Measuring temporal bias in sequential numerosity comparison.

While several methods have been proposed to assess the influence of continuous visual cues in parallel numerosity estimation, the impact of temporal magnitudes on sequential numerosity judgments has been largely ignored. To overcome this issue, we extend a recently proposed framework that makes it possible to separate the contribution of numerical and non-numerical information in numerosity comparison by introducing a novel stimulus space designed for sequential tasks. Our method systematically varies the temporal magnitudes embedded into event sequences through the orthogonal manipulation of numerosity and two latent factors, which we designate as "duration" and "temporal spacing". This allows us to measure the contribution of finer-grained temporal features on numerosity judgments in several sensory modalities. We validate the proposed method on two different experiments in both visual and auditory modalities: results show that adult participants discriminated sequences primarily by relying on numerosity, with similar acuity in the visual and auditory modality. However, participants were similarly influenced by non-numerical cues, such as the total duration of the stimuli, suggesting that temporal cues can significantly bias numerical processing. Our findings highlight the need to carefully consider the continuous properties of numerical stimuli in a sequential mode of presentation as well, with particular relevance in multimodal and cross-modal investigations. We provide the complete code for creating sequential stimuli and analyzing participants' responses.

Don't plan, just do it: Cognitive and sensorimotor contributions to manual dexterity.

Manual dexterity is referred to as the skill to perform fine motor movements and it has been assumed to be associated to the cognitive domain, as well as the sensorimotor one. In this work, we investigated with functional near-infrared spectroscopy the cortical activations elicited by the execution of the 9-HPT, i.e., a standard test evaluating manual dexterity in which nine pegs were taken, placed into and then removed from nine holes on a board as quickly as possible. For comparison, we proposed a new active control task mainly involving the sensorimotor domain, in which the pegs must be placed and removed using the same single hole (1-HPT). Behaviorally, we found two distinct groups based on the difference between the execution time of the 9-HPT and the 1-HPT (ΔHPT). Cortical areas belonging to the network controlling reaching and grasping movements were active in both groups; however, participants showing a large ΔHPT presented significantly higher activation in prefrontal cortical areas (right BA10 and BA11) during 9-HPT and 1-HPT performance with respect to the participants with a small ΔHPT, who showed a deactivation in BA10. Unexpectedly, we observed a significant linear relationship between ΔHPT and right BA10 activity. This suggested that participants performing the 9-HPT more slowly than the 1-HPT recruited prefrontal areas implicitly exploiting the cognitive skills of planning, perhaps in search of a motor strategy to solve the test activating attentional and cognitive control processes, but this resulted not efficient and instead increased the time to accomplish a manual dexterity task.

Moral judgment, decision times and emotional salience of a new developed set of sacrificial manual driving dilemmas.

The growing interest in the subject of moral judgment in driver and autonomous vehicle behavior highlights the importance of investigating the suitability of sacrificial dilemmas as experimental tools in the context of traffic psychology. To this aim a set of validated sacrificial trolley problems and a new set of trolley-like driving dilemmas were compared through an online survey experiment, providing normative values for rates of participants' choices; decision times; evaluation of emotional valence and arousal experienced during the decision process; and ratings of the moral acceptability. Results showed that while both sets of dilemmas led to a more frequent selection of utilitarian outcomes, the driving-type dilemmas seemed to enhance faster decisions mainly based on the utilitarian moral code. No further differences were observed between the two sets, confirming the reliability of the moral dilemma tool in the investigation of moral driving behaviors. We suggest that as moral judgments and behaviors become more lifelike, the individual's moral inclination emerge more automatically and effectively. This new driving-type dilemma set may help researchers who work in traffic psychology and moral decision-making to approach the complex task of developing realistic moral scenarios more easily in the context of autonomous and nonautonomous transportation.

fNIRS & e-drum: An ecological approach to monitor hemodynamic and behavioural effects of rhythmic auditory cueing training.

Converging evidence suggests a beneficial effect of rhythmic music-therapy in easing motor dysfunctions. Nevertheless, the neural systems underpinning both the direct effect and the influence of rhythm on movement control and execution during training in ecological settings are still largely unknown. In this study, we propose an ecological approach to monitor brain activity and behavioural performance during rhythmic auditory cueing short-term training. Our approach envisages the combination of functional near-infrared spectroscopy (fNIRS), which is a non-invasive neuroimaging technique that allows unconstrained movements of participants, with electronic drum (e-drum), which is an instrument able to collect behavioural tapping data in real time. The behavioural and brain effects of this short-term training were investigated on a group of healthy participants, who well tolerated the experimental settings, since none of them withdrew from the study. The rhythmic auditory cueing short-term training improved beat regularity and decreased group variability. At the group level, the training resulted in a reduction of brain activity primarily in premotor areas. Furthermore, participants with the highest behavioural improvement during training showed the smallest reduction in brain activity. Overall, we conclude that our study could pave the way towards translating the proposed approach to clinical settings.

Reward motivation and neurostimulation interact to improve working memory performance in healthy older adults: A simultaneous tDCS-fNIRS study.

Several studies have evaluated the effect of anodal transcranial direct current stimulation (tDCS) over the prefrontal cortex (PFC) for the enhancement of working memory (WM) performance in healthy older adults. However, the mixed results obtained so far suggest the need for concurrent brain imaging, in order to more directly examine tDCS effects. The present study adopted a continuous multimodal approach utilizing functional near-infrared spectroscopy (fNIRS) to examine the interactive effects of tDCS combined with manipulations of reward motivation. Twenty-one older adults (mean age = 69.7 years; SD = 5.05) performed an experimental visuo-spatial WM task before, during and after the delivery of 1.5 mA anodal tDCS/sham over the left prefrontal cortex (PFC). During stimulation, participants received performance-contingent reward for every fast and correct response during the WM task. In both sessions, hemodynamic activity of the bilateral frontal, motor and parietal areas was recorded across the entire duration of the WM task. Cognitive functions and reward sensitivity were also assessed with standard measures. Results demonstrated a significant impact of tDCS on both WM performance and hemodynamic activity. Specifically, faster responses in the WM task were observed both during and after anodal tDCS, while no differences were found under sham control conditions. However, these effects emerged only when taking into account individual visuo-spatial WM capacity. Additionally, during and after the anodal tDCS, increased hemodynamic activity relative to sham was observed in the bilateral PFC, while no effects of tDCS were detected in the motor and parietal areas. These results provide the first evidence of tDCS-dependent functional changes in PFC activity in healthy older adults during the execution of a WM task. Moreover, they highlight the utility of combining reward motivation with prefrontal anodal tDCS, as a potential strategy to improve WM efficiency in low performing healthy older adults.

On the Role of the Inferior Intraparietal Sulcus in Visual Working Memory for Lateralized Single-feature Objects.

A consolidated practice in cognitive neuroscience is to explore the properties of human visual working memory through the analysis of electromagnetic signals using cued change detection tasks. Under these conditions, EEG/MEG activity increments in the posterior parietal cortex scaling with the number of memoranda are often reported in the hemisphere contralateral to the objects' position in the memory array. This highly replicable finding clashes with several reported failures to observe compatible hemodynamic activity modulations using fMRI or fNIRS in comparable tasks. Here, we reconcile this apparent discrepancy by acquiring fMRI data on healthy participants and employing a cluster analysis to group voxels in the posterior parietal cortex based on their functional response. The analysis identified two distinct subpopulations of voxels in the intraparietal sulcus (IPS) showing a consistent functional response among participants. One subpopulation, located in the superior IPS, showed a bilateral response to the number of objects coded in visual working memory. A different subpopulation, located in the inferior IPS, showed an increased unilateral response when the objects were displayed contralaterally. The results suggest that a cluster of neurons in the inferior IPS is a candidate source of electromagnetic contralateral responses to working memory load in cued change detection tasks.

Atypical right hemisphere response to slow temporal modulations in children with developmental dyslexia.

Phase entrainment of neuronal oscillations is thought to play a central role in encoding speech. Children with developmental dyslexia show impaired phonological processing of speech, proposed theoretically to be related to atypical phase entrainment to slower temporal modulations in speech (<10Hz). While studies of children with dyslexia have found atypical phase entrainment in the delta band (~2Hz), some studies of adults with developmental dyslexia have shown impaired entrainment in the low gamma band (~35-50Hz). Meanwhile, studies of neurotypical adults suggest asymmetric temporal sensitivity in auditory cortex, with preferential processing of slower modulations by right auditory cortex, and faster modulations processed bilaterally. Here we compared neural entrainment to slow (2Hz) versus faster (40Hz) amplitude-modulated noise using fNIRS to study possible hemispheric asymmetry effects in children with developmental dyslexia. We predicted atypical right hemisphere responding to 2Hz modulations for the children with dyslexia in comparison to control children, but equivalent responding to 40Hz modulations in both hemispheres. Analyses of HbO concentration revealed a right-lateralised region focused on the supra-marginal gyrus that was more active in children with dyslexia than in control children for 2Hz stimulation. We discuss possible links to linguistic prosodic processing, and interpret the data with respect to a neural 'temporal sampling' framework for conceptualizing the phonological deficits that characterise children with developmental dyslexia across languages.

Number-space interactions in the human parietal cortex: Enlightening the SNARC effect with functional near-infrared spectroscopy.

Interactions between numbers and space have become a major issue in cognitive neuroscience, because they suggest that numerical representations might be deeply rooted in cortical networks that also subserve spatial cognition. The spatial-numerical association of response codes (SNARC) is the most robust and widely replicated demonstration of the link between numbers and space: in magnitude comparison or parity judgments, participants' reaction times to small numbers are faster with left than right effectors, whereas the converse is found for large numbers. However, despite the massive body of research on number-space interactions, the nature of the SNARC effect remains controversial and no study to date has identified its hemodynamic correlates. Using functional near-infrared spectroscopy, we found a hemodynamic signature of the SNARC effect in the bilateral intraparietal sulcus, a core region for numerical magnitude representation, and left angular gyrus (ANG), a region implicated in verbal number processing. Activation of intraparietal sulcus was also modulated by numerical distance. Our findings point to number semantics as cognitive locus of number-space interactions, thereby revealing the intrinsic spatial nature of numerical magnitude representation. Moreover, the involvement of left ANG is consistent with the mediating role of verbal/cultural factors in shaping interactions between numbers and space.

Are the neural correlates of subitizing and estimation dissociable? An fNIRS investigation.

Human performance in visual enumeration tasks typically shows two distinct patterns as a function of set size. For small sets, usually up to 4 items, numerosity judgments are extremely rapid, precise and confident, a phenomenon known as subitizing. When this limit is exceeded and serial counting is precluded, exact enumeration gives way to estimation: performance becomes error-prone and more variable. Surprisingly, despite the importance of subitizing and estimation in numerical cognition, only few neuroimaging studies have examined whether the neural activity related to these two phenomena can be dissociated. In the present work, we used multi-channel near-infrared spectroscopy (fNIRS) to measure hemodynamic activity of the bilateral parieto-occipital cortex during a visual enumeration task. Participants had to judge the numerosity of dot arrays and indicate it by means of verbal response. We observed a different hemodynamic pattern in the parietal cortex, both in terms of amplitude modulation and temporal profile, for numerosities below and beyond the subitizing range. Crucially, the neural dissociation between subitizing and estimation was strongest at the level of right IPS. The present findings confirm that fNIRS can be successfully used to detect subtle temporal differences in hemodynamic activity and to produce inferences on the neural mechanisms underlying cognitive functions.

A semi-immersive virtual reality incremental swing balance task activates prefrontal cortex: a functional near-infrared spectroscopy study.

Previous functional near-infrared spectroscopy (fNIRS) studies indicated that the prefrontal cortex (PFC) is involved in the maintenance of the postural balance after external perturbations. So far, no studies have been conducted to investigate the PFC hemodynamic response to virtual reality (VR) tasks that could be adopted in the field of functional neurorehabilitation. The aim of this fNIRS study was to assess PFC oxygenation response during an incremental and a control swing balance task (ISBT and CSBT, respectively) in a semi-immersive VR environment driven by a depth-sensing camera. It was hypothesized that: i) the PFC would be bilaterally activated in response to the increase of the ISBT difficulty, as this cortical region is involved in the allocation of attentional resources to maintain postural control; and ii) the PFC activation would be greater in the right than in the left hemisphere considering its dominance for visual control of body balance. To verify these hypotheses, 16 healthy male subjects were requested to stand barefoot while watching a 3 dimensional virtual representation of themselves projected onto a screen. They were asked to maintain their equilibrium on a virtual blue swing board susceptible to external destabilizing perturbations (i.e., randomizing the forward-backward direction of the impressed pulse force) during a 3-min ISBT (performed at four levels of difficulty) or during a 3-min CSBT (performed constantly at the lowest level of difficulty of the ISBT). The center of mass (COM), at each frame, was calculated and projected on the floor. When the subjects were unable to maintain the COM over the board, this became red (error). After each error, the time required to bring back the COM on the board was calculated (returning time). An eight-channel continuous wave fNIRS system was employed for measuring oxygenation changes (oxygenated-hemoglobin, O2Hb; deoxygenated-hemoglobin, HHb) related to the PFC activation (Brodmann Areas 10, 11 and 46). The results have indicated that the errors increased between the first and the second level of difficulty of the ISBT, then decreased and remained constant; the returning time progressively increased during the first three levels of difficulty and then remained constant. During the CSBT, the errors and the returning time did not change. In the ISBT, the increase of the first three levels of difficulty was accompanied by a progressive increase in PFC O2Hb and a less consistent decrease in HHb. A tendency to plateau was observable for PFC O2Hb and HHb changes in the fourth level of difficulty of the ISBT, which could be partly explained by a learning effect. A right hemispheric lateralization was not found. A lower amplitude of increase in O2Hb and decrease in HHb was found in the PFC in response to the CSBT with respect to the ISBT. This study has demonstrated that the oxygenation increased over the PFC while performing an ISBT in a semi-immersive VR environment. These data reinforce the involvement of the PFC in attention-demanding balance tasks. Considering the adaptability of this virtual balance task to specific neurological disorders, the absence of motion sensing devices, and the motivating/safe semi-immersive VR environment, the ISBT adopted in this study could be considered valuable for diagnostic testing and for assessing the effectiveness of functional neurorehabilitation.

Motion artifacts in functional near-infrared spectroscopy: a comparison of motion correction techniques applied to real cognitive data.

Motion artifacts are a significant source of noise in many functional near-infrared spectroscopy (fNIRS) experiments. Despite this, there is no well-established method for their removal. Instead, functional trials of fNIRS data containing a motion artifact are often rejected completely. However, in most experimental circumstances the number of trials is limited, and multiple motion artifacts are common, particularly in challenging populations. Many methods have been proposed recently to correct for motion artifacts, including principle component analysis, spline interpolation, Kalman filtering, wavelet filtering and correlation-based signal improvement. The performance of different techniques has been often compared in simulations, but only rarely has it been assessed on real functional data. Here, we compare the performance of these motion correction techniques on real functional data acquired during a cognitive task, which required the participant to speak aloud, leading to a low-frequency, low-amplitude motion artifact that is correlated with the hemodynamic response. To compare the efficacy of these methods, objective metrics related to the physiology of the hemodynamic response have been derived. Our results show that it is always better to correct for motion artifacts than reject trials, and that wavelet filtering is the most effective approach to correcting this type of artifact, reducing the area under the curve where the artifact is present in 93% of the cases. Our results therefore support previous studies that have shown wavelet filtering to be the most promising and powerful technique for the correction of motion artifacts in fNIRS data. The analyses performed here can serve as a guide for others to objectively test the impact of different motion correction algorithms and therefore select the most appropriate for the analysis of their own fNIRS experiment.

Moral reasoning behind the veil of ignorance: An investigation into perspective-taking accessibility in the context of autonomous vehicles.

Perspective-taking (PT) accessibility has been recognized as an important factor in affecting moral reasoning, also playing a non-trivial role in moral investigation towards autonomous vehicles (AVs). A new proposal to deepen this effect leverages the principles of the veil of ignorance (VOI), as a moral reasoning device aimed to control self-interested decisions by limiting the access to specific perspectives and to potentially biased information. Throughout two studies, we deepen the role of VOI reasoning in the moral perception of AVs, disclosing personal and contingent information progressively throughout the experiment. With the use of the moral trilemma paradigm, two different VOI conditions were operationalized, inspired by the Original Position theory by John Rawls and the Equiprobability Model by John Harsanyi. Evidence suggests a significant role of VOI reasoning in affecting moral reasoning, which seems not independent from the order in which information is revealed. Coherently, a detrimental effect of self-involvement on utilitarian behaviours was detected. These results highlight the importance of considering PT accessibility and self-involvement when investigating moral attitudes towards AVs, since it can help the intelligibility of general concerns and hesitations towards this new technology.

A reference-channel based methodology to improve estimation of event-related hemodynamic response from fNIRS measurements.

Functional near-infrared spectroscopy (fNIRS) uses near-infrared light to measure cortical concentration changes in oxygenated (HbO) and deoxygenated hemoglobin (HbR) held to be correlated with cognitive activity. Providing a parametric depiction of such changes in the classic form of stimulus-evoked hemodynamic responses (HRs) can be attained with this technique only by solving two problems. One problem concerns the separation of informative optical signal from structurally analogous noise generated by a variety of spurious sources, such as heart beat, respiration, and vasomotor waves. Another problem pertains to the inherent variability of HRs, which is notoriously contingent on the type of experiment, brain region monitored, and human phenotype. A novel method was devised in the present context to solve both problems based on a two-step algorithm combining the treatment of noise-only data extrapolated from a reference-channel and a Bayesian filter applied on a per-trial basis. The present method was compared to two current methods based on conventional averaging, namely, a typical averaging method and an averaging method implementing the use of a reference-channel. The result of the comparison, carried out both on artificial and real data, revealed a sensitive accuracy improvement in HR estimation using the present method relative to each of the other methods.

Time-on-task in children with ADHD: an ex-Gaussian analysis.

Although it is widely known that high intra-individual variability (IIV) is a key characteristic of attention deficit/hyperactivity disorder (ADHD), a detailed exploration of the IIV pattern during the time course of a cognitive task has never been carried out. In this study, 30 children with ADHD and 30 controls, were administered the Conners' Continuous Performance Task (CPT-II). The across-block individual performance of the groups was analyzed using an ex-Gaussian approach, which enabled a clearer understanding of how individual response times (RTs) fluctuate during a task in comparison with conventional measures of central tendency. While the conventional measures showed a significant group effect on mean RTs but similar RT trends across blocks between the two groups, the ex-Gaussian results revealed no actual differences between the two groups in the normally distributed component of mean RTs (mu). In contrast to the control group, the children with ADHD showed a steep increase in the exponentially distributed component of RTs (tau) across blocks, thereby indicating that extremely long RTs progressively increased soon after the beginning of the task. Taken together, the results demonstrated that sustained attention deficit in ADHD can be detected by analyzing the IIV in the first few task blocks.

Living on the edge: strategic and instructed slowing in the stop signal task.

The stop signal task is widely adopted to assess motor inhibition performance in both clinical and non-clinical populations. Several recent studies explored the influence of strategic approaches to the task. In particular, response slowing seems to be a strategic approach commonly adopted to perform the task. In the present study, we compared a standard version with a strategic version of the task, in which participants were explicitly instructed to slow down responses. Results showed that the instructed slowing did not affect the main inhibition measure, thus confirming the robustness of the stop signal index. On the other hand, it apparently changed the nature of the task, as shown by the lack of correlation between the standard and the strategic versions. In addition, we found a specific influence of individual characteristics on slowing strategies. In the standard version, adherence to task instructions was positively correlated with compliant traits of personality. Despite instructions to maximize response speed, non-compliant participants preferred to adopt a slowing strategy in the standard version of the task, up to a speed level similar to the strategic version, where slowing was required by task instructions. Understanding the role of individual approach to the task seems to be crucial to properly identify how participants cope with task instructions.

Time-of-day influences resting-state functional cortical connectivity.

Time-of-day is rarely considered during experimental protocols investigating motor behavior and neural activity. The goal of this work was to investigate differences in functional cortical connectivity at rest linked to the time of the day using functional Near-Infrared Spectroscopy (fNIRS). Since resting-state brain is shown to be a succession of cognitive, emotional, perceptual, and motor processes that can be both conscious and nonconscious, we studied self-generated thought with the goal to help in understanding brain dynamics. We used the New-York Cognition Questionnaire (NYC-Q) for retrospective introspection to explore a possible relationship between the ongoing experience and the brain at resting-state to gather information about the overall ongoing experience of subjects. We found differences in resting-state functional connectivity in the inter-hemispheric parietal cortices, which was significantly greater in the morning than in the afternoon, whilst the intra-hemispheric fronto-parietal functional connectivity was significantly greater in the afternoon than in the morning. When we administered the NYC-Q we found that the score of the question 27 ("during RS acquisition my thoughts were like a television program or film") was significantly greater in the afternoon with respect to the morning. High scores in question 27 point to a form of thought based on imagery. It is conceivable to think that the unique relationship found between NYC-Q question 27 and the fronto-parietal functional connectivity might be related to a mental imagery process during resting-state in the afternoon.

How tool use and arm position affect peripersonal space representation.

Experiment 1 investigated whether tool use can expand the peripersonal space into the very far extrapersonal space. Healthy participants performed line bisection in peripersonal and extrapersonal space using wooden sticks up to a maximum of 240 cm. Participants misbisected to the left of the true midpoint, both for lines presented in peripersonal and for those presented in extrapersonal space, confirming a peripersonal space expansion up to a distance of 240 cm. Experiment 2 investigated whether arm position could influence the perception of peripersonal and extrapersonal space during tool use. Participants performed line bisection in the peripersonal and in the extrapersonal space (up to a maximum of 120 cm) using wooden sticks in two different conditions: either with the arm bent or with the arm stretched. Results showed stronger pseudoneglect in the stretched arm condition.

Exploring the role of primary and supplementary motor areas in simple motor tasks with fNIRS.

Studies employing functional magnetic resonance imaging (fMRI) have highlighted a covariation between the amplitude of hemodynamic responses recorded in primary and supplementary motor areas (M1 and SMA) and the duration of a motor task. A subset of these studies have hinted to a possible functional dissociation between processing carried out in these areas, with SMA primarily involved in action preparation, while M1 involved in action execution. This proposed functional dissociation was explored in the present study using a different technique--functional near-infrared spectroscopy--which enabled a finer-grained monitoring of the temporal characteristics of the hemodynamic response compared to fMRI. Here, hemodynamic responses in M1 and SMA were recorded in 7 participants during a right-finger-tapping task of short (1 s) or long (3 s) duration. Hemodynamic responses of larger amplitude were recorded from both contralateral M1 and SMA during long-duration than short-duration tapping. Furthermore, the analysis of the temporal profiles of these responses revealed a more sustained and prolonged activity for long-duration versus short-duration tapping in M1, but not in SMA. Rather than functionally dissociable areas, the present results are more compatible with the hypothesis that M1 and SMA subserve different, though strongly interacting, functional subroutines subtended in motor task preparation and execution.

Space coding for sensorimotor transformations can emerge through unsupervised learning.

The posterior parietal cortex (PPC) is fundamental for sensorimotor transformations because it combines multiple sensory inputs and posture signals into different spatial reference frames that drive motor programming. Here, we present a computational model mimicking the sensorimotor transformations occurring in the PPC. A recurrent neural network with one layer of hidden neurons (restricted Boltzmann machine) learned a stochastic generative model of the sensory data without supervision. After the unsupervised learning phase, the activity of the hidden neurons was used to compute a motor program (a population code on a bidimensional map) through a simple linear projection and delta rule learning. The average motor error, calculated as the difference between the expected and the computed output, was less than 3°. Importantly, analyses of the hidden neurons revealed gain-modulated visual receptive fields, thereby showing that space coding for sensorimotor transformations similar to that observed in the PPC can emerge through unsupervised learning. These results suggest that gain modulation is an efficient coding strategy to integrate visual and postural information toward the generation of motor commands.

Influences of hand action on the processing of symbolic numbers: A special role of pointing?

Embodied and grounded cognition theories state that cognitive processing is built upon sensorimotor systems. In the context of numerical cognition, support to this framework comes from the interactions between numerical processing and the hand actions of reaching and grasping documented in skilled adults. Accordingly, mechanisms for the processing of object size and location during reach and grasp actions might scaffold the development of mental representations of numerical magnitude. The present study exploited motor adaptation to test the hypothesis of a functional overlap between neurocognitive mechanisms of hand action and numerical processing. Participants performed repetitive grasping of an object, repetitive pointing, repetitive tapping, or passive viewing. Subsequently, they performed a symbolic number comparison task. Importantly, hand action and number comparison were functionally and temporally dissociated, thereby minimizing context-based effects. Results showed that executing the action of pointing slowed down the responses in number comparison. Moreover, the typical distance effect (faster responses for numbers far from the reference as compared to close ones) was not observed for small numbers after pointing, while it was enhanced by grasping. These findings confirm the functional link between hand action and numerical processing, and suggest new hypotheses on the role of pointing as a meaningful gesture in the development and embodiment of numerical skills.