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Embodied approaches to language understanding hold that comprehension of linguistic material entails a situated simulation of the situation described. Some recent studies have shown that implicit, explicit, and relational properties of objects implied in a sentence are part of this simulation. However, the issue concerning the extent to which language sensorimotor specificity expressed by linguistic constituents of a sentence, contributes to situating the simulation process has not yet been adequately addressed. To fill this gap, we combined a concrete action verb with a noun denoting a graspable or non-graspable object, to form a sensible or non-sensible sentence. Verbs could express a specific action with low degrees of freedom (DoF) or an action with high DoF. Participants were asked to respond indicating whether the sentences were sensible or not. We found that simulation was active in understanding both sensible and non-sensible sentences. Moreover, the simulation was more situated with sentences containing a verb referring to an action with low DoF. Language sensorimotor specificity expressed by the noun, played a role in situating the simulation, only when the noun was preceded by a verb denoting an action with high DoF in sensible sentences. The simulation process in understanding non-sensible sentences evoked both the representations related to the verb and to the noun, these remaining separated rather than being integrated as in sensible sentences. Overall our findings are in keeping with embodied approaches to language understanding and suggest that the language sensorimotor specificity of sentence constituents affects the extent to which the simulation is situated., (Copyright © 2011 Elsevier Srl. All rights reserved.)

Evolution and the brain have done a marvelous job solving many tricky problems in action control, including problems of learning, hierarchical control over serial behavior, continuous recalibration, and fluency in the face of slow feedback. Given that evolution tends to be conservative, it should not be surprising that these solutions are exploited to solve other tricky problems, such as the design of a communication system. We propose that a mechanism of motor control, paired controller/predictor models, has been exploited for language learning, comprehension, and production. Our account addresses the development of grammatical regularities and perspective, as well as how linguistic symbols become meaningful through grounding in perception, action, and emotional systems., (Copyright © 2011 Elsevier Srl. All rights reserved.)

In the last few years, the insula has been the focus of many brain-imaging studies, mostly devoted to clarify its role in emotions and social communication. Physiological data, however, on which one may ground these correlative findings are almost totally lacking. Here, we investigated the functional properties of the insular cortex in behaving monkeys using intracortical microstimulation. Behavioral responses and heart rate changes were recorded. The results showed that the insula is functionally formed by two main subdivisions: (i) a sensorimotor field occupying the caudal-dorsal portion of the insula and appearing as an extension of the parietal lobe; and (ii) a mosaic of orofacial motor programs located in the anterior and centroventral insula sector. These programs show a progressive shift from dorsally located nonemotional motor programs (ingestive activity) to ventral ones laden with emotional and communicative content. The relationship between ingestive and other behaviors is discussed in an evolutionary perspective.

We challenge the classic experience/expression dichotomous account of emotions, according to which experiencing and expressing an emotion are two independent processes. By endorsing Dewey's and Mead's accounts of emotions, and capitalizing upon recent empirical findings, we propose that expression is part of the emotional experience. This proposal partly challenges the purely constructivist approach endorsed by the authors of the target article.

Schizophrenia is often associated with deficits in the domain of language, which are thought to be closely related to deficits in the structure of semantic knowledge. The main aim of the present study was to behaviorally investigate whether semantic impairments in schizophrenia are present also at the very basic level of action verb processing, in particular at the level of motor simulation. We used a go-no go paradigm both for a semantic decision task (with either an early, EGD, or a delayed go-signal delivery, DGD) and for a lexical decision task (control task). Only the first task requires motor simulation to be solved. We found that first-episode schizophrenia (FES) patients, like healthy control (HC) participants, use motor simulation as a basic strategy to semantically judge action verbs. In the EGD condition, both motor simulation and action verb understanding seem to be preserved in FES. However, differently from HC participants, FES patients kept on using the simulation strategy also with the DGD condition, whereas, simultaneously, task performance during this condition appeared to be less efficient and sensitive. Voxel-based morphometry analysis suggested that this altered performance in FES patients could be related to structural brain abnormalities in the right dorsolateral prefrontal cortex. We propose that a prolonged motor simulation in FES may serve as a compensatory strategy for impairments in the selection of action representation and/or for memory deficits disclosed by the DGD condition during the semantic decision task investigated in the present study., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

The aim of this study was to investigate, using eye-tracking technique, the influence of bottom-up and top-down processes on visual behavior while subjects, naïve to art criticism, were presented with representational paintings. Forty-two subjects viewed color and black and white paintings (Color) categorized as dynamic or static (Dynamism) (bottom-up processes). Half of the images represented natural environments and half human subjects (Content); all stimuli were displayed under aesthetic and movement judgment conditions (Task) (top-down processes). Results on gazing behavior showed that content-related top-down processes prevailed over low-level visually-driven bottom-up processes when a human subject is represented in the painting. On the contrary, bottom-up processes, mediated by low-level visual features, particularly affected gazing behavior when looking at nature-content images. We discuss our results proposing a reconsideration of the definition of content-related top-down processes in accordance with the concept of embodied simulation in art perception.

Background: Children with Autistic Spectrum Disorders (ASD) are frequently hampered by motor impairment, with difficulties ranging from imitation of actions to recognition of motor intentions. Such a widespread inefficiency of the motor system is likely to interfere on the ontogeny of both motor planning and understanding of the goals of actions, thus delivering its ultimate effects on the emergence of social cognition., Methodology/principal Findings: We investigate the organization of action representation in 15 high functioning ASD (mean age: 8.11) and in two control samples of typically developing (TD) children: the first one, from a primary school, was matched for chronological age (CA), the second one, from a kindergarten, comprised children of much younger age (CY). We used nine newly designed behavioural motor tasks, aiming at exploring three domains of motor cognition: 1) imitation of actions, 2) production of pantomimes, and 3) comprehension of pantomimes. The findings reveal that ASD children fare significantly worse than the two control samples in each of the inspected components of the motor representation of actions, be it the imitation of gestures, the self-planning of pantomimes, or the (verbal) comprehension of observed pantomimes. In the latter task, owing to its cognitive complexity, ASD children come close to the younger TD children's level of performance; yet they fare significantly worse with respect to their age-mate controls. Overall, ASD children reveal a profound damage to the mechanisms that control both production and pre-cognitive "comprehension" of the motor representation of actions., Conclusions/significance: Our findings suggest that many of the social cognitive impairments manifested by ASD individuals are likely rooted in their incapacity to assemble and directly grasp the intrinsic goal-related organization of motor behaviour. Such impairment of motor cognition might be partly due to an early damage of the Mirror Neuron Mechanism (MNM).

Even though a growing body of research has shown that the processing of action language affects the planning and execution of motor acts, several aspects of this interaction are still hotly debated. The directionality (i.e. does understanding action-related language induce a facilitation or an interference with the corresponding action?), the time course, and the nature of the interaction (i.e. under what conditions does the phenomenon occur?) are largely unclear. To further explore this topic we exploited a go/no-go paradigm in which healthy participants were required to perform arm reaching movements toward a target when verbs expressing either hand or foot actions were shown, and to refrain from moving when abstract verbs were presented. We found that reaction times (RT) and percentages of errors increased when the verb involved the same effector used to give the response. This interference occurred very early, when the interval between verb presentation and the delivery of the go signal was 50 ms, and could be elicited until this delay was about 600 ms. In addition, RTs were faster when subjects used the right arm than when they used the left arm, suggesting that action-verb understanding is left-lateralized. Furthermore, when the color of the printed verb and not its meaning was the cue for movement execution the differences between RTs and error percentages between verb categories disappeared, unequivocally indicating that the phenomenon occurs only when the semantic content of a verb has to be retrieved. These results are compatible with the theory of embodied language, which hypothesizes that comprehending verbal descriptions of actions relies on an internal simulation of the sensory-motor experience of the action, and provide a new and detailed view of the interplay between action language and motor acts.

Two experiments were conducted to investigate whether physical and observational practice in task-sharing entail comparable implicit motor learning. To this end, the social-transfer-of-learning (SToL) effect was assessed when both participants performed the joint practice task (Experiment 1--complete task-sharing), or when one participant observed the other performing half of the practice task (Experiment 2--evocative task-sharing). Since the inversion of the spatial relations between responding agent and stimulus position has been shown to prevent SToL, in the present study we assessed it in both complete and evocative task-sharing conditions either when spatial relations were kept constant or changed from the practice to the transfer session. The same pattern of results was found for both complete and evocative task-sharing, thus suggesting that implicit motor learning in evocative task-sharing is equivalent to that obtained in complete task-sharing. We conclude that this motor learning originates from the simulation of the complementary (rather than the imitative) action.

In this study we investigated whether objects and their name evoke the activation of the same motor programs. In the first experiment participants had to make speeded responses based on the category of an object. They had to signal whether an object, presented visually, either within or outside their reachable space, was natural or manufactured, by making reach-to-precision or reach-to-power grasp responses. We found a compatibility effect between the response required by task, and the grip evoked by the object, for reachable space only. Nevertheless, this finding holds for artefacts and not for natural objects. In the second experiment, participants had to make reach-to-precision or reach-to-power grasp responses when deciding whether an object, presented either within or outside their reachable space, was congruent with a previously displayed word. In this case we found a compatibility effect between the response required by task and the grip evoked by the object's name, however this effect was not limited by participants' reaching range. Our data suggest that objects and objects' name likely correspond to different motor representations. That is, while the former seem to house both stable (i.e., shape and size) and temporary (i.e., orientation and distance with respect to the perceiver) action-relevant information, the latter seem to house only stable action-relevant information., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Simulation theories of social cognition abound in the literature, but it is often unclear what simulation means and how it works. The discovery of mirror neurons, responding both to action execution and observation, suggested an embodied approach to mental simulation. Over the past few years this approach has been hotly debated and alternative accounts have been proposed. We discuss these accounts and argue that they fail to capture the uniqueness of embodied simulation (ES). ES theory provides a unitary account of basic social cognition, demonstrating that people reuse their own mental states or processes represented with a bodily format in functionally attributing them to others., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Previous evidence has shown that active tool-use remaps agents' reaching space with far objects being perceived as reachable and graspable. To date, however, there is no evidence that tool-use observation might also be effective in reaching space remapping. The present six experiments show that not only performing but also observing tool actions may extend the representation of reaching space, useful for grasping objects. In addition, like active tool-use, tool-use observation also impacts on visual distance judgment. Interestingly, these effects only occurred when observers shared the same action potentialities with the agent, i.e., while passively holding a tool compatible with the goal and the spatial range of the observed action. The present findings demonstrate that observing someone else acting with a tool may actually shape the way we map the objects and the space around us, suggesting that such a mapping could provide us with a keystone for coordinating and integrating our actions with those of others., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Previous studies suggested that the observation of other individuals' somatosensory experiences also activates brain circuits processing one's own somatosensory experiences. However, it is unclear whether cortical regions involved with the elementary stages of touch processing are also involved in the automatic coding of the affective consequences of observed touch and to which extent they show overlapping activation for somatosensory experiences of self and others. In order to investigate these issues, in the present fMRI study, healthy participants either experienced touch or watched videos depicting other individuals' inanimate and animate/social touch experiences. Essentially, a distinction can be made between exteroceptive and interoceptive components of touch processing, involved with physical stimulus characteristics and internal feeling states, respectively. Consistent with this distinction, a specific negative modulation was found in the posterior insula by the mere visual perception of other individuals' social or affective cutaneous experiences, compared to neutral inanimate touch. On the other hand, activation in secondary somatosensory and posterior superior temporal regions, strongest for the most intense stimuli, seemed more dependent on the observed physical stimulus characteristics. In contrast to the detected vicarious activation in somatosensory regions, opposite activation patterns for the experience (positive modulation) and observation (negative modulation) of touch suggest that the posterior insula does not reflect a shared representation of self and others' experiences. Embedded in a distributed network of brain regions underpinning a sense of the bodily self, the posterior insula rather appears to differentiate between self and other conditions when affective experiences are implicated.

Impaired understanding of others' sensations and emotions as well as abnormal experience of their own emotions and sensations is frequently reported in individuals with Autism Spectrum Disorder (ASD). It is hypothesized that these abnormalities are based on altered connectivity within "shared" neural networks involved in emotional awareness of self and others. The insula is considered a central brain region in a network underlying these functions, being located at the transition of information about bodily arousal and the physiological state of the body to subjective feelings. The present study investigated the intrinsic functional connectivity properties of the insula in 14 high-functioning participants with ASD (HF-ASD) and 15 typically developing (TD) participants in the age range between 12 and 20 years by means of "resting state" or "nontask" functional magnetic resonance imaging. Essentially, a distinction was made between anterior and posterior regions of the insular cortex. The results show a reduced functional connectivity in the HF-ASD group, compared with the TD group, between anterior as well as posterior insula and specific brain regions involved in emotional and sensory processing. It is suggested that functional abnormalities in a network involved in emotional and interoceptive awareness might be at the basis of altered emotional experiences and impaired social abilities in ASD, and that these abnormalities are partly based on the intrinsic functional connectivity properties of such a network., (Copyright © 2010 Wiley-Liss, Inc.)

We tested the hypothesis that the body self-advantage, i.e., the facilitation in discriminating self versus other people's body-effectors, is the expression of an implicit and body-specific knowledge, based mainly on the sensorimotor representation of one's own body-effectors. Alternatively, the body self-advantage could rely on visual recognition of pictorial cues. According to the first hypothesis, using gray-scale pictures of body-parts, the body self-advantage should emerge when self-body recognition is implicitly required and should be specific for body-effectors and not for inanimate-objects. In contrast, if the self-advantage is due to a mere visual-perceptual facilitation, it should be independent of the implicit or explicit request (and could be extended also to objects). To disentangle these hypotheses, healthy participants were implicitly or explicitly required to recognize either their own body-effectors or inanimate-objects. Participants were more accurate in the implicit task with self rather than with others' body-effectors. In contrast, the self-advantage was not found when an explicit recognition of one's own body-effectors was required, suggesting that the body self-advantage relies upon a sensorimotor, rather than a mere visual representation of one's own body. Moreover, the absence of both self/other and implicit/explicit effects, when processing inanimate-objects, underlines the differences between the body and other objects.

Previous studies demonstrated the human ability to implicitly recognize their own body. When submitted to a visual matching task, participants showed the so-called self-advantage, that is, a better performance with self rather than others' body or body parts. Here, we investigated whether the body self-advantage relies upon a motor representation of one's body. Participants were submitted to a laterality judgment of self and others' hands (Experiment 1 and 3), which involves a sensory-motor mental simulation. Moreover, to investigate whether the self-advantage emerges also when an explicit self processing is required, the same participants were submitted to an explicit self-body recognition task (Experiment 2). Participants showed the self-advantage when performing the laterality judgment, but not when self-recognition was explicitly required. Thus, implicit and explicit recognition of the bodily self dissociate and only an implicit recognition of the bodily self, mapped in motor terms, allows the self-advantage to emerge.

Evidence from a large number of brain imaging studies has shown that, in humans, the insula, and especially its anterior part, is involved in emotions and emotion recognition. Typically, however, these studies revealed that, besides the insula, a variety of other cortical and subcortical areas are also active. Brain imaging studies are correlative in nature, and, as such, they cannot give indications about the necessary contribution of the different centers involved in emotions. In the present study, we aimed to define more clearly the role of the insula in emotional and social behavior of the monkey by stimulating it electrically. Using this technique, one may determine whether direct activation of the insula can produce specific emotional or social behaviors and exactly which parts of this structure are responsible for these behaviors. The results showed that two emotional behaviors, a basic one (disgust) and a social one (affiliative state), were easily elicited by electrical stimulation of specific parts of the insula. Both behaviors were characterized by specific motor and vegetative responses and by a dramatic change in the monkey's responsiveness to external stimuli., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Background: Newborns come into the world wired to socially interact. Is a propensity to socially oriented action already present before birth? Twin pregnancies provide a unique opportunity to investigate the social pre-wiring hypothesis. Although various types of inter-twins contact have been demonstrated starting from the 11(th) week of gestation, no study has so far investigated the critical question whether intra-pair contact is the result of motor planning rather then the accidental outcome of spatial proximity., Methodology/principal Findings: Kinematic profiles of movements in five pairs of twin foetuses were studied by using four-dimensional ultrasonography during two separate recording sessions carried out at the 14(th) and 18(th) week of gestation. We demonstrate that by the 14th week of gestation twin foetuses do not only display movements directed towards the uterine wall and self-directed movements, but also movements specifically aimed at the co-twin, the proportion of which increases between the 14(th) and 18(th) gestational week. Kinematic analysis revealed that movement duration was longer and deceleration time was prolonged for other-directed movements compared to movements directed towards the uterine wall. Similar kinematic profiles were observed for movements directed towards the co-twin and self-directed movements aimed at the eye-region, i.e. the most delicate region of the body., Conclusions/significance: We conclude that performance of movements towards the co-twin is not accidental: already starting from the 14th week of gestation twin foetuses execute movements specifically aimed at the co-twin.

Many authors have proposed that facial expressions, by conveying emotional states of the person we are interacting with, influence the interaction behavior. We aimed at verifying how specific the effect is of the facial expressions of emotions of an individual (both their valence and relevance/specificity for the purpose of the action) with respect to how the action aimed at the same individual is executed. In addition, we investigated whether and how the effects of emotions on action execution are modulated by participants' empathic attitudes. We used a kinematic approach to analyze the simulation of feeding others, which consisted of recording the "feeding trajectory" by using a computer mouse. Actors could express different highly arousing emotions, namely happiness, disgust, anger, or a neutral expression. Response time was sensitive to the interaction between valence and relevance/specificity of emotion: disgust caused faster response. In addition, happiness induced slower feeding time and longer time to peak velocity, but only in blocks where it alternated with expressions of disgust. The kinematic profiles described how the effect of the specificity of the emotional context for feeding, namely a modulation of accuracy requirements, occurs. An early acceleration in kinematic relative-to-neutral feeding profiles occurred when actors expressed positive emotions (happiness) in blocks with specific-to-feeding negative emotions (disgust). On the other hand, the end-part of the action was slower when feeding happy with respect to neutral faces, confirming the increase of accuracy requirements and motor control. These kinematic effects were modulated by participants' empathic attitudes. In conclusion, the social dimension of emotions, that is, their ability to modulate others' action planning/execution, strictly depends on their relevance and specificity to the purpose of the action. This finding argues against a strict distinction between social and nonsocial emotions.

The main aim of the present study was to explore, by means of high-density EEG, the intensity and the temporal pattern of event-related sensory-motor alpha desynchronization (ERD) during the observation of different types of hand motor acts and gestures. In particular, we aimed to investigate whether the sensory-motor ERD would show a specific modulation during the observation of hand behaviors differing for goal-relatedness (hand grasping of an object and meaningless hand movements) and social relevance (communicative hand gestures and grasping within a social context). Time course analysis of alpha suppression showed that all types of hand behaviors were effective in triggering sensory-motor alpha ERD, but to a different degree depending on the category of observed hand motor acts and gestures. Meaningless gestures and hand grasping were the most effective stimuli, resulting in the strongest ERD. The observation of social hand behaviors such as social grasping and communicative gestures, triggered a more dynamic time course of ERD compared to that driven by the observation of simple grasping and meaningless gestures. These findings indicate that the observation of hand motor acts and gestures evoke the activation of a motor resonance mechanism that differs on the basis of the goal-relatedness and the social relevance of the observed hand behavior.

Mirror neurons are a distinct class of neurons that discharge both during the execution of a motor act and during observation of the same or similar motor act performed by another individual. However, the extent to which mirror neurons coding a motor act with a specific goal (e.g., grasping) might also respond to the observation of a motor act having the same goal, but achieved with artificial effectors, is not yet established. In the present study, we addressed this issue by recording mirror neurons from the ventral premotor cortex (area F5) of two monkeys trained to grasp objects with pliers. Neuron activity was recorded during the observation and execution of grasping performed with the hand, with pliers and during observation of an experimenter spearing food with a stick. The results showed that virtually all neurons responding to the observation of hand grasping also responded to the observation of grasping with pliers and, many of them to the observation of spearing with a stick. However, the intensity and pattern of the response differed among conditions. Hand grasping observation determined the earliest and the strongest discharge, while pliers grasping and spearing observation triggered weaker responses at longer latencies. We conclude that F5 grasping mirror neurons respond to the observation of a family of stimuli leading to the same goal. However, the response pattern depends upon the similarity between the observed motor act and the one executed by the hand, the natural motor template.

Background: The humanoid robot WE4-RII was designed to express human emotions in order to improve human-robot interaction. We can read the emotions depicted in its gestures, yet might utilize different neural processes than those used for reading the emotions in human agents., Methodology: Here, fMRI was used to assess how brain areas activated by the perception of human basic emotions (facial expression of Anger, Joy, Disgust) and silent speech respond to a humanoid robot impersonating the same emotions, while participants were instructed to attend either to the emotion or to the motion depicted., Principal Findings: Increased responses to robot compared to human stimuli in the occipital and posterior temporal cortices suggest additional visual processing when perceiving a mechanical anthropomorphic agent. In contrast, activity in cortical areas endowed with mirror properties, like left Broca's area for the perception of speech, and in the processing of emotions like the left anterior insula for the perception of disgust and the orbitofrontal cortex for the perception of anger, is reduced for robot stimuli, suggesting lesser resonance with the mechanical agent. Finally, instructions to explicitly attend to the emotion significantly increased response to robot, but not human facial expressions in the anterior part of the left inferior frontal gyrus, a neural marker of motor resonance., Conclusions: Motor resonance towards a humanoid robot, but not a human, display of facial emotion is increased when attention is directed towards judging emotions., Significance: Artificial agents can be used to assess how factors like anthropomorphism affect neural response to the perception of human actions.

Intersubjectivity entails the taking of another's perspective in order to understand their experience of the world. This perspective taking capacity extends to the intra-individual sharing of tactile experience. Previous studies have shown modulation of motor cortex excitability in response to the observation of aversive tactile stimulation to the hand of another person. Here we used transcranial magnetic stimulation (TMS) and peripheral stimulation to induce a short-latency afferent inhibition (SAI) effect, which we then sought to modulate via observation of non-noxious tactile stimulation to the hand of a model. Side congruency between the observed (model) and the recorded (participant) hand induced an increase of SAI and this effect was found to hold for motor-evoked potentials (MEPs) recorded from both left and right hands. Inhibition was not found with MEPs evoked using unconditioned pulses of TMS. These results demonstrate a sensorimotor response to observed non-noxious stimulation and suggest an empathic matching system for the tactile experiences of others., (Copyright 2010 Elsevier Ireland Ltd. All rights reserved.)

The aim of our paper is to show that there is a sense of body that is enactive in nature and that enables to capture the most primitive sense of self. We will argue that the body is primarily given to us as source or power for action, i.e., as the variety of motor potentialities that define the horizon of the world in which we live, by populating it with things at hand to which we can be directed and with other bodies we can interact with. We will show that this sense of body as bodily self is, on the one hand, antecedent the distinction between sense of agency and sense of ownership, and, on the other, it enables and refines such distinction, providing a conceptual framework for the coherent interpretation of a variety of behavioral and neuropsychological data. We will conclude by positing that the basic experiences we entertain of our selves as bodily selves are from the very beginning driven by our interactions with other bodies as they are underpinned by the mirror mechanism., (2009 Elsevier Ltd. All rights reserved.)

Recent findings in cognitive neuroscience shed light on the existence of a common neural mechanism that could account for action and intention to understand abilities in humans and non-human primates. Empirical evidence on the neural underpinnings of action goals and on their ontogeny and phylogeny is introduced and discussed. It is proposed that the properties of the mirror neuron system and the functional mechanism describing them, embodied simulation, enabled pre-linguistic forms of action and intention understanding. Basic aspects of social cognition appear to be primarily based on the motor cognition that underpins one's own capacity to act, here defined as motor abstraction. On the basis of this new account of the motor system, it is proposed that intersubjectivity is the best conceived of as intercorporeity.

Social life rests in large part on the capacity to understand the intentions behind the behavior of others. What are the origins of this capacity? How is one to construe its development in ontogenesis? By assuming that action understanding can be explained only in terms of the ability to read the minds of others--that is, to represent mental states--the traditional view claims that a sharp discontinuity occurs in both phylogeny and ontogeny. Over the last few years this view has been challenged by a number of ethological and psychological studies, as well as by several neurophysiological findings. In particular, the functional properties of the mirror neuron system and its direct matching mechanism indicate that action understanding may be primarily based on the motor cognition that underpins one's own capacity to act. This article aims to elaborate and motivate the pivotal role of such motor cognition, providing a biologically plausible and theoretically unitary account for the phylogeny and ontogeny of action understanding and also its impairment, as in the case of autistic spectrum disorder.

Previous studies have shown a shared neural circuitry in the somatosensory cortices for the experience of one's own body being touched and the sight of intentional touch. Using functional magnetic resonance imaging (fMRI), the present study aimed to elucidate whether the activation of a visuotactile mirroring mechanism during touch observation applies to the sight of any touch, that is, whether it is independent of the intentionality of observed touching agent. During fMRI scanning, healthy participants viewed video clips depicting a touch that was intentional or accidental, and occurring between animate or inanimate objects. Analyses showed equal overlapping activation for all the touch observation conditions and the experience of one's own body being touched in the bilateral secondary somatosensory cortex (SII), left inferior parietal lobule (IPL)/supramarginal gyrus, bilateral temporal-occipital junction, and left precentral gyrus. A significant difference between the sight of an intentional touch, compared to an accidental touch, was found in the left primary somatosensory cortex (SI/Brodmann's area [BA] 2). Interestingly, activation in SI/BA 2 significantly correlated with the degree of intentionality of the observed touch stimuli as rated by participants. Our findings show that activation of a visuotactile mirroring mechanism for touch observation might underpin an abstract notion of touch, whereas activation in SI might reflect a human tendency to "resonate" more with a present or assumed intentional touching agent.

What is the evolutionary origin of the human ability to understand and predict the behavior of others? Recent studies suggest that human infants' early capacity for understanding others' goal-directed actions relies on nonmentalistic strategies [1-8]. However, there is no consensus about the nature of the mechanisms underpinning these strategies and their evolutionary history. Comparative studies can shed light on these controversial issues. We carried out three preferential looking-time experiments on macaques, modeled on previous work on human infants [1-5], to test whether macaques are sensitive to the functional efficacy of familiar goal-related hand motor acts performed by an experimenter in a given context and to examine to which extent this sensitivity also is present when observing non-goal-related or unusual goal-related motor acts. We demonstrate that macaque monkeys, similar to human infants, do indeed detect action efficacy by gazing longer at less efficient actions. However, they do so only when the observed behavior is directed to a perceptible and familiar goal. Our results show that the direct detection of the functional fitness of action, in relation to goals that have become familiar through previous experience, is the phylogenetic precursor of intentional understanding.

In primates the gaze conveys important information about what others attend to and about their intentions. The ability to follow the gaze direction of conspecifics has been established for several primate species. It has been proposed to be a precursor for more complex cognitive skills related to mind reading. Studies in humans and other primates have shown that this behavior develops during the period between infancy and adulthood; however, the mechanisms responsible for its emergence are still unknown. In a series of experiments we investigated such mechanisms in macaques (Macaca nemestrina). Results show that juvenile macaques improve their ability to follow the gaze of a human experimenter and that adults' ability to follow gaze is more accurate than that of juveniles. Our data also show that this behavior can emerge as the result of learning processes. The discrepancy between the relatively long period of time needed for the full establishment of the gaze-following behavior and its high sensitivity to conditioning procedures may suggest that social experience and integration of this behavior with other social-cognitive skills are required for its development.

This paper discusses the relevance of the discovery of mirror neurons in monkeys and of the mirror neuron system in humans to a neuroscientific account of primates' social cognition and its evolution. It is proposed that mirror neurons and the functional mechanism they underpin, embodied simulation, can ground within a unitary neurophysiological explanatory framework important aspects of human social cognition. In particular, the main focus is on language, here conceived according to a neurophenomenological perspective, grounding meaning on the social experience of action. A neurophysiological hypothesis--the "neural exploitation hypothesis"--is introduced to explain how key aspects of human social cognition are underpinned by brain mechanisms originally evolved for sensorimotor integration. It is proposed that these mechanisms were later on adapted as new neurofunctional architecture for thought and language, while retaining their original functions as well. By neural exploitation, social cognition and language can be linked to the experiential domain of action.

The neural circuits activated in a person carrying out actions, expressing emotions, and experiencing sensations are activated also, automatically via a mirror neuron system, in the observer of those actions, emotions, and sensations. It is proposed that this finding of shared activation suggests a functional mechanism of "embodied simulation" that consists of the automatic, unconscious, and noninferential simulation in the observer of actions, emotions, and sensations carried out and experienced by the observed. It is proposed also that the shared neural activation pattern and the accompanying embodied simulation constitute a fundamental biological basis for understanding another's mind. The implications of this perspective for psychoanalysis are discussed, particularly regarding unconscious communication, projective identification, attunement, empathy, autism, therapeutic action, and transference-countertransference interactions.

The implications of the discovery of mirroring mechanisms and embodied simulation for empathetic responses to images in general, and to works of visual art in particular, have not yet been assessed. Here, we address this issue and we challenge the primacy of cognition in responses to art. We propose that a crucial element of esthetic response consists of the activation of embodied mechanisms encompassing the simulation of actions, emotions and corporeal sensation, and that these mechanisms are universal. This basic level of reaction to images is essential to understanding the effectiveness both of everyday images and of works of art. Historical, cultural and other contextual factors do not preclude the importance of considering the neural processes that arise in the empathetic understanding of visual artworks.

The automatic translation of folk psychology into newly formed brain modules specifically dedicated to mind-reading and other social cognitive abilities should be carefully scrutinized. Searching for the brain location of intentions, beliefs and desires-as such-might not be the best epistemic strategy to disclose what social cognition really is. The results of neurocognitive research suggest that in the brain of primates, mirror neurons, and more generally the premotor system, play a major role in several aspects of social cognition, from action and intention understanding to language processing. This evidence is presented and discussed within the theoretical frame of an embodied simulation account of social cognition. Embodied simulation and the mirror neuron system underpinning it provide the means to share communicative intentions, meaning and reference, thus granting the parity requirements of social communication.

Developmental and cross-cultural studies show that finger counting represents one of the basic number learning strategies. However, despite the ubiquity of such an embodied strategy, the issue of whether there is a neural link between numbers and fingers in adult, literate individuals remains debated. Here, we used transcranial magnetic stimulation to study changes of excitability of hand muscles of individuals performing a visual parity judgment task, a task not requiring counting, on Arabic numerals from 1 to 9. Although no modulation was observed for the left hand muscles, an increase in amplitude of motor-evoked potentials was found for the right hand muscles. This increase was specific for smaller numbers (1 to 4) as compared to larger numbers (6 to 9). These findings indicate a close relationship between hand/finger and numerical representations.

A direct form of 'experiential understanding' of others is achieved by modelling their behaviours as intentional experiences on the basis of the equivalence between what the others do and feel and what we do and feel. This modelling mechanism is embodied simulation. By means of embodied simulation we do not just 'see' an action, an emotion, or a sensation. Side by side with the sensory description of the observed social stimuli, internal representations of the body states associated with actions, emotions, and sensations are evoked in the observer, as if he/she would be doing a similar action or experiencing a similar emotion or sensation. Mirror neurons are likely the neural correlate of this mechanism. The mirror neuron matching systems map the different intentional relations in a compressed fashion, which is neutral about the specific quality or identity of the agentive/subjective parameter. By means of a shared neural state realized in two different bodies that nevertheless obey to the same functional rules, the 'objectual other' becomes 'another self'.

A direct form of experiential understanding of others, "intentional attunement", is achieved by modeling their behavior as intentional experiences on the basis of the activation of shared neural systems underpinning what the others do an feel and what we do and feel. This modeling mechanism is embodied simulation. In parallel with the detached sensory description of the observed social stimuli, internal representations of the body states associated with actions, emotions, and sensations are evoked in the observer, as if he/she would be doing a similar action or experiencing a similar emotion or sensation. Mirror neuron systems are likely the neural correlate of this mechanism. By means of a shared neural state realized in two different bodies, the "objectual other" becomes "another self". A defective intentional attunement caused by a lack of embodies simulation might cause some of the social impairments of autistic individuals.

We investigated the motor and visual properties of F5 grasping neurons, using a controlled paradigm that allows the study of the neuronal discharge during both observation and grasping of many different three-dimensional objects with and without visual guidance. All neurons displayed a preference for grasping of an object or a set of objects. The same preference was maintained when grasping was performed in the dark without visual feedback. In addition to the motor-related discharge, about half of the neurons also responded to the presentation of an object or a set of objects, even when a grasping movement was not required. Often the object evoking the strongest activity during grasping also evoked optimal activity during its visual presentation. Hierarchical cluster analysis indicated that the selectivity of both the motor and the visual discharge of the F5 neurons is determined not by the object shape but by the grip posture used to grasp the object. Because the same paradigm has been used to study the properties of hand-grasping neurons in the dorsal premotor area F2, and in the anterior intraparietal area (AIP), a comparison of the functional properties of grasping-related neurons in the three cortical areas (F5, F2, AIP) is addressed for the first time.

Transcranial magnetic stimulation (TMS) and a behavioral paradigm were used to assess whether listening to action-related sentences modulates the activity of the motor system. By means of single-pulse TMS, either the hand or the foot/leg motor area in the left hemisphere was stimulated in distinct experimental sessions, while participants were listening to sentences expressing hand and foot actions. Listening to abstract content sentences served as a control. Motor evoked potentials (MEPs) were recorded from hand and foot muscles. Results showed that MEPs recorded from hand muscles were specifically modulated by listening to hand-action-related sentences, as were MEPs recorded from foot muscles by listening to foot-action-related sentences. This modulation consisted of an amplitude decrease of the recorded MEPs. In the behavioral task, participants had to respond with the hand or the foot while listening to actions expressing hand and foot actions, as compared to abstract sentences. Coherently with the results obtained with TMS, when the response was given with the hand, reaction times were slower during listening to hand-action-related sentences, while when the response was given with the foot, reaction times were slower during listening to foot-action-related sentences. The present data show that processing verbally presented actions activates different sectors of the motor system, depending on the effector used in the listened-to action.

Concepts are the elementary units of reason and linguistic meaning. They are conventional and relatively stable. As such, they must somehow be the result of neural activity in the brain. The questions are: Where? and How? A common philosophical position is that all concepts-even concepts about action and perception-are symbolic and abstract, and therefore must be implemented outside the brain's sensory-motor system. We will argue against this position using (1) neuroscientific evidence; (2) results from neural computation; and (3) results about the nature of concepts from cognitive linguistics. We will propose that the sensory-motor system has the right kind of structure to characterise both sensory-motor and more abstract concepts. Central to this picture are the neural theory of language and the theory of cogs, according to which, brain structures in the sensory-motor regions are exploited to characterise the so-called "abstract" concepts that constitute the meanings of grammatical constructions and general inference patterns.

Understanding the intentions of others while watching their actions is a fundamental building block of social behavior. The neural and functional mechanisms underlying this ability are still poorly understood. To investigate these mechanisms we used functional magnetic resonance imaging. Twenty-three subjects watched three kinds of stimuli: grasping hand actions without a context, context only (scenes containing objects), and grasping hand actions performed in two different contexts. In the latter condition the context suggested the intention associated with the grasping action (either drinking or cleaning). Actions embedded in contexts, compared with the other two conditions, yielded a significant signal increase in the posterior part of the inferior frontal gyrus and the adjacent sector of the ventral premotor cortex where hand actions are represented. Thus, premotor mirror neuron areas-areas active during the execution and the observation of an action-previously thought to be involved only in action recognition are actually also involved in understanding the intentions of others. To ascribe an intention is to infer a forthcoming new goal, and this is an operation that the motor system does automatically.

Observing actions made by others activates the cortical circuits responsible for the planning and execution of those same actions. This observation-execution matching system (mirror-neuron system) is thought to play an important role in the understanding of actions made by others. In an fMRI experiment, we tested whether this system also becomes active during the processing of action-related sentences. Participants listened to sentences describing actions performed with the mouth, the hand, or the leg. Abstract sentences of comparable syntactic structure were used as control stimuli. The results showed that listening to action-related sentences activates a left fronto-parieto-temporal network that includes the pars opercularis of the inferior frontal gyrus (Broca's area), those sectors of the premotor cortex where the actions described are motorically coded, as well as the inferior parietal lobule, the intraparietal sulcus, and the posterior middle temporal gyrus. These data provide the first direct evidence that listening to sentences that describe actions engages the visuomotor circuits which subserve action execution and observation.

We investigated the properties of neurons located in the distal forelimb field of dorsal premotor area F2 of macaque monkey using a behavioral paradigm for studying the neuronal discharge during observation (object fixation condition) and grasping of different 3-dimensional objects with and without visual guidance of the movement (movement in light and movement in dark conditions, respectively). The main result is that almost all studied neurons were selective for both the type of prehension and the wrist orientation required for grasping an object. Three categories of neurons were found: purely motor, visually modulated, and visuomotor neurons. The discharge of purely motor neurons was not affected by either object presentation or by the visual feedback of the hand approaching to and interacting with the object. Visually modulated neurons presented a different discharge in the 2 movement conditions, this determining a decrease in selectivity for the grip and wrist orientation in the movement in dark condition. Visuomotor neurons typically discharged during the object fixation task even in the absence of any grasping movement. Nine of them also displayed a different discharge rate between the 2 movement conditions. Congruence was observed between the neuron response during the most effective type of prehension and the neuron response during observation of the object requiring that particular prehension. These results indicate an important role of F2 in the control of goal-related hand movements.

Watching the movie scene in which a tarantula crawls on James Bond's chest can make us literally shiver--as if the spider crawled on our own chest. What neural mechanisms are responsible for this "tactile empathy"? The observation of the actions of others activates the premotor cortex normally involved in the execution of the same actions. If a similar mechanism applies to the sight of touch, movies depicting touch should automatically activate the somatosensory cortex of the observer. Here we found using fMRI that the secondary but not the primary somatosensory cortex is activated both when the participants were touched and when they observed someone or something else getting touched by objects. The neural mechanisms enabling our own sensation of touch may therefore be a window also to our understanding of touch.

To have an ontology is to interpret a world. In this paper we argue that the brain, viewed as a representational system aimed at interpreting our world, possesses an ontology too. It creates primitives and makes existence assumptions. It decomposes target space in a way that exhibits a certain invariance, which in turn is functionally significant. We will investigate which are the functional regularities guiding this decomposition process, by answering to the following questions: What are the explicit and implicit assumptions about the structure of reality, which at the same time shape the causal profile of the brain's motor output and its representational deep structure, in particular of the conscious mind arising from it (its "phenomenal output")? How do they constrain high-level phenomena like conscious experience, the emergence of a first-person perspective, or social cognition? By reviewing a series of neuroscientific results and integrating them with a wider philosophical perspective, we will emphasize the contribution the motor system makes to this process. As it will be shown, the motor system constructs goals, actions, and intending selves as basic constituents of the world it interprets. It does so by assigning a single, unified causal role to them. Empirical evidence demonstrates that the brain models movements and action goals in terms of multimodal representations of organism-object-relations. Under a representationalist analysis, this process can be conceived of as an internal, dynamic representation of the intentionality-relation itself. We will show how such a complex form of representational content, once it is in place, can later function as a functional building block for social cognition and for a more complex, consciously experienced representation of the first-person perspective as well.

What neural mechanism underlies the capacity to understand the emotions of others? Does this mechanism involve brain areas normally involved in experiencing the same emotion? We performed an fMRI study in which participants inhaled odorants producing a strong feeling of disgust. The same participants observed video clips showing the emotional facial expression of disgust. Observing such faces and feeling disgust activated the same sites in the anterior insula and to a lesser extent in the anterior cingulate cortex. Thus, as observing hand actions activates the observer's motor representation of that action, observing an emotion activates the neural representation of that emotion. This finding provides a unifying mechanism for understanding the behaviors of others.

Abstraction denotes the cognitive process by means of which general concepts are formed. The dominant view of abstraction considers it not only as a complex and sophisticated cognitive activity, but also as a distinctive hallmark of mankind. The distinctiveness of abstract thought has indeed been closely related to another feature peculiar to our species: language. Following this perspective, the possibility to entertain conceptual representations is thus precluded to animals devoid of full-blown language. I challenge this view and propose that the representational dynamic of the brain is conceivable as a type of self-organization, in which action plays a crucial part. My aim will be to investigate whether, and to what extent, conceptual knowledge can be attributed to non-linguistic animal species, with particular emphasis on nonhuman primates. I therefore introduce the notion of semantic content as a type of 'relational specification'. A review of recent neurophysiological data on the neural underpinnings of action end-states in the macaque monkey brain is presented. On the basis of this evidence, I propose that conceptual representations can be conceived as the expression of a coherent internal world model. This model decomposes the 'outer' space inhabited by things in a meaningful way only to the extent that it accords to biologically constrained, embodied invariance. Finally, I discuss how the 'comparative' neuroscientific approach to abstraction proposed here may shed some light on its nature and its evolutionary origin.