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1.  Learning letter identification in peripheral vision 
Vision research  2005;45(11):1399-1412.
Performance for a variety of visual tasks improves with practice. The purpose of this study was to determine the nature of the processes underlying perceptual learning of identifying letters in peripheral vision. To do so, we tracked changes in contrast thresholds for identifying single letters presented at 10° in the inferior visual field, over a period of six consecutive days. The letters (26 lowercase Times-Roman letters, subtending 1.7°) were embedded within static two-dimensional Gaussian luminance noise, with rms contrast ranging from 0% (no noise) to 20%. We also measured the observers’ response consistency using a double-pass method on days 1, 3 and 6, by testing two additional blocks on each of these days at luminance noise of 3% and 20%. These additional blocks were the exact replicates of the corresponding block at the same noise contrast that was tested on the same day. We analyzed our results using both the linear amplifier model (LAM) and the perceptual template model (PTM). Our results showed that following six days of training, the overall reduction (improvement across all noise levels) in contrast threshold for our seven observers averaged 21.6% (range: 17.2–31%). Despite fundamental differences between LAM and PTM, both models show that learning leads to an improvement of the perceptual template (filter) such that the template is more capable of extracting the crucial information from the signal. Results from both the PTM analysis and the double-pass experiment imply that the stimulus-dependent component of the internal noise does not change with learning.
PMCID: PMC2741315  PMID: 15743610
Perceptual learning; Training; Letter identification; Peripheral vision
2.  Identification of contrast-defined letters benefits from perceptual learning in adults with amblyopia 
Vision research  2006;46(22):3853-3861.
Amblyopes show specific deficits in processing second-order spatial information (e.g. Wong, Levi, & McGraw (2001). Is second-order spatial loss in amblyopia explained by the loss of first-order spatial input? Vision Research, 41, 2951–2960). Recent work suggests there is a significant degree of plasticity in the visual pathway that processes first-order spatial information in adults with amblyopia. In this study, we asked whether or not there is similar plasticity in the ability to process second-order spatial information in adults with amblyopia. Ten adult observers with amblyopia (five strabismic, four anisometropic and one mixed) were trained to identify contrast-defined (second-order) letters using their amblyopic eyes. Before and after training, we determined observers’ contrast thresholds for identifying luminance-defined (first-order) and contrast-defined letters, separately for the non-amblyopic and amblyopic eyes. Following training, eight of the 10 observers showed a significant reduction in contrast thresholds for identifying contrast-defined letters with the amblyopic eye. Five of these observers also showed a partial transfer of improvement to their fellow untrained non-amblyopic eye for identifying contrast-defined letters. There was a small but statistically significant transfer to the untrained task of identifying luminance-defined letters in the same trained eye. Similar to first-order spatial tasks, adults with amblyopia benefit from perceptual learning for identifying contrast-defined letters in their amblyopic eyes, suggesting a sizeable degree of plasticity in the visual pathway for processing second-order spatial information.
PMCID: PMC1852540  PMID: 16930666
Amblyopia; Letter recognition; Perceptual learning; Second-order
3.  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
4.  Letter-recognition and reading speed in peripheral vision benefit from perceptual learning 
Vision research  2004;44(7):695-709.
Visual-span profiles are plots of letter-recognition accuracy as a function of letter position left or right of the midline. Previously, we have shown that contraction of these profiles in peripheral vision can account for slow reading speed in peripheral vision. In this study, we asked two questions: (1) can we modify visual-span profiles through training on letter-recognition, and if so, (2) are these changes accompanied by changes in reading speed? Eighteen normally sighted observers were randomly assigned to one of three groups: training at 10° in the upper visual field, training at 10° in the lower visual field and a no-training control group. We compared observers’ characteristics of reading (maximum reading speed and critical print size) and visual-span profiles (peak amplitude and bits of information transmitted) before and after training, and at trained and untrained retinal locations (10° upper and lower visual fields). Reading speeds were measured for six print sizes at each retinal location, using the rapid serial visual presentation paradigm. Visual-span profiles were measured using a trigram letter-recognition task, for a letter size equivalent to 1.4 × the critical print size for reading. Training consisted of the repeated measurement of 20 visual-span profiles (over four consecutive days) in either the upper or lower visual field. We also tracked the changes in performance in a sub-group of observers for up to three months following training. We found that the visual-span profiles can be expanded (bits of information transmitted increased by 6 bits) through training with a letter-recognition task, and that there is an accompanying increase (41%) in the maximum reading speed. These improvements transferred, to a large extent, from the trained to an untrained retinal location, and were retained, to a large extent, for at least three months following training. Our results are consistent with the view that the visual span is a bottleneck on reading speed, but a bottleneck that can be increased with practice.
PMCID: PMC2729075  PMID: 14751554
Reading; Letter-recognition; Peripheral vision; Perceptual learning; Low vision; Visual rehabilitation
5.  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.
PMCID: PMC2134936  PMID: 17928026
crowding; perceptual learning; training; reading
6.  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
7.  Context and Crowding in Perceptual Learning on a Peripheral Contrast Discrimination Task: Context-Specificity in Contrast Learning 
PLoS ONE  2013;8(5):e63278.
Perceptual learning is an improvement in sensitivity due to practice on a sensory task and is generally specific to the trained stimuli and/or tasks. The present study investigated the effect of stimulus configuration and crowding on perceptual learning in contrast discrimination in peripheral vision, and the effect of perceptual training on crowding in this task. 29 normally-sighted observers were trained to discriminate Gabor stimuli presented at 9° eccentricity with either identical or orthogonally oriented flankers with respect to the target (ISO and CROSS, respectively), or on an isolated target (CONTROL). Contrast discrimination thresholds were measured at various eccentricities and target-flanker separations before and after training in order to determine any learning transfer to untrained stimulus parameters. Perceptual learning was observed in all three training stimuli; however, greater improvement was obtained with training on ISO-oriented stimuli compared to CROSS-oriented and unflanked stimuli. This learning did not transfer to untrained stimulus configurations, eccentricities or target-flanker separations. A characteristic crowding effect was observed increasing with viewing eccentricity and decreasing with target-flanker separation before and after training in both configurations. The magnitude of crowding was reduced only at the trained eccentricity and target-flanker separation; therefore, learning for contrast discrimination and for crowding in the present study was configuration and location specific. Our findings suggest that stimulus configuration plays an important role in the magnitude of perceptual learning in contrast discrimination and suggest context-specificity in learning.
PMCID: PMC3655984  PMID: 23696807
8.  Noise masking reveals channels for second-order letters 
Vision research  2005;46(8-9):1493-1506.
We investigate the channels underlying identification of second-order letters using a critical-band masking paradigm. We find that observers use a single 1–1.5 octave-wide channel for this task. This channel’s best spatial frequency (c/letter) did not change across different noise conditions (indicating the inability of observers to switch channels to improve signal-to-noise ratio) or across different letter sizes (indicating scale invariance), for a fixed carrier frequency (c/letter). However, the channel’s best spatial frequency does change with stimulus carrier frequency (both in c/letter); one is proportional to the other. Following Majaj et al. (Majaj, N. J., Pelli, D. G., Kurshan, P., & Palomares, M. (2002). The role of spatial frequency channels in letter identification. Vision Research, 42, 1165–1184), we define “stroke frequency” as the line frequency (strokes/deg) in the luminance image. That is, for luminance-defined letters, stroke frequency is the number of lines (strokes) across each letter divided by letter width. For second-order letters, letter texture stroke frequency is the number of carrier cycles (luminance lines) within the letter ink area divided by the letter width. Unlike the nonlinear dependence found for first-order letters (implying scale-dependent processing), for second-order letters the channel frequency is half the letter texture stroke frequency (suggesting scale-invariant processing).
PMCID: PMC2760253  PMID: 16203023
Letter identification; Second-order vision; Critical-band masking; Scale invariance; Channel switching
9.  Overt use of a tactile-kinesthetic strategy shifts to covert processing in rehabilitation of letter-by-letter reading 
Aphasiology  2010;24(11):1424-1442.
Letter-by-letter readers identify each letter of the word they are reading serially in left to right order before recognizing the word. When their letter naming is also impaired, letter-by-letter reading is inaccurate and can render even single word reading very poor. Tactile and/or kinesthetic strategies have been reported to improve reading in these patients, but only under certain conditions or for a limited set of stimuli.
The primary aim of the current study was to determine whether a tactile/kinesthetic treatment could significantly improve reading specifically under normal reading conditions, i.e. reading untrained words presented in free vision and read without overt use of the strategy.
Methods & Procedures
Three chronic letter-by-letter readers participated in a tactile/kinesthetic treatment aimed at first improving letter naming accuracy (phase 1) and then letter-by-letter reading speed (phase 2). In a multiple case series design, accuracy and speed of reading untrained words without overt use of the trained tactile/kinesthetic strategy was assessed before phase 1, after phase 1 and again after phase 2.
Outcomes & Results
All three patients significantly improved both their speed and accuracy reading untrained words without overt use of the trained tactile/kinesthetic strategy. All three patients required the additional practice in phase 2 to achieve significant improvement. Treatment did not target sentence level reading, yet two of the three patients became so adept that they could read entire sentences.
This study replicates previous findings on the efficacy of tactile/kinesthetic treatment for letter-by-letter readers with poor letter naming. It further demonstrates that this treatment can alter cognitive processing such that words never specifically trained can be read in free vision without overtly using the trained strategy. The data suggest that an important element in achieving this level of generalization is continuing training beyond the point of initial mastery (i.e. accurate letter naming).
PMCID: PMC3002229  PMID: 21170161
aphasia; cognitive rehabilitation; Speech-Language Pathology; pure alexia; letter-by-letter reading; generalization
10.  Effect of luminance noise on the object frequencies mediating letter identification 
Purpose: To determine if the same object frequency information mediates letter contrast threshold in the presence and absence of additive luminance noise (i.e., “noise-invariant processing”) for letters of different size.
Methods: Contrast thresholds for Sloan letters ranging in size from 0.9 to 1.8 log MAR were obtained from three visually normal observers under three paradigms: (1) high- and low-pass Gaussian filtered letters were presented against a uniform adapting field; (2) high- and low-pass Gaussian filtered letters were presented in additive white luminance noise; and (3) unfiltered letters were presented in high- and low-pass Gaussian filtered luminance noise. A range of high- and low-pass filter cutoffs were used to limit selectively the object frequency content of the letters (paradigms 1 and 2) or noise (paradigm 3). The object frequencies mediating letter identification under each paradigm were derived from plots of log contrast threshold vs. log filter cutoff frequency.
Results: The object frequency band mediating letter identification systematically shifted to higher frequencies with increasing log MAR letter size under all three paradigms. However, the relationship between object frequency and letter size depended on the paradigm under which the measurements were obtained. The largest difference in object frequency among the paradigms was observed at 1.8 log MAR, where the addition of white noise nearly doubled the center frequency of the band of object frequencies mediating letter identification, compared to measurements made in the absence of noise.
Conclusion: Noise can affect the object frequency band mediating letter contrast threshold, particularly for large letters, an effect that is likely due to strong masking of the low frequency letter components by low frequency noise checks. This finding indicates that noise-invariant processing cannot necessarily be assumed for large letters presented in white noise.
PMCID: PMC4080385  PMID: 25071637
visual noise; letter identification; contrast sensitivity; optotype; object spatial frequency; retinal spatial frequency
11.  Crowding between first- and second-order letter stimuli in normal foveal and peripheral vision 
Journal of vision  2007;7(2):10.1-1013.
Evidence that the detection of first- and second-order visual stimuli is processed by separate pathways abounds. This study asked whether first- and second-order stimuli remain independent at the stage of processing where crowding occurs. We measured thresholds for identifying a first-order (luminance defined) or second-order (contrast defined) target letter in the presence of two second- or first-order flanking letters. For comparison, we also measured thresholds when the target and flanking letters were all first or second order. Contrast of the flankers was 1.6 times their respective contrast thresholds. Measurements were obtained at the fovea and 10° in the lower visual field of four normally sighted observers. Two observers were also tested at 10° nasal visual field. As expected, in both the fovea and periphery, the magnitude of crowding (threshold elevation) was maximal at the closest letter separation and decreased as letter separation increased. The magnitude of crowding was greater for second- than for first-order target letters, independent of the order type of flankers; however, the critical distance for crowding was similar for first- and second-order letters. Substantial crossover crowding occurred when the target and flanking letters were of different order type. Our finding of substantial interaction between first- and second-order stimuli suggests that the processing of these stimuli is not independent at the stage of processing at which crowding occurs.
PMCID: PMC2747649  PMID: 18217825
crowding; first order; second order; peripheral vision; letter identification
12.  Detection and Identification of Crowded Mirror-Image Letters in Normal Peripheral Vision 
Vision research  2009;50(3):337.
Performance for discriminating single mirror -image letters in peripheral vision can be as good as that in central vision, provided that letter size is scaled appropriately (Higgins, Arditi & Knoblauch, 1996 Vision Research). In this study, we asked whether or not there is a reduction in performance for discriminating mirror -image letters when the letters are flanked closely by other letters, compared with unflanked (single) letters; and if so, whether or not this effect is greater in peripheral than in central vision. We compared contrast thresholds for detecting and identifying mirror -image letters “b” and “d” for a range of letter separations, at the fovea and 10° eccentricity, for letters that were scaled in size. For comparison, thresholds were also determined for a pair of non - mirror-image letters “o” and “x”. Our principal finding is that there is an additional loss in sensitivity for identifying mirror -image letters (“bd”), compared with non - mirror-image letters (“ox”), when the letters are flanked closely by other letters. The effect is greater in peripheral than central vision. An auxiliary experiment comparing thresholds for letters “d” and “q” vs. “b” and “d” shows that the additional loss in sensitivity for identifying crowded mirror -image letters cannot be attributed to the similarity in letter features between the two letters, but instead, is specific to the axis of symmetry. Our results suggest that in the presence of proximal objects, there is a specific loss in sensitivity for processing broad -band left-right mirror images in peripheral vision.
PMCID: PMC2822048  PMID: 19961868
Crowding; letter identification; mirror images
13.  Learning to identify crowded letters: Does the learning depend on the frequency of training? 
Vision research  2012;77:41-50.
Performance for many visual tasks improves with training. The magnitude of improvement following training depends on the training task, number of trials per training session and the total amount of training. Does the magnitude of improvement also depend on the frequency of training sessions? In this study, we compared the learning effect for three groups of normally sighted observers who repeatedly practiced the task of identifying crowded letters in the periphery for six sessions (1000 trials per session), according to three different training schedules — one group received one session of training everyday, the second group received a training session once a week and the third group once every two weeks. Following six sessions of training, all observers improved in their performance of identifying crowded letters in the periphery. Most importantly, the magnitudes of improvement were similar across the three training groups. The improvement was accompanied by a reduction in the spatial extent of crowding, an increase in the size of visual span and a reduction in letter-size threshold. The magnitudes of these accompanied improvements were also similar across the three training groups. Our finding that the effectiveness of visual perceptual learning is similar for daily, weekly and biweekly training has significant implication for adopting perceptual learning as an option to improve visual functions for clinical patients.
PMCID: PMC3538889  PMID: 23206551
perceptual learning; crowding; letter identification; peripheral vision
14.  Efficient integration across spatial frequencies for letter identification in foveal and peripheral vision 
Journal of vision  2008;8(13):3.1-320.
Objects in natural scenes are spatially broadband; in contrast, feature detectors in the early stages of visual processing are narrowly tuned in spatial frequency. Earlier studies of feature integration using gratings suggested that integration across spatial frequencies is suboptimal. Here we re-examined this conclusion using a letter identification task at the fovea and at 10 deg in the lower visual field. We found that integration across narrow-band (1-octave) spatial frequency components of letter stimuli is optimal in the fovea. Surprisingly, this optimality is preserved in the periphery, even though feature integration is known to be deficient in the periphery from studies of other form-vision tasks such as crowding. A model that is otherwise a white-noise ideal observer except for a limited spatial resolution defined by the human contrast sensitivity function and using internal templates slightly wider in bandwidth than the stimuli is able to account for the human data. Our findings suggest that deficiency in feature integration found in peripheral vision is not across spatial frequencies.
PMCID: PMC2635099  PMID: 19146333
spatial frequency channels; summation; letter identification; fovea; periphery
15.  The Effect of Letter-stroke Boldness on Reading Speed in Central and Peripheral Vision 
Vision research  2013;84:33-42.
People with central vision loss often prefer boldface print over normal print for reading. However, little is known about how reading speed is influenced by the letter-stroke boldness of font. In this study, we examined the reliance of reading speed on stroke boldness, and determined whether this reliance differs between the normal central and peripheral vision. Reading speed was measured using the rapid serial visual presentation paradigm, where observers with normal vision read aloud short single sentences presented on a computer monitor, one word at a time. Text was rendered in Courier at six levels of boldness, defined as the stroke-width normalized to that of the standard Courier font: 0.27, 0.72, 1, 1.48, 1.89 and 3.04× the standard. Testings were conducted at the fovea and 10° in the inferior visual field. Print sizes used were 0.8× and 1.4× the critical print size (smallest print size that can be read at the maximum reading speed). At the fovea, reading speed was invariant for the middle four levels of boldness, but dropped by 23.3% for the least and the most bold text. At 10° eccentricity, reading speed was virtually the same for all boldness <1, but showed a poorer tolerance to bolder text, dropping by 21.5% for 1.89x boldness and 51% for the most bold (3.04x) text. These results could not be accounted for by the changes in print size or the RMS contrast of text associated with changes in stroke boldness. Our results suggest that contrary to the popular belief, reading speed does not benefit from bold text in the normal fovea and periphery. Excessive increase in stroke boldness may even impair reading speed, especially in the periphery.
PMCID: PMC3642228  PMID: 23523572
Reading; stroke boldness; peripheral vision
16.  Metamorphopsia and letter recognition 
Journal of Vision  2014;14(14):1.
Acuity is the most commonly used measure of visual function, and reductions in acuity are associated with most eye diseases. Metamorphopsia—a perceived distortion of visual space—is another common symptom of visual impairment and is currently assessed qualitatively using Amsler (1953) charts. In order to quantify the impact of metamorphopsia on acuity, we measured the effect of physical spatial distortion on letter recognition. Following earlier work showing that letter recognition is tuned to specific spatial frequency (SF) channels, we hypothesized that the effect of distortion might depend on the spatial scale of visual distortion just as it depends on the spatial scale of masking noise. Six normally sighted observers completed a 26 alternate forced choice (AFC) Sloan letter identification task at five different viewing distances, and the letters underwent different levels of spatial distortion. Distortion was controlled using spatially band-pass filtered noise that spatially remapped pixel locations. Noise was varied over five spatial frequencies and five magnitudes. Performance was modeled with logistic regression and worsened linearly with increasing distortion magnitude and decreasing letter size. We found that retinal SF affects distortion at midrange frequencies and can be explained with the tuning of a basic contrast sensitivity function, while object-centered distortion SF follows a similar pattern of letter object recognition sensitivity and is tuned to approximately three cycles per letter (CPL). The interaction between letter size and distortion makes acuity an unreliable outcome for metamorphopsia assessment.
PMCID: PMC4249589  PMID: 25453116
letter recognition; visual acuity; spatial vision; metamorphopsia; clinical vision; macular degeneration
17.  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
18.  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
19.  Asymmetric Transfer of Auditory Perceptual Learning 
Perceptual skills can improve dramatically even with minimal practice. A major and practical benefit of learning, however, is in transferring the improvement on the trained task to untrained tasks or stimuli, yet the mechanisms underlying this process are still poorly understood. Reduction of internal noise has been proposed as a mechanism of perceptual learning, and while we have evidence that frequency discrimination (FD) learning is due to a reduction of internal noise, the source of that noise was not determined. In this study, we examined whether reducing the noise associated with neural phase locking to tones can explain the observed improvement in behavioral thresholds. We compared FD training between two tone durations (15 and 100 ms) that straddled the temporal integration window of auditory nerve fibers upon which computational modeling of phase locking noise was based. Training on short tones resulted in improved FD on probe tests of both the long and short tones. Training on long tones resulted in improvement only on the long tones. Simulations of FD learning, based on the computational model and on signal detection theory, were compared with the behavioral FD data. We found that improved fidelity of phase locking accurately predicted transfer of learning from short to long tones, but also predicted transfer from long to short tones. The observed lack of transfer from long to short tones suggests the involvement of a second mechanism. Training may have increased the temporal integration window which could not transfer because integration time for the short tone is limited by its duration. Current learning models assume complex relationships between neural populations that represent the trained stimuli. In contrast, we propose that training-induced enhancement of the signal-to-noise ratio offers a parsimonious explanation of learning and transfer that easily accounts for asymmetric transfer of learning.
PMCID: PMC3502074  PMID: 23181045
perceptual learning; transfer of learning; frequency discrimination; internal noise; phase locking; integration time; auditory; modeling
20.  Effect of kapalabhati on performance of six-letter cancellation and digit letter substitution task in adults 
International Journal of Yoga  2013;6(2):128-130.
Attention and concentration are valuable skills for all fields of human activity. Training to improve these skills is described in ancient hatha yoga texts.
To study the effect of 1-min Kapalabhati (KB1) and 5-min Kapalabhati (KB5) practice of the Yoga rapid breathing exercise, Kapalabhati (KB), on psychomotor performance, as measured by the six-letter cancellation task (SLCT) and digit-letter substitution task (DLST).
Materials and Methods:
Thirty-six subjects, 21 male (mean age 25.71 years, SD 2.10), 15 female (mean age 24.13 years, SD 2.23) participated in the study. All were participating in a 3-month pranayama training program, part of residential degree courses at Swami Vivekananda Yoga Anusandhana Samsthana, Yoga University. The subjects were divided into two groups, and assessed on the SLCT and DLST, immediately before and after KB on two successive days. The first group did KB1 on day 1, and KB5 on day 2. For the second group, the order was reversed.
There were no significant differences on SLCT and DLST on Total and Net Scores between sessions for the same group, and between groups for the same session i.e. the effects of KB1 and KB5 were not distinguishable. However, both groups made more errors on DLST after the interventions, 525% after KB1 and 562.5% after KB5, P < 0.018 and P < 0.041, respectively (Wilcoxon Signed Rank Test). In contrast, scores on SLCT remained completely unchanged.
Both KB1 and KB5 found no change on both SLCT and DLST. But, this kind of breathing practices leads to increases error score.
PMCID: PMC3734639  PMID: 23930032
DLST; high frequency yoga breathing; kapalabhati; pranayama; SLCT
21.  Grouping in object recognition: The role of a Gestalt law in letter identification 
Cognitive neuropsychology  2009;26(1):36-49.
The Gestalt psychologists reported a set of laws describing how vision groups elements to recognize objects. The Gestalt laws “prescribe for us what we are to recognize ‘as one thing’.” (Köhler, 1920). Were they right? Does object recognition involve grouping? Tests of the laws of grouping have been favorable, but mostly assessed only detection, not identification, of the compound object. The grouping of elements seen in the detection experiments with lattices and ‘snakes in the grass’ is compelling, but falls far short of the vivid everyday experience of recognizing a familiar, meaningful, named thing, which mediates the ordinary identification of an object. Thus, after nearly a century, there is hardly any evidence that grouping plays a role in ordinary object recognition. To assess grouping in object recognition, we made letters out of grating patches and measured threshold contrast for identifying these letters in visual noise as a function of perturbation of grating orientation, phase, and offset. We define a new measure, “wiggle,” to characterize the degree to which these various perturbations violate the Gestalt law of good continuation. We find that efficiency for letter identification is inversely proportional to wiggle, and is wholly determined by wiggle, independent of how the wiggle was produced. Thus the effects of three different kinds of shape perturbation on letter identifiability are predicted by a single measure of goodness of continuation. This shows that letter identification obeys the Gestalt law of good continuation, and may be the first confirmation of the original Gestalt claim that object recognition involves grouping.
PMCID: PMC2679997  PMID: 19424881
Gestalt; grouping; contour integration; good continuation; letter identification; object recognition; features; snake in the grass; snake letters; dot lattice
22.  Grouping in object recognition: The role of a Gestalt law in letter identification 
Cognitive Neuropsychology  2009;26(1):36-49.
The Gestalt psychologists reported a set of laws describing how vision groups elements to recognize objects. The Gestalt laws “prescribe for us what we are to recognize ‘as one thing’” (Köhler, 1920). Were they right? Does object recognition involve grouping? Tests of the laws of grouping have been favourable, but mostly assessed only detection, not identification, of the compound object. The grouping of elements seen in the detection experiments with lattices and “snakes in the grass” is compelling, but falls far short of the vivid everyday experience of recognizing a familiar, meaningful, named thing, which mediates the ordinary identification of an object. Thus, after nearly a century, there is hardly any evidence that grouping plays a role in ordinary object recognition. To assess grouping in object recognition, we made letters out of grating patches and measured threshold contrast for identifying these letters in visual noise as a function of perturbation of grating orientation, phase, and offset. We define a new measure, “wiggle”, to characterize the degree to which these various perturbations violate the Gestalt law of good continuation. We find that efficiency for letter identification is inversely proportional to wiggle and is wholly determined by wiggle, independent of how the wiggle was produced. Thus the effects of three different kinds of shape perturbation on letter identifiability are predicted by a single measure of goodness of continuation. This shows that letter identification obeys the Gestalt law of good continuation and may be the first confirmation of the original Gestalt claim that object recognition involves grouping.
PMCID: PMC2679997  PMID: 19424881
Gestalt; Grouping; Contour integration; Good continuation; Letter identification; Object recognition; Features; Snake in the grass; Snake letters; Dot lattice
23.  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
24.  Involuntary attention enhances identification accuracy for unmasked low contrast letters using non-predictive peripheral cues 
Vision research  2013;89:10.1016/j.visres.2013.06.010.
There is controversy regarding whether or not involuntary attention improves response accuracy at a cued location when the cue is non-predictive and if these cueing effects are dependent on backward masking. Various perceptual and decisional mechanisms of performance enhancement have been proposed, such as signal enhancement, noise reduction, spatial uncertainty reduction, and decisional processes. Herein we review a recent report of mask-dependent accuracy improvements with low contrast stimuli and demonstrate that the experiments contained stimulus artifacts whereby the cue impaired perception of low contrast stimuli, leading to an absence of improved response accuracy with unmasked stimuli. Our experiments corrected these artifacts by implementing an isoluminant cue and increasing its distance relative to the targets. The results demonstrate that cueing effects are robust for unmasked stimuli presented in the periphery, resolving some of the controversy concerning cueing enhancement effects from involuntary attention and mask dependency. Unmasked low contrast and/or short duration stimuli as implemented in these experiments may have a short enough iconic decay that the visual system functions similarly as if a mask were present leading to improved accuracy with a valid cue.
PMCID: PMC3813696  PMID: 23872240
25.  Letters in the forest: global precedence effect disappears for letters but not for non-letters under reading-like conditions 
Normally skilled reading involves special processing strategies for letters, which are habitually funneled into an abstract letter code. On the basis of previous studies we argue that this habit leads to the preferred usage of an analytic strategy for the processing of letters, while non-letters are preferably processed via a holistic strategy. The well-known global precedence effect (GPE) seems to contradict to this assumption, since, with compound, hierarchical figures, including letter items, faster responses are observed to the global than to the local level of the figure, as well as an asymmetric interference effect from global to local level. We argue that with letters these effects depend on presentation conditions; only when they elicit the processing strategies automatized for reading, an analytic strategy for letters in contrast to non-letters is to be expected. We compared the GPE for letters and non-letters in central viewing, with the global stimulus size close to the functional visual field in whole word reading (6.5° of visual angle) and local stimuli close to the critical size for fluent reading of individual letters (0.5° of visual angle). Under these conditions, the GPE remained robust for non-letters. For letters, however, it disappeared: letters showed no overall response time advantage for the global level and symmetric congruence effects (local-to-global as well as global-to-local interference). We interpret these results as according to the view that reading is based on resident analytic visual processing strategies for letters.
PMCID: PMC4102249  PMID: 25101012
reading acquisition; global advantage effect; analytic processing; holistic processing; literacy; developmental dyslexia; congruence effect

Results 1-25 (1151862)