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Results from whole-brain analyses of training effects for the TPW word set. A, B, DIFF-1L pretraining DIFF-1L posttraining (A) and 1L-SAME DIFF-1L posttraining (B). In both analyses, the VWFA region is the only consistent location that shows a change in word selectivity as a result of training. C, Overlap (black) of the ROI from A (red) and B (blue). Color bar indicates t-value; horizontal line across the color bar shows the threshold that was applied for the analysis (see Materials and Methods).
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The nature of orthographic representations in the human brain is still subject of much debate. Recent reports have claimed that the visual word form area (VWFA) in left occipitotemporal cortex contains an orthographic lexicon based on neuronal representations highly selective for individual written real words (RWs). This theory predicts that learni...
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... At the brain level, the two routes are thought to share an initial stage of visual word form processing in left occipitotemporal cortex (Dehaene and Cohen, 2011), with only small topological changes depending on whether the task favors lexical or phonological processing (Bouhali et al., 2019). More anteriorly, ventral temporal cortex is strongly implicated as mediating the lexical route, with mid-fusiform cortex (mFus) likely functioning as the orthographic lexicon, a region where familiar letter strings are mapped onto known words (Nobre et al., 1994;Kronbichler et al., 2004;Glezer et al., 2015;Hirshorn et al., 2016;Lochy et al., 2018;White et al., 2019;Liu et al., 2021;Woolnough et al., 2021Woolnough et al., , 2022. mFus is sensitive to lexicality and word frequency (Kronbichler et al., 2004;White et al., 2019;Woolnough et al., 2021), and its activity is modulated by visual word learning (Taylor et al., 2014a;Glezer et al., 2015). ...
... More anteriorly, ventral temporal cortex is strongly implicated as mediating the lexical route, with mid-fusiform cortex (mFus) likely functioning as the orthographic lexicon, a region where familiar letter strings are mapped onto known words (Nobre et al., 1994;Kronbichler et al., 2004;Glezer et al., 2015;Hirshorn et al., 2016;Lochy et al., 2018;White et al., 2019;Liu et al., 2021;Woolnough et al., 2021Woolnough et al., , 2022. mFus is sensitive to lexicality and word frequency (Kronbichler et al., 2004;White et al., 2019;Woolnough et al., 2021), and its activity is modulated by visual word learning (Taylor et al., 2014a;Glezer et al., 2015). Conversely, the sublexical route, essential for articulating novel words, is thought to engage the inferior parietal lobe (IPL). ...
Reading words aloud is a fundamental aspect of literacy. The rapid rate at which multiple distributed neural substrates are engaged in this process can only be probed via techniques with high spatiotemporal resolution. We probed this with direct intracranial recordings covering most of the left hemisphere in 46 humans (26 male, 20 female) as they read aloud regular, exception and pseudo-words. We used this to create a spatiotemporal map of word processing and to derive how broadband γ activity varies with multiple word attributes critical to reading speed: lexicality, word frequency, and orthographic neighborhood. We found that lexicality is encoded earliest in mid-fusiform (mFus) cortex, and precentral sulcus, and is represented reliably enough to allow single-trial lexicality decoding. Word frequency is first represented in mFus and later in the inferior frontal gyrus (IFG) and inferior parietal sulcus (IPS), while orthographic neighborhood sensitivity resides solely in IPS. We thus isolate the neural correlates of the distributed reading network involving mFus, IFG, IPS, precentral sulcus, and motor cortex and provide direct evidence for parallel processes via the lexical route from mFus to IFG, and the sublexical route from IPS and precentral sulcus to anterior IFG.Significance Statement:Reading aloud depends on multiple complex cerebral computations: mapping from a written letter string on a page to a sequence of spoken sound representations. Here, we used direct intracranial recordings in a large cohort while they read aloud known and novel words, to track, across space and time, the progression of neural representations of behaviorally relevant factors that govern reading speed. We find, concordant with cognitive models of reading, that known and novel words are differentially processed through a lexical route, sensitive to frequency of occurrence of known words in natural language, and a sublexical route, performing letter-by-letter construction of novel words.
... Although previous research revealed the involvement of the left FG for visual word recognition Bruno et al., 2008;Glezer et al., 2009Glezer et al., , 2015Baeck et al., 2015;Lochy et al., 2018), the levels of orthographic structure for the left FG involvement have not been clearly elucidated (Kuo et al., 2004;Liu et al., 2008;Price and Devlin, 2011). Our results showed that the left occipitotemporal cortex preferentially responds to orthographically legal characters (i.e., RWs and PWs), which were consistent to previous findings (Price et al., 1996;Cohen et al., 2002;Ben-Shachar et al., 2007;Vinckier et al., 2007;Chan et al., 2009;Tian et al., 2020;Liu et al., 2021). ...
Visual word recognition has been proposed to have a functional and spatial organization corresponding to hierarchical language-like word forms in the left fusiform gyrus (FG) during visual word recognition in alphabetic languages. However, it is still unclear whether the similar functional gradients of word-like representation exist during Chinese character recognition. In this study, we adopted univariate activation analysis and representational similarity analysis (RSA) methods to investigate the functional organization in the FG for Chinese character recognition using task fMRI data. Native Chinese readers were visually presented with four types of character-like stimuli (i.e., real characters, pseudo-characters, false characters, and stroke combinations). After analysis, we observed a posterior-to-anterior functional gradient in the left FG corresponding to the degree of likeness of stimuli to character. Additionally, distinct subregions of the left FG harbor different orthographic codes. The middle part of the left FG was involved in abstract orthographic processing, while the anterior part of the left FG was involved in lexical orthographic processing (i.e., mapping orthography onto phonology or semantics). Notably, for the right FG, we did not find similar coding pattern for selectivity to character likeness, indicating the asymmetry of the functional hierarchical organization in favor of the left hemisphere. In conclusion, our findings revealed that the left FG presents a posterior-to-anterior gradient functional processing for Chinese character recognition, which expands our understanding of the psychological, neural, and computational theories of word reading.
... Of particular significance is that the participant began to verbalize words. This is of interest to these researchers because other researchers have suggested that for some children, reading acquisition may develop before language, and teaching whole language may be more effective than isolated phonics instruction (Glezer at al., 2015;Manning & Kamii, 2000). Manning and Kamii (2000) found that children who were taught whole language made more progress in both reading and writing, and with more developmental coherence. ...
Many developmental screeners focus heavily on receptive and expressive language skills, and the extent to which an infant can maneuver their environment. Research with young children typically involve motor skills, language, and occasionally simple procedural or problem solving tasks. The current study explores skills infants are expected to attain, and other skills that have never been tested in an infant who is considered “at-risk” due to moderate developmental delays. Researchers collected data via specialized VCA software, video recordings, and the Vineland-3 pre- and post-study. The participant improved in all areas measured by the Vineland-3. Additionally, despite the participant being introduced to novel and progressively more difficult tasks, his average attention span throughout the entirety of the study was significantly longer than previous research suggests for infants. Researchers also implemented the detour box as a gross measure of frontal function. The participant successfully completed the detour task and multi-step problem solving.
... Hence, reading acquisition would 5 In the remainder of the text, we will typically use the term 'letter strings' as a general term that encompasses words (e.g., familiar written forms), non-words (unmeaningful strings of letters), or even consonant strings. The distinction is relevant given the debate of whether the visual word form area (VWFA) processes only pre-lexical (McCandliss et al. 2003) or also lexical representations (Glezer et al. 2009(Glezer et al. , 2015Kronbichler et al. 2004). 6 Note that the spatial frequency hypothesis is not supported by training studies on reading acquisition with atypical visual shapes (such as faces or houses) differing greatly from the natural script characteristics, but also revealing the left middle fusiform gyrus as the site of learning effects (Moore et al. 2014;Hirshorn et al. 2016;Martin et al. 2019). ...
... Moreover, depending on the contrasted material (checkerboards, pseudofonts, or symbols), the location of the area with greater selectivity for letter strings can vary substantially (e.g., Fig. 2) from the "classical" VWFA with posterior coordinates (Talairach atlas: y = − 54; Cohen et al. 2000) to more anterior VOTC regions (y = − 42; Olulade et al. 2013;Turkeltaub et al. 2003; see also Martin et al. 2015). Further, depending on the tasks and contrasts used, this region is sometimes considered as being exclusively prelexical (Vinckier et al. 2007;Vogel et al. 2012) or lexical (Glezer et al. 2009(Glezer et al. , 2015, while its true selectivity to letter strings over object shapes can even be debated (see Fig. 3 above, from Dehaene et al. 2010, in which the "VWFA" response to pictures of tools is larger than to letter strings in all groups tested; see also Price and Devlin 2003;Starrfelt and Gerlach 2007). Future studies addressing a lateralized neural competition hypothesis should clarify these issues and develop more standard protocols, preferably with implicit tasks, in terms of the visual stimuli that are used to define selectivity, both for faces and letter strings. ...
The right hemispheric lateralization of face recognition, which is well documented and appears to be specific to the human species, remains a scientific mystery. According to a long-standing view, the evolution of language, which is typically substantiated in the left hemisphere, competes with the cortical space in that hemisphere available for visuospatial processes, including face recognition. Over the last decade, a specific hypothesis derived from this view according to which neural competition in the left ventral occipito-temporal cortex with selective representations of letter strings causes right hemispheric lateralization of face recognition, has generated considerable interest and research in the scientific community. Here, a systematic review of studies performed in various populations (infants, children, literate and illiterate adults, left-handed adults) and methodologies (behavior, lesion studies, (intra)electroencephalography, neuroimaging) offers little if any support for this reading lateralized neural competition hypothesis. Specifically, right-lateralized face-selective neural activity already emerges at a few months of age, well before reading acquisition. Moreover, consistent evidence of face recognition performance and its right hemispheric lateralization being modulated by literacy level during development or at adulthood is lacking. Given the absence of solid alternative hypotheses and the key role of neural competition in the sensory–motor cortices for selectivity of representations, learning, and plasticity, a revised language-related neural competition hypothesis for the right hemispheric lateralization of face recognition should be further explored in future research, albeit with substantial conceptual clarification and advances in methodological rigor.
... However, our results go further to show that familiarity with specific orientations does not lead to greater errors in part-based models, as might be expected if familiarity led to the formation of specialized whole object detectors. Rather, our results are consistent with the possibility that familiarity leads to sharpening and perceptual tuning of the parts, leading to more separable representations of the whole object (Agrawal et al., 2020;Glezer et al., 2015). We speculate that viewing familiar letters biases the preparation and selection stages of visual search by modifying the underlying feature representations (Eimer, 2014). ...
Fluent reading is an important milestone in education, but we lack a clear understanding of why children vary so widely in attaining it. Language-related factors such as rapid automatized naming (RAN) and phonological awareness have been identified as important factors that explain reading fluency. However, whether any aspects of visual orthographic processing also explain reading fluency beyond phonology is unclear. To investigate these issues, we tested primary school children (n = 68) on four tasks: two reading fluency tasks (word reading and passage reading), a RAN task to measure naming speed, and a visual search task using letters and bigrams. Bigram processing in visual search was accurately explained by single-letter discrimination, and error patterns were unrelated to fluency or bigram frequency, ruling out the contribution of specialized bigram detectors. As expected, the RAN score was strongly correlated with reading fluency. Importantly, there was a highly specific association between reading fluency and upright bigram processing in visual search. This association was specific to upright but not inverted bigrams and to bigrams with normal but not large letter spacing. It was explained by increased letter discrimination across bigrams and reduced interactions between letters within bigrams. Thus, fluent reading is accompanied by specialized changes in letter processing within bigrams. (PsycInfo Database Record (c) 2022 APA, all rights reserved).
... The current study mainly focused on the VWFA, located in the left occipitotemporal sulcus and involved in the visual and prelexical processing of words (Cohen et al., 2000;Dehaene and Cohen, 2011). Previous studies have already shown significant RS in the VWFA both by neuroimaging (e.g., Barton et al., 2010;Glezer et al., 2015) and electrophysiological methods (e.g., Cao et al., 2015;Li et al., 2019;Mercure et al., 2011). Literate humans are considered as experts in the recognition of words of their mother tongue. ...
The magnitude of repetition suppression (RS), measured by fMRI, is modulated by the probability of repetitions (P(rep)) for various sensory stimulus categories. It has been suggested that for visually presented simple letters this P(rep) effect depends on the prior practices of the participants with the stimuli. Here we tested further if previous experiences affect the neural mechanisms of RS, leading to the modulatory effects of stimulus P(rep), for more complex lexical stimuli as well. We measured the BOLD signal in the Visual Word Form Area (VWFA) of native Chinese and German participants and estimated the P(rep) effects for Chinese characters and German words. The results showed a significant P(rep) effect for stimuli of the mother tongue in both participant groups. Interestingly, Chinese participants, learning German as a second language, also showed a significant P(rep) modulation of RS for German words while the German participants who had no prior experiences with the Chinese characters showed no such effects. Our findings suggest that P(rep) effects on RS are manifest for visual word processing as well, but only for words of a language with which participants are highly familiar. These results support further the idea that predictive processes, estimated by P(rep) modulations of RS, require prior experiences.
... These direct neural recordings agree with a growing body of fMRI studies suggesting that the VWFA (likely VWFA-2) encodes word identity (Glezer et al. 2009(Glezer et al. , 2015Riesenhuber & Glezer 2017). Taken together, these studies suggest that learning new written words creates lexical entries that are stored in the VWFA and can be differentiated from letter strings that differ by even a www.annualreviews.org ...
The scientific study of reading has a rich history that spans disciplines from vision science to linguistics, psychology, cognitive neuroscience, neurology, and education. The study of reading can elucidate important general mechanisms in spatial vision, attentional control, object recognition, and perceptual learning, as well as the principles of plasticity and cortical topography. However, literacy also prompts the development of specific neural circuits to process a unique and artificial stimulus. In this review, we describe the sequence of operations that transforms visual features into language, how the key neural circuits are sculpted by experience during development, and what goes awry in children for whom learning to read is a struggle.
Expected final online publication date for the Annual Review of Vision Science, Volume 7 is September 2021. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.
... Because of the known role of Exner's area in handwriting, we created a 5-mm radius spherical ROI centered at the previously reported coordinates in the left PMd (−24, −4, 52) 23 . We also used a 5-mm radius ROI for the left occipitotemporal VWFA associated with orthographic codes of written words (−46, −56, −12) 35,68 , since this region may also play a role in novel word learning in adults 69,70 . For each ROI, neural effects of writing and group were assessed with voxel-level p < 0.05, corrected for multiple comparisons with familywise error across the search volume. ...
Handwriting is thought to impede vocabulary learning in sighted adults because the motor execution of writing interferes with efficient audiovisual processing during encoding. However, the motor memory of writing may facilitate adult word learning when visual sensory inputs are severely restricted. Using functional MRI, we show that late-blind participants, but not sighted participants, learned novel words by recruiting the left dorsal premotor cortex known as Exner’s writing area and its functional coupling with the left hippocampus. During later recall, the phonological and semantic contents of these words are represented in the activation patterns of the left hippocampus as well as in those of left frontotemporal language areas. These findings suggest that motor codes of handwriting help blind participants maintain word-form representations during learning and retrieval. We propose that such reliance on the motor system reflects a broad architecture of the cerebral language network which encompasses the limb motor system as a hardwired component.
... Our model, incorporating just word frequency and length, explained 73% of the variance of mid-fusiform activation. This central role of the mid-fusiform has also recently been suggested by selective haemodynamic changes following training to incorporate new words into the lexicon 43,44 . ...
Reading is a rapid, distributed process that engages multiple components of the ventral visual stream. To understand the neural constituents and their interactions that allow us to identify written words, we performed direct intra-cranial recordings in a large cohort of humans. This allowed us to isolate the spatiotemporal dynamics of visual word recognition across the entire left ventral occipitotemporal cortex. We found that mid-fusiform cortex is the first brain region sensitive to lexicality, preceding the traditional visual word form area. The magnitude and duration of its activation are driven by the statistics of natural language. Information regarding lexicality and word frequency propagates posteriorly from this region to visual word form regions and to earlier visual cortex, which, while active earlier, show sensitivity to words later. Further, direct electrical stimulation of this region results in reading arrest, further illustrating its crucial role in reading. This unique sensitivity of mid-fusiform cortex to sub-lexical and lexical characteristics points to its central role as the orthographic lexicon—the long-term memory representations of visual word forms.
... For hearing readers, it is often not even possible to identify the VWFA in the right hemisphere (the VWFA is typically identified through a separate localizer scan which contrasts words with other visual stimuli). For example, only two out of 20 participants demonstrated a right VWFA in Baker et al. (2007) and only 14 out of 34 in Glezer et al. (2015). The right VWFA was identified in 10 out of the 12 skilled deaf readers in the Glezer et al. (2018) study. ...
Deaf individuals have unique sensory and linguistic experiences that influence how they read and become skilled readers. This review presents our current understanding of the neurocognitive underpinnings of reading skill in deaf adults. Key behavioural and neuroimaging studies are integrated to build a profile of skilled adult deaf readers and to examine how changes in visual attention and reduced access to auditory input and phonology shape how they read both words and sentences. Crucially, the behaviours, processes, and neural circuity of deaf readers are compared to those of hearing readers with similar reading ability to help identify alternative pathways to reading success. Overall, sensitivity to orthographic and semantic information is comparable for skilled deaf and hearing readers, but deaf readers rely less on phonology and show greater engagement of the right hemisphere in visual word processing. During sentence reading, deaf readers process visual word forms more efficiently and may have a greater reliance on and altered connectivity to semantic information compared to their hearing peers. These findings highlight the plasticity of the reading system and point to alternative pathways to reading success.
