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...age (Ben-Shachar et al., 2007; Mahon and Caramazza, 2011; Pinel and Dehaene, 2010). In order to examine the underlying functional connectivity in the blind, we investigated the intrinsic (rest state; =-=Biswal et al., 1995-=-) functional connectivity in the blind from a small seed region focused on the canonical VWFA (for details see Supplemental Experimental Procedures). We found that the VWFA of the blind showed highly ...

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....05 corrected for multiple comparisons, using the spatial extent method based on the theory 650 Neuron 76, 640–652, November 8, 2012 ª2012 Elsevier Inc.of Gaussian random fields (Forman et al., 1995; =-=Friston et al., 1993-=-). This method takes the data contiguity of neighboring voxels directly into account and corrects for the false-positive rate of continuous clusters (a set-level statistical inference correction). Thi...

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...esponses to letters as compared to different visual object categories (such as faces, houses, and objects; Cohen and Dehaene, 2004; Dehaene and Cohen, 2011; Dehaene et al., 2010; Hasson et al., 2002; =-=Puce et al., 1996-=-; Szwed et al., 2011; Tsapkini and Rapp, 2010), similar to the preferential activation of the neighboring regions for faces, scenes, objects, and body shapes (Kanwisher, 2010). Can full category selec...

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...eld (Bolger et al., 2005; NeuronVWFA Visual Selectivity in the Blind Using Sounds2003), which converts visual images to auditory ‘‘soundscapes’’ using a predetermined consistent algorithm (The vOICe; =-=Meijer, 1992-=-). This enabled the blind to perceive high-resolution visual information (Striem-Amit et al., 2012b) and, in this case, to learn to read, with sounds topographically representing visual images of lett...

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...trols for both auditory stimulation and semantic content; see Figure S1B). One area of activation is ofparticular interest given theories on mental imagery originally framed in the context of vision (=-=Kosslyn et al., 1999-=-): we found robust activation to Braille imagery as compared to the semantic control in the hand area of S1 (Figure S1C; t = 6.5, p < 0.000001). This mental imagery reactivation was specific to the re...

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...ual input were restored medically in adulthood. This VWFA Visual Selectivity in the Blind Using Soundsclaim was supported by early studies of a critical period for developing normal sight in animals (=-=Wiesel and Hubel, 1963-=-) and humans (Lewis and Maurer, 2005). It was also supported by the poor functional outcomes observed after rare cases of sight restoration in humans, especially in ventral stream tasks show that comb...

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...honemes, similar to learning a novel script via vision in a literate person (Hashimoto and Sakai, 2004; Maurer et al., 2010; Xue et al., 2006). Bayesian learning principles (Ernst and Bulthoff, 2004; =-=Tenenbaum et al., 2011-=-) enable the extraction of abstract schemas behind superficially different inputs, including sensory modalities. By learning to extract the abstract interpretation of a sound input as a two-dimensiona...

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... (Mahon and Caramazza, 2009; Pinel and Dehaene, 2010) and (2) a sensitivity to the visual features that characterize scripts, such as reliance on line junctions (Szwed et al., 2011), foveal position (=-=Hasson et al., 2002-=-), and high spatial frequencies (Woodhead et al., 2011). How dependent is VWFA selectivity on such visual sensory features? It was recently shown (Rauschecker et al., 2011) that reading activates VWFA...

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...elve images from the same category were presented in each epoch; each image was presented for 800 ms and was followed by a 200 ms blank screen (similar to standard visual localizer experiments; e.g., =-=Hasson et al., 2003-=-). A central red fixation point was present throughout the experiment. The subjects were instructed to covertly classify and identify the displayed objects, as in the main experiment. Imagery Control ...

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...ll-width at half-maximum Gaussian). Group analyses were conducted for the main experiment and visual localizer experiment using a general linear model (GLM) in a hierarchical random-effects analysis (=-=Friston et al., 1999-=-). For the imagery control experiment and the case study, the data were grouped using GLM in a fixed-effects analysis. All GLM contrastsbetween two conditions included comparison of the first term of ...

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...-Canada, Faculty of Medicine lpêtrière, IFR 70, 75013 Paris, France , 75013 Paris, France euroimaging Unit, 91191 Gif sur Yvette, France of Bioimaging, Neurospin, 91191 Gif sur Yvette, France 2000; =-=Dehaene and Cohen, 2011-=-; Schlaggar and McCandliss, 2007) or left ventral occipito-temporal cortex (vOT; Price, 2012; Price and Devlin, 2011; Wandell, 2011). Extensive research has demonstrated the specialization of this reg...

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...al cortex of the blind ‘‘switches tasks’’ and processes nonvisual functions that differ considerably from those typical of the sighted brain, such as memory and language in the primary visual cortex (=-=Amedi et al., 2003-=-; Bedny et al., 2011; see reviews in Frasnelli et al., 2011; Merabet and Pascual-Leone, 2010; Striem-Amit et al., 2011). Here we show that when relevant stimuli and tasks are introduced, the ventral v...

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... (DeWitt and Rauschecker, 2012; Talairachcoordinates 56, 16, 2; statistics from this ROI; t = 11.2, p < 0.000001; see Figure S3), as well as to more posterior areas in the auditory ventral stream (=-=Rauschecker and Scott, 2009-=-), which may correspond to the phoneme-processing network (DeWitt and Rauschecker, 2012). The VWFA of the blind also 648 Neuron 76, 640–652, November 8, 2012 ª2012 Elsevier Inc.showed functional conne...

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...0; Hasson et al., 2002; Puce et al., 1996; Szwed et al., 2011; Tsapkini and Rapp, 2010), similar to the preferential activation of the neighboring regions for faces, scenes, objects, and body shapes (=-=Kanwisher, 2010-=-). Can full category selectivity in the VWFA also emerge without visual experience and by using auditory sensory substi-Figure 1. ‘‘Visual’’ Performance in Blind Users of ‘‘The vOICe’’ Sensory Substit...

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... Similar to the activation in the sighted for letters relative to the baseigure 2A), the congenitally blind group nsive activation of the occipito-temporal (see Figure 2B, as seen previously in blind =-=Burton et al., 2002-=-; Reich et al., 2011). We ditory cortex activation (including A1/ blind for this contrast, given the auditory s the VWFA is characterized not only by Neuronactivation to letters but mostly by its sele...

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...od and improves over many years of practice (Aghababian and Nazir, 2000). Recent evidence shows that plasticity of the ventral visual cortex extends into adolescence and beyond (Dehaene et al., 2010; =-=Golarai et al., 2007-=-). However, does the VWFA show selectivity for script over other object categories when it is trained to read using an SSD in the fully developed adult brain and with quantitatively limited practice? ...

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...es of visual objects (Cohen and Dehaene, 2004; Dehaene and Cohen, 2011; Dehaene et al., 2010; Szwed et al., 2011), its invariance to changes in visual scripts, fonts, or location in the visual field (=-=Bolger et al., 2005-=-; NeuronVWFA Visual Selectivity in the Blind Using Sounds2003), which converts visual images to auditory ‘‘soundscapes’’ using a predetermined consistent algorithm (The vOICe; Meijer, 1992). This enab...

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...to one task over another and fewer yet have investigated the existence in the blind of a critical feature of the ventral visual cortex, namely, its regional selectivity for perceptual categories (see =-=Pietrini et al., 2004-=-; Mahon et al., 2009, who explored large-scale preference patterns). The current study now shows same category selectivity for a specific visual category (letters), as seen in the sighted, in the abse...

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... adulthood. This VWFA Visual Selectivity in the Blind Using Soundsclaim was supported by early studies of a critical period for developing normal sight in animals (Wiesel and Hubel, 1963) and humans (=-=Lewis and Maurer, 2005-=-). It was also supported by the poor functional outcomes observed after rare cases of sight restoration in humans, especially in ventral stream tasks show that combined treatment that includes visual ...

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...visual sensory modality, and further, when reading in this novel modality is learned in adult-selectivity for letters over other visual categories without visual experience, its feature tolerance for =-=Dehaene et al., 2010-=-), as well as its high intersubject anatomicalReading with Sounds: Sensory Substitution Selec the Visual Word Form Area Ella Striem-Amit,1 Laurent Cohen,4,5,6 Stanislas Dehaene,7,8 1Department of Medi...

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...een the groups; p < 0.89). The Tel-Aviv Sourasky Medical Center Ethics Committee approved the experimental procedure and written informed consent was obtained from each subject.(Ackroyd et al., 1974; =-=Fine et al., 2003-=-; Ostrovsky et al., 2009). In the congenitally blind, this may be especially true due to the aforementioned task switching (e.g., for language and memory) that may possibly disturb the visual cortex’s...

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...han sensory-specific computation, possibly linking letter shapes to phonology. INTRODUCTION One key question is what causes the apparent selectivity of the VWFA for letters. This hotly debated issue (=-=Price, 2012-=-; Price and Devlin, 2011) was recently resolved to some extent by an integrative view suggesting that the selectivity of VWFA may arise from a conjunction of two properties that make it optimally appr...

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... nonvisual modalities, how dependent is such specialization on the age and amount of training? Reading in the visual modality is usually learned in childhood and improves over many years of practice (=-=Aghababian and Nazir, 2000-=-). Recent evidence shows that plasticity of the ventral visual cortex extends into adolescence and beyond (Dehaene et al., 2010; Golarai et al., 2007). However, does the VWFA show selectivity for scri...

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...ive view suggesting that the selectivity of VWFA may arise from a conjunction of two properties that make it optimally appropriate for reading: (1) efficient reciprocal projections to language areas (=-=Mahon and Caramazza, 2009-=-; Pinel and Dehaene, 2010) and (2) a sensitivity to the visual features that characterize scripts, such as reliance on line junctions (Szwed et al., 2011), foveal position (Hasson et al., 2002), and h...

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...pecific computation, possibly linking letter shapes to phonology. INTRODUCTION One key question is what causes the apparent selectivity of the VWFA for letters. This hotly debated issue (Price, 2012; =-=Price and Devlin, 2011-=-) was recently resolved to some extent by an integrative view suggesting that the selectivity of VWFA may arise from a conjunction of two properties that make it optimally appropriate for reading: (1)...

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...ation in the sighted for letters relative to the baseigure 2A), the congenitally blind group nsive activation of the occipito-temporal (see Figure 2B, as seen previously in blind Burton et al., 2002; =-=Reich et al., 2011-=-). We ditory cortex activation (including A1/ blind for this contrast, given the auditory s the VWFA is characterized not only by Neuronactivation to letters but mostly by its selectivity for letters ...

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...) is specific to the stimulus-selective cortical location (O’Craven and Kanwisher, 2000), in our case in the VWFA, and (2) is significantly less intense than bottom-up perception of the same stimuli (=-=Amedi et al., 2005-=-; O’Craven and Kanwisher, 2000). Moreover, as in sighted subjects, imagery in the blind can generate activation in the primary sensory cortex related to the stimulus modality and location—in our case ...

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...electivity for letters increases over months and years as a result of schooling and reading practice (Ben-Shachar et al., 2011; Brem et al., 2010; Dehaene et al., 2010; Schlaggar andMcCandliss, 2007; =-=Turkeltaub et al., 2003-=-). In fact, in agreement with the present findings, Brem et al. (2010) also showed that preschoolers may develop a VWFA response for visual letters after less than 4 hr of training with a reading comp...

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...r and fewer yet have investigated the existence in the blind of a critical feature of the ventral visual cortex, namely, its regional selectivity for perceptual categories (see Pietrini et al., 2004; =-=Mahon et al., 2009-=-, who explored large-scale preference patterns). The current study now shows same category selectivity for a specific visual category (letters), as seen in the sighted, in the absence of visual experi...

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... scripts; Bitan et al., 2005; Hashimoto and Sakai, 2004; Xue et al., 2006; Xue and Poldrack, 2007), in exilliterate adults (Dehaene et al., 2010), in effortful reading (e.g., reading a degraded text; =-=Cohen et al., 2008-=-), and in children when initially learning to read (Brem et al., 2010). The same contrast (VR versus VC) caused no activation prior to training, when the shapes of the letters were perceivable but not...

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...006; Xue and Poldrack, 2007), in exilliterate adults (Dehaene et al., 2010), in effortful reading (e.g., reading a degraded text; Cohen et al., 2008), and in children when initially learning to read (=-=Brem et al., 2010-=-). The same contrast (VR versus VC) caused no activation prior to training, when the shapes of the letters were perceivable but not yet associated VWFA Visual Selectivity in the Blind Using Soundsto p...

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...t differ considerably from those typical of the sighted brain, such as memory and language in the primary visual cortex (Amedi et al., 2003; Bedny et al., 2011; see reviews in Frasnelli et al., 2011; =-=Merabet and Pascual-Leone, 2010-=-; Striem-Amit et al., 2011). Here we show that when relevant stimuli and tasks are introduced, the ventral visual cortex displays its normal category-specific function, even with stimulation from an u...

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...e selectivity of VWFA may arise from a conjunction of two properties that make it optimally appropriate for reading: (1) efficient reciprocal projections to language areas (Mahon and Caramazza, 2009; =-=Pinel and Dehaene, 2010-=-) and (2) a sensitivity to the visual features that characterize scripts, such as reliance on line junctions (Szwed et al., 2011), foveal position (Hasson et al., 2002), and high spatial frequencies (...

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...nd ‘‘switches tasks’’ and processes nonvisual functions that differ considerably from those typical of the sighted brain, such as memory and language in the primary visual cortex (Amedi et al., 2003; =-=Bedny et al., 2011-=-; see reviews in Frasnelli et al., 2011; Merabet and Pascual-Leone, 2010; Striem-Amit et al., 2011). Here we show that when relevant stimuli and tasks are introduced, the ventral visual cortex display...

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...vision showing higher or more extensive ventral visual activation in sighted adults reading relatively untrained scripts (artificial or foreign scripts; Bitan et al., 2005; Hashimoto and Sakai, 2004; =-=Xue et al., 2006-=-; Xue and Poldrack, 2007), in exilliterate adults (Dehaene et al., 2010), in effortful reading (e.g., reading a degraded text; Cohen et al., 2008), and in children when initially learning to read (Bre...

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...or location identification in the MOG, and even for the general segregation between the ventral and dorsal visual processing streams (Striem-Amit et al., 2012a; for relevant findings in deafness, see =-=Lomber et al., 2010-=-). This suggests that at least some regions may, despite their bottom-up deafferentation, be sufficiently driven by other innately determined constraints (Mahon and Caramazza, 2011) to develop typical...

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... 0.89). The Tel-Aviv Sourasky Medical Center Ethics Committee approved the experimental procedure and written informed consent was obtained from each subject.(Ackroyd et al., 1974; Fine et al., 2003; =-=Ostrovsky et al., 2009-=-). In the congenitally blind, this may be especially true due to the aforementioned task switching (e.g., for language and memory) that may possibly disturb the visual cortex’s original functions and ...

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...her or more extensive ventral visual activation in sighted adults reading relatively untrained scripts (artificial or foreign scripts; Bitan et al., 2005; Hashimoto and Sakai, 2004; Xue et al., 2006; =-=Xue and Poldrack, 2007-=-), in exilliterate adults (Dehaene et al., 2010), in effortful reading (e.g., reading a degraded text; Cohen et al., 2008), and in children when initially learning to read (Brem et al., 2010). The sam...

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...consistent with studies in vision showing higher or more extensive ventral visual activation in sighted adults reading relatively untrained scripts (artificial or foreign scripts; Bitan et al., 2005; =-=Hashimoto and Sakai, 2004-=-; Xue et al., 2006; Xue and Poldrack, 2007), in exilliterate adults (Dehaene et al., 2010), in effortful reading (e.g., reading a degraded text; Cohen et al., 2008), and in children when initially lea...

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...of the development of such circuitry is probably reciprocal anatomical and functional connectivity with higher-order cortical regions involved in the processing of language (Ben-Shachar et al., 2007; =-=Mahon and Caramazza, 2011-=-; Pinel and Dehaene, 2010). In order to examine the underlying functional connectivity in the blind, we investigated the intrinsic (rest state; Biswal et al., 1995) functional connectivity in the blin...

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..., which converts visual images to auditory ‘‘soundscapes’’ using a predetermined consistent algorithm (The vOICe; Meijer, 1992). This enabled the blind to perceive high-resolution visual information (=-=Striem-Amit et al., 2012-=-b) and, in this case, to learn to read, with sounds topographically representing visual images of letters (see Figure 1). Moreover, subjects also learned to recognize soundscapes of other visually com...

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...no age difference between the groups; p < 0.89). The Tel-Aviv Sourasky Medical Center Ethics Committee approved the experimental procedure and written informed consent was obtained from each subject.(=-=Ackroyd et al., 1974-=-; Fine et al., 2003; Ostrovsky et al., 2009). In the congenitally blind, this may be especially true due to the aforementioned task switching (e.g., for language and memory) that may possibly disturb ...

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... ventral network is consistent with studies in vision showing higher or more extensive ventral visual activation in sighted adults reading relatively untrained scripts (artificial or foreign scripts; =-=Bitan et al., 2005-=-; Hashimoto and Sakai, 2004; Xue et al., 2006; Xue and Poldrack, 2007), in exilliterate adults (Dehaene et al., 2010), in effortful reading (e.g., reading a degraded text; Cohen et al., 2008), and in ...

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...cal functions resulting from sensory deprivation. Specifically, they suggest that the VWFA performs a highly flexible task-specific reading-related operation that can be sensory modality independent (=-=Reich et al., 2012-=-). We suggest that this operation is the learned link between letter shapes and their associated phonological content. This category and task selectivity is maintained in the congenital absence of vis...

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...) and (2) a sensitivity to the visual features that characterize scripts, such as reliance on line junctions (Szwed et al., 2011), foveal position (Hasson et al., 2002), and high spatial frequencies (=-=Woodhead et al., 2011-=-). How dependent is VWFA selectivity on such visual sensory features? It was recently shown (Rauschecker et al., 2011) that reading activates VWFA even when the shape of the letters is derived from at...

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... Bioimaging, Neurospin, 91191 Gif sur Yvette, France 2000; Dehaene and Cohen, 2011; Schlaggar and McCandliss, 2007) or left ventral occipito-temporal cortex (vOT; Price, 2012; Price and Devlin, 2011; =-=Wandell, 2011-=-). Extensive research has demonstrated the specialization of this region for the visual representation of letters, its category selectivity manifested in its preference for letters over other types of...

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...cludes visual training Neuron(along with patching of the nonamblyopic eye) can trigger adult plasticity and greatly improve the perceptual outcome, thus reopening the sensitive period for plasticity (=-=Maurer and Hensch, 2012-=-; see also Bavelier et al., 2010). We suggest that in cases of more profound blindness, such rehabilitation may involve, for example, learning to process complex images using SSDs, as done here, or us...

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... brain may be relatively quickly reconfigured to map a novel set of symbols to the same set of phonemes, similar to learning a novel script via vision in a literate person (Hashimoto and Sakai, 2004; =-=Maurer et al., 2010-=-; Xue et al., 2006). Bayesian learning principles (Ernst and Bulthoff, 2004; Tenenbaum et al., 2011) enable the extraction of abstract schemas behind superficially different inputs, including sensory ...

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..., which converts visual images to auditory ‘‘soundscapes’’ using a predetermined consistent algorithm (The vOICe; Meijer, 1992). This enabled the blind to perceive high-resolution visual information (=-=Striem-Amit et al., 2012-=-b) and, in this case, to learn to read, with sounds topographically representing visual images of letters (see Figure 1). Moreover, subjects also learned to recognize soundscapes of other visually com...

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...ry and task selectivity is maintained in the congenital absence of vision, despite otherwise extreme plasticity for other functions and input types shown previously in the blind brain (see reviews in =-=Frasnelli et al., 2011-=-; Merabetand Pascual-Leone, 2010; Striem-Amit et al., 2011). This implies the presence of innately determined constraints (Striem-Amit et al., 2012a) on the emergence of VWFA selectivity for reading. ...

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...d et al., 2011), foveal position (Hasson et al., 2002), and high spatial frequencies (Woodhead et al., 2011). How dependent is VWFA selectivity on such visual sensory features? It was recently shown (=-=Rauschecker et al., 2011-=-) that reading activates VWFA even when the shape of the letters is derived from atypical features such as themovement or the luminance of sets of dots. This suggests that within vision there is remar...

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...in a variety of visual dimensions, including uppercase/ lowercase (Dehaene et al., 2001), printed/handwriting style (Qiao et al., 2010), location in the visual field (Cohen et al., 2002; but also see =-=Rauschecker et al., 2012-=-, who recently challenged this to some extent), or type of shape-defining visual feature (Rauschecker et al., 2011). A key finding in the present study is that this feature tolerance extends beyond th...

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...absence of vision, despite otherwise extreme plasticity for other functions and input types shown previously in the blind brain (see reviews in Frasnelli et al., 2011; Merabetand Pascual-Leone, 2010; =-=Striem-Amit et al., 2011-=-). This implies the presence of innately determined constraints (Striem-Amit et al., 2012a) on the emergence of VWFA selectivity for reading. Furthermore, in the context of visual rehabilitation, this...

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...s of stimulus type and input modality. Supporting this dissociation, we found no activation for SSD letters in the auditory parallel of the VWFA, the auditory word form area in the left anterior STG (=-=DeWitt and Rauschecker, 2012-=-; see Figures 2E, 2F, and 3; but functional connectivity between these two areaswas found, see below), although vOICe letters are conveyed through audition. Furthermore, our results cannot be readily ...

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...erent visual object categories (such as faces, houses, and objects; Cohen and Dehaene, 2004; Dehaene and Cohen, 2011; Dehaene et al., 2010; Hasson et al., 2002; Puce et al., 1996; Szwed et al., 2011; =-=Tsapkini and Rapp, 2010-=-), similar to the preferential activation of the neighboring regions for faces, scenes, objects, and body shapes (Kanwisher, 2010). Can full category selectivity in the VWFA also emerge without visual...