Early development of letter specialization in left fusiform is associated with better word reading and smaller fusiform face area

A functional region of left fusiform gyrus termed “the visual word form area” (VWFA) develops during reading acquisition to respond more strongly to printed words than to other visual stimuli. Here, we examined responses to letters among 5‐ and 6‐year‐old early kindergarten children (N = 48) with little or no school‐based reading instruction who varied in their reading ability. We used functional magnetic resonance imaging (fMRI) to measure responses to individual letters, false fonts, and faces in left and right fusiform gyri. We then evaluated whether signal change and size (spatial extent) of letter‐sensitive cortex (greater activation for letters versus faces) and letter‐specific cortex (greater activation for letters versus false fonts) in these regions related to (a) standardized measures of word‐reading ability and (b) signal change and size of face‐sensitive cortex (fusiform face area or FFA; greater activation for faces versus letters). Greater letter specificity, but not letter sensitivity, in left fusiform gyrus correlated positively with word reading scores. Across children, in the left fusiform gyrus, greater size of letter‐sensitive cortex correlated with lesser size of FFA. These findings are the first to suggest that in beginning readers, development of letter responsivity in left fusiform cortex is associated with both better reading ability and also a reduction of the size of left FFA that may result in right‐hemisphere dominance for face perception.     

Visual experiences during letter production contribute to the development of the neural systems supporting letter perception

Letter production through handwriting creates visual experiences that may be important for the development of visual letter perception. We sought to better understand the neural responses to different visual percepts created during handwriting at different levels of experience. Three groups of participants, younger children, older children, and adults, ranging in age from 4.5 to 22 years old, were presented with dynamic and static presentations of their own handwritten letters, static presentations of an age‐matched control's handwritten letters, and typeface letters during fMRI. First, data from each group were analyzed through a series of contrasts designed to highlight neural systems that were most sensitive to each visual experience in each age group. We found that younger children recruited ventral‐temporal cortex during perception and this response was associated with the variability present in handwritten forms. Older children and adults also recruited ventral‐temporal cortex; this response, however, was significant for typed letter forms but not variability. The adult response to typed letters was more distributed than in the children, including ventral‐temporal, parietal, and frontal motor cortices. The adult response was also significant for one's own handwritten letters in left parietal cortex. Second, we compared responses among age groups. Compared to older children, younger children demonstrated a greater fusiform response associated with handwritten form variability. When compared to adults, younger children demonstrated a greater response to this variability in left parietal cortex. Our results suggest that the visual perception of the variability present in handwritten forms that occurs during handwriting may contribute to developmental changes in the neural systems that support letter perception.     

视觉词汇加工的动态神经网络及其形成

揭示大脑加工的神经网络机制成为认知神经科学研究的最新取向.本研究以视觉词汇加工脑区(VWFA)的神经功能作为切入点,探讨视觉词汇加工神经网络的动态机制及其形成.研究一考察VWFA在刺激驱动和任务调节下的动态激活,及其与语音、语义脑区所组成神经网络的动态机制.研究二通过跨文化对比以及儿童阅读发展研究,阐明语言经验对视觉词汇加工网络的塑造作用.研究三对比功能网络、静息网络以及白质纤维束联结,探讨视觉词汇加工网络的动态联结及其形成.研究结果有助于建构视觉词汇加工的神经生理模型,为基于脑科学的阅读教学和阅读障碍矫治奠定理论基础,为认知神经科学研究提供了新的思路.     

Familiarity affects visual processing of words.

While previous research has demonstrated that words can be processed more rapidly and/or more accurately than random strings of letters, it has not been convincingly demonstrated that the superior processing of words is a visual effect. In the present experiment, the cases of letters were manipulated in letter strings that were to be compared on the basis of physical identity. Mean response time was shorter for words than for nonwords even for pairs of letter strings that differed only in case (e.g., site-site). This finding implies that the advantage of words over nonwords (the familiarity effect) typically observed in the simultaneous matching task is not due solely to comparison of either the word names or the letter names and, thus, that at least part of the familiarity effect must be due to more rapid formation and/or comparison of visual representations of the two letter strings when they are words. Further analysis failed to reveal a significant involvement of phonemic or lexical codes in the comparison judgments.     

Conflicting strategies and hemispheric suppression in a lexical decision task

The research tests the prediction of the inhibitory-interaction hypothesis that experience with a task accentuates the functional imbalance between the hemispheres. Right-handed males who were experienced readers were presented a letter string to the centre visual field for lexical decision. The string was or was not accompanied by a blinking light to the left or right visual field. It was predicted that asymmetry would be greatest for strings that spelled words, less for strings that were orthographically correct (pseudowords) and least for strings that were orthographically incorrect (nonwords) because efficient adult readers have more experience with letter strings that do than do not spell a word, and have more experience with orthographically correct than incorrect letter strings. The analysis of response times supported the prediction. Moreover, in the nonword and in early trials of the pseudoword conditions, response times were faster when one or other hemisphere was distracted than when both were engaged suggesting the hemispheres use strategies that conflict when suppression has not been accentuated by practice. As well, as the trials progressed in the pseudoword condition, asymmetry reversed before increasing suggesting that the hemispheres reduce conflict by competing for and then strengthening suppression.     

Flank to the left,flank to the right: Testing the modified receptive field hypothesis of letter-specific crowding

The present study tested for effects of number of flankers positioned to the left and to the right of target characters as a function of visual field and stimulus type (letters or shapes). On the basis of the modified receptive field hypothesis (Chanceaux & Grainger, 2012), we predicted that the greatest effects of flanker interference would occur for leftward flankers with letter targets in the left visual field. Target letters and simple shape stimuli were briefly presented and accompanied by either 1, 2, or 3 flankers of the same category either to the left or to the right of the target, and in all conditions with a single flanker on the opposite side. Targets were presented in the left or right visual field at a fixed eccentricity, such that targets and flankers always fell into the same visual field. Results showed greatest interference for leftward flankers associated with letter targets in the left visual field, as predicted by the modified receptive field hypothesis.     

Rapid processing of letters,digits and symbols: what purely visual‐attentional deficit in developmental dyslexia?

Visual‐attentional theories of dyslexia predict deficits for dyslexic children not only for the perception of letter strings but also for non‐alphanumeric symbol strings. This prediction was tested in a two‐alternative forced‐choice paradigm with letters, digits, and symbols. Children with dyslexia showed significant deficits for letter and digit strings but not for symbol strings. This finding is difficult to explain for visual‐attentional theories of dyslexia which postulate identical deficits for letters, digits and symbols. Moreover, dyslexics showed normal W‐shaped serial position functions for letter and digit strings, which suggests that their deficit is not due to an abnormally small attentional window. Finally, the size of the deficit was identical for letters and digits, which suggests that poor letter perception is not just a consequence of the lack of reading. Together then, our results show that symbols that map onto phonological codes are impaired (i.e. letters and digits), whereas symbols that do not map onto phonological codes are not impaired. This dissociation suggests that impaired symbol‐sound mapping rather than impaired visual‐attentional processing is the key to understanding dyslexia.     

An early electrophysiological response associated with expertise in letter perception

Expertise with print is likely to optimize visual processes for recognizing characters of a familiar writing system. Although brain activations have been identified for words and letter strings in contrast with other stimuli, relatively little work has focused on the neural basis of single-letter perception. English readers and Chinese-English bilinguals participated in an ERP study and performed a 1-back identity judgment on Roman letters, Chinese characters, pseudofonts, and their string versions. The Chinese-English bilinguals showed an enhanced N170 for both Roman letters and Chinese characters relative to pseudofonts. For the non-Chinese readers, the N170 amplitude was larger for Roman letters relative to Chinese characters and pseudofonts. Our results suggest that changes in relatively early visual processes underlie expert letter perception.     

Functional specificity of the visual word form area: general activation for words and symbols but specific network activation for words

The functional specificity of the brain region known as the Visual Word Form Area (VWFA) was examined using fMRI. We explored whether this area serves a general role in processing symbolic stimuli, rather than being selective for the processing of words. Brain activity was measured during a visual 1-back task to English words, meaningful symbols (e.g., $, %), digits, words in an unfamiliar language (Hebrew), and geometric control stimuli. Mean activity in the functionally defined VWFA, as well as a pattern of whole-brain activity identified using a multivariate technique, did not differ for words and symbols, but was distinguished from that seen with other stimuli. However, functional connectivity analysis of this region identified a network of regions that was specific to words, including the left hippocampus, left lateral temporal, and left prefrontal cortex. Results support the hypothesis that activity in the VWFA plays a general role in processing abstract stimuli; however, the left VWFA is part of a unique network of brain regions active only during the word condition. These findings suggest that it is the neural "context" of the VWFA, i.e., the broader activity distributed in the brain that is correlated with VWFA, that is specific for visual word representation, not activity in this brain region per se.     

Effects of concurrent spatial and verbal memory loads on serial position functions of laterally presented letter strings

In this study the influence of visuospatial and verbal memory loads on the identification of laterally presented letter strings was investigated. In the asymmetry task without concurrent loads, a clear right visual field advantage for letter identification was obtained. Hemisphere-specific effects due to concurrent loads were particularly observed when the difficulty of the load tasks was increased. The effects of visuospatial loads were found to be sex related, suggesting that under heavy load conditions mental rotation selectively overloads the processing capacity of the right hemisphere in males, while in females capacity limitations were observed in both hemispheres. Concurrent visuospatial loads produced more facilitation (or less interference) for letters in the outermost positions of each visual field than for letters in the innermost positions of each visual field. The results of the verbal memory load tasks revealed that an easy verbal load task facilitated performance which was particularly manifest for the right-most letter of both the left visual field and the right visual field. A difficult verbal memory load task interfered with recognition accuracy of letters which was most marked for the center letter in the right visual field. Letter position effects obtained in this study were interpreted in terms of various processing mechanisms influencing the serial position curves.     

The role of visual spatial attention in adult developmental dyslexia

The present study investigated the nature of visual spatial attention deficits in adults with developmental dyslexia, using a partial report task with five-letter, digit, and symbol strings. Participants responded by a manual key press to one of nine alternatives, which included other characters in the string, allowing an assessment of position errors as well as intrusion errors. The results showed that the dyslexic adults performed significantly worse than age-matched controls with letter and digit strings but not with symbol strings. Both groups produced W-shaped serial position functions with letter and digit strings. The dyslexics' deficits with letter string stimuli were limited to position errors, specifically at the string-interior positions 2 and 4. These errors correlated with letter transposition reading errors (e.g., reading slat as “salt”), but not with the Rapid Automatized Naming (RAN) task. Overall, these results suggest that the dyslexic adults have a visual spatial attention deficit; however, the deficit does not reflect a reduced span in visual–spatial attention, but a deficit in processing a string of letters in parallel, probably due to difficulty in the coding of letter position.     

Selective attention and interhemispheric response competition in the split-brain

The interfering effect of an unattended stimulus on processing of an attended item was studied in a single split-brain participant (LB) and in normal controls. Pairs of letters were presented to the left visual field (LVF), right visual field (RVF), or bilaterally. Participants attended to the rightmost letter while attempting to ignore the leftmost letter. Responses associated with the attended and to-be-ignored letters could be compatible or incompatible. Manual response latencies were generally slower on Response Incompatible compared to Response Compatible trials. Notably, LB displayed this effect on Bilateral trials, where target and distractor were presented to opposite visual fields. LB was unable to perform a same-different matching task with bilateral letter stimuli, but was able to name bilateral letters accurately. Hence, in the bilateral condition, the ability to cross-compare letters was dissociated from attentional interference and from letter naming. Implications of these findings are discussed.     

Left hemisphere superiority for pronounceable nonwords,but not for unpronounceable letter strings

Right-handed adults were asked to identify bilaterally presented linguistic stimuli under three experimental conditions. In Condition A, stimuli were three-letter pronounceable nonwords (such as TUP), and subjects were asked to report them by naming them. In Condition B, stimuli were three-letter pronounceable nonwords, and subjects were asked to report them as strings of letters. In Condition C, stimuli were more or less unpronounceable letter strings (such as UTP) created by rearranging the letters of pronounceable nonwords, and subjects reported them as strings of letters. Pronounceable nonwords were found to be better identified from the right visual hemifield irrespective of the way in which they were reported. Unpronounceable letter strings did not produce any visual hemifield difference. Nonwords are of interest because they can be seen as potential words that lack both specific semantic properties and entries in the subject's internal lexicon. The results of the experiment are consistent with the view that both the left and right cerebral hemispheres are able to identify letters but the left hemisphere is more sensitive to the pronounceability of the nonwords. This may happen either because the left hemisphere can make better use of resemblances to real words or because it has access to spelling to sound correspondence rules.     

Sensori‐motor experience leads to changes in visual processing in the developing brain

Since Broca’s studies on language processing, cortical functional specialization has been considered to be integral to efficient neural processing. A fundamental question in cognitive neuroscience concerns the type of learning that is required for functional specialization to develop. To address this issue with respect to the development of neural specialization for letters, we used functional magnetic resonance imaging (fMRI) to compare brain activation patterns in pre‐school children before and after different letter‐learning conditions: a sensori‐motor group practised printing letters during the learning phase, while the control group practised visual recognition. Results demonstrated an overall left‐hemisphere bias for processing letters in these pre‐literate participants, but, more interestingly, showed enhanced blood oxygen‐level‐dependent activation in the visual association cortex during letter perception only after sensori‐motor (printing) learning. It is concluded that sensori‐motor experience augments processing in the visual system of pre‐school children. The change of activation in these neural circuits provides important evidence that ‘learning‐by‐doing’ can lay the foundation for, and potentially strengthen, the neural systems used for visual letter recognition.     

Parafoveal and foveal processing of abbreviations during eye fixations in reading: making a case for case

The processing of abbreviations in reading was examined with an eye movement experiment. Abbreviations were of 2 distinct types: acronyms (abbreviations that can be read with the normal grapheme-phoneme correspondence [GPC] rules, such as NASA) and initialisms (abbreviations in which the GPCs are letter names, such as NCAA). Parafoveal and foveal processing of these abbreviations was assessed with the use of the boundary change paradigm (K. Rayner, 1975). Using this paradigm, previews of the abbreviations were either identical to the abbreviation (NASA or NCAA), orthographically legal (NUSO or NOBA), or illegal (NRSB or NRBA). The abbreviations were presented as capital letter strings within normal, predominantly lowercase sentences and also sentences in all capital letters such that the abbreviations would not be visually distinct. The results indicate that acronyms and initialisms undergo different processing during reading and that readers can modulate their processing based on low-level visual cues (distinct capitalization) in parafoveal vision. In particular, readers may be biased to process capitalized letter strings as initialisms in parafoveal vision when the rest of the sentence is normal, lowercase letters.     

Whole-word units are used before orthographic knowledge in perceptual development

A visual search task for target letters in multiletter displays was used to investigate information-processing differences between college students and presecond-grade children (mean age = 7 years, 4 months). The stimulus displays consisted of single words, pronounceable pseudowords, and unpronounceable nonwords varying in length from three to five letters. The mean response times for indicating whether or not a target letter occurred in the display increased with the number of display letters for both groups, although there were apparent differences between groups in the rate of search and type of search strategy used. Pre-second-grade children responded faster to word displays than to pseudoword and nonword displays, indicating that familiar letter strings could be processed faster than unfamiliar strings regardless of whether or not the latter were consistent with rules of English orthography. In contrast, college students processed words and pseudowords about equally well, and both resulted in faster responses than nonwords. As reading skills develop, children apparently come to process familiar words differently from other letter strings. Only after a significant sightword vocabulary is established do children seem to recognize the regularities of standard English orthography and make use of this knowledge to facilitate perceptual processes.     

Neural correlates of subliminal and supraliminal letter processing--an event-related fMRI study

One problem of interpreting research on subconscious processing is the possibility that participants are weakly conscious of the stimuli. Here, we compared the fMRI BOLD response in healthy adults to clearly visible single letters (supraliminal presentation) with the response to letters presented in the absence of any behavioural evidence of visibility (subliminal presentation). No letter catch trials served as a control condition. Forced-choice responses did not differ from chance when letter-to-background contrast was low, whereas they were almost 100% correct when contrast was high. A comparison of fMRI BOLD signals for supraliminal and subliminal letters with the control trials revealed a signal increase in left BA 37 (fusiform gyrus). Comparison of supraliminal with subliminal letters showed a significant increase in the right inferior frontal gyrus (BA 44, partly extending to BA 9 and BA 45, as well as BA 46). Finally, a comparison of subliminal with supraliminal letters showed increases in the left middle temporal gyrus (BA 21) and the right extrastriate cortex (BA 19).     

The time course of visual influences in letter recognition

This study builds on a specific characteristic of letters of the Roman alphabet—namely, that each letter name is associated with two visual formats, corresponding to their uppercase and lowercase versions. Participants had to read aloud the names of single letters, and event-related potentials (ERPs) for six pairs of visually dissimilar upper- and lowercase letters were recorded. Assuming that the end product of processing is the same for upper- and lowercase letters sharing the same vocal response, ERPs were compared backward, starting from the onset of articulatory responses, and the first significant divergence was observed 120 ms before response onset. Given that naming responses were produced at around 414 ms, on average, these results suggest that letter processing is influenced by visual information until 294 ms after stimulus onset. This therefore provides new empirical evidence regarding the time course and interactive nature of visual letter perception processes.     

Developmental changes in neural letter‐selectivity: A 1‐year follow‐up of beginning readers

The developmental course of neural tuning to visual letter strings is unclear. Here we tested 39 children longitudinally, at the beginning of grade 1 (6.45 ± 0.33 years old) and 1 year after, with fast periodic visual stimulation in electroencephalography to assess the evolution of selective neural responses to letter strings and their relationship with emerging reading abilities. At both grades, frequency‐tagged letter strings were discriminated from pseudofont strings (i.e. letter‐selectivity) over the left occipito‐temporal cortex, with effects observed at the individual level in 62% of children. However, visual words were not discriminated from pseudowords (lexical access) at either grade. Following 1 year of schooling, letter‐selective responses showed a specific increase in amplitude, a more complex pattern of harmonics, and were located more anteriorly over the left occipito‐temporal cortex. Remarkably, at both grades, neural responses were highly significant at the individual level and correlated with individual reading scores. The amplitude increase in letter‐selective responses between grades was not found for discrimination responses of familiar keyboard symbols from pseudosymbols, and was not related to a general increase in visual stimulation responses. These findings demonstrate a rapid onset of left hemispheric letter selectivity, with 1 year of reading instruction resulting in increased emerging reading abilities and a clear quantitative and qualitative evolution within left hemispheric neural circuits for reading.     

Response latency in visual search with redundancy in the visual display

Two experiments were run to investigate the effects of redundant display items upon response latency in a visual search task. In the first study, Ss searched five-letter displays for a predesignated critical letter. Both critical and noncritical letters could be repeated in the displays. Mean response latency decreased markedly with increasing redundancy in the critical letter and was affected to a lesser extent by redundancy in the noncritical letters. In the second study, Ss were required to detect the presence of redundant letters in displays of from two to five letters, first with no information as to what letter might be repeated, then with knowledge of which letter would be repeated if the display contained a redundant letter. Response latencies in the former case were much slower than in the latter. The implications of these findings for current views of visual information processing were discussed.