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1.  Crowding - an essential bottleneck for object recognition: a mini-review 
Vision research  2008;48(5):635-654.
Crowding, generally defined as the deleterious influence of nearby contours on visual discrimination, is ubiquitous in spatial vision. Crowding impairs the ability to recognize objects in clutter. It has been extensively studied over the last 80 years or so, and much of the renewed interest is the hope that studying crowding may lead to a better understanding of the processes involved in object recognition. Crowding also has important clinical implications for patients with macular degeneration, amblyopia and dyslexia.
There is no shortage of theories for crowding–from low-level receptive field models to high-level attention. The current picture is that crowding represents an essential bottleneck for object perception, impairing object perception in peripheral, amblyopic and possibly developing vision. Crowding is neither masking nor surround suppression. We can localize crowding to the cortex, perhaps as early as V1; however, there is a growing consensus for a two-stage model of crowding in which the first stage involves the detection of simple features (perhaps in V1), and a second stage is required for the integration or interpretation of the features as an object beyond V1. There is evidence for top-down effects in crowding, but the role of attention in this process remains unclear. The strong effect of learning in shrinking the spatial extent of crowding places strong constraints on possible models for crowding and for object recognition.
The goal of this review is to try to provide a broad, balanced and succinct review that organizes and summarizes the diverse and scattered studies of crowding, and also helps to explain it to the non-specialist. A full understanding of crowding may allow us to understand this bottleneck to object recognition and the rules that govern the integration of features into objects.
PMCID: PMC2268888  PMID: 18226828
Crowding; contour interaction; contour integration; contour segmentation; attentional resolution; object recognition; surround suppression; masking; peripheral vision; amblyopia
2.  Perceptual Learning Reduces Crowding in Amblyopia and in the Normal Periphery 
Amblyopia is a developmental visual disorder of cortical origin, characterized by crowding and poor acuity in central vision of the affected eye. Crowding refers to the adverse effects of surrounding items on object identification, common only in normal peripheral but not central vision. We trained a group of adult human amblyopes on a crowded letter identification task to assess whether the crowding problem can be ameliorated. Letter size was fixed well above the acuity limit, and letter spacing was varied to obtain spacing thresholds for central target identification. Normally sighted observers practiced the same task in their lower peripheral visual field. Independent measures of acuity were taken in flanked and unflanked conditions before and after training to measure crowding ratios at three fixed letter separations. Practice improved the letter spacing thresholds of both groups on the training task, and crowding ratios were reduced after posttest. The reductions in crowding in amblyopes were associated with improvements in standard measures of visual acuity. Thus, perceptual learning reduced the deleterious effects of crowding in amblyopia and in the normal periphery. The results support the effectiveness of plasticity-based approaches for improving vision in adult amblyopes and suggest experience-dependent effects on the cortical substrates of crowding.
PMCID: PMC3428833  PMID: 22238083
3.  The Effect of Aging on Crowded Letter Recognition in the Peripheral Visual Field 
Crowding describes the increased difficulty in identifying a target object when it is surrounded by nearby objects (flankers). A recent study investigated the effect of age on visual crowding and found equivocal results: Although crowded visual acuity was worse in older participants, crowding expressed as a ratio did not change with age. However, the spatial extent of crowding is a better index of crowding effects and remains unknown. In the present study, we used established psychophysical methods to characterize the effect of age on visual crowding (magnitude and extent) in a letter recognition task.
Letter recognition thresholds were determined for three different flanker separations in 54 adults (aged 18–76 years) with normal vision. Additionally, the spatial extent of crowding was established by measuring spacing thresholds: the flanker-to-target separation required to produce a given reduction in performance. Uncrowded visual acuity, crowded visual acuity, and spacing thresholds were expressed as a function of age, avoiding arbitrary categorization of young and old participants.
Our results showed that uncrowded and crowded visual acuities do not change significantly as a function of age. Furthermore, spacing thresholds did not change with age and approximated Bouma's law (half eccentricity).
These data show that crowding in adults is unaffected by senescence and provide additional evidence for distinct neural mechanisms mediating surround suppression and visual crowding, since the former shows a significant age effect. Finally, our data suggest that the well-documented age-related decline in peripheral reading ability is not due to age-related changes in visual crowding.
Crowding for a peripheral letter recognition task was measured as a function of age. Both the magnitude and spatial extent of crowding did not change with age. The results have important implications for the visual rehabilitation of patients with central vision loss.
PMCID: PMC4132554  PMID: 24985476
crowding; aging; visual acuity; critical spacing; reading
4.  A Neurophysiologically Plausible Population Code Model for Feature Integration Explains Visual Crowding 
PLoS Computational Biology  2010;6(1):e1000646.
An object in the peripheral visual field is more difficult to recognize when surrounded by other objects. This phenomenon is called “crowding”. Crowding places a fundamental constraint on human vision that limits performance on numerous tasks. It has been suggested that crowding results from spatial feature integration necessary for object recognition. However, in the absence of convincing models, this theory has remained controversial. Here, we present a quantitative and physiologically plausible model for spatial integration of orientation signals, based on the principles of population coding. Using simulations, we demonstrate that this model coherently accounts for fundamental properties of crowding, including critical spacing, “compulsory averaging”, and a foveal-peripheral anisotropy. Moreover, we show that the model predicts increased responses to correlated visual stimuli. Altogether, these results suggest that crowding has little immediate bearing on object recognition but is a by-product of a general, elementary integration mechanism in early vision aimed at improving signal quality.
Author Summary
Visual crowding refers to the phenomenon that objects become more difficult to recognize when other objects surround them. Recently there has been an explosion of studies on crowding, driven, in part, by the belief that understanding crowding will help to understand a range of visual behaviours, including object recognition, visual search, reading, and texture recognition. Given the long-standing interest in the topic and its relevance for a wide range of research fields, it is quite surprising that after nearly a century of research the mechanisms underlying crowding are still as poorly understood as they are today. A nearly complete lack of quantitative models seems to be one of the main reasons for this. Here, we present a mathematical, biologically motivated model of feature integration at the level of neuron populations. Using simulations, we demonstrate that several fundamental properties of the crowding effect can be explained as the by-product of an integration mechanism that may have a function in contour integration. Altogether, these results help differentiate between earlier theories about both the neural and functional origin of crowding.
PMCID: PMC2799670  PMID: 20098499
5.  Training-induced recovery of low-level vision followed by mid-level perceptual improvements in developmental object and face agnosia 
Developmental Science  2014;18(1):50-64.
Long-term deprivation of normal visual inputs can cause perceptual impairments at various levels of visual function, from basic visual acuity deficits, through mid-level deficits such as contour integration and motion coherence, to high-level face and object agnosia. Yet it is unclear whether training during adulthood, at a post-developmental stage of the adult visual system, can overcome such developmental impairments. Here, we visually trained LG, a developmental object and face agnosic individual. Prior to training, at the age of 20, LG's basic and mid-level visual functions such as visual acuity, crowding effects, and contour integration were underdeveloped relative to normal adult vision, corresponding to or poorer than those of 5–6 year olds (Gilaie-Dotan, Perry, Bonneh, Malach & Bentin, 2009). Intensive visual training, based on lateral interactions, was applied for a period of 9 months. LG's directly trained but also untrained visual functions such as visual acuity, crowding, binocular stereopsis and also mid-level contour integration improved significantly and reached near-age-level performance, with long-term (over 4 years) persistence. Moreover, mid-level functions that were tested post-training were found to be normal in LG. Some possible subtle improvement was observed in LG's higher-order visual functions such as object recognition and part integration, while LG's face perception skills have not improved thus far. These results suggest that corrective training at a post-developmental stage, even in the adult visual system, can prove effective, and its enduring effects are the basis for a revival of a developmental cascade that can lead to reduced perceptual impairments.
PMCID: PMC4309467  PMID: 24698161
6.  Training improves visual processing speed and generalizes to untrained functions 
Scientific Reports  2014;4:7251.
Studies show that manipulating certain training features in perceptual learning determines the specificity of the improvement. The improvement in abnormal visual processing following training and its generalization to visual acuity, as measured on static clinical charts, can be explained by improved sensitivity or processing speed. Crowding, the inability to recognize objects in a clutter, fundamentally limits conscious visual perception. Although it was largely considered absent in the fovea, earlier studies report foveal crowding upon very brief exposures or following spatial manipulations. Here we used GlassesOff's application for iDevices to train foveal vision of young participants. The training was performed at reading distance based on contrast detection tasks under different spatial and temporal constraints using Gabor patches aimed at testing improvement of processing speed. We found several significant improvements in spatio-temporal visual functions including near and also non-trained far distances. A remarkable transfer to visual acuity measured under crowded conditions resulted in reduced processing time of 81 ms, in order to achieve 6/6 acuity. Despite a subtle change in contrast sensitivity, a robust increase in processing speed was found. Thus, enhanced processing speed may lead to overcoming foveal crowding and might be the enabling factor for generalization to other visual functions.
PMCID: PMC4246693  PMID: 25431233
7.  Music-reading expertise alters visual spatial resolution for musical notation 
Psychonomic Bulletin & Review  2012;19(4):594-600.
Crowding occurs when the perception of a suprathreshold target is impaired by nearby distractors, reflecting a fundamental limitation on visual spatial resolution. It is likely that crowding limits music reading, as each musical note is crowded by adjacent notes and by the five-line staff, similar to word reading, in which letter recognition is reduced by crowding from adjacent letters. Here, we tested the hypothesis that, with extensive experience, music-reading experts have acquired visual skills such that they experience a smaller crowding effect, resulting in higher music-reading fluency. Experts experienced a smaller crowding effect than did novices, but only for musical stimuli, not for control stimuli (Landolt Cs). The magnitude of the crowding effect for musical stimuli could be predicted by individual fluency in music reading. Our results highlight the role of experience in crowding: Visual spatial resolution can be improved specifically for objects associated with perceptual expertise. Music-reading rates are likely limited by crowding, and our results are consistent with the idea that experience alleviates these limitations.
PMCID: PMC3394230  PMID: 22460744
Reading; Object recognition; Music cognition; Perceptual expertise; Crowding; Reading speed
8.  Training shortens search times in children with visual impairment accompanied by nystagmus 
Perceptual learning (PL) can improve near visual acuity (NVA) in 4–9 year old children with visual impairment (VI). However, the mechanisms underlying improved NVA are unknown. The present study compares feature search and oculomotor measures in 4–9 year old children with VI accompanied by nystagmus (VI+nys [n = 33]) and children with normal vision (NV [n = 29]). Children in the VI+nys group were divided into three training groups: an experimental PL group, a control PL group, and a magnifier group. They were seen before (baseline) and after 6 weeks of training. Children with NV were only seen at baseline. The feature search task entailed finding a target E among distractor E's (pointing right) with element spacing varied in four steps: 0.04°, 0.5°, 1°, and 2°. At baseline, children with VI+nys showed longer search times, shorter fixation durations, and larger saccade amplitudes than children with NV. After training, all training groups showed shorter search times. Only the experimental PL group showed prolonged fixation duration after training at 0.5° and 2° spacing, p's respectively 0.033 and 0.021. Prolonged fixation duration was associated with reduced crowding and improved crowded NVA. One of the mechanisms underlying improved crowded NVA after PL in children with VI+nys seems to be prolonged fixation duration.
PMCID: PMC4162385  PMID: 25309473
perceptual learning; fixation duration; visual impairment; visual search
9.  Prominent effects and neural correlates of visual crowding in a neurodegenerative disease population 
Brain  2014;137(12):3284-3299.
Visual crowding is a perceptual phenomenon whereby recognition of a stimulus is disrupted by the presence of flanker stimuli. Yong et al. observe excessive crowding in individuals with a neurodegenerative condition (posterior cortical atrophy) and identify associations between prominent crowding and lower grey matter volume in the right collateral sulcus.
Crowding is a breakdown in the ability to identify objects in clutter, and is a major constraint on object recognition. Crowding particularly impairs object perception in peripheral, amblyopic and possibly developing vision. Here we argue that crowding is also a critical factor limiting object perception in central vision of individuals with neurodegeneration of the occipital cortices. In the current study, individuals with posterior cortical atrophy (n = 26), typical Alzheimer’s disease (n = 17) and healthy control subjects (n = 14) completed centrally-presented tests of letter identification under six different flanking conditions (unflanked, and with letter, shape, number, same polarity and reverse polarity flankers) with two different target-flanker spacings (condensed, spaced). Patients with posterior cortical atrophy were significantly less accurate and slower to identify targets in the condensed than spaced condition even when the target letters were surrounded by flankers of a different category. Importantly, this spacing effect was observed for same, but not reverse, polarity flankers. The difference in accuracy between spaced and condensed stimuli was significantly associated with lower grey matter volume in the right collateral sulcus, in a region lying between the fusiform and lingual gyri. Detailed error analysis also revealed that similarity between the error response and the averaged target and flanker stimuli (but not individual target or flanker stimuli) was a significant predictor of error rate, more consistent with averaging than substitution accounts of crowding. Our findings suggest that crowding in posterior cortical atrophy can be regarded as a pre-attentive process that uses averaging to regularize the pathologically noisy representation of letter feature position in central vision. These results also help to clarify the cortical localization of feature integration components of crowding. More broadly, we suggest that posterior cortical atrophy provides a neurodegenerative disease model for exploring the basis of crowding. These data have significant implications for patients with, or who will go on to develop, dementia-related visual impairment, in whom acquired excessive crowding likely contributes to deficits in word, object, face and scene perception.
PMCID: PMC4240300  PMID: 25351740
crowding; lateral masking; Alzheimer’s disease; posterior cortical atrophy; acquired dyslexia
10.  Peripheral Vision of Youths with Low Vision: Motion Perception, Crowding, and Visual Search 
Effects of low vision on peripheral visual function are poorly understood, especially in children whose visual skills are still developing. The aim of this study was to measure both central and peripheral visual functions in youths with typical and low vision. Of specific interest was the extent to which measures of foveal function predict performance of peripheral tasks.
We assessed central and peripheral visual functions in youths with typical vision (n = 7, ages 10–17) and low vision (n = 24, ages 9–18). Experimental measures used both static and moving stimuli and included visual crowding, visual search, motion acuity, motion direction discrimination, and multitarget motion comparison.
In most tasks, visual function was impaired in youths with low vision. Substantial differences, however, were found both between participant groups and, importantly, across different tasks within participant groups. Foveal visual acuity was a modest predictor of peripheral form vision and motion sensitivity in either the central or peripheral field. Despite exhibiting normal motion discriminations in fovea, motion sensitivity of youths with low vision deteriorated in the periphery. This contrasted with typically sighted participants, who showed improved motion sensitivity with increasing eccentricity. Visual search was greatly impaired in youths with low vision.
Our results reveal a complex pattern of visual deficits in peripheral vision and indicate a significant role of attentional mechanisms in observed impairments. These deficits were not adequately captured by measures of foveal function, arguing for the importance of independently assessing peripheral visual function.
Peripheral visual processing in observers with low vision is characterized by a complex pattern of visual impairments. These deficits are not adequately captured by measures of foveal function, indicating the importance of independently assessing peripheral visual function.
PMCID: PMC3428114  PMID: 22836766
11.  Learning to Identify Near-Acuity Letters, either with or without Flankers, Results in Improved Letter Size and Spacing Limits in Adults with Amblyopia 
PLoS ONE  2012;7(4):e35829.
Amblyopia is a developmental abnormality that results in deficits for a wide range of visual tasks, most notably, the reduced ability to see fine details, the loss in contrast sensitivity especially for small objects and the difficulty in seeing objects in clutter (crowding). The primary goal of this study was to evaluate whether crowding can be ameliorated in adults with amblyopia through perceptual learning using a flanked letter identification task that was designed to reduce crowding, and if so, whether the improvements transfer to untrained visual functions: visual acuity, contrast sensitivity and the size of visual span (the amount of information obtained in one fixation). To evaluate whether the improvements following this training task were specific to training with flankers, we also trained another group of adult observers with amblyopia using a single letter identification task that was designed to improve letter contrast sensitivity, not crowding. Following 10,000 trials of training, both groups of observers showed improvements in the respective training task. The improvements generalized to improved visual acuity, letter contrast sensitivity, size of the visual span, and reduced crowding. The magnitude of the improvement for each of these measurements was similar in the two training groups. Perceptual learning regimens aimed at reducing crowding or improving letter contrast sensitivity are both effective in improving visual acuity, contrast sensitivity for near-acuity objects and reducing the crowding effect, and could be useful as a clinical treatment for amblyopia.
PMCID: PMC3340394  PMID: 22558234
12.  Foveal contour interactions and crowding effects at the resolution limit of the visual system 
Journal of vision  2007;7(2):25.1-2518.
We describe several experiments on contour interactions and crowding effects at the resolution limit of the visual system. As test stimuli we used characters that are often employed in optometric practice for testing visual acuity: Landolt C's, Snellen E's and rectangular gratings. We tested several hypotheses that have been put forward to explain contour interaction and crowding effects. In Experiment 1 and Experiment 2, Landolt C's were the test stimuli, and bars, or Landolt C's, or gratings served as distractors. In Experiment 1, we showed that neither scale invariance, nor spatial frequency selectivity are characteristic of foveal crowding effects. These results allowed us to conclude that mechanisms other than lateral masking contribute to observers' performance in ‘crowded’ tasks. Hess, Dakin & Kappor (2000) suggested that the spatial-frequency band most appropriate for target recognition is shifted by the surrounding bars to higher spatial frequencies that cannot be resolved by observers. Our Experiment 2 rejects this hypothesis as the experimental data do not follow theoretical predictions. In Experiment 3 we employed Snellen E's both as test stimuli and as distractors. The masking functions were similar to those measured in Experiment 1 when the test Landolt C was surrounded by Landolt C's. In Experiment 4 we extended the range of test stimuli to rectangular gratings; same-frequency or high-frequency gratings were distractors. In this case, if the distracting gratings had random orientation from trial to trial, the critical spacing was twice larger than in the first three experiments. If the orientation of the distractors was fixed during the whole experiment, the critical spacing was similar to that measured in the first three experiments. We suggest that the visual system can use different mechanisms for the discrimination of different test stimuli in the presence of particular surround. Different receptive fields with different spatial characteristics can be employed. To explain why crowding-effects at the resolution limit of the visual system are not scale invariant, we suggest that a range of stimuli, slightly varying in size, may all be processed by the same neural channel – the channel with the smallest receptive fields of the visual system.
PMCID: PMC2652120  PMID: 18217840
contour interaction; crowding effect; lateral masking; visual acuity; optotype; Landolt C; Snellen E; rectangular grating; spatial frequency
13.  Effects of Macromolecular Crowding on Protein Conformational Changes 
PLoS Computational Biology  2010;6(7):e1000833.
Many protein functions can be directly linked to conformational changes. Inside cells, the equilibria and transition rates between different conformations may be affected by macromolecular crowding. We have recently developed a new approach for modeling crowding effects, which enables an atomistic representation of “test” proteins. Here this approach is applied to study how crowding affects the equilibria and transition rates between open and closed conformations of seven proteins: yeast protein disulfide isomerase (yPDI), adenylate kinase (AdK), orotidine phosphate decarboxylase (ODCase), Trp repressor (TrpR), hemoglobin, DNA β-glucosyltransferase, and Ap4A hydrolase. For each protein, molecular dynamics simulations of the open and closed states are separately run. Representative open and closed conformations are then used to calculate the crowding-induced changes in chemical potential for the two states. The difference in chemical-potential change between the two states finally predicts the effects of crowding on the population ratio of the two states. Crowding is found to reduce the open population to various extents. In the presence of crowders with a 15 Å radius and occupying 35% of volume, the open-to-closed population ratios of yPDI, AdK, ODCase and TrpR are reduced by 79%, 78%, 62% and 55%, respectively. The reductions for the remaining three proteins are 20–44%. As expected, the four proteins experiencing the stronger crowding effects are those with larger conformational changes between open and closed states (e.g., as measured by the change in radius of gyration). Larger proteins also tend to experience stronger crowding effects than smaller ones [e.g., comparing yPDI (480 residues) and TrpR (98 residues)]. The potentials of mean force along the open-closed reaction coordinate of apo and ligand-bound ODCase are altered by crowding, suggesting that transition rates are also affected. These quantitative results and qualitative trends will serve as valuable guides for expected crowding effects on protein conformation changes inside cells.
Author Summary
The biophysical properties of proteins inside cells can be expected to be quite different from those typically measured by in vitro experiments in dilute solutions. In particular, intracellular macromolecular crowding may significantly affect the equilibria and transition rates between different conformations of a protein, and hence its functions. What are the trends and magnitudes of such crowding effects? We address this question here by applying a recently developed approach for modeling crowding. Seven proteins, each with structures for both an open state and a closed state, are studied. Crowding exerts significant effects on the open-closed equilibria of four proteins and more modest effects on the remaining three. Potentials of mean force along the open-closed reaction coordinate, and hence transition rates, are similarly affected. The extent of conformational changes is the main determinant for the magnitudes of crowding effects, but the protein size also plays an important role. The effects of crowding become stronger as the protein size increases. Conformational transitions of the ribosome, an extremely large complex, during translation are predicted to experience particularly strong effects of intracellular crowding. We conclude that deduction of intracellular behaviors from in vitro experiments requires explicit consideration of crowding effects.
PMCID: PMC2895631  PMID: 20617196
14.  Reducing Crowding by Weakening Inhibitory Lateral Interactions in the Periphery with Perceptual Learning 
PLoS ONE  2011;6(10):e25568.
We investigated whether lateral masking in the near-periphery, due to inhibitory lateral interactions at an early level of central visual processing, could be weakened by perceptual learning and whether learning transferred to an untrained, higher-level lateral masking known as crowding. The trained task was contrast detection of a Gabor target presented in the near periphery (4°) in the presence of co-oriented and co-aligned high contrast Gabor flankers, which featured different target-to-flankers separations along the vertical axis that varied from 2λ to 8λ. We found both suppressive and facilitatory lateral interactions at target-to-flankers distances (2λ - 4λ and 8λ, respectively) that were larger than those found in the fovea. Training reduces suppression but does not increase facilitation. Most importantly, we found that learning reduces crowding and improves contrast sensitivity, but has no effect on visual acuity (VA). These results suggest a different pattern of connectivity in the periphery with respect to the fovea as well as a different modulation of this connectivity via perceptual learning that not only reduces low-level lateral masking but also reduces crowding. These results have important implications for the rehabilitation of low-vision patients who must use peripheral vision to perform tasks, such as reading and refined figure-ground segmentation, which normal sighted subjects perform in the fovea.
PMCID: PMC3204963  PMID: 22065990
15.  Context and Crowding in Perceptual Learning on a Peripheral Contrast Discrimination Task: Context-Specificity in Contrast Learning 
PLoS ONE  2013;8(5):e63278.
Perceptual learning is an improvement in sensitivity due to practice on a sensory task and is generally specific to the trained stimuli and/or tasks. The present study investigated the effect of stimulus configuration and crowding on perceptual learning in contrast discrimination in peripheral vision, and the effect of perceptual training on crowding in this task. 29 normally-sighted observers were trained to discriminate Gabor stimuli presented at 9° eccentricity with either identical or orthogonally oriented flankers with respect to the target (ISO and CROSS, respectively), or on an isolated target (CONTROL). Contrast discrimination thresholds were measured at various eccentricities and target-flanker separations before and after training in order to determine any learning transfer to untrained stimulus parameters. Perceptual learning was observed in all three training stimuli; however, greater improvement was obtained with training on ISO-oriented stimuli compared to CROSS-oriented and unflanked stimuli. This learning did not transfer to untrained stimulus configurations, eccentricities or target-flanker separations. A characteristic crowding effect was observed increasing with viewing eccentricity and decreasing with target-flanker separation before and after training in both configurations. The magnitude of crowding was reduced only at the trained eccentricity and target-flanker separation; therefore, learning for contrast discrimination and for crowding in the present study was configuration and location specific. Our findings suggest that stimulus configuration plays an important role in the magnitude of perceptual learning in contrast discrimination and suggest context-specificity in learning.
PMCID: PMC3655984  PMID: 23696807
16.  Ideal observer analysis of crowding and the reduction of crowding through learning 
Journal of vision  2010;10(5):16.
Crowding is a prominent phenomenon in peripheral vision where nearby objects impede one’s ability to identify a target of interest. The precise mechanism of crowding is not known. We used ideal observer analysis and a noise-masking paradigm to identify the functional mechanism of crowding. We tested letter identification in the periphery with and without flanking letters and found that crowding increases equivalent input noise and decreases sampling efficiency. Crowding effectively causes the signal from the target to be noisier and at the same time reduces the visual system’s ability to make use of a noisy signal. After practicing identification of flanked letters without noise in the periphery for 6 days, subjects’ performance for identifying flanked letters improved (reduction of crowding). Across subjects, the improvement was attributable to either a decrease in crowding-induced equivalent input noise or an increase in sampling efficiency, but seldom both. This pattern of results is consistent with a simple model whereby learning reduces crowding by adjusting the spatial extent of a perceptual window used to gather relevant input features. Following learning, subjects with inappropriately large windows reduced their window sizes; while subjects with inappropriately small windows increased their window sizes. The improvement in equivalent input noise and sampling efficiency persists for at least 6 months.
PMCID: PMC3096759  PMID: 20616136
peripheral vision; crowding; perceptual learning; ideal observer analysis
17.  Uncovering foveal crowding? 
Scientific Reports  2014;4:4067.
Visual crowding, as context modulation, reduce the ability to recognize objects in clutter, sets a fundamental limit on visual perception and object recognition. It's considered that crowding does not exist in the fovea and extensive efforts explored crowding in the periphery revealed various models that consider several aspects of spatial processing. Studies showed that spatial and temporal crowding are correlated, suggesting a tradeoff between spatial and temporal processing of crowding. We hypothesized that limiting stimulus availability should decrease object recognition in clutter. Here we show, for the first time, that robust contour interactions exist in the fovea for much larger target-flanker spacing than reported previously: participants overcome crowded conditions for long presentations times but exhibit contour interaction effects for short presentation times. Thus, by enabling enough processing time in the fovea, contour interactions can be overcome, enabling object recognition. Our results suggest that contemporary models of context modulation should include both time and spatial processing.
PMCID: PMC3921636  PMID: 24518803
18.  The nature of letter crowding as revealed by first- and second-order classification images 
Journal of vision  2007;7(2):5.1-526.
Visual crowding refers to the marked inability to identify an otherwise perfectly identifiable object when it is flanked by other objects. Crowding places a significant limit on form vision in the visual periphery; its mechanism is, however, unknown. Building on the method of signal-clamped classification images (Tjan & Nandy, 2006), we developed a series of first- and second-order classification-image techniques to investigate the nature of crowding without presupposing any model of crowding. Using an “o” versus “x” letter-identification task, we found that (1) crowding significantly reduced the contrast of first-order classification images, although it did not alter the shape of the classification images; (2) response errors during crowding were strongly correlated with the spatial structures of the flankers that resembled those of the erroneously perceived targets; (3) crowding had no systematic effect on intrinsic spatial uncertainty of an observer nor did it suppress feature detection; and (4) analysis of the second-order classification images revealed that crowding reduced the amount of valid features used by the visual system and, at the same time, increased the amount of invalid features used. Our findings strongly support the feature-mislocalization or source-confusion hypothesis as one of the proximal contributors of crowding. Our data also agree with the inappropriate feature-integration account with the requirement that feature integration be a competitive process. However, the feature-masking account and a front-end version of the spatial attention account of crowding are not supported by our data.
PMCID: PMC2635026  PMID: 18217820
crowding; letter identification; peripheral vision; classification images
19.  Crowding during restricted and free viewing 
Vision research  2013;84:50-59.
Crowding impairs the perception of form in peripheral vision. It is likely to be a key limiting factor of form vision in patients without central vision. Crowding has been extensively studied in normally sighted individuals, typically with a stimulus duration of a few hundred milliseconds to avoid eye movements. These restricted testing conditions do not reflect the natural behavior of a patient with central field loss. Could unlimited stimulus duration and unrestricted eye movements change the properties of crowding in any fundamental way? We studied letter identification in the peripheral vision of normally sighted observers in three conditions: (i) a fixation condition with a brief stimulus presentation of 250 ms, (ii) another fixation condition but with an unlimited viewing time, and (iii) an unrestricted eye movement condition with an artificial central scotoma and an unlimited viewing time. In all conditions, contrast thresholds were measured as a function of target-to-flanker spacing, from which we estimated the spatial extent of crowding in terms of critical spacing. We found that presentation duration beyond 250 ms had little effect on critical spacing with stable gaze. With unrestricted eye movements and a simulated central scotoma, we found a large variability in critical spacing across observers, but more importantly, the variability in critical spacing was well correlated with the variability in target eccentricity. Our results assure that the large body of findings on crowding made with briefly presented stimuli remains relevant to conditions where viewing time is unconstrained. Our results further suggest that impaired oculomotor control associated with central vision loss can confound peripheral form vision beyond the limits imposed by crowding.
PMCID: PMC3665516  PMID: 23563172
crowding; form vision; peripheral vision; eye movements
20.  Factors Affecting Crowded Acuity: Eccentricity and Contrast 
Acuity measurement is a fundamental method to assess visual performance in the clinic. Little is known about how acuity measured in the presence of neighboring letters, as in the case of letter charts, changes with contrast and with non-foveal viewing. This information is crucial for acuity measurement using low-contrast charts and when patients cannot use their fovea. In this study, we evaluated how optotype acuity, with and without flankers, is affected by contrast and eccentricity.
Five young adults with normal vision identified the orientation of a Tumbling-E alone or in the presence of four flanking Tumbling-Es. Edge-to-edge letter spacing ranged from 1 to 20 bar widths. Stimuli were presented on a white background for 150 ms with Weber contrast ranging from −2.5% to −99%. Flankers had the same size and contrast as the target. Testings were performed at the fovea, 3, 5 and 10 degrees in the inferior visual field.
When plotted as a function of letter spacing, acuity remains unaffected by the presence of flankers until the flankers are within the critical spacing, which averages an edge-to-edge spacing of 4.4 bar widths at the fovea, and approximately 16 bar widths at all three eccentricities. Critical spacing decreases with a reduction in contrast. When plotted as a function of contrast, acuity only worsens when the contrast falls below approximately 24% at the fovea and 17% in the periphery, for flanked and unflanked conditions alike.
The letter spacing on conventional letter charts exceeds the critical spacing for acuity measurement at the fovea, at all contrast levels. Thus these charts are appropriate for assessing foveal acuity. In the periphery, the critical spacing is larger than the letter spacing on conventional charts. Consequently, these charts may underestimate the acuity measured in the periphery due to the effects of crowding.
PMCID: PMC3837536  PMID: 23770657
acuity; contrast; eccentricity; crowding; periphery; critical spacing; critical contrast
21.  The Glenn A. Fry Award Lecture 2012: Plasticity of the Visual System Following Central Vision Loss 
Following the onset of central vision loss, most patients develop an eccentric retinal location outside the affected macular region, the preferred retinal locus (PRL), as their new reference for visual tasks. The first goal of this paper is to present behavioral evidence showing the presence of experience-dependent plasticity in people with central vision loss. The evidence includes (1) the presence of oculomotor re-referencing of fixational saccades to the PRL; (2) the characteristics of the shape of the crowding zone (spatial region within which the presence of other objects affects the recognition of a target) at the PRL are more “foveal-like” instead of resembling those of the normal periphery; and (3) the change in the shape of the crowding zone at a para-PRL location that includes a component referenced to the PRL. These findings suggest that there is a shift in the referencing locus of the oculomotor and the sensory visual system from the fovea to the PRL for people with central vision loss, implying that the visual system for these individuals is still plastic and can be modified through experiences. The second goal of the paper is to demonstrate the feasibility of applying perceptual learning, which capitalizes on the presence of plasticity, as a tool to improve functional vision for people with central vision loss. Our finding that visual function could improve with perceptual learning presents an exciting possibility for the development of an alternative rehabilitative strategy for people with central vision loss.
PMCID: PMC3673767  PMID: 23670125
central vision loss; low vision; AMD; plasticity; eye movements; acuity; crowding; perceptual learning; rehabilitation
22.  Cortical Reorganization after Long-Term Adaptation to Retinal Lesions in Humans 
The Journal of Neuroscience  2013;33(46):18080-18086.
Single-unit recordings demonstrated that the adult mammalian visual cortex is capable of reorganizing after induced retinal lesions. In humans, whether the adult cortex is capable of reorganizing has only been studied using functional magnetic resonance imaging, with equivocal results. Here, we exploited the phenomenon of visual crowding, a major limitation on object recognition, to show that, in humans with long-standing retinal (macular) lesions that afflict the fovea and thus use their peripheral vision exclusively, the signature properties of crowding are distinctly different from those of the normal periphery. Crowding refers to the inability to recognize objects when the object spacing is smaller than the critical spacing. Critical spacing depends only on the retinal location of the object, scales linearly with its distance from the fovea, and is approximately two times larger in the radial than the tangential direction with respect to the fovea, thus demonstrating the signature radial–tangential anisotropy of the crowding zone. Using retinal imaging combined with behavioral measurements, we mapped out the crowding zone at the precise peripheral retinal locations adopted by individuals with macular lesions as the new visual reference loci. At these loci, the critical spacings are substantially smaller along the radial direction than expected based on the normal periphery, resulting in a lower scaling of critical spacing with the eccentricity of the peripheral locus and a loss in the signature radial–tangential anisotropy of the crowding zone. These results imply a fundamental difference in the substrate of cortical processing in object recognition following long-term adaptation to macular lesions.
PMCID: PMC3828461  PMID: 24227718
23.  A double dissociation of the acuity and crowding limits to letter identification, and the promise of improved visual screening 
Journal of Vision  2014;14(5):3.
Here, we systematically explore the size and spacing requirements for identifying a letter among other letters. We measure acuity for flanked and unflanked letters, centrally and peripherally, in normals and amblyopes. We find that acuity, overlap masking, and crowding each demand a minimum size or spacing for readable text. Just measuring flanked and unflanked acuity is enough for our proposed model to predict the observer's threshold size and spacing for letters at any eccentricity.
We also find that amblyopia in adults retains the character of the childhood condition that caused it. Amblyopia is a developmental neural deficit that can occur as a result of either strabismus or anisometropia in childhood. Peripheral viewing during childhood due to strabismus results in amblyopia that is crowding limited, like peripheral vision. Optical blur of one eye during childhood due to anisometropia without strabismus results in amblyopia that is acuity limited, like blurred vision. Furthermore, we find that the spacing:acuity ratio of flanked and unflanked acuity can distinguish strabismic amblyopia from purely anisometropic amblyopia in nearly perfect agreement with lack of stereopsis. A scatter diagram of threshold spacing versus acuity, one point per patient, for several diagnostic groups, reveals the diagnostic power of flanked acuity testing. These results and two demonstrations indicate that the sensitivity of visual screening tests can be improved by using flankers that are more tightly spaced and letter like.
Finally, in concert with Strappini, Pelli, Di Pace, and Martelli (submitted), we jointly report a double dissociation between acuity and crowding. Two clinical conditions—anisometropic amblyopia and apperceptive agnosia—each selectively impair either acuity A or the spacing:acuity ratio S/A, not both. Furthermore, when we specifically estimate crowding, we find a double dissociation between acuity and crowding. Models of human object recognition will need to accommodate this newly discovered independence of acuity and crowding.
PMCID: PMC4021854  PMID: 24799622
amblyopia; crowding; strabismic; anisometropic; acuity; screening; spacing:acuity ratio; critical spacing; threshold spacing; legibility; overlap masking; letter identification; object recognition
24.  Grouping and Crowding Affect Target Appearance over Different Spatial Scales 
PLoS ONE  2013;8(8):e71188.
Crowding is the impairment of peripheral target perception by nearby flankers. A number of recent studies have shown that crowding shares many features with grouping. Here, we investigate whether effects of crowding and grouping on target perception are related by asking whether they operate over the same spatial scale. A target letter T had two sets of flanking Ts of varying orientations. The first set was presented close to the target, yielding strong crowding. The second set was either close enough to cause crowding on their own or too far to cause crowding on their own. The Ts of the second set had the same orientation that either matched the target’s orientation (Grouped condition) or not (Ungrouped condition). In Experiment 1, the Grouped flankers reduced crowding independently of their distance from the target, suggesting that grouping operated over larger distances than crowding. In Experiments 2 and 3 we found that grouping did not affect sensitivity but produced a strong bias to report that the grouped orientation was present at the target location whether or not it was. Finally, we investigated whether this bias was a response or perceptual bias, rejecting the former in favor of a perceptual grouping explanation. We suggest that the effect of grouping is to assimilate the target to the identity of surrounding flankers when they are all the same, and that this shape assimilation effect differs in its spatial scale from the integration effect of crowding.
PMCID: PMC3742795  PMID: 23967164
25.  Modeling Magnification and Anisotropy in the Primate Foveal Confluence 
PLoS Computational Biology  2010;6(1):e1000651.
A basic organizational principle of the primate visual system is that it maps the visual environment repeatedly and retinotopically onto cortex. Simple algebraic models can be used to describe the projection from visual space to cortical space not only for V1, but also for the complex of areas V1, V2 and V3. Typically a conformal (angle-preserving) projection ensuring local isotropy is regarded as ideal and primate visual cortex is often regarded as an approximation of this ideal. However, empirical data show systematic deviations from this ideal that are especially relevant in the foveal projection. The aims of this study were to map the nature of anisotropy predicted by existing models, to investigate the optimization targets faced by different types of retino-cortical maps, and finally to propose a novel map that better models empirical data than other candidates. The retino-cortical map can be optimized towards a space-conserving homogenous representation or a quasi-conformal mapping. The latter would require a significantly enlarged representation of specific parts of the cortical maps. In particular it would require significant enlargement of parafoveal V2 and V3 which is not supported by empirical data. Further, the recently published principal layout of the foveal singularity cannot be explained by existing models. We suggest a new model that accurately describes foveal data, minimizing cortical surface area in the periphery but suggesting that local isotropy dominates the most foveal part at the expense of additional cortical surface. The foveal confluence is an important example of the detailed trade-offs between the compromises required for the mapping of environmental space to a complex of neighboring cortical areas. Our models demonstrate that the organization follows clear morphogenetic principles that are essential for our understanding of foveal vision in daily life.
Author Summary
Cortical areas V1, V2 and V3 each contain a complete and orderly representation of the visual environment, and their detailed layout is often studied. Here we discuss and compare algebraic mapping functions providing a transformation from visual field to cortical maps. Such algebraic models allow prediction of the layout in individuals from sparse data and advance our understanding of the mapping principles guiding the cortical organization in this important brain region. We show the deficiencies in previous work and provide the first plausible account of the combined layout of the first three visual maps, deriving an optimal compromise between the desirable principles of local isotropy of the maps and the extent of cortex that they occupy. This formal mapping scheme has only a few parameters but accurately predicts the layout of cortical areas, particularly the confluence of the three retinotopic areas in the central fovea, which we have only recently resolved. This scheme highlights the result that V2 and V3 have larger cortical representations than V1 in the central fovea, explaining the release from acuity crowding in this region. In summary, our novel model describes the high-resolution retinotopic mapping data to the level of their existing empirical accuracy.
PMCID: PMC2813258  PMID: 20126528

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