Mauro Pesenti

Neuroanatomical Substrates of Arabic Number Processing, Numerical Comparison, and Simple Addition: A PET Study

Positron emission tomography was used to localize the cerebral networks specifically involved in three basic numerical processes: arabic numeral processing, numerical magnitude comparison, and retrieval of simple addition facts. Relative cerebral blood flow changes were measured while normal volunteers were resting with eyes closed, making physical judgment on nonnumerical characters or arabic digits, comparing, or adding the same digits. Processing arabic digits bilaterally produced a large nonspecific activation of occipito-parietal areas, as well as a specific activation of the right anterior insula. Comparison and simple addition fact retrieval revealed a fronto-parietal network involving mainly the left intraparietal sulcus, the superior parietal lobule and the precentral gyrus. Comparison also activated, but to a lesser extent, the right superior parietal lobe, whereas addition also activated the orbito-frontal areas and the anterior insula in the right hemisphere. Implications for current anatomo-functional models of numerical cognition are drawn.

Neural Correlates of Simple and Complex Mental Calculation

Some authors proposed that exact mental calculation is based on linguistic representations and relies on the perisylvian language cortices, while the understanding of proximity relations between numerical quantities implicates the parietal cortex. However, other authors opposed developmental arguments to suggest that number sense emerges from nonspecific visuospatial processing areas in the parietal cortex. Within this debate, the present study aimed at revealing the functional anatomy of the two basic resolution strategies involved in mental calculation, namely arithmetical fact retrieval and actual computation, questioning in particular the respective role of language and/or visuospatial cerebral areas. Regional cerebral blood flow was measured with positron emission tomography while subjects were at rest (Rest), read digits (Read), retrieved simple arithmetic facts from memory (i.e., 2 x 4, Retrieve), and performed mental complex calculation (i.e., 32 x 24, Compute). Compared to Read, Retrieve engaged a left parieto-premotor circuit representing a developmental trace of a finger-counting representation that mediates, by extension, the numerical knowledge in adult. Beside this basic network, Retrieve involved a naming network, including the left anterior insula and the right cerebellar cortex, while it did not engage the perisylvian language areas, which were deactivated as compared to Rest. In addition to this retrieval network, Compute specifically involved two functional networks: a left parieto-frontal network in charge of the holding of the multidigit numbers in visuospatial working memory and a bilateral inferior temporal gyri related to the visual mental imagery resolution strategy. Overall, these results provide strong evidence of the involvement of visuospatial representations in different levels of mental calculation.

Mental calculation in a prodigy is sustained by right prefrontal and medial temporal areas

Calculating prodigies are individuals who are exceptional at quickly and accurately solving complex mental calculations. With positron emission tomography (PET), we investigated the neural bases of the cognitive abilities of an expert calculator and a group of non-experts, contrasting complex mental calculation to memory retrieval of arithmetic facts. We demonstrated that calculation expertise was not due to increased activity of processes that exist in non-experts; rather, the expert and the non-experts used different brain areas for calculation. We found that the expert could switch between short-term effort-requiring storage strategies and highly efficient episodic memory encoding and retrieval, a process that was sustained by right prefrontal and medial temporal areas.

Finger–digit compatibility in Arabic numeral processing

Finger–digit response compatibility was tested by asking participants to identify Arabic digits by pressing 1 of 10 keys with all 10 fingers. The direction of the finger–digit mapping was varied by manipulating the global direction of the hand–digit mapping as well as the direction of the finger–digit mapping within each hand (in each case, from small to large digits, or the reverse). The hypothesis of a left-to-right mental number line predicted that a complete left-to-right mapping should be easier whereas the hypothesis of a representation based on finger counting predicted that a counting-congruent mapping should be easier. The results show that when all 10 fingers are used to answer, a mapping congruent with the prototypical finger-counting strategy reported by the participants leads to better performance than does a mapping congruent with a left-to-right oriented mental number line, both in palm-down and palm-up postures of the hands, and they demonstrate that finger-counting strategies influence the way that numerical information is mentally represented and processed.

Dissociation of numerosity and duration processing in the left intraparietal sulcus: a transcranial magnetic stimulation study.

A possible dissociation of duration and numerosity processing was tested in an off-line repetitive transcranial magnetic stimulation (rTMS) design. Participants had to compare the numerosity of flashed dot sequences or the duration of single dot displays before and after 15 min of 1 Hz rTMS over one of three sites (the left or right intraparietal sulcus (IPS), or the vertex chosen as a control site). Compared to the control site, performance was only slowed down for the numerosity comparison task after the left IPS stimulation, whereas it was not affected for the duration comparison task for any of the parietal sites. These results show that the parietal area critically involved in numerosity processing is not involved in duration processing, revealing at least one cerebral site where duration and numerosity comparison processes dissociate.

Selective impairment as evidence for mental organisation of arithmetical facts: BB, a case of preserved subtraction?

We report here the case of a patient (BB), suffering from a precocious evolving dementia with impaired arithmetic performance, who showed specific and theoretical pertinent dissociations in basic mental arithmetic. First, in a task involving production of answers to simple arithmetic problems, a strong dissociation was found among operation: while multiplication was severely impaired, addition was moderately and subtraction only slightly impaired. A second dissociation was found between problems potentially solvable by rules and the others, with the former being better preserved. Finally, in multiplication verification tasks, the rate and distribution of errors among problems were not different from those observed in the multiplication production task. This pattern of performance like the one presented by the patient RG (Dagenbach and McCloskey, 1992), suggests first that stored arithmetical fact representations are segregated by arithmetic operation and second that a distinction has to be drawn between arithmetical rules and arithmetical facts. Last, the parallelism of performance observed here in verification and production tasks suggests that the same deficit(s) is (are) responsible for errors in both tasks.

Role of distinct parietal areas in arithmetic: An fMRI-guided TMS study

Although several parietal areas are known to be involved in number processing, their possible role in arithmetic operations remains debated. It has been hypothesized that the horizontal segment of the intraparietal sulcus (hIPS) and the posterior superior parietal lobule (PSPL) contribute to operations solved by calculation procedures, such as subtraction, but whether these areas are also involved in operations solved by memory retrieval, such as multiplication, is controversial. In the present study, we first identified the parietal areas involved in subtraction and multiplication by means of functional magnetic resonance imaging (fMRI) and we found an increased activation, bilaterally, in the hIPS and PSPL during both arithmetic operations. In order to test whether these areas are causally involved in subtraction and multiplication, we used transcranial magnetic stimulation (TMS) to create, in each participant, a virtual lesion of either the hIPS or PSPL, over the sites corresponding to the peaks of activation gathered in fMRI. When compared to a control site, we found an increase in response latencies in both operations after a virtual lesion of either the left or right hIPS, but not of the PSPL. Moreover, TMS over the hIPS increased the error rate in the multiplication task. The present results indicate that even operations solved by memory retrieval, such as multiplication, rely on the hIPS. In contrast, the PSPL seems to underlie processes that are nonessential to solve basic subtraction and multiplication problems.

Number magnitude potentiates action judgements.

Motor actions can be simulated and generated through the perception of objects and their characteristics. Such functional characteristics of objects with given action capabilities are called affordances. Here we report an interaction between the perception of affordances and the processing of numerical magnitude, and we show that the numerical information calibrates the judgement of action even when no actual action is required. In Experiment 1, participants had to judge whether they would be able to grasp a rod lengthways between their thumb and index finger. The presentation of the rod was preceded by a number or a non-numerical symbol. When a small number preceded the rod, participants overestimated their grasp; conversely, when a large number preceded the rods, they underestimated their grasp. In Experiment 2, participants were requested to judge if two successive rods had the same length, a judgement that did not involve any grasping. The numerical primes had no effect on this judgement, showing that the magnitude/affordance interaction was not due to a simple perceptual effect. Finally, Experiment 3 showed that the interaction was not present with a non-numerical ordered sequence, thereby eliminating sequence order as a potentially confounding variable.

Cross-modal interactions between human faces and voices involved in person recognition

Faces and voices are key features of human recognition but the way the brain links them together is still unknown. In this study, we measured brain activity using functional magnetic resonance imaging (fMRI) while participants were recognizing previously learned static faces, voices and voice-static face associations. Using a subtraction method between bimodal and unimodal conditions, we observed that voice-face associations activated both unimodal visual and auditory areas, and specific multimodal regions located in the left angular gyrus and the right hippocampus. Moreover, a functional connectivity analysis confirmed the connectivity of the right hippocampus with the unimodal areas. These findings demonstrate that binding faces and voices rely on a cerebral network sustaining different aspects of integration such as sensory inputs processing, attention and memory.

Creating number semantics through finger movement perception

Communication, language and conceptual knowledge related to concrete objects may rely on the sensory-motor systems from which they emerge. How abstract concepts can emerge from these systems is however still unknown. Here we report a functional interaction between a specific meaningful finger movement, such as a finger grip closing, and a concept as abstract as numerical magnitude. Participants were presented with Arabic digits to recall before or after they perceived a biological or non-biological hand movement. The results show that perceiving a grip closing slows down the processing of large magnitude numbers. Importantly, we show that this motor-to-semantic interaction differs from the reverse semantic-to-motor interaction, and that it does not result from a general movement amplitude processing as it is only observed for biological hand movements. These results demonstrate the functional link between number meaning and goal-directed finger movements, and show how abstract concept semantics can emerge from the sensory-motor circuits of the brain.