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1.  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.
doi:10.3389/fpsyg.2012.00154
PMCID: PMC3356879  PMID: 22661961
letter-string processing; letter-position encoding; letter-identity encoding; letter transposition; letter substitution; reading acquisition; dyslexic children
2.  Overt use of a tactile-kinesthetic strategy shifts to covert processing in rehabilitation of letter-by-letter reading 
Aphasiology  2010;24(11):1424-1442.
Background
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.
Aims
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.
Conclusions
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).
doi:10.1080/02687030903580333
PMCID: PMC3002229  PMID: 21170161
aphasia; cognitive rehabilitation; Speech-Language Pathology; pure alexia; letter-by-letter reading; generalization
3.  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.
doi:10.3389/fpsyg.2013.00119
PMCID: PMC3595509  PMID: 23494610
letter perception; crowding; non-adjacent flankers; number of flankers; spatial attention
4.  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.
doi:10.1167/7.2.2
PMCID: PMC2729067  PMID: 18217817
visual span; reading speed; letter spacing; visual crowding
5.  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.
doi:10.3389/neuro.09.056.2009
PMCID: PMC2796900  PMID: 20046826
word reading; ERPs; visual cortex; visual orthography
6.  Eye Movements When Reading Transposed Text: The Importance of Word-Beginning Letters 
Participants’ eye movements were recorded as they read sentences with words containing transposed adjacent letters. Transpositions were either external (e.g., problme, rpoblem) or internal (e.g., porblem, probelm) and at either the beginning (e.g., rpoblem, porblem) or end (e.g., problme, probelm) of words. The results showed disruption for words with transposed letters compared to the normal baseline condition, and the greatest disruption was observed for word-initial transpositions. In Experiment 1, transpositions within low frequency words led to longer reading times than when letters were transposed within high frequency words. Experiment 2 demonstrated that the position of word-initial letters is most critical even when parafoveal preview of words to the right of fixation is unavailable. The findings have important implications for the roles of different letter positions in word recognition and the effects of parafoveal preview on word recognition processes.
doi:10.1037/0096-1523.34.5.1261
PMCID: PMC2662926  PMID: 18823209
reading; eye movements; word recognition; transposed letters; parafoveal processing
7.  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.
doi:10.1016/j.bandl.2012.12.009
PMCID: PMC3612367  PMID: 23454070
visual-word recognition; position coding; ERPs; word-frequency; transposed-letter effects
8.  The impact of letter detection on eye movement patterns during reading: Reconsidering lexical analysis in connected text as a function of task 
A comparison was made between reading tasks performed with and without the additional requirement of detecting target letters. At issue was whether eye movement measures are affected by the additional requirement of detection. Global comparisons showed robust effects of task type with longer fixations and fewer word skippings when letter detection was required. Detailed analyses of target words, however, further showed that reading with and without letter detection yielded virtually identical effects of word class and text predictability for word-skipping rate and similar effects for different word viewing duration measures. The overall oculomotor pattern suggested that detection does not substantially shift normal reading movements in response to lexical cues and thereby indicated that detection tasks are informative about word and specifically word class processing in normal reading.
doi:10.1080/17470210600654776
PMCID: PMC2694497  PMID: 16885138
9.  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.


doi:10.1136/jnnp.68.6.782
PMCID: PMC1736980  PMID: 10811708
10.  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.
doi:10.3389/fpsyg.2013.00452
PMCID: PMC3715728  PMID: 23882249
RSVP; visual perception; hemispheric asymmetry; hemispheric specialization; lateralization; left visual-field advantage
11.  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.
doi:10.1162/jocn_a_00085
PMCID: PMC3357212  PMID: 21736455
12.  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 http://compbio.cs.utoronto.ca/varid
Contact: varid@cs.toronto.edu
doi:10.1093/bioinformatics/btq184
PMCID: PMC2881369  PMID: 20529926
13.  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.
doi:10.1080/13546800802550134
PMCID: PMC2679997  PMID: 19424881
Gestalt; Grouping; Contour integration; Good continuation; Letter identification; Object recognition; Features; Snake in the grass; Snake letters; Dot lattice
14.  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.
doi:10.1080/13546800802550134
PMCID: PMC2679997  PMID: 19424881
Gestalt; grouping; contour integration; good continuation; letter identification; object recognition; features; snake in the grass; snake letters; dot lattice
15.  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.
doi:10.1016/j.visres.2004.11.021
PMCID: PMC2741315  PMID: 15743610
Perceptual learning; Training; Letter identification; Peripheral vision
16.  A Common Left Occipito-Temporal Dysfunction in Developmental Dyslexia and Acquired Letter-By-Letter Reading? 
PLoS ONE  2010;5(8):e12073.
Background
We used fMRI to examine functional brain abnormalities of German-speaking dyslexics who suffer from slow effortful reading but not from a reading accuracy problem. Similar to acquired cases of letter-by-letter reading, the developmental cases exhibited an abnormal strong effect of length (i.e., number of letters) on response time for words and pseudowords.
Results
Corresponding to lesions of left occipito-temporal (OT) regions in acquired cases, we found a dysfunction of this region in our developmental cases who failed to exhibit responsiveness of left OT regions to the length of words and pseudowords. This abnormality in the left OT cortex was accompanied by absent responsiveness to increased sublexical reading demands in phonological inferior frontal gyrus (IFG) regions. Interestingly, there was no abnormality in the left superior temporal cortex which—corresponding to the onological deficit explanation—is considered to be the prime locus of the reading difficulties of developmental dyslexia cases.
Conclusions
The present functional imaging results suggest that developmental dyslexia similar to acquired letter-by-letter reading is due to a primary dysfunction of left OT regions.
doi:10.1371/journal.pone.0012073
PMCID: PMC2920311  PMID: 20711448
17.  Representation of Letter Position in Spelling: Evidence from Acquired Dysgraphia 
Cognition  2010;115(3):466-490.
The graphemic representations that underlie spelling performance must encode not only the identities of the letters in a word, but also the positions of the letters. This study investigates how letter position information is represented. We present evidence from two dysgraphic individuals, CM and LSS, who perseverate letters when spelling: that is, letters from previous spelling responses intrude into subsequent responses. The perseverated letters appear more often than expected by chance in the same position in the previous and subsequent responses. We used these errors to address the question of how letter position is represented in spelling. In a series of analyses we determined how often the perseveration errors produced maintain position as defined by a number of alternative theories of letter position encoding proposed in the literature. The analyses provide strong evidence that the grapheme representations used in spelling encode letter position such that position is represented in a graded manner based on distance from both edges of the word.
doi:10.1016/j.cognition.2010.03.013
PMCID: PMC2953246  PMID: 20378104
letter position coding; dysgraphia; spelling; letter perseveration errors; orthographic processing
18.  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.
doi:10.1016/j.bandl.2008.04.001
PMCID: PMC2599869  PMID: 18829095
Schizophrenia and language; Lateralization; Lexical-semantic processing; Imaging; Effective Connectivity; Modeling
19.  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.
doi:10.1371/journal.pone.0068460
PMCID: PMC3699680  PMID: 23844204
20.  Faces in the mist: illusory face and letter detection 
i-Perception  2011;2(5):458-476.
We report three behavioral experiments on the spatial characteristics evoking illusory face and letter detection. False detections made to pure noise images were analyzed using a modified reverse correlation method in which hundreds of observers rated a modest number of noise images (480) during a single session. This method was originally developed for brain imaging research, and has been used in a number of fMRI publications, but this is the first report of the behavioral classification images. In Experiment 1 illusory face detection occurred in response to scattered dark patches throughout the images, with a bias to the left visual field. This occurred despite the use of a fixation cross and expectations that faces would be centered. In contrast, illusory letter detection (Experiment 2) occurred in response to centrally positioned dark patches. Experiment 3 included an oval in all displays to spatially constrain illusory face detection. With the addition of this oval the classification image revealed an eyes/nose/mouth pattern. These results suggest that face detection is triggered by a minimal face-like pattern even when these features are not centered in visual focus.
doi:10.1068/i0421
PMCID: PMC3485785  PMID: 23145238
vision; face perception; reverse correlation; letter perception; top down; false detection
21.  Letter case and text legibility in normal and low vision 
Vision research  2007;47(19):2499-2505.
It is thought by cognitive scientists and typographers alike, that lower-case text is more legible than upper-case. Yet lower-case letters are, on average, smaller in height and width than upper-case characters, which suggests an upper-case advantage. Using a single unaltered font and all upper-, all lower-, and mixed-case text, we assessed size thresholds for words and random strings, and reading speeds for text with normal and visually impaired participants. Lower-case thresholds were roughly 0.1 log unit higher than upper-. Reading speeds were higher for upper- than for mixed-case text at sizes twice acuity size; at larger sizes, the upper-case advantage disappeared. Results suggest that upper-case is more legible than the other case styles, especially for visually-impaired readers, because smaller letter sizes can be used than with the other case styles, with no diminution of legibility.
doi:10.1016/j.visres.2007.06.010
PMCID: PMC2016788  PMID: 17675131
22.  Neural Correlates of Letter Reversal in Children and Adults 
PLoS ONE  2014;9(5):e98386.
Children often make letter reversal errors when first learning to read and write, even for letters whose reversed forms do not appear in normal print. However, the brain basis of such letter reversal in children learning to read is unknown. The present study compared the neuroanatomical correlates (via functional magnetic resonance imaging) and the electrophysiological correlates (via event-related potentials or ERPs) of this phenomenon in children, ages 5–12, relative to young adults. When viewing reversed letters relative to typically oriented letters, adults exhibited widespread occipital, parietal, and temporal lobe activations, including activation in the functionally localized visual word form area (VWFA) in left occipito-temporal cortex. Adults exhibited significantly greater activation than children in all of these regions; children only exhibited such activation in a limited frontal region. Similarly, on the P1 and N170 ERP components, adults exhibited significantly greater differences between typical and reversed letters than children, who failed to exhibit significant differences between typical and reversed letters. These findings indicate that adults distinguish typical and reversed letters in the early stages of specialized brain processing of print, but that children do not recognize this distinction during the early stages of processing. Specialized brain processes responsible for early stages of letter perception that distinguish between typical and reversed letters may develop slowly and remain immature even in older children who no longer produce letter reversals in their writing.
doi:10.1371/journal.pone.0098386
PMCID: PMC4032318  PMID: 24859328
23.  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.
doi:10.1016/j.visres.2003.09.028
PMCID: PMC2729075  PMID: 14751554
Reading; Letter-recognition; Peripheral vision; Perceptual learning; Low vision; Visual rehabilitation
24.  Which Children Benefit from Letter Names in Learning Letter Sounds? 
Cognition  2007;106(3):1322-1338.
Typical U.S. children use their knowledge of letters' names to help learn the letters' sounds. They perform better on letter sound tests with letters that have their sounds at the beginnings of their names, such as v, than with letters that have their sounds at the ends of their names, such as m, and letters that do not have their sounds in their names, such as h. We found this same pattern among children with speech sound disorders, children with language impairments as well as speech sound disorders, and children who later developed serious reading problems. Even children who scored at chance on rhyming and sound matching tasks performed better on the letter sound task with letters such as v than with letters such as m and h. Our results suggest that a wide range of children use the names of letters to help learn the sounds and that phonological awareness, as conventionally measured, is not required in order to do so.
doi:10.1016/j.cognition.2007.06.006
PMCID: PMC2267370  PMID: 17692304
25.  The nature of letter crowding as revealed by first- and second-order classification images 
Journal of vision  2007;7(2):5.1-526.
Visual crowding refers to the marked inability to identify an otherwise perfectly identifiable object when it is flanked by other objects. Crowding places a significant limit on form vision in the visual periphery; its mechanism is, however, unknown. Building on the method of signal-clamped classification images (Tjan & Nandy, 2006), we developed a series of first- and second-order classification-image techniques to investigate the nature of crowding without presupposing any model of crowding. Using an “o” versus “x” letter-identification task, we found that (1) crowding significantly reduced the contrast of first-order classification images, although it did not alter the shape of the classification images; (2) response errors during crowding were strongly correlated with the spatial structures of the flankers that resembled those of the erroneously perceived targets; (3) crowding had no systematic effect on intrinsic spatial uncertainty of an observer nor did it suppress feature detection; and (4) analysis of the second-order classification images revealed that crowding reduced the amount of valid features used by the visual system and, at the same time, increased the amount of invalid features used. Our findings strongly support the feature-mislocalization or source-confusion hypothesis as one of the proximal contributors of crowding. Our data also agree with the inappropriate feature-integration account with the requirement that feature integration be a competitive process. However, the feature-masking account and a front-end version of the spatial attention account of crowding are not supported by our data.
doi:10.1167/7.2.5
PMCID: PMC2635026  PMID: 18217820
crowding; letter identification; peripheral vision; classification images

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