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1.  The Temporal Dynamics of Implicit Processing of Non-Letter, Letter, and Word-Forms in the Human Visual Cortex 
The decoding of visually presented line segments into letters, and letters into words, is critical to fluent reading abilities. Here we investigate the temporal dynamics of visual orthographic processes, focusing specifically on right hemisphere contributions and interactions between the hemispheres involved in the implicit processing of visually presented words, consonants, false fonts, and symbolic strings. High-density EEG was recorded while participants detected infrequent, simple, perceptual targets (dot strings) embedded amongst a of character strings. Beginning at 130 ms, orthographic and non-orthographic stimuli were distinguished by a sequence of ERP effects over occipital recording sites. These early latency occipital effects were dominated by enhanced right-sided negative-polarity activation for non-orthographic stimuli that peaked at around 180 ms. This right-sided effect was followed by bilateral positive occipital activity for false-fonts, but not symbol strings. Moreover the size of components of this later positive occipital wave was inversely correlated with the right-sided ROcc180 wave, suggesting that subjects who had larger early right-sided activation for non-orthographic stimuli had less need for more extended bilateral (e.g., interhemispheric) processing of those stimuli shortly later. Additional early (130–150 ms) negative-polarity activity over left occipital cortex and longer-latency centrally distributed responses (>300 ms) were present, likely reflecting implicit activation of the previously reported ‘visual-word-form’ area and N400-related responses, respectively. Collectively, these results provide a close look at some relatively unexplored portions of the temporal flow of information processing in the brain related to the implicit processing of potentially linguistic information and provide valuable information about the interactions between hemispheres supporting visual orthographic processing.
PMCID: PMC2796900  PMID: 20046826
word reading; ERPs; visual cortex; visual orthography
2.  Sensory factors limiting horizontal and vertical visual span for letter recognition 
Journal of Vision  2014;14(9):23.
Reading speed for English text is slower for text oriented vertically than horizontally. Yu, Park, Gerold, and Legge (2010) showed that slower reading of vertical text is associated with a smaller visual span (the number of letters recognized with high accuracy without moving the eyes). Three possible sensory determinants of the size of the visual span are: resolution (decreasing acuity at letter positions farther from the midline), mislocations (uncertainty about the relative position of letters in strings), and crowding (interference from flanking letters in recognizing the target letter). In the present study, we asked which of these factors is most important in determining the size of the visual span, and likely in turn in determining the horizontal/vertical difference in reading when letter size is above the critical print size for reading. We used a decomposition analysis to represent constraints due to resolution, mislocations, and crowding as losses in information transmitted (in bits) about letter recognition. Across vertical and horizontal conditions, crowding accounted for 75% of the loss in information, mislocations accounted for 19% of the loss, and declining acuity away from fixation accounted for only 6%. We conclude that crowding is the major factor limiting the size of the visual span, and that the horizontal/vertical difference in the size of the visual span is associated with stronger crowding along the vertical midline.
PMCID: PMC4144171
visual span; crowding; reading; vertical text; acuity; mislocation
3.  Effect of letter spacing on visual span and reading speed 
Journal of vision  2007;7(2):2.1-210.
S. T. L. Chung (2002) has shown that rapid serial visual presentation (RSVP) reading speed varies with letter spacing, peaking near the standard letter spacing for text and decreasing for both smaller and larger spacings. In this study, we tested the hypothesis that the dependence of reading speed on letter spacing is mediated by the size of the visual span—the number of letters recognized with high accuracy without moving the eyes. If so, the size of the visual span and reading speed should show a similar dependence on letter spacing. We tested this prediction for RSVP reading and asked whether it generalizes to the reading of blocks of text requiring eye movements. We measured visual-span profiles and reading speeds as a function of letter spacing. Visual-span profiles, measured with trigrams (strings of three random letters), are plots of letter-recognition accuracy as a function of letter position left or right of fixation. Size of the visual span was quantified by a measure of the area under the visual-span profile. Reading performance was measured using two presentation methods: RSVP and flashcard (a short block of text on four lines). We found that the size of the visual span and the reading speeds measured by the two presentation methods showed a qualitatively similar dependence on letter spacing and that they were highly correlated. These results are consistent with the view that the size of the visual span is a primary visual factor that limits reading speed.
PMCID: PMC2729067  PMID: 18217817
visual span; reading speed; letter spacing; visual crowding
4.  Are All Letters Really Processed Equally and in Parallel? Further Evidence of a Robust First Letter Advantage 
Acta psychologica  2013;144(2):10.1016/j.actpsy.2013.07.018.
This present study examined accuracy and response latency of letter processing as a function of position within a horizontal array. In a series of 4 Experiments, target-strings were briefly (33 ms for Experiment 1 to 3, 83 ms for Experiment 4) displayed and both forward and backward masked. Participants then made a two alternative forced choice. The two alternative responses differed just in one element of the string, and position of mismatch was systematically manipulated. In Experiment 1, words of different lengths (from 3 to 6 letters) were presented in separate blocks. Across different lengths, there was a robust advantage in performance when the alternative response was different for the letter occurring at the first position, compared to when the difference occurred at any other position. Experiment 2 replicated this finding with the same materials used in Experiment 1, but with words of different lengths randomly intermixed within blocks. Experiment 3 provided evidence of the first position advantage with legal nonwords and strings of consonants, but did not provide any first position advantage for non-alphabetic symbols. The lack of a first position advantage for symbols was replicated in Experiment 4, where target-strings were displayed for a longer duration (83 ms). Taken together these results suggest that the first position advantage is a phenomenon that occurs specifically and selectively for letters, independent of lexical constraints. We argue that the results are consistent with models that assume a processing advantage for coding letters in the first position, and are inconsistent with the commonly held assumption in visual word recognition models that letters are equally processed in parallel independent of letter position.
PMCID: PMC3859888  PMID: 24012723
letter-processing; serial position; visual word recognition
5.  Neural Dissociation of Number from Letter Recognition and Its Relationship to Parietal Numerical Processing 
The visual recognition of letters dissociates from the recognition of numbers at both the behavioral and neural level. In this article, using fMRI, we investigate whether the visual recognition of numbers dissociates from letters, thereby establishing a double dissociation. In Experiment 1, participants viewed strings of consonants and Arabic numerals. We found that letters activated the left midfusiform and inferior temporal gyri more than numbers, replicating previous studies, whereas numbers activated a right lateral occipital area more than letters at the group level. Because the distinction between letters and numbers is culturally defined and relatively arbitrary, this double dissociation provides some of the strongest evidence to date that a neural dissociation can emerge as a result of experience. We then investigated a potential source of the observed neural dissociation. Specifically, we tested the hypothesis that lateralization of visual number recognition depends on lateralization of higher-order numerical processing. In Experiment 2, the same participants performed addition, subtraction, and counting on arrays of nonsymbolic stimuli varying in numerosity, which produced neural activity in and around the intraparietal sulcus, a region associated with higher-order numerical processing. We found that individual differences in the lateralization of number activity in visual cortex could be explained by individual differences in the lateralization of numerical processing in parietal cortex, suggesting a functional relationship between the two regions. Together, these results demonstrate a neural double dissociation between letter and number recognition and suggest that higher-level numerical processing in parietal cortex may influence the neural organization of number processing in visual cortex.
PMCID: PMC3357212  PMID: 21736455
6.  VARiD: A variation detection framework for color-space and letter-space platforms 
Bioinformatics  2010;26(12):i343-i349.
Motivation: High-throughput sequencing (HTS) technologies are transforming the study of genomic variation. The various HTS technologies have different sequencing biases and error rates, and while most HTS technologies sequence the residues of the genome directly, generating base calls for each position, the Applied Biosystem's SOLiD platform generates dibase-coded (color space) sequences. While combining data from the various platforms should increase the accuracy of variation detection, to date there are only a few tools that can identify variants from color space data, and none that can analyze color space and regular (letter space) data together.
Results: We present VARiD—a probabilistic method for variation detection from both letter- and color-space reads simultaneously. VARiD is based on a hidden Markov model and uses the forward-backward algorithm to accurately identify heterozygous, homozygous and tri-allelic SNPs, as well as micro-indels. Our analysis shows that VARiD performs better than the AB SOLiD toolset at detecting variants from color-space data alone, and improves the calls dramatically when letter- and color-space reads are combined.
Availability: The toolset is freely available at
PMCID: PMC2881369  PMID: 20529926
7.  Effect of Orthographic Processes on Letter Identity and Letter-Position Encoding in Dyslexic Children 
The ability to identify letters and encode their position is a crucial step of the word recognition process. However and despite their word identification problem, the ability of dyslexic children to encode letter identity and letter-position within strings was not systematically investigated. This study aimed at filling this gap and further explored how letter identity and letter-position encoding is modulated by letter context in developmental dyslexia. For this purpose, a letter-string comparison task was administered to French dyslexic children and two chronological age (CA) and reading age (RA)-matched control groups. Children had to judge whether two successively and briefly presented four-letter strings were identical or different. Letter-position and letter identity were manipulated through the transposition (e.g., RTGM vs. RMGT) or substitution of two letters (e.g., TSHF vs. TGHD). Non-words, pseudo-words, and words were used as stimuli to investigate sub-lexical and lexical effects on letter encoding. Dyslexic children showed both substitution and transposition detection problems relative to CA-controls. A substitution advantage over transpositions was only found for words in dyslexic children whereas it extended to pseudo-words in RA-controls and to all type of items in CA-controls. Letters were better identified in the dyslexic group when belonging to orthographically familiar strings. Letter-position encoding was very impaired in dyslexic children who did not show any word context effect in contrast to CA-controls. Overall, the current findings point to a strong letter identity and letter-position encoding disorder in developmental dyslexia.
PMCID: PMC3356879  PMID: 22661961
letter-string processing; letter-position encoding; letter-identity encoding; letter transposition; letter substitution; reading acquisition; dyslexic children
8.  A comparison of Lea Symbols versus ETDRS letter distance visual acuity in a population of young children with a high prevalence of astigmatism 
To compare visual acuity results obtained using the Lea Symbols chart with results obtained using Early Treatment Diabetic Retinopathy Study (ETDRS) charts in young children who are members of a population with a high prevalence of astigmatism.
Subjects were 438 children aged 5 through 7 years who were enrolled in kindergarten or 1st grade on the Tohono O’odham Reservation: 241 (55%) had astigmatism ≥1.00 D in one or both eyes (range, 0.00 to 6.75 D). While wearing best correction, each child had right eye visual acuity tested with the 62 cm by 65 cm Lea Symbols chart at 3 m and with the 62 cm by 65 cm ETDRS chart at 4 m. Visual acuity was scored as the smallest optotype size at which the child correctly identified 3 out of a maximum of 5 optotypes. ETDRS visual acuity was also scored based on the total number of letters that the child correctly identified.
Correlation between Lea Symbols visual acuity and ETDRS visual acuity was 0.78 (p < 0.001). Mean Lea Symbols visual acuity was one-half line (0.04 to 0.06 logMAR) better than mean ETDRS visual acuity (p < 0.001). Difference between Lea Symbols and ETDRS visual acuity was not correlated with the mean of the Lea Symbols and ETDRS visual acuity scores, which ranged from −0.3 logMAR (20/10) to 0.74 logMAR (20/110).
In this population of young children, in whom the primary source of reduced visual acuity is astigmatism-related amblyopia, the Lea Symbols chart produced visual acuity scores that were about 0.5 line better than visual acuity scores obtained with ETDRS charts.
PMCID: PMC2764356  PMID: 19345596
9.  Reading Embossed Capital Letters: An fMRI Study in Blind and Sighted Individuals 
Human brain mapping  2006;27(4):325-339.
Reading Braille activates visual cortex in blind people [Burton et al., J Neurophysiol 2002;87: 589-611; Sadato et al., Nature 1996;380:526-528; Sadato et al., Brain 1998;121:1213-1229]. Because learning Braille requires extensive training, we had sighted and blind people read raised block capital letters to determine whether all groups engage visual cortex similarly when reading by touch. Letters were passively rubbed across the right index finger at 30 mm/s using an MR-compatible drum stimulator. Age-matched sighted, early blind (lost sight 0–5 years), and late blind (lost sight >5.5 years) volunteers performed three tasks: stating an identified letter, stating a verb containing an identified letter, and feeling a moving smooth surface. Responses were voiced immediately after the drum stopped moving across the fingertip. All groups showed increased activity in visual areas V1 and V2 during both letter identification tasks. Blind compared to sighted participants showed greater activation increases predominantly in the parafoveal-peripheral portions of visuotopic areas and posterior parts of BA 20 and 37. Sighted participants showed suppressed activity in most of the same areas except for small positive responses bilaterally in V1, left V5/MT+, and bilaterally in BA 37/20. Blind individuals showed suppression of the language areas in the frontal cortex, while sighted individuals showed slight positive responses. Early blind showed a more extensive distribution of activity in superior temporal sulcal multisensory areas. These results show cross-modal reorganization of visual cortex and altered response dynamics in nonvisual areas that plausibly reflect mechanisms for adaptive plasticity in blindness.
PMCID: PMC3684958  PMID: 16142777
blindness; human; magnetic resonance imaging; visual cortex; physiology
10.  Evidence for Letter-Specific Position Coding Mechanisms 
PLoS ONE  2013;8(7):e68460.
The perceptual matching (same-different judgment) paradigm was used to investigate precision in position coding for strings of letters, digits, and symbols. Reference and target stimuli were 6 characters long and could be identical or differ either by transposing two characters or substituting two characters. The distance separating the two characters was manipulated such that they could either be contiguous, separated by one intervening character, or separated by two intervening characters. Effects of type of character and distance were measured in terms of the difference between the transposition and substitution conditions (transposition cost). Error rates revealed that transposition costs were greater for letters than for digits, which in turn were greater than for symbols. Furthermore, letter stimuli showed a gradual decrease in transposition cost as the distance between the letters increased, whereas the only significant difference for digit and symbol stimuli arose between contiguous and non-contiguous changes, with no effect of distance on the non-contiguous changes. The results are taken as further evidence for letter-specific position coding mechanisms.
PMCID: PMC3699680  PMID: 23844204
11.  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
12.  Constraints on Letter-in-String Identification in Peripheral Vision: Effects of Number of Flankers and Deployment of Attention 
Effects of non-adjacent flanking elements on crowding of letter stimuli were examined in experiments manipulating the number of flanking elements and the deployment of spatial attention. To this end, identification accuracy of single letters was compared with identification of letter targets surrounded by two, four, or six flanking elements placed symmetrically left and right of the target. Target stimuli were presented left or right of a central fixation, and appeared either unilaterally or with an equivalent number of characters in the contralateral visual field (bilateral presentation). Experiment 1A tested letter targets with random letter flankers, and Experiments 1B and 2 tested letter targets with Xs as flanking stimuli. The results revealed a number of flankers effect that extended beyond standard two-flanker crowding. Flanker interference was stronger with random letter flankers compared with homogeneous Xs, and performance was systematically better under unilateral presentation conditions compared with bilateral presentation. Furthermore, the difference between the zero-flanker and two-flanker conditions was significantly greater under bilateral presentation, whereas the difference between two-flankers and four-flankers did not differ across unilateral and bilateral presentation. The complete pattern of results can be captured by the independent contributions of excessive feature integration and deployment of spatial attention to letter-in-string visibility.
PMCID: PMC3595509  PMID: 23494610
letter perception; crowding; non-adjacent flankers; number of flankers; spatial attention
13.  The nature of letter crowding as revealed by first- and second-order classification images 
Journal of vision  2007;7(2):5.1-526.
Visual crowding refers to the marked inability to identify an otherwise perfectly identifiable object when it is flanked by other objects. Crowding places a significant limit on form vision in the visual periphery; its mechanism is, however, unknown. Building on the method of signal-clamped classification images (Tjan & Nandy, 2006), we developed a series of first- and second-order classification-image techniques to investigate the nature of crowding without presupposing any model of crowding. Using an “o” versus “x” letter-identification task, we found that (1) crowding significantly reduced the contrast of first-order classification images, although it did not alter the shape of the classification images; (2) response errors during crowding were strongly correlated with the spatial structures of the flankers that resembled those of the erroneously perceived targets; (3) crowding had no systematic effect on intrinsic spatial uncertainty of an observer nor did it suppress feature detection; and (4) analysis of the second-order classification images revealed that crowding reduced the amount of valid features used by the visual system and, at the same time, increased the amount of invalid features used. Our findings strongly support the feature-mislocalization or source-confusion hypothesis as one of the proximal contributors of crowding. Our data also agree with the inappropriate feature-integration account with the requirement that feature integration be a competitive process. However, the feature-masking account and a front-end version of the spatial attention account of crowding are not supported by our data.
PMCID: PMC2635026  PMID: 18217820
crowding; letter identification; peripheral vision; classification images
14.  On Coding Non-Contiguous Letter Combinations 
Starting from the hypothesis that printed word identification initially involves the parallel mapping of visual features onto location-specific letter identities, we analyze the type of information that would be involved in optimally mapping this location-specific orthographic code onto a location-invariant lexical code. We assume that some intermediate level of coding exists between individual letters and whole words, and that this involves the representation of letter combinations. We then investigate the nature of this intermediate level of coding given the constraints of optimality. This intermediate level of coding is expected to compress data while retaining as much information as possible about word identity. Information conveyed by letters is a function of how much they constrain word identity and how visible they are. Optimization of this coding is a combination of minimizing resources (using the most compact representations) and maximizing information. We show that in a large proportion of cases, non-contiguous letter sequences contain more information than contiguous sequences, while at the same time requiring less precise coding. Moreover, we found that the best predictor of human performance in orthographic priming experiments was within-word ranking of conditional probabilities, rather than average conditional probabilities. We conclude that from an optimality perspective, readers learn to select certain contiguous and non-contiguous letter combinations as information that provides the best cue to word identity.
PMCID: PMC3122073  PMID: 21734901
bigrams; visual word recognition; information; optimization
15.  Differences between visual hemifields in identifying rapidly presented target stimuli: letters and digits, faces, and shapes 
The right hemisphere has been shown to play a dominant role in processing of visuo-spatial information. Recently, this role has been studied in the two-stream rapid serial visual presentation task. In this task, two alphanumerical targets are embedded in left and right simultaneous streams of rapidly changing letters. The second target (T2) is identified better in the left than in the right visual field. This difference has been interpreted as advantage of the right hemisphere (RH). However, a disadvantage of the left hemisphere (LH) could not be excluded so far. The LH, specialized for processing of verbal stimuli, might be overloaded due to constant input of letters from both visual fields. In the present study, this overload hypothesis was tested by reducing demands on verbal processing (Experiment 1), and by overloading the RH with non-verbal stimuli: faces (Experiment 2) and irregular shapes (Experiment 3). The left visual field advantage proved to be largely independent from the level of verbal load and from stimulus type. Therefore, although not entirely disproving the overload hypothesis, these results suggest as the most parsimonious explanation this asymmetry reflects a RH advantage, presumably in perceptual and attentional processing, rather than a LH disadvantage caused by verbal overload.
PMCID: PMC3715728  PMID: 23882249
RSVP; visual perception; hemispheric asymmetry; hemispheric specialization; lateralization; left visual-field advantage
16.  Dual-modality impairment of implicit learning of letter-strings versus color-patterns in patients with schizophrenia 
Implicit learning was reported to be intact in schizophrenia using artificial grammar learning. However, emerging evidence indicates that artificial grammar learning is not a unitary process. The authors used dual coding stimuli and schizophrenia clinical symptom dimensions to re-evaluate the effect of schizophrenia on various components of artificial grammar learning.
Letter string and color pattern artificial grammar learning performances were compared between 63 schizophrenic patients and 27 comparison subjects. Four symptom dimensions derived from a Chinese Positive and Negative Symptom Scale ratings were correlated with patients' artificial grammar implicit learning performances along the two stimulus dimensions. Patients' explicit memory performances were assessed by verbal paired associates and visual reproduction subtests of the Wechsler Memory Scales Revised Version to provide a contrast to their implicit memory function.
Schizophrenia severely hindered color pattern artificial grammar learning while the disease affected lexical string artificial grammar learning to a lesser degree after correcting the influences from age, education and the performance of explicit memory function of both verbal and visual modalities. Both learning performances correlated significantly with the severity of patients' schizophrenic clinical symptom dimensions that reflect poor abstract thinking, disorganized thinking, and stereotyped thinking.
The results of this study suggested that schizophrenia affects various mechanisms of artificial grammar learning differently. Implicit learning, knowledge acquisition in the absence of conscious awareness, is not entirely intact in patients with schizophrenia. Schizophrenia affects implicit learning through an impairment of the ability of making abstractions from rules and at least in part decreasing the capacity for perceptual learning.
PMCID: PMC1334227  PMID: 16343344
17.  The Overlap Model: A Model of Letter Position Coding 
Psychological review  2008;115(3):577-600.
Recent research has shown that letter identity and letter position are not integral perceptual dimensions (e.g., jugde primes judge in word-recognition experiments). Most comprehensive computational models of visual word recognition (e.g., the interactive activation model, J. L. McClelland & D. E. Rumelhart, 1981, and its successors) assume that the position of each letter within a word is perfectly encoded. Thus, these models are unable to explain the presence of effects of letter transposition (trial-trail), letter migration (beard-bread), repeated letters (moose-mouse), or subset/superset effects (faulty-faculty). The authors extend R. Ratcliff's (1981) theory of order relations for encoding of letter positions and show that the model can successfully deal with these effects. The basic assumption is that letters in the visual stimulus have distributions over positions so that the representation of one letter will extend into adjacent letter positions. To test the model, the authors conducted a series of forced-choice perceptual identification experiments. The overlap model produced very good fits to the empirical data, and even a simplified 2-parameter model was capable of producing fits for 104 observed data points with a correlation coefficient of .91.
PMCID: PMC2597794  PMID: 18729592
lexical process; letter position coding; word recognition; modeling; perceptual matching
18.  Letter representations in writing: an fMRI adaptation approach 
Behavioral and neuropsychological research in reading and spelling has provided evidence for the role of the following types of orthographic representations in letter writing: letter shapes, letter case, and abstract letter identities. We report on the results of an fMRI investigation designed to identify the neural substrates of these different representational types. Using an fMRI adaptation paradigm we examined the neural distribution of inhibition and release from inhibition in a letter-writing task in which, on every trial, participants produced three repetitions of the same letter and a fourth letter that was either identical to (no-change trial) or different from the previous three (change trial). Change trials involved a change in the shape, case, and/or identity of the letter. After delineating the general letter writing network by identifying areas that exhibited significant neural adaptation effects on no-change trials, we used deconvolution analysis to examine this network for effects of release from inhibition on change trials. In this way we identified regions specifically associated with the representation of letter shape (in the left SFS and SFG/pre-CG) and letter identity [in the left fusiform gyrus (FG)] or both [right cerebellum, left post-central gyrus (post-CG), and left middle frontal gyrus (MFG)]. No regions were associated with the representation of letter case. This study showcases an investigational approach that allows for the differentiation of the neurotopography of the representational types that are key to our ability to produce written language.
PMCID: PMC3809555  PMID: 24194724
letter; writing; fMRI; fMRI adaptation; neural habituation; letter shape; letter identity; letter case
19.  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
20.  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
21.  Fixation-Related fMRI Analysis in the Domain of Reading Research: Using Self-Paced Eye Movements as Markers for Hemodynamic Brain Responses During Visual Letter String Processing 
Cerebral Cortex (New York, NY)  2013;24(10):2647-2656.
The present study investigated the feasibility of using self-paced eye movements during reading (measured by an eye tracker) as markers for calculating hemodynamic brain responses measured by functional magnetic resonance imaging (fMRI). Specifically, we were interested in whether the fixation-related fMRI analysis approach was sensitive enough to detect activation differences between reading material (words and pseudowords) and nonreading material (line and unfamiliar Hebrew strings). Reliable reading-related activation was identified in left hemisphere superior temporal, middle temporal, and occipito-temporal regions including the visual word form area (VWFA). The results of the present study are encouraging insofar as fixation-related analysis could be used in future fMRI studies to clarify some of the inconsistent findings in the literature regarding the VWFA. Our study is the first step in investigating specific visual word recognition processes during self-paced natural sentence reading via simultaneous eye tracking and fMRI, thus aiming at an ecologically valid measurement of reading processes. We provided the proof of concept and methodological framework for the analysis of fixation-related fMRI activation in the domain of reading research.
PMCID: PMC4153805  PMID: 23645718
cerebrum; functional magnetic resonance imaging; language; visual word form area; visual word recognition
22.  Intact verbal description of letters with diminished awareness of their forms 
Visual processing and its conscious awareness can be dissociated. To examine the extent of dissociation between ability to read characters or words and to be consciously aware of their forms, reading ability and conscious awareness for characters were examined using a tachistoscope in an alexic patient. A right handed woman with 14 years of education presented with incomplete right hemianopia, alexia with kanji (ideogram) agraphia, anomia, and amnesia. Brain MRI disclosed cerebral infarction limited to the left lower bank of the calcarine fissure, lingual and parahippocampal gyri, and an old infarction in the right medial frontal lobe. Tachistoscopic examination disclosed that she could read characters aloud in the right lower hemifield when she was not clearly aware of their forms and only noted their presence vaguely. Although her performance in reading kanji was better in the left than the right field, she could read kana (phonogram) characters and Arabic numerals equally well in both fields. By contrast, she claimed that she saw only a flash of light in 61% of trials and noticed vague forms of stimuli in 36% of trials. She never recognised a form of a letter in the right lower field precisely. She performed judgment tasks better in the left than right lower hemifield where she had to judge whether two kana characters were the same or different. Although dissociation between performance of visual recognition tasks and conscious awareness of the visual experience was found in patients with blindsight or residual vision, reading (verbal identification) of characters without clear awareness of their forms has not been reported in clinical cases. Diminished awareness of forms in our patient may reflect incomplete input to the extrastriate cortex.

PMCID: PMC1736980  PMID: 10811708
23.  ERP correlates of letter identity and letter position are modulated by lexical frequency 
Brain and language  2013;125(1):11-27.
The encoding of letter position is a key aspect in all recently proposed models of visual-word recognition. We analyzed the impact of lexical frequency on letter position assignment by examining the temporal dynamics of lexical activation induced by pseudowords extracted from words of different frequencies. For each word (e.g., BRIDGE), we created two pseudowords: A transposed-letter (TL: BRIGDE) and a replaced-letter pseudoword (RL: BRITGE). ERPs were recorded while participants read words and pseudowords in two tasks: Semantic categorization (Experiment 1) and lexical decision (Experiment 2). For high-frequency stimuli, similar ERPs were obtained for words and TL-pseudowords, but the N400 component to words was reduced relative to RL-pseudowords, indicating less lexical/semantic activation. In contrast, TL- and RL-pseudowords created from low-frequency stimuli elicited similar ERPs. Behavioral responses in the lexical decision task paralleled this asymmetry. The present findings impose constraints on computational and neural models of visual-word recognition.
PMCID: PMC3612367  PMID: 23454070
visual-word recognition; position coding; ERPs; word-frequency; transposed-letter effects
24.  Effect of yoga relaxation techniques on performance of digit–letter substitution task by teenagers 
International Journal of Yoga  2009;2(1):30-34.
Memory and selective attention are important skills for academic and professional performance. Techniques to improve these skills are not taught either in education or company training courses. Any system which can systematically improve these skills will be of value in schools, universities, and workplaces. Aims:To investigate possible improvements in memory and selective attention, as measured by the Digit–Letter Substitution Task (DLST), due to practice of Cyclic Meditation (CM), a yoga relaxation technique, as compared to Supine Rest (SR).
Materials and Methods
Subjects consisted of 253 school students, 156 boys, 97 girls, in the age range 13–16 years, who were attending a 10-day yoga training course during summer vacation. The selected subjects had English as their medium of instruction in school and they acted as their own controls. They were allocated to two groups, and tested on the DLST, immediately before and after 22.5 minutes practice of CM on one day, and immediately before and after an equal period of SR on the other day. The first group performed CM on day 9 and SR on day 10. For the second group, the order was reversed.
Within each group pre-post test differences were significant for both the relaxation techniques. The magnitude of net score improvement was greater after SR (7.85%) compared to CM (3.95%). Significance levels were P < 0.4 × 10-9for SR and P < 0.1 × 10-3 for CM. The number of wrong attempts also increased significantly on both interventions, even after removing two outlier data points on day 1 in the SR group.
Both CM and SR lead to improvement in performance on the DLST. However, these relaxation techniques lead to more wrong cancellation errors.
PMCID: PMC3017965  PMID: 21234213
DLST; yoga; relaxation; meditation
25.  Word and Letter String Processing Networks in Schizophrenia: Evidence for Anomalies and Compensation 
Brain and language  2008;107(2):158-166.
Imaging studies show that in normal language correlated activity between anterior and posterior brain regions increases as the linguistic and semantic content (i.e., from false fonts, letter strings, pseudo words, to words) of stimuli increase. In schizophrenia however, disrupted functional connectivity between frontal and posterior brain regions has been frequently reported and these disruptions may change the nature of language organization. We characterized basic linguistic operations in word and letter string processing in a region-of-interest network using structural equation modeling (SEM). Healthy volunteers and volunteers with schizophrenia performed an fMRI one-back matching task with real words and consonant letter strings. We hypothesized that left hemisphere network dysfunction in schizophrenia would be present during processes dealing with linguistic/semantic content. The modeling results suggest aberrant left hemisphere function in schizophrenia, even in tasks requiring minimal access to language. Alternative mechanisms included increases in right hemisphere involvement and increased top-down influence from frontal to posterior regions.
PMCID: PMC2599869  PMID: 18829095
Schizophrenia and language; Lateralization; Lexical-semantic processing; Imaging; Effective Connectivity; Modeling

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