Verb generation for presurgical mapping: Gaining specificity.

Verb generation is among the most frequently used tasks in presurgical mapping. Because this task involves many processes, the overall brain effects are not specific. While it is necessary to identify the whole network involving noun comprehension or semantic retrieval and lexical selection to produce the verb, isolation of those components is also crucial. Here, we present data from four patients undergoing presurgical brain mapping. The study implied a reanalysis of magnetoencephalography data with a recategorization of the used items. It aimed to extract the task component that relies on the inferior frontal gyrus (IFG). The task could be applied with higher specificity when targeting frontal areas. For that, we based item classification on the selection demands imposed by the noun. It is a robust finding that the IFG carries out this selection and that a quantitative index can be calculated for each noun, which depends on the selection effort (Proceedings of the National Academy of Sciences of the United States of America, 1997; 94(26):14792-14797, Proceedings of the National Academy of Sciences of the United States of America, 1998; 95(26):15855-15860). Data showed focality and specificity, with a correlation between this derived index and source activations in the inferior frontal gyrus for all patients. Strikingly, we detected when the right-hemisphere homologue area was involved in the selection process in two patients showing reorganization or language right lateralization. The present data are a step towards a dissection of broad specific tasks frequently used in presurgical protocols.

The brain lateralization and development of math functions: progress since Sperry, 1974.

In 1974, Roger Sperry, based on his seminal studies on the split-brain condition, concluded that math was almost exclusively sustained by the language dominant left hemisphere. The right hemisphere could perform additions up to sums less than 20, the only exception to a complete left hemisphere dominance. Studies on lateralized focal lesions came to a similar conclusion, except for written complex calculation, where spatial abilities are needed to display digits in the right location according to the specific requirements of calculation procedures. Fifty years later, the contribution of new theoretical and instrumental tools lead to a much more complex picture, whereby, while left hemisphere dominance for math in the right-handed is confirmed for most functions, several math related tasks seem to be carried out in the right hemisphere. The developmental trajectory in the lateralization of math functions has also been clarified. This corpus of knowledge is reviewed here. The right hemisphere does not simply offer its support when calculation requires generic space processing, but its role can be very specific. For example, the right parietal lobe seems to store the operation-specific spatial layout required for complex arithmetical procedures and areas like the right insula are necessary in parsing complex numbers containing zero. Evidence is found for a complex orchestration between the two hemispheres even for simple tasks: each hemisphere has its specific role, concurring to the correct result. As for development, data point to right dominance for basic numerical processes. The picture that emerges at school age is a bilateral pattern with a significantly greater involvement of the right-hemisphere, particularly in non-symbolic tasks. The intraparietal sulcus shows a left hemisphere preponderance in response to symbolic stimuli at this age.

Two dissociable semantic mechanisms predict naming errors and their responsive brain sites in awake surgery. DO80 revisited.

How do we choose words, and what affects the selection of a specific term? Naming tests such as the DO80 are frequently used to assess language function during brain mapping in awake surgery. The present study aimed to explore whether specific semantic errors become more probable under the stimulation of specific brain areas. Moreover, it meant to determine whether specific semantic characteristics of the items may evoke specific types of error. A corpus-based qualitative semantic analysis of the DO80 items, and the emitted naming errors to those items during direct cortical electrostimulation (DCE) revealed that the number of hyperonyms (i.e. 'vehicle' for car') of an item predicted the emission of a synonym ('automobile' for 'car'). This association occurred mainly in frontal tumor patients, which was corroborated by behavior to lesion analyses. In contrast, the emission of co-hyponyms was associated with tumors located in temporal areas. These two behavior-lesion associations thus dissociated, and were also dependent on item semantic characteristics. Co-hyponym errors might generate from the disruption in a temporal semantic-to-lexical process, and the production of synonyms could be the result of an impairment in a frontal lexical-selection mechanism. A hypothesis on the lexical selection mechanisms exerted by the inferior frontal gyrus is proposed. Crucially, the present data suggest the need for more restrictive naming tasks, with items conditioned by tumor location.

Neurofunctional Components of Simple Calculation: A Magnetoencephalography Study.

Our ability to calculate implies more than the sole retrieval of the correct solution. Essential processes for simple calculation are related to the spreading of activation through arithmetic memory networks. There is behavioral and electrophysiological evidence for these mechanisms. Their brain location is, however, still uncertain. Here, we measured magnetoencephalographic brain activity during the verification of simple multiplication problems. Following the operands, the solutions to verify could be preactivated correct solutions, preactivated table-related incorrect solutions, or unrelated incorrect solutions. Brain source estimation, based on these event-related fields, revealed 3 main brain networks involved in simple calculation: 1) bilateral inferior frontal areas mainly activated in response to correct, matching solutions; 2) a left-lateralized frontoparietal network activated in response to incorrect table-related solutions; and (3) a strikingly similar frontoparietal network in the opposite hemisphere activated in response to unrelated solutions. Directional functional connectivity analyses revealed a bidirectional causal loop between left parietal and frontal areas for table-related solutions, with frontal areas explaining the resolution of arithmetic competition behaviorally. Hence, this study isolated at least 3 neurofunctional networks orchestrated between hemispheres during calculation.

A MEG Study on the Processing of Time and Quantity: Parietal Overlap but Functional Divergence.

A common magnitude system for the processing of time and numerosity, supported by areas in the posterior parietal cortex, has been proposed by some authors. The present study aims to investigate possible intersections between the neural processing of non-numerical (time) and numerical magnitudes in the posterior parietal lobe. Using Magnetoencephalography for the comparison of brain source activations during the processing of duration and numerosity contrasts, we demonstrate parietal overlap as well as dissociations between these two dimensions. Within the parietal cortex, the main areas of overlap were bilateral precuneus, bilateral intraparietal sulci, and right supramarginal gyrus. Interestingly, however, these regions did not equivalently correlated with the behavior for the two dimensions: left and right precuneus together with the right supramarginal gyrus accounted functionally for durational judgments, whereas numerosity judgments were accounted by the activation pattern in the right intraparietal sulcus. Present results, indeed, demonstrate an overlap between the neural substrates for processing duration and quantity. However, the functional relevance of parietal overlapping areas for each dimension is not the same. In fact, our data indicates that the same parietal sites rule differently non-numerical and numerical dimensions, as parts of broader networks.

Oscillatory brain activity reveals linguistic prints in the quantity code.

Number representations change through education, although it is currently unclear whether and how language could impact the magnitude representation that we share with other species. The most prominent view is that language does not play any role in modulating the core numeric representation involved in the contrast of quantities. Nevertheless, possible cultural hints on the numerical magnitude representation are currently on discussion focus. In fact, the acquisition of number words provides linguistic input that the quantity system may not ignore. Bilingualism offers a window to the study of this question, especially in bilinguals where the two number wording systems imply also two different numerical systems, such as in Basque-Spanish bilinguals. The present study evidences linguistic prints in the core number representational system through the analysis of EEG oscillatory activity during a simple number comparison task. Gamma band synchronization appears when Basque-Spanish bilinguals compare pairs of Arabic numbers linked through the Basque base-20 wording system, but it does not if the pairs are related through the base-10 system. Crucially, this gamma activity, originated in a left fronto-parietal network, only appears in bilinguals who learned math in Basque and not in equivalent proficiency bilinguals who learned math in Spanish. Thus, this neural index reflected in gamma band synchrony appears to be triggered by early learning experience with the base-20 numerical associations in Basque number words.

Functional mapping of left parietal areas involved in simple addition and multiplication. A single-case study of qualitative analysis of errors.

All electrostimulation studies on arithmetic have so far solely reported general errors. Nonetheless, a classification of the errors during stimulation can inform us about underlying arithmetic processes. The present electrostimulation study was performed in a case of left parietal glioma. The patient's erroneous responses suggested that calculation was mainly applied for addition and a combination of retrieval and calculation was mainly applied for multiplication. The findings of the present single-case study encourage follow up with further data collection with the same paradigm.

Core number representations are shaped by language.

Language and math have been predominantly related through exact calculation. In the present study we investigated a more fundamental link between language and math: whether the most basic quantity representation used for the contrast of numerosities could be shaped by language. We selected two groups of balanced, equally proficient Basque-Spanish bilinguals. Crucially, the two groups differed with respect to the language in which math had been learned at the point of earliest formal instruction in mathematics (Language of learning Math - LL(math)). They performed a simple comparison task between pairs of Arabic digits related through the decimal system or through the vigesimal system. The vigesimal system is retained in Basque for the naming of certain numerals, while for other numerals the decimal system is used, just as for all Spanish number words. Event-related potential (ERP) distance effects were taken as the dependent variable, indexing the activation of quantity. Results showed an N1-P2 distance effect during the comparison of digit pairs related through the base-10 system in both groups. Importantly, this N1-P2 effect appeared only for the group whose LL(math) was Basque when base-20 related digits were compared, even if both groups were perfectly fluent in Basque. Thus the early N1-P2 component appears to be sensitive to verbal components contained in quantity representation. Since the task did not contain any verbal input, the present data suggest that quantity representation may have verbal traces inherited from early learning. In turn, LL(math) should be the optimal medium for numerical communication.

Right parietal cortex and calculation processing: intraoperative functional mapping of multiplication and addition in patients affected by a brain tumor.

OBJECT: The role of parietal areas in number processing is well known. The significance of intraoperative functional mapping of these areas has been only partially explored, however, and only a few discordant data are available in the surgical literature with regard to the right parietal lobe. The purpose of this study was to evaluate the clinical impact of simple calculation in cortical electrostimulation of right-handed patients affected by a right parietal brain tumor. METHODS: Calculation mapping in awake surgery was performed in 3 right-handed patients affected by high-grade gliomas located in the right parietal lobe. Preoperatively, none of the patients presented with calculation deficits. In all 3 cases, after sensorimotor and language mapping, cortical and intraparietal sulcus areas involved in single-digit multiplication and addition calculations were mapped using bipolar electrostimulation. RESULTS: In all patients, different sites of the right parietal cortex, mainly in the inferior lobule, were detected as being specifically related to calculation (multiplication or addition). In 2 patients the intraparietal sulcus was functionally specific for multiplication. No functional sites for language were detected. All sites functional for calculation were spared during tumor resection, which was complete in all cases without postoperative neurological deficits. CONCLUSIONS: These findings provide intraoperative data in support of an anatomofunctional organization for multiplication and addition within the right parietal area. Furthermore, the study shows the potential clinical relevance of intraoperative mapping of calculation in patients undergoing surgery in the right parietal area. Further and larger studies are needed to confirm these data and assess whether mapped areas are effectively essential for function.

Neural correlates of visual versus abstract letter processing in Roman and Arabic scripts.

In alphabetic orthographies, letter identification is a critical process during the recognition of visually presented words. In the present experiment, we examined whether and when visual form influences letter processing in two very distinct alphabets (Roman and Arabic). Disentangling visual versus abstract letter representations was possible because letters in the Roman alphabet may look visually similar/dissimilar in lowercase and uppercase forms (e.g., c-C vs. r-R) and letters in the Arabic alphabet may look visually similar/dissimilar, depending on their position within a word (e.g., [Formula: see text] - [Formula: see text] vs. [Formula: see text] - [Formula: see text]). We employed a masked priming same-different matching task while ERPs were measured from individuals who had learned the two alphabets at an early age. Results revealed a prime-target relatedness effect dependent on visual form in early components (P/N150) and a more abstract relatedness effect in a later component (P300). Importantly, the pattern of data was remarkably similar in the two alphabets. Thus, these data offer empirical support for a universal (i.e., across alphabets) hierarchical account of letter processing in which the time course of letter processing in different scripts follows a similar trajectory from visual features to visual form independent of abstract representations.

Early learning shapes the memory networks for arithmetic: evidence from brain potentials in bilinguals.

Language and math are intertwined during children's learning of arithmetic concepts, but the importance of language in adult arithmetic processing is less clear. To determine whether early learning plays a critical role in the math-language connection in adults, we tested retrieval of simple multiplication in adult bilinguals who learned arithmetic in only one language. We measured electrophysiological and behavioral responses during correctness judgments for problems presented as digits or as number words in Spanish or English. Problems presented in the language in which participants learned arithmetic elicited larger, more graded, and qualitatively different brain responses than did problems presented in participants' other language, and these responses more closely resembled responses for digits, even when participants' other language was more dominant. These findings suggest that the memory networks for simple multiplication are established when arithmetic concepts are first learned and are independent of language dominance in adulthood.

Single pulse TMS induced disruption to right and left parietal cortex on addition and multiplication.

Whether or not mathematical operations are dependent on verbal codes in left hemisphere areas - particularly the left intraparietal sulcus - remains an issue of intense debate. Using single pulse transcranial magnetic stimulation directed at horizontal and ventral regions of the left and right intraparietal sulcus, we examined disruption to reaction times in simple addition and multiplication. Results indicate that these two operations differ in the pattern of lateralization across time for the two areas studied. These show that computational efficiency is not specifically dependent on left hemisphere regions and, in particular, that efficiency in multiplication is dependent on the ventral region of the intraparietal sulcus in the right hemisphere considered to be critical for motion representation and automatization.

Numbers in the blind's "eye".

BACKGROUND: Although lacking visual experience with numerosities, recent evidence shows that the blind perform similarly to sighted persons on numerical comparison or parity judgement tasks. In particular, on tasks presented in the auditory modality, the blind surprisingly show the same effect that appears in sighted persons, demonstrating that numbers are represented through a spatial code, i.e. the Spatial-Numerical Association of Response Codes (SNARC) effect. But, if this is the case, how is this numerical spatial representation processed in the brain of the blind? PRINCIPAL FINDINGS: Here we report that, although blind and sighted people have similarly organized numerical representations, the attentional shifts generated by numbers have different electrophysiological correlates (sensorial N100 in the sighted and cognitive P300 in the blind). CONCLUSIONS: These results highlight possible differences in the use of spatial representations acquired through modalities other than vision in the blind population.

Leftward motion restores number space in neglect.

In the present study, a group of patients with left-sided neglect performed a number comparison task that co-occurred either with coherent motion in different directions or with random motion. Their performance was compared to that of a healthy control group and to a group of patients with right hemisphere damage (RHD) but no signs of neglect. The presence of leftward motion alleviated the difficulties that neglect patients typically show for a number smaller than the reference number 5 (i.e., number 4). Moreover, the standard distance effect was only present when the task co-occurred with leftward motion. These effects were not present in a group of participants with RHD without neglect or in a control group. The present data extend the effects of optokinetic stimulation (OKS) to representational neglect, suggesting that an external redirection of attention by the perception of motion may restore the altered access to the representation of the mental number line in neglect.

Motion on numbers: transcranial magnetic stimulation on the ventral intraparietal sulcus alters both numerical and motion processes.

It has often been proposed that there is a close link between representation of number and space. In the present work, single-pulse transcranial magnetic stimulation (TMS) was applied to the ventral intraparietal sulcus (VIPS) to determine effects on performance in motion detection and number comparison tasks. Participants' reaction times and thresholds for perception of laterally presented coherent motion in random dot kinematograms increased significantly when the contralateral VIPS was stimulated in contrast to the interhemispheric sulcus (Experiment 1) and to the ipsilateral VIPS (Experiment 2). In number comparison tasks, participants compared the magnitude of the laterally presented numbers 1-9 with the number 5. Again, reaction times significantly increased when TMS was applied to the contralateral VIPS in contrast to control sites. The finding that VIPS-directed TMS results in impaired efficiency in both motion perception and number comparison suggests that these processes share a common neural substrate.

Behavioural and neurophysiological evidence of semantic interaction between iconic gestures and words.

We report two experiments that provide converging behavioural and neurophysiological evidence on the relationship between the meaning of iconic gestures and words. Experiment 1 exploited a semantic priming paradigm and revealed interference between gestures and words when they were not related in meaning, but no facilitation when they were. This result was confirmed in Experiment 2, where ERPs were recorded during silent word reading with the same paradigm. The analysis showed a negative deflection peaking near 400 ms (N400) and, in the left hemisphere, greater negative values for verbs than for nouns. Differently from the classical distribution obtained with verbal stimuli, we found an N400 that spread more over central-anterior areas of the scalp, suggesting that the meaning systems of gesture and language do not overlap completely. These results are consistent with the view that the meaning systems for gesture and speech are tightly integrated.

Sensory and cognitive processes of shifts of spatial attention induced by numbers: an ERP study.

The relationship between space and number has become a focus of intensive investigation (Hubbard et al., 2005; Walsh, 2003). The present paper aims to explore the nature of attentional shifts induced by the perception of irrelevant numbers as it was shown by Fischer et al. (2003). We measured the event related potentials induced by the perception of visual lateralized targets cued by numbers that differed in their magnitude. Congruent trials were defined as those where a target presented in the Right Visual Field (RVF) followed a large number and those where a target presented in the Left Visual Field (LVF) followed a small number. Numbers generate a modulation of evoked potentials on targets as soon as 80 msec after the presentation of the target: congruency of the target determined the amplitude on perceptual P100 and cognitive P300 in both sides of presentation of the target. Although a typical distribution of the components was found, effects of congruency were distributed around anterior and Centro-Parietal sites. Due to the functional properties of the mentioned components, the present data suggest that, in fact, perception of numbers does affect the location of attention to external space. Moreover, the distribution of the congruency effect signals so that the representational nature of numbers makes a difference with respect to the stimuli classically used in cueing studies of visual attention to location. The role of top-down control generated by numbers is discussed.