Balancing the 2 Hemispheres in Simple Calculation: Evidence From Direct Cortical Electrostimulation

Published: 24 September 2016How do the parietal lobes contribute to simple calculation? Clinical and neuroimaging methods, which are based mainly on correlational evidence, have provided contrasting results so far. Here we used direct cortical electrostimulation during brain surgery to causally infer the role of the left and right parietal lobes in simple calculation. Stimulation provoked errors for addition and multiplication in different parietal areas on both hemispheres. Crucially, an innovative qualitative error analysis unveiled the functional contrast of the 2 parietal lobes. Right or left stimulation led to different types of substitution errors in multiplication, unveiling the function of the more active hemisphere. While inhibition of the left hemisphere led mainly to approximation errors, right hemisphere inhibition enhanced retrieval within a stored repertory. These results highlight the respective roles of each hemisphere in the network: rote retrieval of possible solutions by the left parietal areas and approximation to the correct solution by the right hemisphere. The bilateral orchestration between these functions guarantees precise calculation.This work was supported by the Italian Ministry of Health (grant RF-2009-1530973); by the University of Padua (Grant Progetto d’Ateneo CPDA131328 and Progetto strategico NEURAT) to C.S., and by the Spanish Ministry of Science and Innovation (grant PSI2014-53351) to E.S. and by financial assistance as a Severo Ochoa Center of Excellence SEV-2015-0490 to the BCBL (E.S.)

Subjective discomfort of TMS predicts reaction times differences in published studies

Transcranial magnetic stimulation (TMS) was developed thirty years ago, in part to decrease the peripheral side-effects associated with transcranial electrical stimulation (Barker, 1991). TMS has been effective in that aim, and great advances have been made over the past 30 years. TMS can still be uncomfortable and painful, however, as it stimulates excitable superficial tissue including scalp muscles and peripheral nerves (Maizey et al., 2013). This causes annoyance, pain, and muscle twitches (i.e., discomfort) that vary systematically across the scalp (Meteyard & Holmes, 2018). For this Opinion, we investigated whether the TMS-related discomfort measured in our previous work could predict the reported differences in RT in studies published in the last 10 years. For single-pulse TMS studies, differences in RT between TMS and control conditions were significantly correlated with both the mean of median ratings of muscle twitches, and the mean effect of TMS on RT from Meteyard & Holmes (2018)

More than skin deep: body representation beyond primary somatosensory cortex

The neural circuits underlying initial sensory processing of somatic information are relatively well understood. In contrast, the processes that go beyond primary somatosensation to create more abstract representations related to the body are less clear. In this review, we focus on two classes of higher-order processing beyond somatosensation. Somatoperception refers to the process of perceiving the body itself, and particularly of ensuring somatic perceptual constancy. We review three key elements of somatoperception: (a) remapping information from the body surface into an egocentric reference frame (b) exteroceptive perception of objects in the external world through their contact with the body and (c) interoceptive percepts about the nature and state of the body itself. Somatorepresentation, in contrast, refers to the essentially cognitive process of constructing semantic knowledge and attitudes about the body, including: (d) lexical-semantic knowledge about bodies generally and one’s own body specifically, (e) configural knowledge about the structure of bodies, (f) emotions and attitudes directed towards one’s own body, and (g) the link between physical body and psychological self. We review a wide range of neuropsychological, neuroimaging and neurophysiological data to explore the dissociation between these different aspects of higher somatosensory function

Methods and models for brain connectivity assessment across levels of consciousness

The human brain is one of the most complex and fascinating systems in nature. In the last decades, two events have boosted the investigation of its functional and structural properties. Firstly, the emergence of novel noninvasive neuroimaging modalities, which helped improving the spatial and temporal resolution of the data collected from in vivo human brains. Secondly, the development of advanced mathematical tools in network science and graph theory, which has recently translated into modeling the human brain as a network, giving rise to the area of research so called Brain Connectivity or Connectomics. In brain network models, nodes correspond to gray-matter regions (based on functional or structural, atlas-based parcellations that constitute a partition), while links or edges correspond either to structural connections as modeled based on white matter fiber-tracts or to the functional coupling between brain regions by computing statistical dependencies between measured brain activity from different nodes. Indeed, the network approach for studying the brain has several advantages: 1) it eases the study of collective behaviors and interactions between regions; 2) allows to map and study quantitative properties of its anatomical pathways; 3) gives measures to quantify integration and segregation of information processes in the brain, and the flow (i.e. the interacting dynamics) between different cortical and sub-cortical regions. The main contribution of my PhD work was indeed to develop and implement new models and methods for brain connectivity assessment in the human brain, having as primary application the analysis of neuroimaging data coming from subjects at different levels of consciousness. I have here applied these methods to investigate changes in levels of consciousness, from normal wakefulness (healthy human brains) or drug-induced unconsciousness (i.e. anesthesia) to pathological (i.e. patients with disorders of consciousness)

Causal role of the posterior parietal cortex for two-digit mental subtraction and addition: A repetitive TMS study

Although parietal areas of the left hemisphere are known to be involved in simple mental calculation, the possible role of the homologue areas of the right hemisphere in mental complex calculation remains debated. In the present study, we tested the causal role of the posterior parietal cortex of both hemispheres in two-digit mental addition and subtraction by means of neuronavigated repetitive TMS (rTMS), investigating possible hemispheric asymmetries in specific parietal areas. In particular, we performed two rTMS experiments, which differed only for the target sites stimulated, on independent samples of participants. rTMS was delivered over the horizontal and ventral portions of the intraparietal sulcus (HIPS and VIPS, respectively) of each hemisphere in Experiment 1, and over the angular and supramarginal gyri (ANG and SMG, respectively) of each hemisphere in Experiment 2. First, we found that each cerebral area of the posterior parietal cortex is involved to some degree in the two-digit addition and subtraction. Second, in Experiment 1, we found a stronger pattern of hemispheric asymmetry for the involvement of HIPS in addition compared to subtraction. In particular, results showed a greater involvement of the right HIPS than the left one for addition. Moreover, we found less asymmetry for the VIPS. Taken together, these results suggest that two-digit mental addition is more strongly associated with the use of a spatial mapping compared to subtraction. In support of this view, in Experiment 2, a greater role of left and right ANG was found for addition needed in verbal processing of numbers and in visuospatial attention processes, respectively. We also revealed a greater involvement of the bilateral SMG in two-digit mental subtraction, in response to greater working memory load required to solve this latter operation compared to addition

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

Impaired Arithmetic Fact Retrieval in an Adult with Developmental Dyscalculia: Evidence from Behavioral and Functional Brain Imaging Data

Developmental dyscalculia (DD) is a developmental disorder characterized by arithmetic difficulties. Recently, it has been suggested that the neural networks supporting procedure-based calculation (e.g., in subtraction) and left-hemispheric verbal arithmetic fact retrieval (e.g., in multiplication) are partially distinct. Here we compared the neurofunctional correlates of subtraction and multiplication in a 19-year-old student (RM) with DD to 18 age-matched controls. Behaviorally, RM performed significantly worse than controls in multiplication, while subtraction was unaffected. Neurofunctional differences were most pronounced regarding multiplication: RM showed significantly stronger activation than controls not only in left angular gyrus but also in a fronto-parietal network (including left intraparietal sulcus and inferior frontal gyrus) typically activated during procedure-based calculation. Region-of-interest analyses indicated group differences in multiplication only, which, however, did not survive correction for multiple comparisons. Our results are consistent with dissociable and processing-specific, but not operation-specific neurofunctional networks. Procedure-based calculation is not only associated with subtraction but also with (untrained) multiplication facts. Only after rote learning, facts can be retrieved quasi automatically from memory. We suggest that this learning process and the associated shift in activation patterns has not fully occurred in RM, as reflected in her need to resort to procedure-based strategies to solve multiplication facts

Mirror Activity in the Macaque Motor System

Mirror neurons (MirNs) within ventral premotor cortex (PMv) and primary motor cortex (M1), including pyramidal tract neurons (PTNs) projecting to the spinal cord, modulate their activity during both the execution and observation of motor acts. However, movement is not produced in the latter condition, and mirror responses cannot be explained by lowlevel muscle activity. Relatively reduced activity in M1 during observation may help to suppress movement. Here, we examined the extent to which activity at different stages of action observation reflects grasp representation and suppression of movement across multiple levels of the mirror system in monkeys and humans. We recorded MirNs in M1 and F5 (rostral PMv), including identified PTNs, in two macaque monkeys as they performed, observed, and withheld reach-to-grasp actions. Time-varying population activity was more distinct between execution and observation in M1 than in F5, and M1 activity in the lead-up to the observation of movement onset shared parallels with movement withholding activity. In separate experiments, modulation of short-latency responses evoked in hand muscles by pyramidal tract stimulation revealed modest grasp-specific facilitation at the spinal level during grasp observation. This contrasted with a relative suppression of excitability prior to observed movement onset or when monkeys simply withheld movement. Additional cortical recording experiments examined how contextual factors, such as observing to imitate, observing while engaged in action, or observation with reduced visual information, modulated mirror activity in M1 and F5. Finally, single-pulse transcranial magnetic stimulation (TMS) in healthy human volunteers was used to examine changes in corticospinal excitability (CSE) during action observation and withholding. Overall, the results reveal distinctions in the profile of mirror activity across premotor and motor areas. While F5 maintains a more abstract representation of grasp independent of the acting agent, a balance of excitation and inhibition in motor cortex and spinal circuitry during action observation may support a flexible dissociation between initiation of grasping actions and representation of observed grasp

Disruption of spatio-temporal processing in human vision using transcranial magnetic stimulation

Transcranial magnetic stimulation (TMS) is a non-invasive technique used to reversibly modulate the activity of cortical neurons using time-varying magnetic fields. Recently TMS has been used to demonstrate the functional necessity of human cortical areas to visual tasks. For example, it has been shown that delivering TMS over human visual area V5/MT selectively disrupts global motion perception. The temporal resolution of TMS is considered to be one of its main advantages as each pulse has a duration of less than 1 ms. Despite this impressive temporal resolution, however, the critical period(s) during which TMS of area V5/MT disrupts performance on motion-based tasks is still far from clear. To resolve this issue, the influence of TMS on direction discrimination was measured for translational global motion stimuli and components of optic flow (rotational and radial global motion). The results of these experiments provide evidence that there are two critical periods during which delivery of TMS over V5/MT disrupts performance on global motion tasks: an early temporal window centred at 64 ms prior to and a late temporal window centred at 146 ms post global motion onset. Importantly, the early period cannot be explained by a TMS-induced muscular artefact. The onset of the late temporal window was contrast-dependent, consistent with longer neural activation latencies associated with lower contrasts. The theoretical relevance of the two epochs is discussed in relation to feedforward and feedback pathways known to exist in the human visual system, and the first quantitative model of the effects of TMS on global motion processing is presented. A second issue is that the precise mechanism behind TMS disruption of visual perception is largely unknown. For example, one view is that the “virtual lesion” paradigm reduces the effective signal strength, which can be likened to a reduction in perceived target visibility. Alternatively, other evidence suggests that TMS induces neural noise, thereby reducing the signal-to-noise ratio, which results in an overall increase in threshold. TMS was delivered over the primary visual cortex (area V1) to determine whether its influence on orientation discrimination could be characterised as a reduction in the visual signal strength, or an increase in TMS-induced noise. It was found that TMS produced a uniform reduction in perceived stimulus visibility for all observers. In addition, an overall increase in threshold (JND) was also observed for some observers, but this effect disappeared when TMS intensity was reduced. Importantly, susceptibility to TMS, defined as an overall increase in JND, was not dependent on observers’ phosphene thresholds. It is concluded that single-pulse TMS can both reduce signal strength (perceived visibility) and induce task-specific noise, but these effects are separable, dependent on TMS intensity and individual susceptibility