). The perceptive field (PF) is the fundamental processing unit of human vision and both masking and crowding depend on its size. An increased PF size is found in amblyopic populations or in young children. We hypothesized that the PF's size would be asymmetric only for the phoric group (larger along the HM). We estimated the PF's size using two different methods (LM with equal-phase and opposite-phase flankers). Phoric subjects exhibited a larger binocular PF size, only for the HM, confirming our hypothesis of an asymmetric PF size. However, the monocular PF size of phoric and control subjects was similar. Phoria affects the PF's size similarly to meridional amblyopia but without being attributed to abnormal refraction. We suggest that these findings could help explain the inter-observer variability found in the masking literature and the reading difficulties often encountered in subjects with high heterophoria. Since perceptual learning can reduce the PF's size, further investigation of training may provide a novel therapy to reduce some symptoms related to heterophoria.

Binocular fusion disorders impair basic visual processing.

In an era of increasing screen consumption, the requirement for binocular vision is demanding, leading to the emergence of syndromes such as the computer vision syndrome (CVS) or visual discomfort reported by virtual reality (VR) users. Heterophoria (phoria) is a latent eye misalignment (with a prevalence up to 35%) that appears in conditions that disrupt binocular vision and may affect the quality of binocular fusion. Collinear facilitation (CF), the mechanism for grouping contour elements, is a process that reveals lateral interactions by improving the visibility of a target by flankers placed collinearly. An abnormal pattern of CF has been observed in strabismic amblyopia. We hypothesize that phoria may affect CF in the horizontal meridian (HM) due to latent eye misalignment and its impact on binocular fusion. Fully corrected participants (phoria group and controls) completed a standard CF experiment for horizontal and vertical meridians during binocular and monocular viewing. Phoric observers exhibited (1) an asymmetry and an abnormal pattern of CF only for the HM, during both monocular and binocular viewing, (2) poor binocular summation between the monocular inputs, and (3) no binocular advantage of the CF. Phoria affects the CF in a way that is reminiscent of meridional amblyopia without being attributed to abnormal refraction. The abnormal pattern of CF in monocular viewing suggests that phoria could be a binocular developmental disorder that affects monocular spatial interactions. We suggest that the results could contribute to explain the visual discomfort experienced with VR users or symptoms when presenting CVS.

Nearby contours abolish the binocular advantage.

That binocular viewing confers an advantage over monocular viewing for detecting isolated low luminance or low contrast objects, has been known for well over a century; however, the processes involved in combining the images from the two eyes are still not fully understood. Importantly, in natural vision, objects are rarely isolated but appear in context. It is well known that nearby contours can either facilitate or suppress detection, depending on their distance from the target and the global configuration. Here we report that at close distances collinear (but not orthogonal) flanking contours suppress detection more under binocular compared to monocular viewing, thus completely abolishing the binocular advantage, both at threshold and suprathreshold levels. In contrast, more distant flankers facilitate both monocular and binocular detection, preserving a binocular advantage up to about four times the detection threshold. Our results for monocular and binocular viewing, for threshold contrast discrimination without nearby flankers, can be explained by a gain control model with uncertainty and internal multiplicative noise adding additional constraints on detection. However, in context with nearby flankers, both contrast detection threshold and suprathreshold contrast appearance matching require the addition of both target-to-target and flank-to-target interactions occurring before the site of binocular combination. To test an alternative model, in which the interactions occur after the site of binocular combination, we performed a dichoptic contrast matching experiment, with the target presented to one eye, and the flanks to the other eye. The two models make very different predictions for abutting flanks under dichoptic conditions. Interactions after the combination site predict that the perceived contrast of the flanked target will be strongly suppressed, while interactions before the site predict the perceived contrast will be more or less veridical. The data are consistent with the latter model, strongly suggesting that the interactions take place before the site of binocular combination.

Binocular summation is affected by crowding and tagging.

In perceptual crowding, a letter easily recognized on its own, becomes unrecognizable if it is surrounded by other letters, an effect that confers a limit on the visual processing. Models assume that crowding is a hallmark of the periphery but that it is almost absent in the fovea. However, recently it was shown that crowding occurs in the fovea of people with an abnormal development of functional vision (amblyopia), when the stimulus is presented for a very short time. When targets and flankers are dissimilar, the crowding is reduced (tagging). Since a combination of binocular inputs increases the processing load, we investigated whether color tagging the target reduces crowding in the fovea of subjects with normal vision and determined how crowding is combined with binocular vision. The crowding effect at the fovea was significantly reduced by tagging with a color target. Interestingly, whereas binocular summation for a single letter was expected to be about 40%, it was significantly reduced and almost absent under crowding conditions. Our results are consistent with the notion that the crowding effect produces a high processing load on visual processing, which interferes with other processes such as binocular summation. We assume that the tagging effect in our experiment improved the subject's abilities (sensitivity and RT) by creating a "segmentation", i.e., a visual simulated separation between the target letter and the background. Interestingly, tagging the target with a distinct color can eliminate or reduce the crowding effect and consequently, binocular summation recovers.

Development of global visual processing: From the retina to the perceptive field.

Young children exhibit poorer visual performance than adults due to immaturity of the fovea and of the fundamental processing of visual functions such as masking and crowding. Recent studies suggest that masking and crowding are closely related to the size of the fundamental processing unit-the perceptive field (PF). However, while it is known that the retina and basic visual functions develop throughout childhood, it is not clear whether and how changes in the size of the PF affect masking and crowding. Furthermore, no retinal and perceptual development data have been collected from the same cohort and time. Here we explored the developmental process of the PF and the basic visual functions. Psychophysical and imaging methods were used to test visual functions and foveal changes in participants ranging from 3-17 years old. Lateral masking, crowding and contrast sensitivity were tested using computerized tasks. Foveal measurements were obtained from spectral-domain optical coherence tomography (OCT). The children patterns below 6 years exhibited high crowding, while the expected facilitation was found only at a larger target-flanker distance than required for children above 6 years, who exhibited the typical adult. Foveal thickness and macular volume for the children below 6 years were significantly lower than for the older group. Significant correlation was found for contrast sensitivity, foveal thickness and macular volume with age and between contrast sensitivity and foveal thickness. Our data suggest that the developmental processes at the retina and visual cortex occur in the same age range. Thus, in parallel to maturation of the PF, which enables reduction in crowding, foveal development contributes to increasing contrast sensitivity.

Distorted optical input affects human perception.

Collinear facilitation, the mechanism for grouping contour elements, is a process involving lateral interactions that improve the detectability of a target by the presence of collinear flankers. It was shown that the development of collinear facilitation is experience dependent and that it may be impaired when the visual input is distorted in one meridian (meridional amblyopia). In oblique astigmatism, the blurring is on the opposite oblique meridian in both eyes, resulting in two conflicting images, which may affect the development of binocular vision. We hypothesized that the collinear facilitation of adults with oblique astigmatism is reminiscent of the abnormal development of the lateral facilitation of meridional amblyopia. We explored the perception of binocular vision and collinear facilitation in cases of both distorted and non-distorted vision. Fully corrected participants that tested for the target contrast detection of Gabor patches and two collinear flankers, presented for 80 ms, were positioned at different orientations (0° (180°), 45°, 90°, and 135°) and for different eyes (monocular, binocular). The results show a significant anisotropy for monocular collinear facilitation between the blured and the clear meridians, being lower in the blurriest meridian than in the clearest meridian, resembling the meridional amblyopia results. Collinear facilitation results in poor binocular summation between the monocular channels. Our results indicate that the perceptual behavior was similar to that of meridional amblyopic subjects having an anisotropy of collinear facilitation between cardinal meridians in oblique astigmatic subjects.

Monocular and Binocular Temporal Visual Perception of Infantile Nystagmus.

Contrast sensitivity is mostly used as a tool for testing aspects of visual functions. Infantile nystagmus is a pathological phenomenon that affects the spatial-temporal visual functions due to spontaneous oscillating movements of the eyes. We examined the spatial-temporal aspects of nystagmus perception, aiming to investigate the mechanisms underlying the deterioration of their visual performance. We tested the monocular and binocular contrast sensitivity of nystagmus and normally sighted subjects by measuring contrast detection of a Gabor target with spatial frequencies slightly above the cutoff threshold of each subject (nystagmus ~3; controls = 9cpd; presentation times 60-480 ms). The dominant eye of nystagmus revealed large differences over the non-dominant eye, highlighting the superiority of the dominant over the non-dominant eye in nystagmus. In addition, binocular summation mechanism was impaired in majority of the nystagmus subjects. Furthermore, these differences are not attributed to differences in visual acuity. Moreover, the visual performance in nystagmus continue to improve for longer presentation time compared with controls and was longer in the poor eye. Since the results are not due to differences in eye movements and strabismus, we suggest that the differences are due to developmental impairment in the visual system during the critical period.

Spatial visual function in anomalous trichromats: Is less more?

Color deficiency is a common inherited disorder affecting 8% of Caucasian males with anomalous trichromacy (AT); it is the most common type of inherited color vision deficiency. Anomalous trichromacy is caused by alteration of one of the three cone-opsins' spectral sensitivity; it is usually considered to impose marked limitations for daily life as well as for choice of occupation. Nevertheless, we show here that anomalous trichromat subjects have superior basic visual functions such as visual acuity (VA), contrast sensitivity (CS), and stereo acuity, compared with participants with normal color vision. Both contrast sensitivity and stereo acuity performance were correlated with the severity of color deficiency. We further show that subjects with anomalous trichromacy exhibit a better ability to detect objects camouflaged in natural gray scale figures. The advantages of color-deficient subjects in spatial vision performance could explain the relatively high prevalence of color-vision polymorphism in humans.

Vision improvement in pilots with presbyopia following perceptual learning.

Israeli Air Force (IAF) pilots continue flying combat missions after the symptoms of natural near-vision deterioration, termed presbyopia, begin to be noticeable. Because modern pilots rely on the displays of the aircraft control and performance instruments, near visual acuity (VA) is essential in the cockpit. We aimed to apply a method previously shown to improve visual performance of presbyopes, and test whether presbyopic IAF pilots can overcome the limitation imposed by presbyopia. Participants were selected by the IAF aeromedical unit as having at least initial presbyopia and trained using a structured personalized perceptual learning method (GlassesOff application), based on detecting briefly presented low-contrast Gabor stimuli, under the conditions of spatial and temporal constraints, from a distance of 40 cm. Our results show that despite their initial visual advantage over age-matched peers, training resulted in robust improvements in various basic visual functions, including static and temporal VA, stereoacuity, spatial crowding, contrast sensitivity and contrast discrimination. Moreover, improvements generalized to higher-level tasks, such as sentence reading and aerial photography interpretation (specifically designed to reflect IAF pilots' expertise in analyzing noisy low-contrast input). In concert with earlier suggestions, gains in visual processing speed are plausible to account, at least partially, for the observed training-induced improvements.

Temporal asynchrony and spatial perception.

Collinear facilitation is an enhancement in the visibility of a target by laterally placed iso-oriented flankers in a collinear (COL) configuration. Iso-oriented flankers placed in a non-collinear configuration (side-by-side, SBS) produce less facilitation. Surprisingly, presentation of both configurations simultaneously (ISO-CROSS) abolishes the facilitation rather than increases it - a phenomenon that can't be fully explained by the spatial properties of the target and flankers. Based on our preliminary data and recent studies, we hypothesized that there might be a novel explanation based on the temporal properties of the excitation and inhibition, resulting in asynchrony between the lateral inputs received from COL and SBS, leading to cancelation of the facilitatory component in ISO-CROSS. We explored this effect using a detection task in humans. The results replicated the previous results showing that the preferred facilitation for COL and SBS was abolished for the ISO-CROSS configuration. However, presenting the SBS flankers, but not the COL flankers 20 msec before ISO-CROSS restored the facilitatory effect. We propose a novel explanation that the perceptual advantage of collinear facilitation may be cancelled by the delayed input from the sides; thus, the final perception is determined by the overall spatial-temporal integration of the lateral interactions.

Gains following perceptual learning are closely linked to the initial visual acuity.

The goal of the present study was to evaluate the dependence of perceptual learning gains on initial visual acuity (VA), in a large sample of subjects with a wide range of VAs. A large sample of normally sighted and presbyopic subjects (N = 119; aged 40 to 63) with a wide range of uncorrected near visual acuities (VA, -0.12 to 0.8 LogMAR), underwent perceptual learning. Training consisted of detecting briefly presented Gabor stimuli under spatial and temporal masking conditions. Consistent with previous findings, perceptual learning induced a significant improvement in near VA and reading speed under conditions of limited exposure duration. Our results show that the improvements in VA and reading speed observed following perceptual learning are closely linked to the initial VA, with only a minor fraction of the observed improvement that may be attributed to the additional sessions performed by those with the worse VA.

Space and time in masking and crowding.

Masking and crowding are major phenomena associated with contextual modulations, but the relationship between them remains unclear. We have recently shown that crowding is apparent in the fovea when the time available for processing is limited, pointing to the strong relationship between crowding in the spatial and temporal domains. Models of crowding emphasize the size (acuity) of the target and the spacing between the target and flankers as the main determinants that predict crowding. Our model, which is based on lateral interactions, posits that masking and crowding are related in the spatial and temporal domains at the fovea and periphery and that both can be explained by the increasing size of the human perceptive field (PF) with increasing eccentricity. We explored the relations between masking and crowding using letter identification and contrast detection by correlating the crowding effect with the estimated size of the PF and with masking under different spatiotemporal conditions. We found that there is a large variability in PF size and crowding effects across observers. Nevertheless, masking and crowding were both correlated with the estimated size of the PF in the fovea and periphery under a specific range of spatiotemporal parameters. Our results suggest that under certain conditions, crowding and masking share common neural mechanisms that underlie the spatiotemporal properties of these phenomena in both the fovea and periphery. These results could explain the transfer of training gains from spatiotemporal Gabor masking to letter acuity, reading, and reduced crowding.

Crowding is proportional to visual acuity in young and aging eyes.

Spatial crowding decreases object recognition and conscious visual perception in clutter. In a previous study we used brief presentation times to reveal the effects of a crowded presentation in the fovea. Here we aimed to test the relationships between varying visual acuity (VA) and crowding in the fovea, under the assumption that in uncorrected presbyopia, the processing is relatively normal, whereas the retinal input is blurred. We tested whether normal participants whose near VA is gradually reduced due to age-related deterioration (presbyopia, or "aging eye") will show an acuity-dependent increase in foveal crowding. We used brief presentations and acuity-threshold letter targets in order to magnify the crowding-effect amplitude in the fovea. A total of 195 participants with an age range of 20-68 years and an average of 44.3 ± 11.7 years (M ± SD) were divided into four age groups, all without any optical correction for the near distance. Our findings show that crowding is proportional to VA. This proportionality is affected by VA-age dependency, with a nonlinear S-shaped pattern: A steep VA reduction begins to develop, which is compatible with the normal onset age of presbyopia symptoms and a saturation in the VA-age dependency in the oldest age group, for which we propose a VA-eccentricity account. Finally, there is a high variance in the crowding amplitude in the young, even before the onset age of presbyopia symptoms, suggesting crowding conditions with limited presentation times as a highly sensitive measure of VA, which predicts visual performance in complex tasks, such as reading.

Training on spatiotemporal masking improves crowded and uncrowded visual acuity.

Spatial crowding impairs conscious visual perception and object recognition in clutter.Short presentation times produce crowding in the normal fovea, in young participants and in uncorrected presbyopes ("aging eye"), measured as reduced visual acuity (VA). On the other hand, perceptual learning improves near VA in healthy young adults, in presbyopia, and in amblyopia. Here we aimed at exploring the effects of perceptual training on crowded VA in uncorrected presbyopes with naturally decreased VA, with two specific objectives: (a) to objectively measure crowded VA, taking advantage of the natural deterioration of near visual acuity from being normal or better than normal (i.e., 20/20 or better) in young participants to naturally decreasing in uncorrected presbyopes; and (b) to explore whether perceptual training previously shown to improve visual functions as complex as reading will affect crowded VA. Visual acuity was measured under crowded and uncrowded conditions by having subjects identify letters presented for short durations ranging from 34 to 116 msec. Training consisted of detecting brief Gabor stimuli under spatial and temporal masking conditions, using the GlassesOff mobile application (GlassesOff, Inc., New York, NY)on iOS devices from a distance of 40 cm. Before training, a robust reduction in crowded VA was found in the fovea of presbyopes. Training resulted in significant improvement of letter identification under both crowded and uncrowded VA conditions for all stimulus durations. After training, the crowded condition threshold reached the level of the uncrowded threshold measured before training. Thus, training enabled the subjects to overcome the effect of reduced VA under the crowded condition. We suggest that more efficient spatial and temporal processing induced by perceptual learning allows one to improve crowded VA, including that found on naturally reduced near VA, and that this effect may transfer to improve complex visual functions, such as reading and object recognition.