Emotion words constitute a special class of verbal stimuli which can quickly activate the limbic system outside the left-hemisphere language network. Such fast response to emotion words may arise independently of the left occipitotemporal area involved in visual word-form analysis and rely on a distinct amygdala-dependent emotion circuit involved in fearful face processing. Using a hemifield priming paradigm with fMRI, we explored how the left and right amygdala systems interact with the reading network during emotion word processing. On each trial, participants viewed a centrally presented target which was preceded by a masked prime flashed either to the left or right visual field. Primes and targets, each denoting negative or positive nouns, could be either affectively congruent or incongruent with each other. We observed that affective congruency produced parallel changes in neural priming between the left frontal and parietotemporal regions and the bilateral amygdala. However, we also found that the left, but not right, amygdala exhibited significant change in functional connectivity with the neural components of reading as a function of affective congruency. Collectively, these results suggest that emotion words activate the bilateral amygdala during early stages of emotion word processing, whereas only the left amygdala exerts a long-distance regulatory influence over the reading network via its strong within-hemisphere connectivity.

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Do the neural circuits for reading vary across culture? Reading of visually complex writing systems such as Chinese has been proposed to rely on areas outside the classical left-hemisphere network for alphabetic reading. Here, however, we show that, once potential confounds in cross-cultural comparisons are controlled for by presenting handwritten stimuli to both Chinese and French readers, the underlying network for visual word recognition may be more universal than previously suspected. Using functional magnetic resonance imaging in a semantic task with words written in cursive font, we demonstrate that two universal circuits, a shape recognition system (reading by eye) and a gesture recognition system (reading by hand), are similarly activated and show identical patterns of activation and repetition priming in the two language groups. These activations cover most of the brain regions previously associated with culture-specific tuning. Our results point to an extended reading network that invariably comprises the occipitotemporal visual wordform system, which is sensitive to well-formed static letter strings, and a distinct left premotor region, Exner's area, which is sensitive to the forward or backward direction with which cursive letters are dynamically presented. These findings suggest that cultural effects in reading merely modulate a fixed set of invariant macroscopic brain circuits, depending on surface features of orthographies.cross-cultural invariance | functional magnetic resonance imaging | neuronal recycling | masked priming A pproximately one-fifth of today's world population is still unable to read and write (1). The acquisition of written language does not rely on a specific innate ability but is an education-dependent skill resulting from the learning of mapping rules linking written codes, speech sounds, and word meanings. At the neural level, literacy acquisition imposes various structural and functional changes to the human brain, particularly in the visual cortex where responses become attuned to a specific script, but also in other areas of the temporal and parietal lobes (2, 3).The issue that we raise here is whether those changes vary considerably from one culture to another or whether they consistently engage a universal and largely invariant brain network. Past research indicated that skilled reading universally relies on a posterior left-hemisphere network, including the lateral occipitotemporal visual word-form area (VWFA) for perceptual analysis of written words (4, 5), the inferior parietal and superior temporal cortices involved in print-to-sound translation (6, 7), and lateral temporal cortices involved in access to word meaning (8-10). Reading of alphabetic scripts engages this multicomponent system with only small cultural variation depending on the degree of transparency (11) and grain size (12) of the orthographic system. However, beyond this shared left posterior network, several previous studies with normal (8,13,14) and dyslexic (15, 16) Chinese participants defended a cu...

Abstract& Recent evidence has suggested that the human occipitotemporal region comprises several subregions, each sensitive to a distinct processing level of visual words. To further explore the functional architecture of visual word recognition, we employed a subliminal priming method with functional magnetic resonance imaging (fMRI) during semantic judgments of words presented in two different Japanese scripts, Kanji and Kana. Each target word was preceded by a subliminal presentation of either the same or a different word, and in the same or a different script. Behaviorally, word repetition produced significant priming regardless of whether the words were presented in the same or different script. At the neural level, this cross-script priming was associated with repetition suppression in the left inferior temporal cortex anterior and dorsal to the visual word form area hypothesized for alphabetical writing systems, suggesting that cross-script convergence occurred at a semantic level. fMRI also evidenced a shared visual occipito-temporal activation for words in the two scripts, with slightly more mesial and right-predominant activation for Kanji and with greater occipital activation for Kana. These results thus allow us to separate script-specific and script-independent regions in the posterior temporal lobe, while demonstrating that both can be activated subliminally. &

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Humans and primates can quickly recognize mirror images of previously exposed pictures. This spontaneous mirror invariance, though advantageous for visual recognition, makes it difficult to distinguish the orientation of letters (e.g. to differentiate a "b" from a "d"), and may result in classical mirror reading and writing errors in preschoolers. Mirror invariance must therefore be overcome during reading acquisition. The Visual Word Form Area (VWFA), a region in the ventral stream that develops with reading expertise, was previously shown to discriminate words from their mirror images in literate adults. Here we investigate whether this region underlies mirror-image discrimination at the most elementary level of the orthographic code, the single-letter level. Using an fMRI priming paradigm, we demonstrate that the VWFA distinguishes the left-right orientation of single letters in skilled readers, and yet exhibits mirror invariance for simple pictures of matched complexity. These results clarify how letter shapes, after reading acquisition, escape the process of mirror invariance which is a basic property of the ventral visual shape recognition pathway.

1. We examined single-neuronal activity in the temporal pole of monkeys, including the anterior ventromedial temporal (VMT) cortex (the temporopolar cortex, area 36, area 35, and the entorhinal cortex) and the anterior inferotemporal (IT) cortex, during a visual recognition memory task. In the task, a trial began when the monkey pressed a lever. After a waiting period, a visual sample stimulus (S) was presented one to four times on a monitor with an interstimulus delay. Thereafter, a new stimulus (R) was presented. The monkeys were trained to remember S during the delay period and to release the lever in response to R. Colored photographs of natural objects were used as visual stimuli. 2. About 70% of the recorded neurons (225 of 311) responded to at least one of the Ss tested. Thirty percent of these neurons (68 of 225) continued to fire during the subsequent delay periods. In 75% of these neurons (51 of 68), the firing during the delay period strongly correlated with the response to S. 3. The discharge rate during the delay period did not correlate with the monkey's eye movements, pressing or releasing of the lever, or the reaction time. 4. If the monkey erroneously released the lever in response to S or during the delay period, the firing disappeared after the erroneous lever release. If the monkey failed to release the lever in response to R, the firing persisted even after R was withdrawn. The discharge rate in incorrect trials was comparable with that in correct trials. The neurons were considered to fire for as long as the memory of S was necessary. 5. Firing persisted even when an achromatic version or half (even a portion) of S was presented, indicating that the color, a particular portion, or the entire shape of S was not always necessary to elicit firing. 6. An S that elicited firing during the delay period invariably elicited a visual response. Neurons that fired during the delay period showed a higher stimulus selectivity than other visually responsive neurons in the anterior VMT cortex. Thus neurons that fire during the delay period represent a subgroup of visually responsive neurons that are selectively tuned to a certain stimulus. 7. More neurons fired during the delay period in the anterior VMT cortex than in the anterior IT cortex. 8. We conclude that firing during the delay period by neurons in the temporal pole reflects the short-term storage of visual information regarding a particular S.

Learning to read requires the acquisition of an efficient visual procedure for quickly recognizing fine print. Thus, reading practice could induce a perceptual learning effect in early vision. Using functional magnetic resonance imaging (fMRI) in literate and illiterate adults, we previously demonstrated an impact of reading acquisition on both high-and low-level occipitotemporal visual areas, but could not resolve the time course of these effects. To clarify whether literacy affects early vs. late stages of visual processing, we measured event-related potentials to various categories of visual stimuli in healthy adults with variable levels of literacy, including completely illiterate subjects, early-schooled literate subjects, and subjects who learned to read in adulthood (ex-illiterates). The stimuli included written letter strings forming pseudowords, on which literacy is expected to have a major impact, as well as faces, houses, tools, checkerboards, and false fonts. To evaluate the precision with which these stimuli were encoded, we studied repetition effects by presenting the stimuli in pairs composed of repeated, mirrored, or unrelated pictures from the same category. The results indicate that reading ability is correlated with a broad enhancement of early visual processing, including increased repetition suppression, suggesting better exemplar discrimination, and increased mirror discrimination, as early as ∼100-150 ms in the left occipitotemporal region. These effects were found with letter strings and false fonts, but also were partially generalized to other visual categories. Thus, learning to read affects the magnitude, precision, and invariance of early visual processing.reading | brain plasticity | education R eading is a cultural activity in which contemporary humans have considerable training. Fluently accessing the sounds and meanings of written words requires very fast and efficient visual recognition of letter strings, at rates exceeding 100 words/ min. Neuroimaging studies have begun to show how learning to read modulates the functioning of the visual system, from early retinotopic areas (1, 2) to extrastriate occipital and temporal cortex (1, 3, 4). In particular, a restricted region of the left occipitotemporal cortex, the visual word form area (VWFA), is robustly activated when orthographic stimuli are presented to literate subjects.This VWFA activation is reproducible across participants and writing systems (5, 6), even when orthographic stimuli are flashed unconsciously (7). Orthographic processing in the VWFA is thought to be very fast, peaking at ∼170-200 ms (8-10), and is colateralized to the dominant hemisphere for language (11,12). Reading practice enhances activation of the VWFA (1, 13, 14), even in dyslexic children (15). Reading also modulates nonvisual circuits, such as the spoken language network (1,14,16,17).In addition to these positive effects of learning to read, the theory of neuronal recycling (18) proposes that literacy acquisition also may have a negative "unlearning" e...