Biomedical subjects
Publications and source records attributed to S Dehaene.
Recent computational models, or mathematical realizations of neurobiological theories, are providing insights into the organization and workings of the association cortex. Such models concern the construction of cortical maps, the neural basis of cognitive functions such as visual perception, reward-motivated learning and some aspects of consciousness.
Neuropsychological studies have revealed different subtypes of dyscalculia, including dissociations between exact calculation and approximation abilities, and an impact of number size on performance. To understand the origins of these effects, we measured cerebral activity with functional MRI at 3 Tesla and event-related potentials while healthy volunteers performed exact and approximate calculation tasks with small and large numbers. Bilateral intraparietal, precentral, dorsolateral and superior prefrontal regions showed greater activation during approximation, while the left inferior prefrontal cortex and the bilateral angular regions were more activated during exact calculation. Increasing number size during exact calculation led to increased activation in the same bilateral intraparietal regions as during approximation, as well the left inferior and superior frontal gyri. Event-related potentials gave access to the temporal dynamics of calculation processes, showing that effects of task and of number size could be found as early as 200-300 ms following problem presentation. Altogether, the results reveal two cerebral networks for number processing. Rote arithmetic operations with small numbers have a greater reliance on left-lateralized regions, presumably encoding numbers in verbal format. Approximation and exact calculation with large numbers, however, put heavier emphasis on the left and right parietal cortices, which may encode numbers in a non-verbal quantity format. Subtypes of dyscalculia can be explained by lesions disproportionately affecting only one of these networks.
A standard model of word reading postulates that visual information is initially processed by occipitotemporal areas contralateral to the stimulated hemifield, from whence it is subsequently transferred to the visual word form (VWF) system, a left inferior temporal region specifically devoted to the processing of letter strings. For stimuli displayed in the left visual field, this transfer proceeds from the right to the left hemisphere through the posterior portion of the corpus callosum. In order to characterize the spatial and temporal organization of these processes, reading tasks with split-field presentation were performed by five control subjects and by two patients suffering from left hemialexia following posterior callosal lesions. The subjects' responses were studied using behavioural measures and functional brain imaging techniques, providing both high spatial resolution (functional MRI, fMRI) and high temporal resolution (high-density event-related potentials, ERPs). Early visual processing was revealed as activations contralateral to stimulation, located by fMRI in the inferior occipitotemporal region and presumably coincident with area V4. A negative wave occurring 150-160 ms post-stimulus, also strictly contralateral to stimulation, was recorded over posterior electrodes. In contrast with these hemifield-dependent effects, the VWF system was revealed as a strictly left-hemispheric activation which, in control subjects, was identical for stimuli presented in the left or in the right hemifield and was located in the middle portion of the left fusiform gyrus. The electrical signature of the VWF system consisted of a unilateral sharp negativity, recorded 180-200 ms post-stimulus over left inferior temporal electrodes. In callosal patients, due to the inability of visual information to pass across the posterior part of the corpus callosum, the VWF system was activated only by stimuli presented in the right visual field. Similarly, a significant influence of the word/non-word status on ERPs recorded over the left hemisphere was discernible for either hemifield in controls, while it affected only right-hemifield stimuli in callosal patients. These findings provide direct support for the main components of the classical model of reading and help specify their timing and cerebral substrates.
Cerebral activity during number comparison was studied with functional magnetic resonance imaging using an event-related design. We identified an extended network of task-related areas that showed a phasic activation following each trial, including anterior cingulate, bilateral sensorimotor areas, inferior occipito-temporal cortices, posterior parietal cortices, inferior and dorsolateral prefrontal cortices, and thalami. We then tested which of these areas were affected by number notation, numerical distance and response side, three variables that specifically target processes of visual identification, quantity manipulation and motor response in a serial-stage model of the number comparison task. Our results confirm the role of the right fusiform gyrus in digit identification processes, and of the inferior parietal lobule in the internal manipulation of numerical quantities.
Does the human capacity for mathematical intuition depend on linguistic competence or on visuo-spatial representations? A series of behavioral and brain-imaging experiments provides evidence for both sources. Exact arithmetic is acquired in a language-specific format, transfers poorly to a different language or to novel facts, and recruits networks involved in word-association processes. In contrast, approximate arithmetic shows language independence, relies on a sense of numerical magnitudes, and recruits bilateral areas of the parietal lobes involved in visuo-spatial processing. Mathematical intuition may emerge from the interplay of these brain systems.
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We measured cerebral activation with functional magnetic resonance imaging at 3 Tesla while eight healthy volunteers performed various number processing tasks known to be dissociable in brain-lesioned patients: naming, comparing, multiplying, or subtracting single digits. The results revealed the activation of a circuit comprising bilateral intraparietal, prefrontal, and anterior cingulate components. The extension and lateralization of this circuit was modulated by task demands. The intraparietal and prefrontal activation was more important in the right hemisphere during the comparison task and in the left hemisphere during the multiplication task and was intensely bilateral during the subtraction task. Thus, partially distinct cerebral circuits with the dorsal parietal pathway underlie distinct arithmetic operations.
A minimal hypothesis is proposed concerning the brain processes underlying effortful tasks. It distinguishes two main computational spaces: a unique global workspace composed of distributed and heavily interconnected neurons with long-range axons, and a set of specialized and modular perceptual, motor, memory, evaluative, and attentional processors. Workspace neurons are mobilized in effortful tasks for which the specialized processors do not suffice. They selectively mobilize or suppress, through descending connections, the contribution of specific processor neurons. In the course of task performance, workspace neurons become spontaneously coactivated, forming discrete though variable spatio-temporal patterns subject to modulation by vigilance signals and to selection by reward signals. A computer simulation of the Stroop task shows workspace activation to increase during acquisition of a novel task, effortful execution, and after errors. We outline predictions for spatio-temporal activation patterns during brain imaging, particularly about the contribution of dorsolateral prefrontal cortex and anterior cingulate to the workspace.
Visual words that are masked and presented so briefly that they cannot be seen may nevertheless facilitate the subsequent processing of related words, a phenomenon called masked priming. It has been debated whether masked primes can activate cognitive processes without gaining access to consciousness. Here we use a combination of behavioural and brain-imaging techniques to estimate the depth of processing of masked numerical primes. Our results indicate that masked stimuli have a measurable influence on electrical and haemodynamic measures of brain activity. When subjects engage in an overt semantic comparison task with a clearly visible target numeral, measures of covert motor activity indicate that they also unconsciously apply the task instructions to an unseen masked numeral. A stream of perceptual, semantic and motor processes can therefore occur without awareness.
The introduction, in the late sixties, of the concepts and methods of molecular biology to the study of the nervous system had a profound impact on the field, primarily through the identification of its basic molecular components. These structures include, for example, the elementary units of the synapse: neurotransmitters, neuropeptides and their receptors, but also ionic channels, intracellular second messengers and the relevant enzymes, cell surface adhesion molecules, or growth and trophic factors [21,78,81, 52,79]. Attempts to establish appropriate causal relationships between these molecular components, the actual organisation of neural networks, and a defined behavior, nevertheless, still must overcome many difficulties. A first problem is the recognition of the minimum levels of organisation, from the molecular, cellular, or multicellular (circuit) to the higher cognitive levels, that determine the given physiological and/or behavioral performance under investigation. A common difficulty (and potential source of errors of interpretation) is to relate a cognitive function to a network organization which does not possess the required structural complexity and vice-versa. Another problem is to distinguish, among the components of the system, those which are actually necessary and those which, taken together, suffice for a given behavior to take place. Identification of such a minimal set of building blocks may receive decisive insights from the elaboration of neurally plausible formal models that bring together, within a single and coherent 'artificial organism', the neuronal network, the circulating activity, and the behavior they determine (see [42,43,45,72,30]). In this communication, we shall attempt, still in a preliminary fashion, to bring together: (1) our recent knowledge on the molecular biology of brain nicotinic receptors (nAChRs) and their allosteric properties and (2) integrated behaviors, such as cognitive learning, investigated for instance with delayed-response or passive avoidance tasks that are likely to involve nAChRs in particular at the level of reinforcement (or reward) mechanisms (see [18,29,135]).
There is evidence to suggest that animals, young infants and adult humans possess a biologically determined, domain-specific representation of number and of elementary arithmetic operations. Behavioral studies in infants and animals reveal number perception, discrimination and elementary calculation abilities in non-verbal organisms. Lesion and brain-imaging studies in humans indicate that a specific neural substrate, located in the left and right intraparietal area, is associated with knowledge of numbers and their relations ('number sense'). The number domain is a prime example where strong evidence points to an evolutionary endowment of abstract domain-specific knowledge in the brain because there are parallels between number processing in animals and humans.The numerical distance effect, which refers to the finding that the ability to discriminate between two numbers improves as the numerical distance between them increases, has been demonstrated in humans and animals, as has the number size effect,which refers to the finding that for equal numerical distance,discrimination of two numbers worsens as their numerical size increases.
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Functional imaging methods show differences in the pattern of cerebral activation associated with the subject's native language (L1) compared with a second language (L2). In a recent PET investigation on bilingualism we showed that auditory processing of stories in L1 (Italian) engages the temporal lobes and temporoparietal cortex more extensively than L2 (English). However, in that study the Italian subjects learned L2 late and attained a fair, but not an excellent command of this language (low proficiency, late acquisition bilinguals). Thus, the different patterns of activation could be ascribed either to age of acquisition or to proficiency level. In the current study we use a similar paradigm to evaluate the effect of early and late acquisition of L2 in highly proficient bilinguals. We studied a group of Italian-English bilinguals who acquired L2 after the age of 10 years (high proficiency, late acquisition bilinguals) and a group of Spanish-Catalan bilinguals who acquired L2 before the age of 4 years (high proficiency, early acquisition bilinguals). The differing cortical responses we had observed when low proficiency volunteers listened to stories in L1 and L2 were not found in either of the high proficiency groups in this study. Several brain areas, similar to those observed for L1 in low proficiency bilinguals, were activated by L2. These findings suggest that, at least for pairs of L1 and L2 languages that are fairly close, attained proficiency is more important than age of acquisition as a determinant of the cortical representation of L2.
Perseveration consists of the inappropriate repetition of a preceding behaviour when a new adapted response is expected. We have developed statistical tools that make it possible to reveal such perseverations, assess their significance and study their finer characteristics, such as their temporal course and impaired processing level. This approach is illustrated and evaluated through analyses of naming errors produced by three patients with impairments affecting different stages of the processing chain leading from visual perception to speech production. These examples of perseverations include the intrusion not only of whole words (patient R.A.V.) but also of isolated phonemes (patient D.U.M.) or of visual features (patient Y.M.) from previous trials. In all cases, the probability that an error is a perseveration from a previous trial is an exponentially decreasing function of the lag between the two trials considered. This suggests that perseverations reflect a decaying internal variable, such as an internal level of activation of previous utterances. Based on these empirical results, we put forward a tentative mechanism for the generation of perseverations: whenever a given processing level is deprived of its normal input, persistent activity inherited from previous trials is no longer overcome by current input, and is revealed in the form of perseverations.
Functional magnetic resonance imaging was used to assess inter-subject variability in the cortical representation of language comprehension processes. Moderately fluent French-English bilinguals were scanned while they listened to stories in their first language (L1 = French) or in a second language (L2 = English) acquired at school after the age of seven. In all subjects, listening to L1 always activated a similar set of areas in the left temporal lobe, clustered along the left superior temporal sulcus. Listening to L2, however, activated a highly variable network of left and right temporal and frontal areas, sometimes restricted only to right-hemispheric regions. These results support the hypothesis that first language acquisition relies on a dedicated left-hemispheric cerebral network, while late second language acquisition is not necessarily associated with a reproducible biological substrate. The postulated contribution of the right hemisphere to L2 comprehension is found to hold only on average, individual subjects varying from complete right lateralization to standard left lateralization for L2.
Planning a goal-directed sequence of behavior is a higher function of the human brain that relies on the integrity of prefrontal cortical areas. In the Tower of London test, a puzzle in which beads sliding on pegs must be moved to match a designated goal configuration, patients with lesioned prefrontal cortex show deficits in planning a goal-directed sequence of moves. We propose a neuronal network model of sequence planning that passes this test and, when lesioned, fails in a way that mimics prefrontal patients' behavior. Our model comprises a descending planning system with hierarchically organized plan, operation, and gesture levels, and an ascending evaluative system that analyzes the problem and computes internal reward signals that index the correct/erroneous status of the plan. Multiple parallel pathways connecting the evaluative and planning systems amend the plan and adapt it to the current problem. The model illustrates how specialized hierarchically organized neuronal assemblies may collectively emulate central executive or supervisory functions of the human brain.
We describe two acalculic patients, one with a left subcortical lesion and the other with a right inferior parietal lesion and Gerstmann's syndrome. Both suffered from "pure anarithmetia": they could read arabic numerals and write them to dictation, but experienced a pronounced calculation deficit. On closer analysis, however, distinct deficits were found. The subcortical case suffered from a selective deficit of rote verbal knowledge, including but not limited to arithmetic tables, while her semantic knowledge of numerical quantities was intact. Conversely the inferior parietal case suffered from a category-specific impairment of quantitative numerical knowledge, particularly salient in subtraction and number bissection tasks, with preserved knowledge of rote arithmetic facts. This double dissociation suggests that numerical knowledge is processed in different formats within distinct cerebral pathways. We suggest that a left subcortical network contributes to the storage and retrieval of rote verbal arithmetic facts, while a bilateral inferior parietal network is dedicated to the mental manipulation of numerical quantities.