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1.  Prentice Award Lecture 2011: Removing the Brakes on Plasticity in the Amblyopic Brain 
Optometry and Vision Science  2012;89(6):827-838.
Experience-dependent plasticity is closely linked with the development of sensory function. Beyond this sensitive period, developmental plasticity is actively limited; however, new studies provide growing evidence for plasticity in the adult visual system. The amblyopic visual system is an excellent model for examining the “brakes” that limit recovery of function beyond the critical period. While amblyopia can often be reversed when treated early, conventional treatment is generally not undertaken in older children and adults. However new clinical and experimental studies in both animals and humans provide evidence for neural plasticity beyond the critical period. The results suggest that perceptual learning and video game play may be effective in improving a range of visual performance measures and importantly the improvements may transfer to better visual acuity and stereopsis. These findings, along with the results of new clinical trials, suggest that it might be time to re-consider our notions about neural plasticity in amblyopia.
PMCID: PMC3369432  PMID: 22581119
amblyopia; stereopsis; perceptual learning; plasticity; video-game play
2.  Vernier perceptual learning transfers to completely untrained retinal locations after double training: A “piggybacking” effect 
Journal of Vision  2014;14(13):12.
Perceptual learning, a process in which training improves visual discrimination, is often specific to the trained retinal location, and this location specificity is frequently regarded as an indication of neural plasticity in the retinotopic visual cortex. However, our previous studies have shown that “double training” enables location-specific perceptual learning, such as Vernier learning, to completely transfer to a new location where an irrelevant task is practiced. Here we show that Vernier learning can be actuated by less location-specific orientation or motion-direction learning to transfer to completely untrained retinal locations. This “piggybacking” effect occurs even if both tasks are trained at the same retinal location. However, piggybacking does not occur when the Vernier task is paired with a more location-specific contrast-discrimination task. This previously unknown complexity challenges the current understanding of perceptual learning and its specificity/transfer. Orientation and motion-direction learning, but not contrast and Vernier learning, appears to activate a global process that allows learning transfer to untrained locations. Moreover, when paired with orientation or motion-direction learning, Vernier learning may be “piggybacked” by the activated global process to transfer to other untrained retinal locations. How this task-specific global activation process is achieved is as yet unknown.
PMCID: PMC4233766  PMID: 25398974
perceptual learning; transfer; double training; Vernier
3.  Perceptual Learning as a potential treatment for amblyopia: a mini-review 
Vision research  2009;49(21):2535-2549.
Amblyopia is a developmental abnormality that results from physiological alterations in the visual cortex and impairs form vision. It is a consequence of abnormal binocular visual experience during the “sensitive period” early in life. While amblyopia can often be reversed when treated early, conventional treatment is generally not undertaken in older children and adults. A number of studies over the last twelve years or so suggest that Perceptual Learning (PL) may provide an important new method for treating amblyopia.
The aim of this mini-review is to provide a critical review and “meta-analysis” of perceptual learning in adults and children with amblyopia, with a view to extracting principles that might make PL more effective and efficient. Specifically we evaluate:
What factors influence the outcome of perceptual learning?Specificity and generalization – two sides of the coin.Do the improvements last?How does PL improve visual function?Should PL be part of the treatment armamentarium?
A review of the extant studies makes it clear that practicing a visual task results in a long-lasting improvement in performance in an amblyopic eye. The improvement is generally strongest for the trained eye, task, stimulus and orientation, but appears to have a broader spatial frequency bandwidth than in normal vision. Importantly, practicing on a variety of different tasks and stimuli seems to transfer to improved visual acuity. Perceptual learning operates via a reduction of internal neural noise and/or through more efficient use of the stimulus information by retuning the weighting of the information. The success of PL raises the question of whether it should become a standard part of the armamentarium for the clinical treatment of amblyopia, and suggests several important principles for effective perceptual learning in amblyopia.
PMCID: PMC2764839  PMID: 19250947
Amblyopia; perceptual learning; sensitive period; critical period; internal noise; template-retuning; occlusion
4.  Perceptual Learning Improves Adult Amblyopic Vision Through Rule-Based Cognitive Compensation 
We investigated whether perceptual learning in adults with amblyopia could be enabled to transfer completely to an orthogonal orientation, which would suggest that amblyopic perceptual learning results mainly from high-level cognitive compensation, rather than plasticity in the amblyopic early visual brain.
Nineteen adults (mean age = 22.5 years) with anisometropic and/or strabismic amblyopia were trained following a training-plus-exposure (TPE) protocol. The amblyopic eyes practiced contrast, orientation, or Vernier discrimination at one orientation for six to eight sessions. Then the amblyopic or nonamblyopic eyes were exposed to an orthogonal orientation via practicing an irrelevant task. Training was first performed at a lower spatial frequency (SF), then at a higher SF near the cutoff frequency of the amblyopic eye.
Perceptual learning was initially orientation specific. However, after exposure to the orthogonal orientation, learning transferred to an orthogonal orientation completely. Reversing the exposure and training order failed to produce transfer. Initial lower SF training led to broad improvement of contrast sensitivity, and later higher SF training led to more specific improvement at high SFs. Training improved visual acuity by 1.5 to 1.6 lines (P < 0.001) in the amblyopic eyes with computerized tests and a clinical E acuity chart. It also improved stereoacuity by 53% (P < 0.001).
The complete transfer of learning suggests that perceptual learning in amblyopia may reflect high-level learning of rules for performing a visual discrimination task. These rules are applicable to new orientations to enable learning transfer. Therefore, perceptual learning may improve amblyopic vision mainly through rule-based cognitive compensation.
Amblyopic perceptual learning is cognitive compensation.
PMCID: PMC3974581  PMID: 24550359
amblyopia; perceptual learning; orientation
5.  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
6.  The response of the amblyopic visual system to noise 
Vision research  2007;47(19):2531-2542.
Visual perception is limited by both the strength of the neural signals, and by the noise in the visual nervous system. Here we use one-dimensional white noise as input, to study the response of amblyopic visual system. We measured the thresholds for detection and discrimination of noise contrast. Using an N-Pass reverse correlation technique, we derived classification images and estimated response consistency.
Our results provide the first report of the sensitivity of the amblyopic visual system to white noise. We show that amblyopes have markedly reduced sensitivity for detecting noise, particularly at high spatial frequencies, and much less loss for discriminating suprathreshold noise contrast. Compensating for the detection loss almost (but not quite) equates performance of the amblyopic and normal visual system.
The classification images suggest that the amblyopic visual system contains adjustable channels for noise, similar to those found in normal vision, but “tuned” to slightly lower spatial frequencies than in normal observers. Our N-pass results show that the predominant factor limiting performance in our task in both normal and amblyopic vision is internal random multiplicative noise. For the detection of white noise the raised thresholds of the amblyopic visual system can be attributed primarily to extra additive noise. However, for the discrimination of suprathreshold white noise contrast, there is surprisingly little additional deficit, after accounting for the visibility of the noise.
PMCID: PMC2099256  PMID: 17697689
white noise; psychophysics; amblyopia; classification images; reverse-correlation; double-pass; internal noise
7.  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
8.  Task relevancy and demand modulate double-training enabled transfer of perceptual learning 
Vision Research  2011;61:33-38.
Location-specific perceptual learning can be rendered transferrable to a new location with double training, in which feature training (e.g., contrast) is accompanied by additional location training at the new location even with an irrelevant task (e.g. orientation). Here we investigated the impact of relevancy (to feature training) and demand of location training tasks on double training enabled learning transfer. We found that location training with an irrelevant task (Gabor vs. letter judgment, or contrast discrimination) limited transfer of Vernier learning to the trained orientation only. However, performing a relevant suprathreshold orthogonal Vernier task prompted additional transfer to an untrained orthogonal orientation. In addition, the amount of learning transfer may depend on the demand of location training as well as the double training procedure. These results characterize how double training potentiates the functional connections between a learned high-level decision unit and visual inputs from an untrained location to enable transfer of learning across retinal locations.
PMCID: PMC3227777  PMID: 21820004
Perceptual learning; double training; location specificity; transfer
9.  A Weber-like law for perceptual learning 
Scientific Reports  2013;3:1158.
What determines how much an organism can learn? One possibility is that the neural factors that limit sensory performance prior to learning, place an upper limit on the amount of learning that can take place. We tested this idea by comparing learning on a sensory task where performance is limited by cortical mechanisms, at two retinal eccentricities. Prior to learning, visual performance at the two eccentricities was either unmatched or equated in two different ways (through spatial scaling or visual crowding). The magnitude of learning was equivalent when initial levels of performance were matched regardless of how performance was equated. The magnitude of learning was a constant proportion of initial performance. This Weber-like law for perceptual learning demonstrates that it should be possible to predict the degree of perceptual improvement and the final level of performance that can be achieved via sensory training, regardless of what cortical constraint limits performance.
PMCID: PMC3557449  PMID: 23362458
10.  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
11.  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.
PMCID: PMC3070834  PMID: 21420894
12.  Reduced sampling efficiency causes degraded Vernier hyperacuity with normal aging: Vernier acuity in position noise 
Scientific Reports  2012;2:300.
Vernier acuity, a form of visual hyperacuity, is amongst the most precise forms of spatial vision. Under optimal conditions Vernier thresholds are much finer than the inter-photoreceptor distance. Achievement of such high precision is based substantially on cortical computations, most likely in the primary visual cortex. Using stimuli with added positional noise, we show that Vernier processing is reduced with advancing age across a wide range of noise levels. Using an ideal observer model, we are able to characterize the mechanisms underlying age-related loss, and show that the reduction in Vernier acuity can be mainly attributed to the reduction in efficiency of sampling, with no significant change in the level of internal position noise, or spatial distortion, in the visual system.
PMCID: PMC3293147  PMID: 22393476
13.  Training the brain to overcome the effect of aging on the human eye 
Scientific Reports  2012;2:278.
Presbyopia, from the Greek for aging eye, is, like death and taxes, inevitable. Presbyopia causes near vision to degrade with age, affecting virtually everyone over the age of 50. Presbyopia has multiple negative effects on the quality of vision and the quality of life, due to limitations on daily activities – in particular, reading. In addition presbyopia results in reduced near visual acuity, reduced contrast sensitivity, and slower processing speed. Currently available solutions, such as optical corrections, are not ideal for all daily activities. Here we show that perceptual learning (repeated practice on a demanding visual task) results in improved visual performance in presbyopes, enabling them to overcome and/or delay some of the disabilities imposed by the aging eye. This improvement was achieved without changing the optical characteristics of the eye. The results suggest that the aging brain retains enough plasticity to overcome the natural biological deterioration with age.
PMCID: PMC3284862  PMID: 22363834
14.  Visual deficits in anisometropia 
Vision research  2010;51(1):48-57.
Amblyopia is usually associated with the presence of anisometropia, strabismus or both early in life. We set out to explore quantitative relationships between the degree of anisometropia and the loss of visual function, and to examine how the presence of strabismus affects visual function in observers with anisometropia. We measured optotype acuity, Pelli-Robson contrast sensitivity and stereoacuity in 84 persons with anisometropia and compared their results with those of 27 persons with high bilateral refractive error (isoametropia) and 101 persons with both strabismus and anisometropia. All subjects participated in a large scale study of amblyopia (McKee, Levi & Movshon, 2003). We found no consistent visual abnormalities in the strong eye, and therefore report only on vision in the weaker, defined as the eye with lower acuity. LogMAR acuity falls off markedly with increasing anisometropia in non-strabismic anisometropes, while contrast sensitivity is much less affected. Acuity degrades rapidly with increases in both hyperopic and myopic anisometropia, but the risk of amblyopia is about twice as great in hyperopic than myopic anisometropes of comparable refractive imbalance. For a given degree of refractive imbalance, strabismic anisometropes perform considerably worse than anisometropes without strabismus – visual acuity for strabismics was on average 2.5 times worse than for non-strabismics with similar anisometropia. For observers with equal refractive error in the two eyes there is very little change in acuity or sensitivity with increasing (bilateral) refractive error except for one extreme individual (bilaterally refractive error of -15 D). Most pure anisometropes with interocular differences less than 4 D retain some stereopsis, and the degree is correlated with the acuity of the weak eye. We conclude that even modest interocular differences in refractive error can influence visual function.
PMCID: PMC3010510  PMID: 20932989
anisometropia; amblyopia; spatial vision; contrast sensitivity; Vernier acuity; binocular vision
15.  Video-Game Play Induces Plasticity in the Visual System of Adults with Amblyopia 
PLoS Biology  2011;9(8):e1001135.
A pilot study suggests that playing video games may enhance a range of spatial vision functions in adults with amblyopia.
Abnormal visual experience during a sensitive period of development disrupts neuronal circuitry in the visual cortex and results in abnormal spatial vision or amblyopia. Here we examined whether playing video games can induce plasticity in the visual system of adults with amblyopia. Specifically 20 adults with amblyopia (age 15–61 y; visual acuity: 20/25–20/480, with no manifest ocular disease or nystagmus) were recruited and allocated into three intervention groups: action videogame group (n = 10), non-action videogame group (n = 3), and crossover control group (n = 7). Our experiments show that playing video games (both action and non-action games) for a short period of time (40–80 h, 2 h/d) using the amblyopic eye results in a substantial improvement in a wide range of fundamental visual functions, from low-level to high-level, including visual acuity (33%), positional acuity (16%), spatial attention (37%), and stereopsis (54%). Using a cross-over experimental design (first 20 h: occlusion therapy, and the next 40 h: videogame therapy), we can conclude that the improvement cannot be explained simply by eye patching alone. We quantified the limits and the time course of visual plasticity induced by video-game experience. The recovery in visual acuity that we observed is at least 5-fold faster than would be expected from occlusion therapy in childhood amblyopia. We used positional noise and modelling to reveal the neural mechanisms underlying the visual improvements in terms of decreased spatial distortion (7%) and increased processing efficiency (33%). Our study had several limitations: small sample size, lack of randomization, and differences in numbers between groups. A large-scale randomized clinical study is needed to confirm the therapeutic value of video-game treatment in clinical situations. Nonetheless, taken as a pilot study, this work suggests that video-game play may provide important principles for treating amblyopia, and perhaps other cortical dysfunctions.
Trial Registration NCT01223716
Author Summary
Early abnormal visual experience disrupts neuronal circuitry in the brain and results in reduced vision, known as amblyopia or “lazy eye,” the most frequent cause of permanent visual loss in childhood. It is generally believed that adult amblyopia is irreversible beyond the sensitive period of brain development during childhood. In this study, we examine whether playing video games, both action and non-action, has an effect on the vision of adults with amblyopia. We assessed visual acuity (visual resolution), positional acuity (the ability to localize object's relative position), spatial attention (the ability to direct visual attention to various locations in the visual field), and stereoacuity (stereo-vision / 3-D depth perception) in a small group of teenagers and adults. We found that they tended to recover vision much faster than we would have expected from the results of conventional occlusion therapy in childhood amblyopia. Additional experiments and modelling suggest that the improvements are a result of decreasing spatial distortion and increasing information processing efficiency in the amblyopic brain. Thus, video games may include essential elements for active vision training to boost visual performance. Most importantly, our findings suggest that video-game play may provide important principles for treating amblyopia, a suggestion that we are pursuing with larger scale clinical trials.
PMCID: PMC3166159  PMID: 21912514
16.  Human efficiency for classifying natural versus random text 
Vision research  2010;50(6):557-563.
Humans are remarkably efficient at processing natural text. We quantified efficiency for discriminating a sample of meaningful text from a sample of random text by disrupting the meaningful sample, and measuring how much disruption human readers can tolerate before the two samples become indistinguishable. We performed these measurements for a wide range of conditions, involving samples of different lengths and containing letters, words or Chinese characters. We then compared human performance to the best possible performance achieved by a Bayesian estimator under the conditions in which we tested our participants, and in so doing we determined their absolute efficiency. Values were mostly in the range 5–40%, in agreement with reported efficiencies for many visual tasks. Although not intended as a veridical model of human processing, we found that the Bayesian model captured some (but not all) aspects of how humans classified text in our tasks and conditions.
PMCID: PMC2832918  PMID: 20079757
17.  Rule-Based Learning Explains Visual Perceptual Learning and Its Specificity and Transfer 
The Journal of Neuroscience  2010;30(37):12323-12328.
Visual perceptual learning models, as constrained by orientation and location specificities, propose that learning either reflects changes in V1 neuronal tuning or reweighting specific V1 inputs in either the visual cortex or higher areas. Here we demonstrate that, with a training-plus-exposure procedure, in which observers are trained at one orientation and either simultaneously or subsequently passively exposed to a second transfer orientation, perceptual learning can completely transfer to the second orientation in tasks known to be orientation-specific. However, transfer fails if exposure precedes the training. These results challenge the existing specific perceptual learning models by suggesting a more general perceptual learning process. We propose a rule-based learning model to explain perceptual learning and its specificity and transfer. In this model, a decision unit in high-level brain areas learns the rules of reweighting the V1 inputs through training. However, these rules cannot be applied to a new orientation/location because the decision unit cannot functionally connect to the new V1 inputs that are unattended or even suppressed after training at a different orientation/location, which leads to specificity. Repeated orientation exposure or location training reactivates these inputs to establish the functional connections and enable the transfer of learning.
PMCID: PMC3842491  PMID: 20844128
18.  Complete Transfer of Perceptual Learning across Retinal Locations Enabled by Double Training 
Current biology : CB  2008;18(24):1922-1926.
Practice improves discrimination of many basic visual features, such as contrast, orientation, positional offset, etc. [1–7]. Perceptual learning of many of these tasks is found to be retinal location specific, in that learning transfers little to an untrained retinal location [1, 6–8]. In most perceptual learning models, this location specificity is interpreted as a pointer to a retinotopic early visual cortical locus of learning [1, 6–11]. Alternatively, an untested hypothesis is that learning could occur in a central site, but it consists of two separate aspects: learning to discriminate a specific stimulus feature (“feature learning”), and learning to deal with stimulus non-specific factors like local noise at the stimulus location (“location learning”) [12]. Therefore, learning is not transferable to a new location that has never been location-trained. To test this hypothesis we developed a novel double-training paradigm that employed conventional feature training (e.g., contrast) at one location, and additional training with an irrelevant feature/task (e.g. orientation) at a second location, either simultaneously or at a different time. Our results showed that this additional location training enabled a complete transfer of feature learning (e.g., contrast) to the second location. This finding challenges location specificity and its inferred cortical retinotopy as central concepts to many perceptual learning models, and suggests perceptual learning involves higher non-retinotopic brain areas that enable location transfer.
PMCID: PMC3045109  PMID: 19062277
19.  Crowding in peripheral vision: why bigger is better 
Current biology : CB  2009;19(23):1988-1993.
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.
PMCID: PMC3045113  PMID: 19853450
20.  Aging and Visual Counting 
PLoS ONE  2010;5(10):e13434.
Much previous work on how normal aging affects visual enumeration has been focused on the response time required to enumerate, with unlimited stimulus duration. There is a fundamental question, not yet addressed, of how many visual items the aging visual system can enumerate in a “single glance”, without the confounding influence of eye movements.
Methodology/Principal Findings
We recruited 104 observers with normal vision across the age span (age 21–85). They were briefly (200 ms) presented with a number of well- separated black dots against a gray background on a monitor screen, and were asked to judge the number of dots. By limiting the stimulus presentation time, we can determine the maximum number of visual items an observer can correctly enumerate at a criterion level of performance (counting threshold, defined as the number of visual items at which ≈63% correct rate on a psychometric curve), without confounding by eye movements. Our findings reveal a 30% decrease in the mean counting threshold of the oldest group (age 61–85: ∼5 dots) when compared with the youngest groups (age 21–40: 7 dots). Surprisingly, despite decreased counting threshold, on average counting accuracy function (defined as the mean number of dots reported for each number tested) is largely unaffected by age, reflecting that the threshold loss can be primarily attributed to increased random errors. We further expanded this interesting finding to show that both young and old adults tend to over-count small numbers, but older observers over-count more.
Here we show that age reduces the ability to correctly enumerate in a glance, but the accuracy (veridicality), on average, remains unchanged with advancing age. Control experiments indicate that the degraded performance cannot be explained by optical, retinal or other perceptual factors, but is cortical in origin.
PMCID: PMC2956663  PMID: 20976149
21.  Stochastic model for detection of signals in noise 
Fifty years ago Birdsall, Tanner, and colleagues made rapid progress in developing signal detection theory into a powerful psychophysical tool. One of their major insights was the utility of adding external noise to the signals of interest. These methods have been enhanced in recent years by the addition of multipass and classification-image methods for opening up the black box. There remain a number of as yet unresolved issues. In particular, Birdsall developed a theorem that large amounts of external input noise can linearize nonlinear systems, and Tanner conjectured, with mathematical backup, that what had been previously thought of as a nonlinear system could actually be a linear system with uncertainty. Recent findings, both experimental and theoretical, have validated Birdsall’s theorem and Tanner’s conjecture. However, there have also been experimental and theoretical findings with the opposite outcome. In this paper we present new data and simulations in an attempt to sort out these issues. Our simulations and experiments plus data from others show that Birdsall’s theorem is quite robust. We argue that uncertainty can serve as an explanation for violations of Birdsall’s linearization by noise and also for reports of stochastic resonance. In addition, we modify present models to better handle detection of signals with both noise and pedestal backgrounds.
PMCID: PMC2942087  PMID: 19884912
22.  Improving the performance of the amblyopic visual system 
Experience-dependent plasticity is closely linked with the development of sensory function; however, there is also growing evidence for plasticity in the adult visual system. This review re-examines the notion of a sensitive period for the treatment of amblyopia in the light of recent experimental and clinical evidence for neural plasticity. One recently proposed method for improving the effectiveness and efficiency of treatment that has received considerable attention is ‘perceptual learning’. Specifically, both children and adults with amblyopia can improve their perceptual performance through extensive practice on a challenging visual task. The results suggest that perceptual learning may be effective in improving a range of visual performance and, importantly, the improvements may transfer to visual acuity. Recent studies have sought to explore the limits and time course of perceptual learning as an adjunct to occlusion and to investigate the neural mechanisms underlying the visual improvement. These findings, along with the results of new clinical trials, suggest that it might be time to reconsider our notions about neural plasticity in amblyopia.
PMCID: PMC2674474  PMID: 19008199
amblyopia; perceptual learning; sensitive periods; plasticity
23.  Learning to Identify Near-Threshold Luminance-Defined and Contrast-Defined Letters in Observers with Amblyopia 
Vision research  2008;48(27):2739-2750.
We assessed whether or not the sensitivity for identifying luminance-defined and contrast-defined letters improved with training in a group of amblyopic observers who have passed the critical period of development. In Experiment 1, we tracked the contrast threshold for identifying luminance-defined letters with training in a group of 11 amblyopic observers. Following training, six observers showed a reduction in thresholds, averaging 20%, for identifying luminance-defined letters. This improvement transferred extremely well to the untrained task of identifying contrast-defined letters (average improvement = 38%) but did not transfer to an acuity measurement. Seven of the 11 observers were subsequently trained on identifying contrast-defined letters in Experiment 2. Following training, five of these seven observers demonstrated a further improvement, averaging 17%, for identifying contrast-defined letters. This improvement did not transfer to the untrained task of identifying luminance-defined letters. Our findings are consistent with predictions based on the locus of learning for first- and second-order stimuli according to the filter-rectifier-filter model for second-order visual processing.
PMCID: PMC2642955  PMID: 18824189
Amblyopia; Perceptual learning; Training; First-order; Second-order; Letter recognition
24.  Prolonged Perceptual Learning of Positional Acuity in Adult Amblyopia 
Amblyopia is a developmental abnormality that results in physiological alterations in the visual cortex and impairs form vision. It is often successfully treated by patching the sound eye in infants and young children, but is generally considered to be untreatable in adults. However, a number of recent studies suggest that repetitive practice of a visual task using the amblyopic eye results in improved performance in both children and adults with amblyopia. These perceptual learning studies have used relatively brief periods of practice; however, clinical studies have shown that the time-constant for successful patching is long. The time-constant for perceptual learning in amblyopia is still unknown. Here we show that the time-constant for perceptual learning depends on the degree of amblyopia. Severe amblyopia requires more than 50 hours (≈35,000 trials) to reach plateau, yielding as much as a five-fold improvement in performance at a rate of ≈1.5% per hour. There is significant transfer of learning from the amblyopic to the dominant eye, suggesting that the learning reflects alterations in higher decision stages of processing. Using a reverse correlation technique, we document, for the first time, a dynamic retuning of the amblyopic perceptual decision template and a substantial reduction in internal spatial distortion. These results show that the mature amblyopic brain is surprisingly malleable, and point to more intensive treatment methods for amblyopia.
PMCID: PMC2765479  PMID: 19109504
plasticity; critical period; visual learning; positional acuity; classification image; amblyopia
25.  Using visual noise to characterize amblyopic letter identification 
Journal of vision  2004;4(10):904-920.
Amblyopia is a much-studied but poorly understood developmental visual disorder that reduces acuity, profoundly reducing contrast sensitivity for small targets. Here we use visual noise to probe the letter identification process and characterize its impairment by amblyopia. We apply five levels of analysis — threshold, threshold in noise, equivalent noise, optical MTF, and noise modeling — to obtain a two-factor model of the amblyopic deficit: substantially reduced efficiency for small letters and negligibly increased cortical noise. Cortical noise, expressed as an equivalent input noise, varies among amblyopes but is roughly 1.4× normal, as though only 1/1.4 the normal number of cortical spikes are devoted to the amblyopic eye. This raises threshold contrast for large letters by a factor of √1.4 = 1.2×, a negligible effect. All 16 amblyopic observers showed near-normal efficiency for large letters (> 4× acuity) and greatly reduced efficiency for small letters: 1/4 normal at 2× acuity and approaching 1/16 normal at acuity. Finding that the acuity loss represents a loss of efficiency rules out all models of amblyopia except those that predict the same sensitivity loss on blank and noisy backgrounds. One such model is the last-channel hypothesis, which supposes that the highest-spatial-frequency channels are missing, leaving the remaining highest-frequency channel struggling to identify the smallest letters. However, this hypothesis is rejected by critical band masking of letter identification, which shows that the channels used by the amblyopic eye have normal tuning for even the smallest letters. Finally, based on these results, we introduce a new “Dual Acuity” chart that promises to be a quick diagnostic test for amblyopia.
PMCID: PMC2751822  PMID: 15595894
amblyopia; noise; efficiency; cortical noise; Pelli-Levi Dual Acuity Chart

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