Playing three-dimensional video games boosts stereo vision.

Playing two-dimensional video games has been shown to result in improvements in a range of visual and cognitive tasks, and these improvements appear to generalize widely1,2,3,4,5,6. Here we report that young adults with healthy vision, surprisingly, showed a dramatic improvement in stereo vision after playing three-dimensional, but not two-dimensional, video games for a relatively short period of time. Intriguingly, neither group showed any significant improvement in binocular contrast sensitivity. This dissociation suggests that the visual enhancement was specific to genuine stereoscopic processing, not indirectly resulting from enhanced contrast processing, and required engaging in a disparity cue-rich three-dimensional environment.

Brief Adaptation to Astigmatism Reduces Meridional Anisotropy in Contrast Sensitivity.

Purpose: To investigate the effect of visual adaptation to orientation-dependent optical blur on meridional contrast sensitivity function in artificially imposed astigmatism. Methods: The study adopted a top-up adapt-test paradigm. During the blur adaptation process, the 18 non-astigmatic young adult participants were briefly presented with natural scene images (first trial, 10 minutes; subsequent trials, 6 seconds). Contrast sensitivities for horizontal and vertical gratings at spatial frequencies ranging from 1 to 8 cycles per degree (cpd) were measured immediately before and after adaptation to +3.00 diopters cylinder (DC) with-the-rule or against-the-rule astigmatism. Meridional anisotropy was measured to quantify the contrast sensitivity difference between the two grating orientations. Results: Adapting to astigmatic blur enhanced contrast sensitivity at the blurred power meridian but reduced contrast sensitivity at the least affected axis meridian. In with-the-rule conditions, contrast sensitivity for horizontal gratings was significantly increased, whereas that for vertical gratings was significantly decreased. Similarly, in against-the-rule conditions, contrast sensitivity for vertical gratings was significantly increased, whereas that for horizontal gratings was significantly decreased. These two factors together resulted in a substantial systematic reduction, averaging 34%, in meridional anisotropy of contrast sensitivity across the spatial frequency spectrum. Conclusions: Astigmatism adaptation occurs in natural scene viewing. Brief exposure to astigmatic blur altered contrast sensitivity in the opposite direction at the two principal meridians, indicating that the mature visual system possesses functional plasticity to recalibrate the response characteristics of orientationally tuned cortical filters and thus promote substantial reductions of meridional anisotropy in astigmatic vision, to some extent counterbalancing the elongated oval shape of astigmatic blur.

Meridional Anisotropy of Foveal and Peripheral Resolution Acuity in Adults With Emmetropia, Myopia, and Astigmatism.

Purpose: To quantify astigmatism-related meridional anisotropy in visual resolution at central, nasal, and inferior visual fields. Methods: Three groups of young adults (range, 18-30 years) with corrected-to-normal visual acuity (logMAR 0) were recruited: (1) myopic astigmats (MA): spherical-equivalent error (SE) < -0.75D, with-the-rule astigmatism ≥ 2.00D, n = 19; (2) simple myopes (SM): SE < -0.75D, astigmatism ≤ 0.50D, n = 20; and (3) emmetropes (EM): SE ± 0.50D, astigmatism ≤ 0.50D, n = 14. Resolution acuity was measured for the horizontal and vertical gratings at central and peripheral visual fields (eccentricity: 15°) using a 3-down 1-up staircase paradigm. On- and off-axis refractive errors were corrected by ophthalmic lenses. Results: The MA group exhibited meridional anisotropy preferring vertical gratings. At the central field, the MA group had better resolution acuity for vertical than horizontal gratings, and their resolution acuity for horizontal gratings was significantly worse than the SM and EM groups. At peripheral visual fields, both the SM and EM groups showed better resolution acuity for the radial (i.e., nasal field: horizontal gratings; inferior field: vertical gratings) than tangential orientation. However, the MA group tended to have better resolution acuity for the tangential orientation (i.e., vertical gratings), and their resolution acuity for horizontal gratings was significantly lower than the SM and EM groups at the nasal field. No significant differences were found in the inferior field among the three groups. Conclusions: This study provided evidence of astigmatism-related meridional anisotropy at the fovea and nasal visual fields, underscoring the significant impact of astigmatism on orientation-dependent visual functions.

Sequential perceptual learning of letter identification and "uncrowding" in normal peripheral vision: Effects of task, training order, and cholinergic enhancement.

Human adults with normal vision are capable of improving performance on visual tasks through repeated practice. Previous work has shown that enhancing synaptic levels of acetylcholine (ACh) in healthy human adults with donepezil (trade name: Aricept) can increase the magnitude and specificity of perceptual learning (PL) for motion direction discrimination in the perifovea. In the current study, we ask whether increasing the synaptic levels of ACh in healthy human adults with donepezil boosts learning of low-contrast isolated letter identification and high-contrast flanked letter identification in normal peripheral vision. Two groups of observers performed sequential training over multiple days while ingesting donepezil. One group trained on isolated low-contrast letters in Phase 1 and crowded high-contrast letters in Phase 2, and the other group performed the reverse sequence, thereby enabling us to differentiate possible effects of drug and training order on PL of letter identification. All testing and training were performed monocularly in peripheral vision, at an eccentricity of 10 degrees along the lower vertical meridian. Our experimental design allowed us to evaluate the effects of sequential training and to ask whether increasing cholinergic signaling boosted learning and/or transfer of low-contrast isolated letter identification and high-contrast flanked letter identification in normal peripheral vision. We found that both groups improved on each of the two tasks. However, our results revealed an effect of training task order on flanked letter identification: Observers who trained on isolated targets first showed rapid early improvement in flanked letter identification but little to no additional improvement after 30 training blocks, while observers who first trained with flanked letters improved gradually on flanked letter identification over the entire 100-block course of training. In addition, we found no effect of donepezil on PL of either isolated or flanked letter identification. In other words, donepezil neither boosted nor blocked learning to identify isolated low-contrast letters or learning to uncrowd in normal peripheral vision.

Donepezil Does Not Enhance Perceptual Learning in Adults with Amblyopia: A Pilot Study.

Amblyopia is a developmental disorder that results in a wide range of visual deficits. One proven approach to recovering vision in adults with amblyopia is perceptual learning (PL). Recent evidence suggests that neuromodulators can enhance adult plasticity. In this pilot study, we asked whether donepezil, a cholinesterase inhibitor, enhances visual PL in adults with amblyopia. Nine adults with amblyopia were first trained on a low-contrast single-letter identification task while taking a daily dose (5 mg) of donepezil throughout training. Following 10,000 trials of training, participants showed improved contrast sensitivity for identifying single letters. However, the magnitude of improvement was no greater than, and the rate of improvement was slower than, that obtained in a previous study in which six adults with amblyopia were trained using an identical task and protocol but without donepezil (Chung et al., 2012). In addition, we measured transfer of learning effects to other tasks and found that for donepezil, the post-pre performance ratios in both a size-limited (acuity) and a spacing-limited (crowding) task were not significantly different from those found in the previous study without donepezil administration. After an interval of several weeks, six participants returned for a second course of training on identifying flanked (crowded) letters, again with concurrent donepezil administration. Although this task has previously been shown to be highly amenable to PL in adults with amblyopia (Chung et al., 2012; Hussain et al., 2012), only one observer in our study showed significant learning over 10,000 trials of training. Auxiliary experiments showed that the lack of a learning effect on this task during donepezil administration was not due to either the order of training of the two tasks or the use of a sequential training paradigm. Our results reveal that cholinergic enhancement with donepezil during training does not improve or speed up PL of single-letter identification in adults with amblyopia, and importantly, it may even halt learning and transfer related to a crowding task. Clinical Trial Registration: This study was registered with ClinicalTrials.gov (NCT03109314).

Monocular blur alters the tuning characteristics of stereopsis for spatial frequency and size.

Our sense of depth perception is mediated by spatial filters at different scales in the visual brain; low spatial frequency channels provide the basis for coarse stereopsis, whereas high spatial frequency channels provide for fine stereopsis. It is well established that monocular blurring of vision results in decreased stereoacuity. However, previous studies have used tests that are broadband in their spatial frequency content. It is not yet entirely clear how the processing of stereopsis in different spatial frequency channels is altered in response to binocular input imbalance. Here, we applied a new stereoacuity test based on narrow-band Gabor stimuli. By manipulating the carrier spatial frequency, we were able to reveal the spatial frequency tuning of stereopsis, spanning from coarse to fine, under blurred conditions. Our findings show that increasing monocular blur elevates stereoacuity thresholds 'selectively' at high spatial frequencies, gradually shifting the optimum frequency to lower spatial frequencies. Surprisingly, stereopsis for low frequency targets was only mildly affected even with an acuity difference of eight lines on a standard letter chart. Furthermore, we examined the effect of monocular blur on the size tuning function of stereopsis. The clinical implications of these findings are discussed.

Sharpening coarse-to-fine stereo vision by perceptual learning: asymmetric transfer across the spatial frequency spectrum.

Neurons in the early visual cortex are finely tuned to different low-level visual features, forming a multi-channel system analysing the visual image formed on the retina in a parallel manner. However, little is known about the potential 'cross-talk' among these channels. Here, we systematically investigated whether stereoacuity, over a large range of target spatial frequencies, can be enhanced by perceptual learning. Using narrow-band visual stimuli, we found that practice with coarse (low spatial frequency) targets substantially improves performance, and that the improvement spreads from coarse to fine (high spatial frequency) three-dimensional perception, generalizing broadly across untrained spatial frequencies and orientations. Notably, we observed an asymmetric transfer of learning across the spatial frequency spectrum. The bandwidth of transfer was broader when training was at a high spatial frequency than at a low spatial frequency. Stereoacuity training is most beneficial when trained with fine targets. This broad transfer of stereoacuity learning contrasts with the highly specific learning reported for other basic visual functions. We also revealed strategies to boost learning outcomes 'beyond-the-plateau'. Our investigations contribute to understanding the functional properties of the network subserving stereovision. The ability to generalize may provide a key principle for restoring impaired binocular vision in clinical situations.

Characteristics of fixational eye movements in amblyopia: Limitations on fixation stability and acuity?

Persons with amblyopia, especially those with strabismus, are known to exhibit abnormal fixational eye movements. In this paper, we compared six characteristics of fixational eye movements among normal control eyes (n=16), the non-amblyopic fellow eyes and the amblyopic eyes of anisometropic (n=14) and strabismic amblyopes (n=14). These characteristics include the frequency, magnitude of landing errors, amplitude and speed of microsaccades, and the amplitude and speed of slow drifts. Fixational eye movements were recorded using retinal imaging while observers monocularly fixated a 1° cross. Eye position data were recovered using a cross-correlation procedure. We found that in general, the characteristics of fixational eye movements are not significantly different between the fellow eyes of amblyopes and controls, and that the strabismic amblyopic eyes are always different from the other groups. Next, we determined the primary factors that limit fixation stability and visual acuity in amblyopic eyes by examining the relative importance of the different oculomotor characteristics, adding acuity (for fixation stability) or fixation stability (for acuity), and the type of amblyopia, as predictive factors in a multiple linear regression model. We show for the first time that the error magnitude of microsaccades, acuity, amplitude and frequency of microsaccades are primary factors limiting fixation stability; while the error magnitude, fixation stability, amplitude of drifts and amplitude of microsaccades are the primary factors limiting acuity. A mediation analysis showed that the effects of error magnitude and amplitude of microsaccades on acuity could be explained, at least in part, by their effects on fixation stability.

Relieving the attentional blink in the amblyopic brain with video games.

Video game play induces a generalized recovery of a range of spatial visual functions in the amblyopic brain. Here we ask whether video game play also alters temporal processing in the amblyopic brain. When visual targets are presented in rapid succession, correct identification of the first target (T1) can interfere with identification of the second (T2). This is known as the "attentional blink". We measured the attentional blink in each eye of adults with amblyopia before and after 40 hours of active video game play, using a rapid serial visual presentation technique. After videogame play, we observed a ~40% reduction in the attentional blink (identifying T2 200 ms after T1) seen through the amblyopic eye and this improvement in performance transferred substantially to the untrained fellow sound eye. Our experiments show that the enhanced performance cannot be simply explained by eye patching alone, or to improved visual acuity, but is specific to videogame experience. Thus, videogame training might have important therapeutic applications for amblyopia and other visual brain disorders.

A Weber-like law for perceptual learning.

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.

Learning to identify near-acuity letters, either with or without flankers, results in improved letter size and spacing limits in adults with amblyopia.

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.

Reduced sampling efficiency causes degraded Vernier hyperacuity with normal aging: Vernier acuity in position noise.

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.

Video-game play induces plasticity in the visual system of adults with amblyopia.

UNLABELLED: 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: ClinicalTrials.gov NCT01223716.

Removing brakes on adult brain plasticity: from molecular to behavioral interventions.

Adult brain plasticity, although possible, remains more restricted in scope than during development. Here, we address conditions under which circuit rewiring may be facilitated in the mature brain. At a cellular and molecular level, adult plasticity is actively limited. Some of these "brakes" are structural, such as perineuronal nets or myelin, which inhibit neurite outgrowth. Others are functional, acting directly upon excitatory-inhibitory balance within local circuits. Plasticity in adulthood can be induced either by lifting these brakes through invasive interventions or by exploiting endogenous permissive factors, such as neuromodulators. Using the amblyopic visual system as a model, we discuss genetic, pharmacological, and environmental removal of brakes to enable recovery of vision in adult rodents. Although these mechanisms remain largely uncharted in the human, we consider how they may provide a biological foundation for the remarkable increase in plasticity after action video game play by amblyopic subjects.

Aging and visual counting.

BACKGROUND: 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. CONCLUSION/SIGNIFICANCE: 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.

Perceptual learning as a potential treatment for amblyopia: a mini-review.

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: 1). What factors influence the outcome of perceptual learning? 2). Specificity and generalization - two sides of the coin. 3). Do the improvements last? 4). How does PL improve visual function? 5). 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.

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.

Prolonged perceptual learning of positional acuity in adult amblyopia: perceptual template retuning dynamics.

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 >50 h (approximately equal to 35,000 trials) to reach plateau, yielding as much as a five-fold improvement in performance at a rate of approximately equal to 1.5%/h. 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.

Learning to identify near-threshold luminance-defined and contrast-defined letters in observers with amblyopia.

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.