Elementary arithmetic requires a complex interplay between several brain regions. The classical view, arising from fMRI, is that the intraparietal sulcus (IPS) and the superior parietal lobe (SPL) are the main hubs for arithmetic calculations. However, recent studies using intracranial electroencephalography have discovered a specific site, within the posterior inferior temporal cortex (pITG), that activates during visual perception of numerals, with widespread adjacent responses when numerals are used in calculation. Here, we reexamined the contribution of the IPS, SPL, and pITG to arithmetic by recording intracranial electroencephalography signals while participants solved addition problems. Behavioral results showed a classical problem size effect: RTs increased with the size of the operands. We then examined how high-frequency broadband (HFB) activity is modulated by problem size. As expected from previous fMRI findings, we showed that the total HFB activity in IPS and SPL sites increased with problem size. More surprisingly, pITG sites showed an initial burst of HFB activity that decreased as the operands got larger, yet with a constant integral over the whole trial, thus making these signals invisible to slow fMRI. Although parietal sites appear to have a more sustained function in arithmetic computations, the pITG may have a role of early identification of the problem difficulty, beyond merely digit recognition. Our results ask for a reevaluation of the current models of numerical cognition and reveal that the ventral temporal cortex contains regions specifically engaged in mathematical processing.

How does reading expertise change the visual system? Here, we explored whether the visual system could develop dedicated perceptual mechanisms in early and intermediate visual cortex under the pressure for fast processing that is particularly strong in reading. We compared fMRI activations in Chinese participants with limited knowledge of French and in French participants with no knowledge of Chinese, exploiting these doubly dissociated reading skills as a tool to study the neural correlates of visual expertise. All participants viewed the same stimuli: words in both languages and matched visual controls, presented at a fast rate comparable with fluent reading. In the Visual Word Form Area, all participants showed enhanced responses to their known scripts. However, group differences were found in occipital cortex. In French readers reading French, activations were enhanced in left-hemisphere visual area V1, with the strongest differences between French words and their controls found at the central and horizontal meridian representations. Chinese participants, who were not expert French readers, did not show these early visual activations. In contrast, Chinese readers reading Chinese showed enhanced activations in intermediate visual areas V3v/hV4, absent in French participants. Together with our previous findings [Szwed, M., Dehaene, S., Kleinschmidt, A., Eger, E., Valabregue, R., Amadon, A., et al. Specialization for written words over objects in the visual cortex. Neuroimage, 56, 330–344, 2011], our results suggest that the effects of extensive practice can be found at the lowest levels of the visual system. They also reveal their cross-script variability: Alphabetic reading involves enhanced engagement of central and right meridian V1 representations that are particularly used in left-to-right reading, whereas Chinese characters put greater emphasis on intermediate visual areas.

When two displays are presented in close temporal succession at the same location, how does the brain assign them to one versus two conscious percepts? We investigate this issue using a novel reading paradigm in which the odd and even letters of a string are presented alternatively at a variable rate. The results reveal a window of temporal integration during reading, with a nonlinear boundary around ∼80 msec of presentation duration. Below this limit, the oscillating stimulus is easily fused into a single percept, with all characteristics of normal reading. Above this limit, reading times are severely slowed and suffer from a word-length effect. ERPs indicate that, even at the fastest frequency, the oscillating stimulus elicits synchronous oscillations in posterior visual cortices, while late ERP components sensitive to lexical status vanish beyond the fusion threshold. Thus, the fusion/segregation dilemma is not resolved by retinal or subcortical filtering, but at cortical level by at most 300 msec. The results argue against theories of visual word recognition and letter binding that rely on temporal synchrony or other fine temporal codes.

Language and arithmetic are both lateralized to the left hemisphere in the majority of right-handed adults. Yet, does this similar lateralization reflect a single overall constraint of brain organization, such an overall “dominance” of the left hemisphere for all linguistic and symbolic operations? Is it related to the lateralization of specific cerebral subregions? Or is it merely coincidental? To shed light on this issue, we performed a “colateralization analysis” over 209 healthy subjects: We investigated whether normal variations in the degree of left hemispheric asymmetry in areas involved in sentence listening and reading are mirrored in the asymmetry of areas involved in mental arithmetic. Within the language network, a region-of-interest analysis disclosed partially dissociated patterns of lateralization, inconsistent with an overall “dominance” model. Only two of these areas presented a lateralization during sentence listening and reading which correlated strongly with the lateralization of two regions active during calculation. Specifically, the profile of asymmetry in the posterior superior temporal sulcus during sentence processing covaried with the asymmetry of calculation-induced activation in the intraparietal sulcus, and a similar colateralization linked the middle frontal gyrus with the superior posterior parietal lobule. Given recent neuroimaging results suggesting a late emergence of hemispheric asymmetries for symbolic arithmetic during childhood, we speculate that these colateralizations might constitute developmental traces of how the acquisition of linguistic symbols affects the cerebral organization of the arithmetic network.

As literate adults, we appreciate numerical values as abstract entities that can be represented by a numeral, a word, a number of lines on a scorecard, or a sequence of chimes from a clock. This abstract, notation-independent appreciation of numbers develops gradually over the first several years of life. Here, using functional magnetic resonance imaging, we examine the brain mechanisms that 6- and 7-year-old children and adults recruit to solve numerical comparisons across different notation systems. The data reveal that when young children compare numerical values in symbolic and nonsymbolic notations, they invoke the same network of brain regions as adults including occipito-temporal and parietal cortex. However, children also recruit inferior frontal cortex during these numerical tasks to a much greater degree than adults. Our data lend additional support to an emerging consensus from adult neuroimaging, nonhuman primate neurophysiology, and computational modeling studies that a core neural system integrates notation-independent numerical representations throughout development but, early in development, higher-order brain mechanisms mediate this process.

Priming effects have been well documented in behavioral psycholinguistics experiments: The processing of a word or a sentence is typically facilitated when it shares lexico-semantic or syntactic features with a previously encountered stimulus. Here, we used fMRI priming to investigate which brain areas show adaptation to the repetition of a sentence's content or syntax. Participants read or listened to sentences organized in series which could or not share similar syntactic constructions and/or lexico-semantic content. The repetition of lexico-semantic content yielded adaptation in most of the temporal and frontal sentence processing network, both in the visual and the auditory modalities, even when the same lexico-semantic content was expressed using variable syntactic constructions. No fMRI adaptation effect was observed when the same syntactic construction was repeated. Yet behavioral priming was observed at both syntactic and semantic levels in a separate experiment where participants detected sentence endings. We discuss a number of possible explanations for the absence of syntactic priming in the fMRI experiments, including the possibility that the conglomerate of syntactic properties defining “a construction” is not an actual object assembled during parsing.

The visual system of literate adults develops a remarkable perceptual expertise for printed words. To delineate the aspects of this competence intrinsic to the occipitotemporal “what” pathway, we studied a patient with bilateral lesions of the occipitoparietal “where” pathway. Depending on critical geometric features of the display (rotation angle, letter spacing, mirror reversal, etc.), she switched from a good performance, when her intact ventral pathway was sufficient to encode words, to severely impaired reading, when her parietal lesions prevented the use of alternative reading strategies as a result of spatial and attentional impairments. In particular, reading was disrupted (a) by rotating word by more than 50°, providing an approximation of the invariance range for words encoding in the ventral pathway; (b) by separating letters with double spaces, revealing the limits of letter grouping into perceptual wholes; (c) by mirror-reversing words, showing that words escape the default mirror-invariant representation of visual objects in the ventral pathway. Moreover, because of her parietal lesions, she was unable to discriminate mirror images of common objects, although she was excellent with reversible pseudowords, confirming that the breaking of mirror symmetry was intrinsic to the occipitotemporal cortex. Thus, charting the display conditions associated with preserved or impaired performance allowed us to infer properties of word coding in the normal ventral pathway and to delineate the roles of the parietal lobes in single-word recognition.

Recent evidence has suggested that the human occipito-temporal 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.

Measuring reaction times (RTs) using the additive-factors method provides information about the sequence of processing stages in a cognitive task. Here, I describe how the simultaneous recording of event-related potentials (ERPs) in the same task can provide complementary information that cannot be obtained using RTs alone. Most notably, ERP data can reveal the absolute activation time and the coarse brain localization of processing stages. RTs and ERPs can also be used to cross-validate a serial-stage model. These notions were applied to a study of the temporal unfolding of brain activations in a number comparison task. ERPs were recorded from 64 scalp electrodes while normal subjects classified numbers as larger or smaller than 5. Specific scalp signatures and timing data were obtained for stages of word and digit identification, magnitude comparison, response programming, and error capture and correction. The observed localizations were compatible with previous neuropsychological and brain imaging data and provided new insights into the cerebral lateralization and timing of number processing.

Despite their lack of language, human infants and several animal species possess some elementary abilities for numerical processing. These include the ability to recognize that a given numerosity is being presented visually or auditorily, and, at a later stage of development, the ability to compare two nume-rosities and to decide which is larger. We propose a model for the development of these abilities in a formal neuronal network. Initially, the model is equipped only with unordered numerosity detectors. It can therefore detect the numerosity of an input set and can be conditioned to react accordingly. In a later stage, the addition of a short-term memory network is shown to be sufficient for number comparison abilities to develop. Our computer simulations account for several phenomena in the numerical domain, including the distance effect and Fechner's law for numbers. They also demonstrate that infants' numerosity detection abilities may be explained without assuming that infants can count. The neurobiological bases of the critical components of the model are discussed.

Both psychologists and neurobiologists have used delayed response (DR), AB, and delayed matching-to-sample (DMS) tasks as tools to study the functions of prefrontal cortex in primates and humans. We describe a simulation model that relates behavioral and electrophysiological-data relevant to these tasks into a minimal neural network. The inputs to the network are two visual objects and a positive or negative reinforcement signal. As the output, the network orients toward one of the two objects. We subdivide the architecture of the network into two levels, both of which embody constraints from neuroanatomy in a simplified form. Level 1 consists of a sensory-motor loop with modifiable synaptic weights and provides a capacity for grasping. Level 2 contains memory and rule-coding units and modulates the lower level 1. When level 1 only is simulated, the network fails to learn the tasks. The errors made by the network resemble those of young monkeys, infants, or adults with prefrontal lesions. In particular, the systematic AB error can be reproduced. With level 2 on top of level 1, the network acquires systematic rules of behavior by mere reinforcement and rapidly adapts to changes in the reinforcement schedule. Learning takes place by selection among a repertoire of possible rules. The properties of the model are discussed in terms of actual behavioral and physiological data, and several critical experimental predictions are presented. In particular, we address the issues of prefrontal functions, “systematicity” in neural networks, and “mental Darwinism.”