Tuning of the human left fusiform gyrus to sublexical orthographic structure
Open accessAbstract
Neuropsychological and neurophysiological evidence point to a role for the left fusiform gyrus in visual word recognition, but the specific nature of this role remains a topic of debate. The aim of this study was to measure the sensitivity of this region to sublexical orthographic structure. We measured blood oxygenation (BOLD) changes in the brain with functional magnetic resonance imaging while fluent readers of English viewed meaningless letter strings. The stimuli varied systematically in their approximation to English orthography, as measured by the probability of occurrence of letters and sequential letter pairs (bigrams) comprising the string. A whole-brain analysis showed a single region in the lateral left fusiform gyrus where BOLD signal increased with letter sequence probability; no other brain region showed this response pattern. The results suggest tuning of this cortical area to letter probabilities as a result of perceptual experience and provide a possible neural correlate for the 'word superiority effect' observed in letter perception research.
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Learning to read tunes the receptive properties of underlying neurons to combinations of letters found within familiar words.
“led to suggestions that learning to read tuned the receptive properties of the underlying neurons to combinations of letters (such as bigrams and trigrams) that are found within familiar words (Cohen and Dehaene, 2004; Binder et al., 2006).”
From A review and synthesis of the first 20years of PET and fMRI studies of heard speech, spoken language and reading (2012) · cites this paper as Binder et al., 2006
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“Previous studies showed that the left fusiform gyrus is responsive to orthographic structure during visual word recognition (Binder et al., 2006), leading researchers to suggest that the role of the left fusiform is to integrate features into elaborate schemes that represent whole words or objects (Starrfelt and Gerla…”
BackgroundThe Interactive Account of ventral occipitotemporal contributions to reading
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“Part of our results overlap with those of a recent study by Binder et al. (2006), who collected fMRI data while subjects were presented with letter strings at four levels of positional bigram frequency, ranging from very low to high.”
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Binder, J. R., Westbury, C. F., McKiernan, K. A., Possing, E. T., & Medler, D. A. (2005) · Journal of Cognitive Neuroscience · doi:10.1162/0898929054021102 · Cited by 643 · see citing works
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“Indeed, in expert alpha-betic readers, the VWFA is organized in a posterior-to-anterior hierarchy 42–44 : posterior parts respond to individual letters 43 (irrespective of case 44 ), whereas anterior parts respond to letter combinations such as bigrams 42,45 .”
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BackgroundMethodInfluentialCracking the orthographic code: An introduction
Grainger, J. (2008) · Language and Cognitive Processes · doi:10.1080/01690960701578013 · Cited by 323 · see citing works
“This conclusion led to the ‘word shape’ hypothesis, according to which some form of supra-letter or holistic information about a printed word (e.g., the word envelope generated by a specific combination of ascending, descending, and neutral letters; Bouma, 1973) is used to identify the word.”
BackgroundMethodA ventral visual stream reading center independent of visual experience
Reich, L., Szwed, M., Cohen, L., & Amedi, A. (2011) · Current Biology · doi:10.1016/j.cub.2011.01.040 · Cited by 295 · see citing works
“Interestingly, previous studies in the sighted have suggested a visual anterior-posterior word processing chain along the left ventral occipitotemporal cortex, with preference for semantics anteriorly and word form posteriorly [31–33].”
Neural Systems for Reading Aloud: A Multiparametric Approach
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“Not only has this area been shown to be positively correlated with graphotactic probability in functional brain-imaging studies of healthy readers (Binder et al. 2006; Vinckier et al. 2007), but it is also one of the major areas that, when damaged, leads to a type of acquired dyslexia known as pure…”
BackgroundResultInfluentialSelf-citationEvidence for Highly Selective Neuronal Tuning to Whole Words in the “Visual Word Form Area”
Glezer, L. S., Jiang, X., & Riesenhuber, M. (2009) · Neuron · doi:10.1016/j.neuron.2009.03.017 · Cited by 293 · see citing works
“In summary, our results therefore provide strong support for theories of experience-driven plasticity of the neural representations underlying reading (Baker et al., 2007; Binder et al., 2006; Dehaene et al., 2005; Vinckier et al., 2007), establishing that this learning does not just apply to lower-level representatio…”
BackgroundResultInfluentialFunctional MRI of language lateralization during development in children
Holland, S. K., Vannest, J. J., Mecoli, M. D., Jacola, L. M., Tillema, J., Karunanayaka, P., Schmithorst, V. J., Yuan, W., Plante, E., & Byars, A. W. (2007) · International Journal of Audiology · doi:10.1080/14992020701448994 · Cited by 272 · see citing works
“The reader is referred to the works of Booth et al (2004); Binder et al (2006); Pugh (2006); and others for further insights about developing neural substrates supporting these facets of language so critical for reading proficiency.”
MethodChildren with dyslexia lack multiple specializations along the visual word-form (VWF) system.
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“Such deficits at multiple levels are in line with converging evidence that the occipitotemporal cortex has more than one function and responds to multiple levels of sublexical orthographical structures (Binder et al., 2006; Dehaene et al., 2002; Vinckier et al., 2007; Wydell et al., 2003).”
Origins of the specialization for letters and numbers in ventral occipitotemporal cortex
Hannagan, T., Amedi, A., Cohen, L., Dehaene‐Lambertz, G., & Dehaene, S. (2015) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2015.05.006 · Cited by 228 · see citing works
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Anatomical Connections of the Visual Word Form Area
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“Considering the positive correlation of fMRI activation in ROI 5 with bigram frequency (Binder et al., 2006), one may suggest that ROI 5 and the posterior MTG constitute an anatomically connected tandem involved in grapheme-to-phoneme mapping.”
BackgroundSpecialization for written words over objects in the visual cortex
Marcin Szwed, A., Dehaene, S., Kleinschmidt, A. K., Eger, E., Valabrègue, R., Amadon, A., & Cohen, L. (2011) · NeuroImage · doi:10.1016/j.neuroimage.2011.01.073 · Cited by 202 · see citing works
“This finding is consistent with consistent with greater activation in the context of more efficient recognition at the whole word level (Binder et al., 2006; Vinckier et al., 2007; Levy et al., 2008; Glezer et al., 2009).”
InfluentialThe Development of Cortical Sensitivity to Visual Word Forms
Ben-Shachar, M., Dougherty, R. F., Deutsch, G. K., & Wandell, B. A. (2011) · Journal of Cognitive Neuroscience · doi:10.1162/jocn.2011.21615 · Cited by 190 · see citing works
Dehaene, S., Nakamura, K., Jobert, A., Kuroki, C., Ogawa, S., & Cohen, L. (2009) · NeuroImage · doi:10.1016/j.neuroimage.2009.09.024 · Cited by 166 · see citing works
“…recognition (knowing that A and a are the same letter), learning of specific letter shapes (e.g. Hebrew versus Roman script), and compiling statistics of letter co-occurrence (bigrams), all of which have now been related to the VWFA (Baker et al., 2007; Binder et al., 2006; Dehaene et al., 2001).”
BackgroundThe Literate Brain: The Relationship between Spelling and Reading
Rapp, B. C., & Lipka, K. (2010) · Journal of Cognitive Neuroscience · doi:10.1162/jocn.2010.21507 · Cited by 160 · see citing works
“This finding is consistent with proposals in which the posterior–anterior axis of the left fusiform is characterized as instantiating increasingly abstract, complex, and eventually modality-independent lexical processing (Vinckier et al, 2007 ; Binder, Medler, Westbury, Liebenthal, & Buchanan, 2006 ; Dehaene, Cohen, S…”
ResultSequential then interactive processing of letters and words in the left fusiform gyrus
Thesen, T., McDonald, C. R., Carlson, C., Doyle, W., Cash, S., Sherfey, J. S., Felsovalyi, O., Girard, H. M., Barr, W., Devinsky, O., et al. (2012) · Nature Communications · doi:10.1038/ncomms2220 · Cited by 157 · see citing works
“Critically, activation in this area to letter-strings increases with their similarity to actual words[6, 7], especially in more anterior VWFA[8], suggested that it actually comprises a succession of detectors responding to progressively more abstract lexico-semantic aspects of the letter-strings.”
Neurobiological Underpinnings of Math and Reading Learning Disabilities
Ashkenazi, S., Black, J. M., Abrams, D. A., Hoeft, F., & Menon, V. (2013) · Journal of Learning Disabilities · doi:10.1177/0022219413483174 · Cited by 157 · see citing works
“…found that within the VOT, the left fusiform gyrus (FG; Table 1) is consistently activated across a wide range of numerical tasks (Arsalidou & Taylor, 2011), consistent with its hypothesized role in processing orthographic structure (Binder, Medler, Westbury, Liebenthal, & Buchanan, 2006).”
BackgroundWernicke’s area revisited: Parallel streams and word processing
DeWitt, I., & Rauschecker, J. P. (2013) · Brain and Language · doi:10.1016/j.bandl.2013.09.014 · Cited by 153 · see citing works
“Identification of this VOT site, analogous to macaque infero-temporal cortex, permitted detailed, mechanistic investigations to proceed, producing a prolific literature (Cohen et al., 2004; Binder et al., 2006; Gaillard et al., 2006; Baker et al., 2007; Vinckier et al., 2007; Turkeltaub et al., 2008; Glezer, Jiang & R…”
BackgroundBitan, T., Cheon, J., Lu, D., Burman, D. D., & Booth, J. R. (2008) · Journal of Cognitive Neuroscience · doi:10.1162/jocn.2009.21065 · Cited by 144 · see citing works
“Four left hemisphere regions were chosen because they were identified in our previous effective connectivity studies as part of the language network involved in the phonological judgment of visually presented words (Bitan et al., 2005, 2006).”
InfluentialThe neurobiological basis of seeing words
Wandell, B. A. (2011) · Annals of the New York Academy of Sciences · doi:10.1111/j.1749-6632.2010.05954.x · Cited by 142 · see citing works
“Some authors entertain the hypothesis that the VOT represents full words (lexical) [81,82,88,92,93] and memory for words [94].”
Wandell, B. A., Rauschecker, A. M., & Yeatman, J. D. (2011) · Annual Review of Psychology · doi:10.1146/annurev-psych-120710-100434 · Cited by 135 · see citing works
“…alexia (Cohen et al. 2003, Gaillard et al. 2006, Leff et al. 2001, Warrington & Shallice 1980), (c) VOT responses are sensitive to bigram frequency (Binder et al. 2006, Vinckier et al. 2007), (d ) there is a word-specific pattern of orthographic priming in VWFA (Dehaene et al. 2001), and (e) VWFA…”
BackgroundChauncey, K., Holcomb, P. J., & Jonathan Grainger, B. (2008) · Language and Cognitive Processes · doi:10.1080/01690960701579839 · Cited by 137 · see citing works
“Orthographic processing would then proceed in a posterior-anterior sweep through left fusiform, known to be sensitive to orthographic structure (e.g., Binder, Medler, Westbury, Liebenthal, & Buchanan, 2006; Cohen et al., 2000; Dehaene et al., 2005).”
BackgroundCortical dynamics of word recognition
Mainy, N., Jung, J., Baciu, M., Kahane, P., Schoendorff, B., Minotti, L., Hoffmann, D., Bertrand, O., & Lachaux, J.-P. (2007) · Human Brain Mapping · doi:10.1002/hbm.20457 · Cited by 135 · see citing works
“…predominantly in the left hemisphere, involving in (a) visual analysis: early visual areas (BA 17, 18, and 19) and midfusiform gyrus (BA 19, 37) [Binder et al., 2006; Cohen et al., 2000; Devlin et al., 2006; Gaillard et al., 2006; James et al., 2005; Mechelli et al., 2007; Vigneau et al.,…”
BackgroundSpeech-Specific Tuning of Neurons in Human Superior Temporal Gyrus
Chan, A. M., Dykstra, A. R., Jayaram, V., Leonard, M. K., Travis, K. E., Gygi, B., Baker, J. M., Eskandar, E. N., Hochberg, L. R., Halgren, E., et al. (2013) · Cerebral Cortex · doi:10.1093/cercor/bht127 · Cited by 136 · see citing works
Auditory Selective Attention to Speech Modulates Activity in the Visual Word Form Area
Yoncheva, Y. N., Zevin, J. D., Maurer, U., & McCandliss, B. D. (2009) · Cerebral Cortex · doi:10.1093/cercor/bhp129 · Cited by 131 · see citing works
“by performance measures), both known to affect activity in the VWFA (Booth et al. 2003; Binder et al. 2006).”
BackgroundInfluentialDecoding and disrupting left midfusiform gyrus activity during word reading
Hirshorn, E. A., Li, Y., Ward, M., Richardson, R. M., Fiez, J. A., & Ghuman, A. S. (2016) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1604126113 · Cited by 127 · see citing works
“We next used techniques from machine learning in iEEG data from P1 and P4 to assess the sensitivity of lmFG to sublexical, orthographic statistics (bigram frequency) that has been hypothesized as an indicator for a visual word form system (16, 21).”
BackgroundMethodSpatiotemporal Dynamics of Orthographic and Lexical Processing in the Ventral Visual Pathway
Woolnough, O., Donos, C., Rollo, P., Forseth, K., Lakretz, Y., Crone, N., Fischer-Baum, S., Dehaene, S., & Tandon, N. (2020) · Nature Human Behaviour · doi:10.1038/s41562-020-00982-w · Cited by 121 · see citing works
Dien, J. (2008) · Neuropsychologia · doi:10.1016/j.neuropsychologia.2008.08.024 · Cited by 122 · see citing works
Interactive Activation and Mutual Constraint Satisfaction in Perception and Cognition
McClelland, J. L., Mirman, D., Bolger, D. J., & Khaitan, P. (2014) · Cognitive Science · doi:10.1111/cogs.12146 · Cited by 122 · see citing works
Dien, J. (2008) · Biological Psychology · doi:10.1016/j.biopsycho.2008.04.013 · Cited by 119 · see citing works
Arithmetic in the Child and Adult Brain
Menon, V. (2014) · Oxford University Press eBooks · doi:10.1093/oxfordhb/9780199642342.013.041 · Cited by 119 · see citing works
Lochy, A., Jacques, C., Maillard, L. G., Colnat‐Coulbois, S., Rossion, B., & Jonas, J. (2018) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1718987115 · Cited by 114 · see citing works
Sensitivity to orthographic familiarity in the occipito-temporal region.
Bruno, J. L., Zumberge, A., Manis, F. R., Lu, Z.-L., & Goldman, J. G. (2008) · NeuroImage · doi:10.1016/j.neuroimage.2007.10.044 · Cited by 114 · see citing works
“The findings of Binder et al. (2006) that the OT region responded to manipulations of orthographic familiarity at the sub-word level necessitated holding bigram frequency constant across conditions. Given that we controlled for bigram frequency, we are able to conclude that OT is sensitive to whole-word orthographic f…”
InfluentialSchurz, M., Sturm, D., Richlan, F., Kronbichler, M., Ladurner, G., & Wimmer, H. (2009) · NeuroImage · doi:10.1016/j.neuroimage.2009.10.082 · Cited by 114 · see citing works
“Recent evidence for prelexical tuning of the VWFA was provided by Binder et al. (2006), who found that OT cortex activation systematically increased with increasingly higher letter sequence probability (i.e., bigram-frequency) of a nonword.”
Attentional Focus During Learning Impacts N170 ERP Responses to an Artificial Script
Yoncheva, Y. N., Blau, V. C., Maurer, U., & McCandliss, B. D. (2010) · Developmental Neuropsychology · doi:10.1080/87565641.2010.480918 · Cited by 110 · see citing works
Levy, J., Pernet, C., Treserras, S., Boulanouar, K., Aubry, F., Démonet, J.-F., & Celsis, P. (2009) · PLoS ONE · doi:10.1371/journal.pone.0006675 · Cited by 107 · see citing works
“However, in recent years more and more experimental evidence is gained over the more general role of the LOT [12] reflecting an operation common to the processing of words [46], word sub-units [19], pictures [47], objects [48], or stored visual forms and structures in general [49].”
BackgroundInfluentialLochy, A., Belle, G. V., & Rossion, B. (2014) · Neuropsychologia · doi:10.1016/j.neuropsychologia.2014.11.007 · Cited by 105 · see citing works
“This region is usually equally activated by real words and readable pseudowords but less by nonwords (consonant strings for instance) and shows, as we found here, a gradually increasing response to alphabetic stimuli with increasing letter probability, thus with higher word-likeness (Binder et al., 2006).”
Tsapkini, K., & Rapp, B. C. (2009) · Cortex · doi:10.1016/j.cortex.2009.02.025 · Cited by 104 · see citing works
“N IH -PA Author M anuscript N IH -PA Author M anuscript N IH -PA Author M anuscript increasingly more “abstract” orthographic functions (Cohen, et al., 2003; Vinckier et al., 2007; Binder et al., 2006).”
BackgroundAlpha and theta brain oscillations index dissociable processes in spoken word recognition
Strauß, A., Kotz, S. A., Scharinger, M., & Obleser, J. (2014) · NeuroImage · doi:10.1016/j.neuroimage.2014.04.005 · Cited by 98 · see citing works
“Binder et al. (2006) gathered evidence that this area is especially sensitive to sublexical bigram frequency—a pivotal element of our study design.”
Weber-Goericke, F., & Muehlhan, M. (2018) · Journal of Affective Disorders · doi:10.1016/j.jad.2018.09.049 · Cited by 97 · see citing works
Effects of Literacy in Early Visual and Occipitotemporal Areas of Chinese and French Readers
Szwed, M., Qiao, E., Jobert, A., Dehaene, S., & Cohen, L. (2014) · Journal of Cognitive Neuroscience · doi:10.1162/jocn_a_00499 · Cited by 98 · see citing works
“Xue and colleagues (2005) found strong left-hemispheric dominance in 12-year-old Chinese children, and Chan and colleagues (2009) found that, in Chinese readers, only the left-hemisphere visual system shows a spatial hierarchical organization of brain responses to increasingly languagelike word forms, as occurs for La…”
BackgroundPark, J., Hebrank, A. C., Polk, T. A., & Park, D. C. (2011) · Journal of Cognitive Neuroscience · doi:10.1162/jocn_a_00085 · Cited by 98 · see citing works
“Activation in VWFA is hierarchically organized so that the strength of activation increases with orthographic regularities (Vinckier et al., 2007) and bigram frequency ( Binder, Medler, Westbury, Liebenthal, & Buchanan, 2006 ).”
The Neural Basis of Visual Word Form Processing: A Multivariate Investigation
Nestor, A., Behrmann, M., & Plaut, D. C. (2012) · Cerebral Cortex · doi:10.1093/cercor/bhs158 · Cited by 93 · see citing works
“This proposal found ground in the equivalent levels of activation detected for words and pseudowords at the level of the vWFA (Dehaene et al. 2002; Binder et al. 2006; Vinckier et al. 2007).”
Mano, Q. R., Humphries, C., Desai, R. H., Seidenberg, M. S., Osmon, D. C., Stengel, B. C., & Binder, J. R. (2012) · Cerebral Cortex · doi:10.1093/cercor/bhs093 · Cited by 87 · see citing works
“This approach removes variation both within and between conditions that can be explained by a linear function of RT (Binder et al. 2006; Graves et al. 2010).”
InfluentialSelf-citationPosition sensitivity in the visual word form area
Rauschecker, A. M., Bowen, R. F., Parvizi, J., & Wandell, B. A. (2012) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1121304109 · Cited by 90 · see citing works
“If no absolute visual field position information is present, the neural circuitry could encode relative position implicitly by representing ordered letter strings (bigrams, trigrams, and so forth) (34, 35).”
BackgroundHauk, O., Davis, M. H., & Friedemann Pulvermüller, A. (2008) · NeuroImage · doi:10.1016/j.neuroimage.2008.05.054 · Cited by 84 · see citing works
“One fMRI study found larger activation for more “word-like” nonwords compared to nonwords with low frequency letter constellations in a letter detection task (Binder et al., 2006).”
Rothlein, D., & Rapp, B. C. (2013) · NeuroImage · doi:10.1016/j.neuroimage.2013.11.054 · Cited by 77 · see citing works
“This region has been strongly associated with orthographic processing in adult reading (Binder et al., 2006; Cohen et al., 2000; Gaillard et al., 2006; Dehaene et al., 2002; Jobard et al., 2003) and spelling (Rapp and Lipka, 2011; Purcell et al.”
BackgroundProblems with visual statistical learning in developmental dyslexia
Sigurdardottir, H. M., Daníelsdóttir, H. B., Gudmundsdottir, M., Hjartarson, K. H., Þórarinsdóttir, E. Á., & Kristjansson, A. K. (2017) · Scientific Reports · doi:10.1038/s41598-017-00554-5 · Cited by 75 · see citing works
Impaired recognition of faces and objects in dyslexia: Evidence for ventral stream dysfunction?
Sigurdardottir, H. M., Ívarsson, E., Kristinsdóttir, K., & Kristjánsson, Á. (2015) · Neuropsychology · doi:10.1037/neu0000188 · Cited by 73 · see citing works
A Comparative Perspective on the Human Temporal Lobe
Bryant, K., & Preuss, T. M. (2017) · Springer eBooks · doi:10.1007/978-4-431-56582-6_16 · Cited by 71 · see citing works
Functional Neuroanatomy of Contextual Acquisition of Concrete and Abstract Words
Mestres-Missé, A., Münte, T. F., & Rodríguez‐Fornells, A. (2008) · Journal of Cognitive Neuroscience · doi:10.1162/jocn.2008.21171 · Cited by 70 · see citing works
“The VWFA has been related to prelexical representation of visual words, responding preferentially to letter strings than to other categories of visual stimuli (for a review, see Cohen & Dehaene, 2004), and to word and word-like (pseudowords) stimuli than consonant strings ( Binder, Medler, Westbury, Liebenthal, & Buch…”
An ERP study of category priming: Evidence of early lexical semantic access
Segalowitz, S. J., & Zheng, X. W. (2008) · Biological Psychology · doi:10.1016/j.biopsycho.2008.04.009 · Cited by 67 · see citing works
“Using fMRI, Binder et al. (2006) found that the brain region that is sensitive to positional bigram frequencies is the left lateral fusiform gyrus, which is the visual word form area (VWFA), a brain region that is particularly responsive to words and pseudowords compared to other types of visual…”
Do orthotactics and phonology constrain the transposed-letter effect?
Perea, M., & Carreiras, M. (2008) · Language and Cognitive Processes · doi:10.1080/01690960701578146 · Cited by 66 · see citing works
“The lexical interpretation would be consistent with the recent findings reported by Binder, Medler, Westbury, Liebenthal, and Buchanan (2006) using an fMRI methodology.”
A mesial-to-lateral dissociation for orthographic processing in the visual cortex
Bouhali, F., Bézagu, Z., Dehaene, S., & Cohen, L. (2019) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1904184116 · Cited by 64 · see citing works
“12–14) and have shown that it probably hosts multiple orthographic representations of letters (15, 16), bigrams (17), and short words (18).”
BackgroundPleisch, G., Karipidis, I. I., Brauchli, C., Röthlisberger, M., Hofstetter, C., Stämpfli, P., Walitza, S., & Brem, S. (2019) · NeuroImage · doi:10.1016/j.neuroimage.2019.01.046 · Cited by 62 · see citing works
“, 2003; Kourtzi and Kanwisher, 2001; Yovel and Kanwisher, 2004), and its crucial role in orthographic processing is widely accepted and has been demonstrated in numerous studies (Baker et al., 2007a; Binder et al., 2006; Bruno et al., 2008; Cohen et al., 2002; Dehaene and Cohen, 2011; Dehaene et al., 2002, 2005; Gleze…”
Reconsidering the role of orthographic redundancy in visual word recognition
Chetail, F. (2015) · Frontiers in Psychology · doi:10.3389/fpsyg.2015.00645 · Cited by 62 · see citing works
“More generally, a critical point regarding orthographic redundancy is that it naturally covaries with other factors known to impact visual word recognition, such as pronunceability (Binder et al., 2006), word frequency (Adams, 1981), or orthographic neighborhood (Andrews, 1992).”
BackgroundDien, J., Brian, E. S., Molfese, D. L., & Gold, B. T. (2013) · Cortex · doi:10.1016/j.cortex.2013.03.008 · Cited by 59 · see citing works
“…the broad expanse of the FG, a region labeled the Visual Word Form Area (VWFA), located at roughly y=−50, is thought to mediate pairwise letter associations, or bigrams (Binder et al., 2006; Dehaene and Cohen, 2011; Dehaene et al., 2005; Dien, 2009a; McCandliss et al., 2003; Polk and Farah, 1997).”
Anatomy and physiology of word-selective visual cortex: from visual features to lexical processing
Caffarra, S., Karipidis, I. I., Yablonski, M., & Yeatman, J. D. (2021) · Brain Structure and Function · doi:10.1007/s00429-021-02384-8 · Cited by 62 · see citing works
Does the visual word form area split in bilingual readers? A millimeter-scale 7-T fMRI study
Zhan, M.-Y., Pallier, C., Agrawal, A., Dehaene, S., & Cohen, L. (2022) · bioRxiv · doi:10.1126/sciadv.adf6140 · Cited by 45 · see citing works
“When written stimuli respect increasingly well the statistics of words in the known language, they elicit increasingly greater activity in the VWFA ( 72 ).”
Kinoshita, S., Wit, B. D., & Norris, D. G. (2016) · Journal of Experimental Psychology Learning Memory and Cognition · doi:10.1037/xlm0000311 · Cited by 58 · see citing works
Byun, J., Kim, H., Kang, H., Cha, K. S., Sunwoo, J., Shin, J.-W., Moon, J., Lee, S., Jung, K., Chu, K., et al. (2020) · Sleep Medicine · doi:10.1016/j.sleep.2020.01.010 · Cited by 57 · see citing works
Left fusiform BOLD responses are inversely related to word-likeness in a one-back task
Wang, X., Yang, J., Hua Shu, A., & Zevin, J. D. (2011) · NeuroImage · doi:10.1016/j.neuroimage.2010.12.062 · Cited by 56 · see citing works
den Broeck, W. V., & Geudens, A. (2012) · Cognitive Psychology · doi:10.1016/j.cogpsych.2012.06.003 · Cited by 55 · see citing works
Decoding levels of representation in reading: A representational similarity approach
Fischer‐Baum, S., Bruggemann, D., Gallego, I. F., Li, D. S. P., & Tamez, E. R. (2017) · Cortex · doi:10.1016/j.cortex.2017.02.017 · Cited by 56 · see citing works
Woollams, A. M., Silani, G., Okada, K., Patterson, K., & Price, C. J. (2011) · Journal of Cognitive Neuroscience · doi:10.1162/jocn.2010.21502 · Cited by 54 · see citing works
“, 2004), frequent bigrams (Vinckier et al., 2007; Binder et al., 2006), high demands on visual discrimination (Rogers et al.”
BackgroundUncovering phonological and orthographic selectivity across the reading network using fMRI-RA
Glezer, L. S., Eden, G. F., Jiang, X., Luetje, M. M., Napoliello, E. M., Kim, J. S., & Riesenhuber, M. (2016) · NeuroImage · doi:10.1016/j.neuroimage.2016.05.072 · Cited by 58 · see citing works
“Specifically, we have shown that rather than processing prelexical information, such as commonly occurring letter combinations (Binder et al., 2006; Cohen and Dehaene, 2004; Dehaene and Cohen, 2011) the VWFA is finely tuned to whole real words (Glezer et al., 2009) AC C EP TE D M AN U SC R IP T and…”
Winter, D., Krause-Utz, A., Lis, S., Chiu, C.-D., Lanius, R. A., Schriner, F., Bohus, M., & Schmahl, C. (2015) · Psychiatry Research: Neuroimaging · doi:10.1016/j.pscychresns.2015.05.018 · Cited by 54 · see citing works
“, 2001) and word recognition (Nobre et al., 1994; Pammer et al., 2004; Binder et al., 2006), among others.”
BackgroundThe Neural Correlates of Reading Fluency Deficits in Children
Langer, N., Benjamin, C. F. A., Minas, J. E., & Gaab, N. (2013) · Cerebral Cortex · doi:10.1093/cercor/bht330 · Cited by 54 · see citing works
Rakhlin, N. V., Mourgues, C., Cardoso-Martins, C., Kornev, A., & Grigorenko, E. (2019) · Contemporary Educational Psychology · doi:10.1016/j.cedpsych.2018.12.002 · Cited by 43 · see citing works
Early Visual Word Processing Is Flexible: Evidence from Spatiotemporal Brain Dynamics
Chen, Y., Davis, M. H., Friedemann Pulvermüller, A., & Hauk, O. (2015) · Journal of Cognitive Neuroscience · doi:10.1162/jocn_a_00815 · Cited by 51 · see citing works
Beyond the visual word form area: The orthography–semantics interface in spelling and reading
Purcell, J. J., Shea, J., & Rapp, B. C. (2014) · Cognitive Neuropsychology · doi:10.1080/02643294.2014.909399 · Cited by 51 · see citing works
Piecemeal recruitment of left-lateralized brain areas during reading: A spatio-functional account
Lévy, J., Pernet, C. R., Treserras, S., Boulanouar, K., Berry, I., Aubry, F., Démonet, J., & Celsis, P. (2008) · NeuroImage · doi:10.1016/j.neuroimage.2008.08.008 · Cited by 51 · see citing works
“…operation common to the processing of words with regular (Kronbichler et al., 2007) or irregular (Bruno et al., 2008) orthographies, word sub-units (Binder et al., 2006), pictures (Starrfelt and Gerlach 2007), objects (Bar et al., 2001), or stored visual forms and structures in general (Joseph and…”
Tsapkini, K., Vindiola, M. M., & Rapp, B. C. (2010) · NeuroImage · doi:10.1016/j.neuroimage.2010.12.024 · Cited by 50 · see citing works
“…hemisphere mid-fusiform gyrus/occipito-temproal sulcus has been identified as an important component of the reading network (Baker et al., 2007; Binder et al., 2003, 2006; Booth et al., 2002; Carreiras et al., 2007; Cohen et al., 2000, 2002, 2004; Dehaene et al., 2002, 2004; Fiez et al., 1999;…”
Hannagan, T., Agrawal, A., Cohen, L., & Dehaene, S. (2021) · Proceedings of the National Academy of Sciences of the United States of America · doi:10.1073/pnas.2104779118 · Cited by 41 · see citing works
Automaticity of phonological and semantic processing during visual word recognition
Pattamadilok, C., Chanoine, V., Pallier, C., Anton, J., Nazarian, B., Belin, P., & Ziegler, J. C. (2017) · NeuroImage · doi:10.1016/j.neuroimage.2017.02.003 · Cited by 48 · see citing works
“Precisely, in the left anterior and middle fusiform gyrus (at the location of the Visual Word Form Area reported in previous studies, e.g., Cohen et al., 2002; Jobard et al., 2003; Binder et al., 2006), precentral gyrus, SMA and IFG tri, the effect of stimulus visibility was significantly (or marginally, p ≤ .”
Processing deficits for familiar and novel faces in patients with left posterior fusiform lesions
Roberts, D. J., Ralph, M. A. L., Kim, E., Tainturier, M., Beeson, P. M., Rapcsak, S. Z., & Woollams, A. M. (2015) · Cortex · doi:10.1016/j.cortex.2015.02.003 · Cited by 47 · see citing works
“Neuroimaging results indicate that the VWFA is sensitive to the familiarity of subword letter combinations like bigrams and trigrams (Vinckier et al., 2007; Binder et al., 2006).”
Graziano, M., & Sigman, M. (2008) · Journal of Vision · doi:10.1167/8.5.9 · Cited by 48 · see citing works
Top-Down Processing of Symbolic Meanings Modulates the Visual Word Form Area
Song, Y., Tian, M., & Jia Liu, B. (2012) · Journal of Neuroscience · doi:10.1523/jneurosci.1874-12.2012 · Cited by 47 · see citing works
“…Baker et al., 2007; Brem et al., 2010; Dehaene et al., 2010) and is not sensitive to low-level features of written words, such as size, position, font, or letter case (Dehaene et al., 2004; Binder et al., 2006; Vinckier et al., 2007; Glezer et al., 2009; Qiao et al., 2010; Braet et al., 2012).”
BackgroundLudersdorfer, P., Wimmer, H., Richlan, F., Schurz, M., Hutzler, F., & Kronbichler, M. (2016) · NeuroImage · doi:10.1016/j.neuroimage.2015.09.039 · Cited by 42 · see citing works
“…have shown that activation in this region ismodulated by the familiarity of sublexical orthographic features (e.g., the frequency of pairs or triplets of letters) with frequent features eliciting higher activation relative to less frequent features (Binder et al., 2006; Vinckier et al., 2007).”
The impact of letter spacing on reading: A test of the bigram coding hypothesis
Vinckier, F., Qiao, E., Pallier, C., Dehaene, S., & Cohen, L. (2011) · Journal of Vision · doi:10.1167/11.6.8 · Cited by 45 · see citing works
Orthographic Contamination of Broca’s Area
Montant, M., Schön, D., Anton, J., & Ziegler, J. C. (2011) · Frontiers in Psychology · doi:10.3389/fpsyg.2011.00378 · Cited by 46 · see citing works
“As a matter of fact, a wealth of research indicates that the vOTC is the brain region that is sensitive to the orthographic familiarity of letter strings (Binder et al., 2006; Dehaene and Cohen, 2011).”
BackgroundVisual Feature-Tolerance in the Reading Network
Rauschecker, A. M., Bowen, R. F., Perry, L. M., Kevan, A., Dougherty, R. F., & Wandell, B. A. (2011) · Neuron · doi:10.1016/j.neuron.2011.06.036 · Cited by 44 · see citing works
“shape representations (Binder et al., 2006; Dehaene et al., 2005; Glezer et al., 2009; Vinckier et al., 2007), independent of the specific visual features that define these shapes.”
BackgroundYablonski, M., Rastle, K., Taylor, J. S. H., & Ben‐Shachar, M. (2018) · Cortex · doi:10.1016/j.cortex.2018.06.011 · Cited by 42 · see citing works
Visual print tuning deficits in dyslexic adolescents under minimized phonological demands
Kronschnabel, J., Schmid, R., Maurer, U., & Brandeis, D. (2013) · NeuroImage · doi:10.1016/j.neuroimage.2013.02.014 · Cited by 41 · see citing works
“…al., 2006; Brem et al., 2009; Maurer et al., 2005, 2006; Xue and Poldrack, 2007), while fine-tuning refers to the sublexical and whole word levels (Binder et al., 2006; Dehaene et al., 2005; Glezer et al., 2009; Kronbichler et al., 2004; Vinckier et al., 2007) as for example reflected by…”
BackgroundAging Influences the Neural Correlates of Lexical Decision but Not Automatic Semantic Priming
Gold, B. T., Andersen, A. H., Jicha, G. A., & Smith, C. D. (2009) · Cerebral Cortex · doi:10.1093/cercor/bhp018 · Cited by 41 · see citing works
“…regions within this network play different roles in LD. Extrastriate regions have been linked with orthographic processes such as visual feature extraction and mapping visual patterns onto lexical representations (Petersen et al. 1990; Binder et al. 2003, 2006; Cohen et al. 2003; Gold et al. 2006).”
Graves, W. W., Binder, J. R., Desai, R. H., Humphries, C., Stengel, B. C., & Seidenberg, M. S. (2014) · Brain and Language · doi:10.1016/j.bandl.2014.03.005 · Cited by 40 · see citing works
Self-citationNeural networks underlying contributions from semantics in reading aloud
Boukrina, O. V., & Graves, W. W. (2013) · Frontiers in Human Neuroscience · doi:10.3389/fnhum.2013.00518 · Cited by 39 · see citing works
“This should be most evident in areas previously implicated in semantic processing such as the ITS (Binder et al., 2009; Graves et al., 2010) and AG (Binder et al., 1997, 2005, 2009; Price and Mechelli, 2005; Price, 2012).”
BackgroundResultProtein Analysis Meets Visual Word Recognition: A Case for String Kernels in the Brain
Hannagan, T., & Jonathan Grainger, B. (2012) · Cognitive Science · doi:10.1111/j.1551-6709.2012.01236.x · Cited by 39 · see citing works
“Turning to Open-bigram coding, Binder et al. (2006) found that activity in the VWFA was correlated with the frequency of (contiguous) bigrams in letter strings presented to subjects.”
BackgroundWhat is special about orthographic processing?
Grainger, J., & Hannagan, T. (2014) · Written Language & Literacy · doi:10.1075/wll.17.2.03gra · Cited by 37 · see citing works
“Furthermore, fMRI research has provided evidence for the existence of such letter combination representations in the VWFA (Binder et al., 2006; Vinckier et al., 2007), and our own computational modeling work has shown how letter combinations can emerge during orthographic learning (Hannagan et al., 2013).”
BackgroundInfluentialIndependence of Early Speech Processing from Word Meaning
Travis, K. E., Leonard, M. K., Chan, A. M., Torres, C., Sizemore, M. L., Qu, Z., Eskandar, E. N., Dale, A. M., Elman, J. L., Cash, S. S., et al. (2012) · Cerebral Cortex · doi:10.1093/cercor/bhs228 · Cited by 38 · see citing works
“Keywords: ECoG, MEG, N400, speech processing”
- Authors
- Binder, J. R., Medler, D. A., Westbury, C. F., Liebenthal, E., & Buchanan, L.
- Affiliations
- Medical College of Wisconsin (US)
- University of Alberta (CA)
- University of Windsor (CA)
- Corresponding author
- Binder, Jeffrey R.
- Venue
- NeuroImage
- Publisher
- Elsevier BV
- Published
- 2006-11
- Volume
- 33
- Issue
- 2
- Pages
- 739-748
- ISSN
- 1053-8119 (ISSN-L)
- PMC
- PMC1634933
- OpenAlex
- W2070993882
- Cited by
- 286
- References
- 74
- Citations by year
- 2026: 2 · 2025: 10 · 2024: 9 · 2023: 9 · 2022: 14 · 2021: 15 · 2020: 9 · 2019: 14 · 2018: 16 · 2017: 15 · 2016: 13 · 2015: 26 · 2014: 16 · 2013: 20 · 2012: 21
- FWCI
- 23.72
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Locations
- NeuroImage (publishedVersion)
- PubMed (publishedVersion)
- Europe PMC (PubMed Central) (acceptedVersion)
- Funders
- National Institute of Mental Health (P01MH51358, P01MH051358)
- National Institute of Neurological Disorders and Stroke (R01NS033576, R01 NS33576)
- National Center for Research Resources (M01-RR00058, M01RR000058)
- Topics
- Social Sciences › Psychology › Developmental and Educational Psychology › Reading and Literacy Development
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neurobiology of Language and Bilingualism
- Life Sciences › Neuroscience › Cognitive Neuroscience › Visual perception and processing mechanisms
- Keywords
- fusiform gyrus, visual word recognition, bigrams, word superiority effect, letter perception, perceptual experience, English orthography, BOLD signal
- MeSH terms
- Adult, Brain Mapping, Cerebrovascular Circulation, Educational Status, Female, Gyrus Cinguli, Humans, Image Processing, Computer-Assisted, Functional Laterality, Magnetic Resonance Imaging, Male, Middle Aged, Oxygen, Reference Values
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