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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.
doi:10.1016/j.visres.2007.12.009
PMCID: PMC2268888  PMID: 18226828
Crowding; contour interaction; contour integration; contour segmentation; attentional resolution; object recognition; surround suppression; masking; peripheral vision; amblyopia
2.  A summary-statistic representation in peripheral vision explains visual crowding 
Journal of vision  2009;9(12):13.1-1318.
Peripheral vision provides a less faithful representation of the visual input than foveal vision. Nonetheless, we can gain a lot of information about the world from our peripheral vision, for example in order to plan eye movements. The phenomenon of crowding shows that the reduction of information available in the periphery is not merely the result of reduced resolution. Crowding refers to visual phenomena in which identification of a target stimulus is significantly impaired by the presence of nearby stimuli, or flankers. What information is available in the periphery? We propose that the visual system locally represents peripheral stimuli by the joint statistics of responses of cells sensitive to different position, phase, orientation, and scale. This “textural” representation by summary statistics predicts the subjective “jumble” of features often associated with crowding. We show that the difficulty of performing an identification task within a single pooling region using this representation of the stimuli is correlated with peripheral identification performance under conditions of crowding. Furthermore, for a simple stimulus with no flankers, this representation can be adequate to specify the stimulus with some position invariance. This provides evidence that a unified neuronal mechanism may underlie peripheral vision, ordinary pattern recognition in central vision, and texture perception. A key component of our methodology involves creating visualizations of the information available in the summary statistics of a stimulus. We call these visualizations “mongrels” and show that they are highly useful in examining how the early visual system represents the visual input. Mongrels enable one to study the “equivalence classes” of our model, i.e., the sets of stimuli that map to the same representation according to the model.
doi:10.1167/9.12.13
PMCID: PMC2923917  PMID: 20053104
peripheral vision; crowding; texture perception; texture synthesis; computational model
3.  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.
doi:10.3758/s13423-012-0242-x
PMCID: PMC3394230  PMID: 22460744
Reading; Object recognition; Music cognition; Perceptual expertise; Crowding; Reading speed
4.  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.
doi:10.1167/7.2.5
PMCID: PMC2635026  PMID: 18217820
crowding; letter identification; peripheral vision; classification images
5.  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.
doi:10.1371/journal.pcbi.1000646
PMCID: PMC2799670  PMID: 20098499
6.  Spatial Resolution of Conscious Visual Perception in Infants 
Psychological science  2010;21(10):1502-1509.
Conscious awareness of objects in the visual periphery is limited. This limit is not entirely the result of reduced visual acuity, but is primarily caused by crowding—the inability to identify an object when surrounded by clutter. Crowding represents a fundamental limitation of the visual system, and has to date been unexplored in infants. Do infants have a fine-grained “spotlight”, similar to adults, or a diffuse “lantern” that sets limits on what they can register in the periphery? An eye-tracking paradigm was designed to psychophysically measure crowding in 6- to 15-month-olds by showing pairs of faces at three eccentricities, in the presence or absence of flankers, and recording infants’ first saccade from central fixation to either face. Results reveal that infants can discriminate faces in the periphery, and flankers impair this ability as close as 3 degrees; the effective spatial resolution of visual perception increased with age but was only half that of adults.
doi:10.1177/0956797610382787
PMCID: PMC3038241  PMID: 20817914
inversion; crowding; attention; peripheral vision; Mooney face
7.  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.
doi:10.1523/JNEUROSCI.3845-11.2012
PMCID: PMC3428833  PMID: 22238083
8.  Peripheral Vision of Youths with Low Vision: Motion Perception, Crowding, and Visual Search 
Purpose.
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.
Methods.
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.
Results.
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.
Conclusions.
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.
doi:10.1167/iovs.12-10350
PMCID: PMC3428114  PMID: 22836766
9.  The Neural Correlates of Crowding-Induced Changes in Appearance 
Current Biology  2012;22(13):1199-1206.
Summary
Object recognition in the peripheral visual field is limited by crowding: the disruptive influence of nearby clutter [1, 2]. Despite its severity, little is known about the cortical locus of crowding. Here, we examined the neural correlates of crowding by combining event-related fMRI adaptation with a change-detection paradigm [3]. Crowding can change the appearance of objects, such that items become perceptually matched to surrounding objects; we used this change in appearance as a signature of crowding and measured brain activity that correlated with the crowded percept. Observers adapted to a peripheral patch of noise surrounded by four Gabor flankers. When crowded, the noise appears oriented and perceptually indistinguishable from the flankers. Consequently, substitution of the noise for a Gabor identical to the flankers (“change-same”) is rarely detected, whereas substitution for an orthogonal Gabor (“change-different”) is rarely missed. We predicted that brain areas representing the crowded percept would show repetition suppression in change-same trials but release from adaptation in change-different trials. This predicted pattern was observed throughout cortical visual areas V1–V4, increasing in strength from early to late visual areas. These results depict crowding as a multistage process, involving even the earliest cortical visual areas, with perceptual consequences that are increasingly influenced by later visual areas.
Highlights
► Crowding influences brain activity throughout the early retinotopic cortex ► Modulation increases from early (V1/V2) to late (V3/V4) visual areas ► The progressive influence of crowding parallels the increase in receptive-field size ► These findings suggest crowding is a multistage process
doi:10.1016/j.cub.2012.04.063
PMCID: PMC3396841  PMID: 22658599
10.  Visual Crowding: a fundamental limit on conscious perception and object recognition 
Trends in cognitive sciences  2011;15(4):160-168.
Crowding, the inability to recognize objects in clutter, sets a fundamental limit on conscious visual perception and object recognition throughout most of the visual field. Despite how widespread and essential it is to object recognition, reading, and visually guided action, a solid operational definition of what crowding is has only recently become clear. The goal of this review is to provide a broad-based synthesis of the most recent findings in this area, to define what crowding is and is not, and to set the stage for future work that will extend crowding well beyond low-level vision. Here we define five diagnostic criteria for what counts as crowding, and further describe factors that both escape and break crowding. All of these lead to the conclusion that crowding occurs at multiple stages in the visual hierarchy.
doi:10.1016/j.tics.2011.02.005
PMCID: PMC3070834  PMID: 21420894
11.  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
12.  Crowding Changes Appearance 
Current Biology  2010;20(6):496-501.
Summary
Crowding is the breakdown in object recognition that occurs in cluttered visual environments [1–4] and the fundamental limit on peripheral vision, affecting identification within many visual modalities [5–9] and across large spatial regions [10]. Though frequently characterized as a disruptive process through which object representations are suppressed [11, 12] or lost altogether [13–15], we demonstrate that crowding systematically changes the appearance of objects. In particular, target patches of visual noise that are surrounded (“crowded”) by oriented Gabor flankers become perceptually oriented, matching the flankers. This was established with a change-detection paradigm: under crowded conditions, target changes from noise to Gabor went unnoticed when the Gabor orientation matched the flankers (and the illusory target percept), despite being easily detected when they differed. Rotation of the flankers (leaving target noise unaltered) also induced illusory target rotations. Blank targets led to similar results, demonstrating that crowding can induce apparent structure where none exists. Finally, adaptation to these stimuli induced a tilt aftereffect at the target location, consistent with signals from the flankers “spreading” across space. These results confirm predictions from change-based models of crowding, such as averaging [16], and establish crowding as a regularization process that simplifies the peripheral field by promoting consistent appearance among adjacent objects.
Graphical Abstract
Highlights
► Patches of visual noise become perceptually oriented when crowded by Gabor elements ► Changes from crowded-noise targets to Gabors go unnoticed when perceptually matched ► Adaptation to crowded change gives tilt aftereffects in the target location ► Crowding promotes consistency by regularizing the peripheral visual field
doi:10.1016/j.cub.2010.01.023
PMCID: PMC2849014  PMID: 20206527
SYSNEURO
13.  Visuomotor Crowding: The Resolution of Grasping in Cluttered Scenes 
Reaching toward a cup of coffee while reading the newspaper becomes exceedingly difficult when other objects are nearby. Although much is known about the precision of visual perception in cluttered scenes, relatively little is understood about acting within these environments – the spatial resolution of visuomotor behavior. When the number and density of objects overwhelm visual processing, crowding results, which serves as a bottleneck for object recognition. Despite crowding, featural information of the ensemble persists, thereby supporting texture perception. While texture is beneficial for visual perception, it is relatively uninformative for guiding the metrics of grasping. Therefore, it would be adaptive if the visual and visuomotor systems utilized the clutter differently. Using an orientation task, we measured the effect of crowding on vision and visually guided grasping and found that the density of clutter similarly limited discrimination performance. However, while vision integrates the surround to compute a texture, action discounts this global information. We propose that this dissociation reflects an optimal use of information by each system.
doi:10.3389/neuro.08.049.2009
PMCID: PMC2783724  PMID: 19949462
crowding; clutter; grasping; kinematics; dual-visual systems; perception-action dissociation
14.  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.
doi:10.1167/7.2.25
PMCID: PMC2652120  PMID: 18217840
contour interaction; crowding effect; lateral masking; visual acuity; optotype; Landolt C; Snellen E; rectangular grating; spatial frequency
15.  The Clinical Course of the Idiopathic Epiretinal Membrane After Surgery 
Purpose
To evaluate the clinical course of visual acuity and foveal thickness in the idiopathic epiretinal membrane (ERM) after a vitrectomy with the use of triamcinolone.
Methods
We retrospectively reviewed the records of 30 patients (30 eyes) with ERM that were treated by vitrectomy from 2004 to 2008. Visual acuity and foveal thickness from optical coherence tomography imaging was obtained preoperatively and at every postoperative follow-up visit.
Results
Visual acuity improved by two or more lines of vision in 30%, 50%, 60%, and 70%, and stayed the same within ±1 line in 47%, 50%, 40%, and 30% at one month, three months, five months, and seven months after surgery. Twenty-three percents of the subjects deteriorated by two or more lines of vision within one month after surgery. None of the subjects had reduced vision three months after surgery. Foveal thickness decreased significantly after surgery. The mean thickness was 409.7±107.9 µm before surgery and 288.6±66.1 µm seven months after surgery. Parameters which were significantly correlated with the final visual acuity included preoperative visual acuity (0.683), preoperative foveal thickness (0.544), and final foveal thickness (0.643) (p<0.005).
Conclusions
Foveal thickness and visual acuity improved until seven months after the vitrectomy in patients with idiopathic ERM. Preoperative visual acuity, foveal thickness, and final foveal thickness had a significant correlation with the final visual acuity.
doi:10.3341/kjo.2009.23.4.249
PMCID: PMC2789947  PMID: 20046683
Epiretinal membrane; Foveal thickness; Optical coherence tomography; Visual acuity; Vitrectomy
16.  Recovery of a crowded object by masking the flankers: Determining the locus of feature integration 
Journal of vision  2009;9(10):4.1-4.9.
Object recognition is a central function of the visual system. As a first step, the features of an object are registered; these independently encoded features are then bound together to form a single representation. Here we investigate the locus of this “feature integration” by examining crowding, a striking breakdown of this process. Crowding, an inability to identify a peripheral target surrounded by flankers, results from “excessive integration” of target and flanker features. We presented a standard crowding display with a target C flanked by four flanker C's in the periphery. We then masked only the flankers (but not the target) with one of three kinds of masks—noise, metacontrast, and object substitution—each of which interferes at progressively higher levels of visual processing. With noise and metacontrast masks (low-level masking), the crowded target was recovered, whereas with object substitution masks (high-level masking), it was not. This places a clear upper bound on the locus of interference in crowding suggesting that crowding is not a low-level phenomenon. We conclude that feature integration, which underlies crowding, occurs prior to the locus of object substitution masking. Further, our results indicate that the integrity of the flankers, but not their identification, is crucial for crowding to occur.
doi:10.1167/9.10.4
PMCID: PMC2766569  PMID: 19810785
Object recognition; feature integration; crowding; masking; target recovery; extrastriate cortex
17.  Crowding by Invisible Flankers 
PLoS ONE  2011;6(12):e28814.
Background
Human object recognition degrades sharply as the target object moves from central vision into peripheral vision. In particular, one's ability to recognize a peripheral target is severely impaired by the presence of flanking objects, a phenomenon known as visual crowding. Recent studies on how visual awareness of flanker existence influences crowding had shown mixed results. More importantly, it is not known whether conscious awareness of the existence of both the target and flankers are necessary for crowding to occur.
Methodology/Principal Findings
Here we show that crowding persists even when people are completely unaware of the flankers, which are rendered invisible through the continuous flash suppression technique. Contrast threshold for identifying the orientation of a grating pattern was elevated in the flanked condition, even when the subjects reported that they were unaware of the perceptually suppressed flankers. Moreover, we find that orientation-specific adaptation is attenuated by flankers even when both the target and flankers are invisible.
Conclusions
These findings complement the suggested correlation between crowding and visual awareness. What's more, our results demonstrate that conscious awareness and attention are not prerequisite for crowding.
doi:10.1371/journal.pone.0028814
PMCID: PMC3237546  PMID: 22194919
18.  Learning to Identify Crowded Letters: Does It Improve Reading Speed? 
Vision research  2007;47(25):3150-3159.
Crowding, the difficulty in identifying a letter embedded in other letters, has been suggested as an explanation for slow reading in peripheral vision. In this study, we asked whether crowding in peripheral vision can be reduced through training on identifying crowded letters, and if so, whether these changes will lead to improved peripheral reading speed. We measured the spatial extent of crowding, and reading speeds for a range of print sizes at 10° inferior visual field before and after training. Following training, averaged letter identification performance improved by 88% at the trained (the closest) letter separation. The improvement transferred to other untrained separations such that the spatial extent of crowding decreased by 38%. However, averaged maximum reading speed improved by a mere 7.2%. These findings demonstrated that crowding in peripheral vision could be reduced through training. Unfortunately, the reduction in the crowding effect did not lead to improved peripheral reading speed.
doi:10.1016/j.visres.2007.08.017
PMCID: PMC2134936  PMID: 17928026
crowding; perceptual learning; training; reading
19.  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.
doi:10.1371/journal.pone.0035829
PMCID: PMC3340394  PMID: 22558234
20.  Dissociable effects of attention and crowding on orientation averaging 
Journal of vision  2009;9(11):28.1-2816.
It has been proposed that visual crowding—the breakdown in recognition that occurs when objects are presented in cluttered scenes—reflects a limit imposed by visual attention. We examined this idea in the context of an orientation averaging task, having subjects judge the mean orientation of a set of oriented signal elements either in isolation, or “crowded” by nearby randomly oriented elements. In some conditions, subjects also had to perform an attentionally demanding secondary task. By measuring performance at different levels of signal orientation variability, we show that crowding increases subjects’ local uncertainty (about the orientation of individual elements) but that diverting attention reduces their global efficiency (the effective number of elements they can average over). Furthermore, performance with the same stimulus-sequence, presented multiple times, reveals that crowding does not induce more stimulus-independent variability (as would be predicted by some accounts based on attention). We conclude that crowding and attentional load have dissociable perceptual consequences for orientation averaging, suggesting distinct neural mechanisms for both. For the task we examined, attention can modulate the effects of crowding by changing the efficiency with which information is analyzed by the visual system but since crowding changes local uncertainty, not efficiency, crowding does not reflect an attentional limit.
doi:10.1167/9.11.28
PMCID: PMC2927104  PMID: 20053091
crowding; attention; orientation; context
21.  Crowding in peripheral vision: why bigger is better 
Current biology : CB  2009;19(23):1988-1993.
Summary
We enjoy the illusion that visual resolution is high across the entire field of vision. However, this illusion can be easily dispelled by trying to identify objects in a cluttered environment out of the corner of your eye. This reflects in part, the well-known decline in visual resolution in peripheral vision, however, the main bottleneck for reading or object recognition in peripheral vision is the crowding. Objects that can be easily identified in isolation seem indistinct and jumbled in clutter. Crowding is thought to reflect inappropriate integration of the target and flankers in peripheral vision1,2. Here we uncover and explain a paradox in peripheral crowding: under certain conditions increasing the size or number of flanking rings results in a paradoxical decrease in the magnitude of crowding – i.e., the bigger or more numerous the flanks, the smaller the crowding. These surprising results are predicted by a model in which crowding is determined by the centroids of ≈ 4 – 8 independent features within ≈ 0.5 times the target eccentricity. These features are then integrated into a texture beyond the stage of feature analysis. We speculate that this process may contribute to the illusion of high resolution across the field of vision.
doi:10.1016/j.cub.2009.09.056
PMCID: PMC3045113  PMID: 19853450
22.  Isolated Foveal Hypoplasia: Tomographic, Angiographic and Autofluorescence Patterns 
Purpose. To report clinical aspects, tomographic, angiographic, and autofluorescence patterns of two cases of isolated foveal hypoplasia. Methods. Foveal hypoplasia was found in a 23-year-old male patient and in a 64-year-old woman with impaired visual acuity of unknown etiology that remained unchanged for years. Results. In the first case, spectral-domain optical coherence tomography (SD-OCT) showed reduced foveal pit and continuity of inner retinal layers in the fovea. Photoreceptor layer had a normal thickness centrally. The foveal avascular zone (FAZ) was absent in the flourescein angiogram (FA). Fundus autofluorescence showed reduced foveal attenuation of autofluorescence. In the second patient, there was the same pattern in SD-OCT, with normal aspect in FA and only a slightly reduced foveal attenuation of autofluorescence. Conclusion. OCT, as a noninvasive and quick method, is helpful in the diagnosis of foveal hypoplasia. FA and fundus autofluorescence were less sensitive.
doi:10.1155/2012/864958
PMCID: PMC3415089  PMID: 22900218
23.  Cholinergic enhancement reduces orientation-specific surround suppression but not visual crowding 
Acetylcholine (ACh) reduces the spatial spread of excitatory fMRI responses in early visual cortex and receptive field size of V1 neurons. We investigated the perceptual consequences of these physiological effects of ACh with surround suppression and crowding, two phenomena that involve spatial interactions between visual field locations. Surround suppression refers to the reduction in perceived stimulus contrast by a high-contrast surround stimulus. For grating stimuli, surround suppression is selective for the relative orientations of the center and surround, suggesting that it results from inhibitory interactions in early visual cortex. Crowding refers to impaired identification of a peripheral stimulus in the presence of flankers and is thought to result from excessive integration of visual features. We increased synaptic ACh levels by administering the cholinesterase inhibitor donepezil to healthy human subjects in a placebo-controlled, double-blind design. In Experiment 1, we measured surround suppression of a central grating using a contrast discrimination task with three conditions: (1) surround grating with the same orientation as the center (parallel), (2) surround orthogonal to the center, or (3) no surround. Contrast discrimination thresholds were higher in the parallel than in the orthogonal condition, demonstrating orientation-specific surround suppression (OSSS). Cholinergic enhancement decreased thresholds only in the parallel condition, thereby reducing OSSS. In Experiment 2, subjects performed a crowding task in which they reported the identity of a peripheral letter flanked by letters on either side. We measured the critical spacing between the targets and flanking letters that allowed reliable identification. Cholinergic enhancement with donepezil had no effect on critical spacing. Our findings suggest that ACh reduces spatial interactions in tasks involving segmentation of visual field locations but that these effects may be limited to early visual cortical processing.
doi:10.3389/fnbeh.2012.00061
PMCID: PMC3444757  PMID: 23049505
acetylcholine; surround suppression; crowding; pharmacology; psychophysics
24.  Stochastic re-calibration: contextual effects on perceived tilt 
The human visual system exaggerates the difference between the tilts of adjacent lines or grating patches. In addition to this tilt illusion, we found that oblique flanks reduced acuity for small changes of tilt in the centre of the visual field. However, no flanks—regardless of their tilts—decreased sensitivity to contrast. Thus, the foveal tilt illusion should not be attributed to orientation-selective lateral inhibition. Nor is it similar to conventional crowding, which typically does not impair letter recognition in the fovea. Our observers behaved as though the reference orientation (horizontal) had a small tilt in the direction of the flanks. We suggest that the extent of this re-calibration varies randomly over trials, and we demonstrate that this stochastic re-calibration can explain flank-induced acuity loss in the fovea.
doi:10.1098/rspb.2006.3634
PMCID: PMC1635463  PMID: 17002955
spatial vision; tilt illusion; crowding; lateral inhibition; orientation preference
25.  Brain visual impairment in childhood: mini review 
Hippokratia  2010;14(4):249-251.
Cerebral visual impairment (CVI) is one of the leading causes of severe visual impairment in childhood. This article was written to highlight any new knowledge related to cerebral visual impairment in childhood.
The international literature was searched to describe the type of visual, oculomotor and / or visuo-perceptual disturbances and to discuss the prognosis. CVI children show a wide range of visual disturbances. These could be either visual, oculomotor, perceptual or a combination of all. The severity of CVI depends on the time, location and extend of the brain damage.
The visual function seems to improve in CVI children, especially in the cortically damaged, mainly due to brain plasticity. The increased survival rate of very premature infants during the last decades has increased the incidence of CVI in childhood. Better understanding of the pathophysiological mechanisms of CVI, early diagnosis and early intervention could lead to a better quality of life of these children.
PMCID: PMC3031318  PMID: 21311632
cerebral visual impairment; hypoxic ischemic injury; periventricular leukomalacia; review

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