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Area 8A has traditionally been considered to be the frontal eye field (FEF), i.e. the area for the motor production of eye movements. However, recent research has shown that the FEF lies posteriorly in premotor area 6. Research in macaque monkeys has demonstrated that, in contrast to premotor area 6, which is involved in the production of motor actions, area 8A is implicated in the cognitive allocation of attention to stimuli based on instruction cues. The aim of the present study was to elucidate the specific cognitive role of area 8A by examining a unique patient with a lesion restricted to area 8A, i.e. sparing the premotor cortex. This right‐handed male patient underwent neuropsychological assessment and testing on two conditional associative‐learning tasks: the motor hand conditional associative‐learning task (MCALT) assessing the selection of motor actions based on instruction cues and the visual conditional associative‐learning task (VCALT) assessing the selection of visual stimuli based on instruction cues. The patient's performance on the VCALT was significantly impaired compared to control subjects, but performance on the MCALT and on an Eye Movement Control Task was preserved. The present study provides evidence that area 8A is critical for the cognitive process regulating the allocation of attention to different stimuli in the environment, but not in the production of eye movements. These findings enhance understanding of the functional organization of the posterior dorsolateral frontal region in the human brain and suggest a specialized role of area 8A in high‐level cognitive processes. [ABSTRACT FROM AUTHOR]

Actions are rarely devoid of emotional content. Thus, a more complete picture of the neural mechanisms underlying the mental simulation of observed actions requires more research using emotion information. The present study used high‐density electroencephalography to investigate mental simulation associated with facial emotion categorisation. Alpha‐mu rhythm modulation was measured at each frequency, from 8 Hz to 13 Hz, to infer the degree of sensorimotor simulation. Results suggest the sensitivity of the sensorimotor activity to emotional information, because (1) categorising static images of neutral faces as happy or sad was associated with stronger suppression in the central region than categorising clearly happy faces, (2) there was preliminary evidence indicating that the strongest suppression in the central region was in response to neutral faces, followed by sad and then happy faces and (3) in the control task, which required categorising images with the head oriented right, left, or forward as right or left, differences between conditions showed a pattern more indicative of task difficulty rather than sensorimotor engagement. Dissociable processing of emotional information in facial expressions and directionality information in head orientations was further captured in beta band activity (14–20 Hz). Stronger mu suppression to neutral faces indicates that sensorimotor simulation extends beyond crude motor mimicry. We propose that mu rhythm responses to facial expressions may serve as a biomarker for empathy circuit activation. Future research should investigate whether atypical or inconsistent mu rhythm responses to facial expressions indicate difficulties in understanding or sharing emotions. [ABSTRACT FROM AUTHOR]

The activation of the Mirror Neuron System (MNS) has been described to reflect visible movements, but not postural, non‐visible, adaptations that accompany the observed movements. Since any motor act is the result of a well‐tailored dialogue between these two components, we decided to investigate whether a motor resonance to nonvisible postural adaptations could be detected. Possible changes in soleus corticospinal excitability were investigated by eliciting the H‐reflex during the observation of three videos, corresponding to three distinct experimental conditions: 'Chest pass', 'Standing' and 'Sitting', and comparing its size with that measured during observation of a control videoclip (a landscape). In the observed experimental conditions, the Soleus muscle has different postural roles: a dynamic role in postural adaptations during the Chest pass; a static role while Standing still; no role while Sitting. The H‐reflex amplitude was significantly enhanced in the 'Chest pass' condition compared to the 'Sitting' and 'Standing' conditions. No significant difference was found between 'Sitting' and 'Standing' conditions. The increased corticospinal excitability of the Soleus during the 'Chest pass' condition suggests that the mirror mechanisms produce a resonance to postural components of an observed action, although they may not be visible. This observation highlights the fact that mirror mechanisms echo non intentional movements as well and points to a novel possible role of mirror neurons in motor recovery. [ABSTRACT FROM AUTHOR]

Autism has been characterized by atypical task-related brain activation and functional connections, coinciding with deficits in sociocommunicative abilities. However, evidence of the brain's experience-dependent plasticity suggests that abnormal activity patterns may be reversed with treatment. In particular, neurofeedback training (NFT), an intervention based on operant conditioning resulting in self-regulation of brain electrical oscillations, has shown increasing promise in addressing abnormalities in brain function and behavior. We examined the effects of ≥ 20 h of sensorimotor mu-rhythm-based NFT in children with high-functioning autism spectrum disorders (ASD) and a matched control group of typically developing children (ages 8-17). During a functional magnetic resonance imaging imitation and observation task, the ASD group showed increased activation in regions of the human mirror neuron system following the NFT, as part of a significant interaction between group (ASD vs. controls) and training (pre- vs. post-training). These changes were positively correlated with behavioral improvements in the ASD participants, indicating that mu-rhythm NFT may be beneficial to individuals with ASD.

The discovery of mirror neurons in the macaque brain in the 1990s triggered investigations on putative human mirror neurons and their potential functionality. The leading proposed function has been action understanding: Accordingly, we understand the actions of others by 'simulating' them in our own motor system through a direct matching of the visual information to our own motor programmes. Furthermore, it has been proposed that this simulation involves the prediction of the sensory consequences of the observed action, similar to the prediction of the sensory consequences of our executed actions. Here, we tested this proposal by quantifying somatosensory attenuation behaviourally during action observation. Somatosensory attenuation manifests during voluntary action and refers to the perception of self‐generated touches as less intense than identical externally generated touches because the self‐generated touches are predicted from the motor command. Therefore, we reasoned that if an observer simulates the observed action and, thus, he/she predicts its somatosensory consequences, then he/she should attenuate tactile stimuli simultaneously delivered to his/her corresponding body part. In three separate experiments, we found a systematic attenuation of touches during executed self‐touch actions, but we found no evidence for attenuation when such actions were observed. Failure to observe somatosensory attenuation during observation of self‐touch is not compatible with the hypothesis that the putative human mirror neuron system automatically predicts the sensory consequences of the observed action. In contrast, our findings emphasize a sharp distinction between the motor representations of self and others. [ABSTRACT FROM AUTHOR]

Rats have elaborate cognitive capacities for playing Hide & Seek. Playing Hide & Seek strongly engages medial prefrontal cortex and the activity of prefrontal cortex neurons reflects the structure of the game. We wondered if prefrontal neurons would also show a mirroring of play‐related neural activity. Specifically, we asked how does the activity in the rat medial prefrontal cortex differ when the animal plays itself versus when it observes others playing. Consistent with our previous work, when the animal plays itself we observed medial prefrontal cortex activity that was sharply locked to game events. Observing play, however, did not lead to a comparable activation of rat medial prefrontal cortex. Firing rates during observing play were lower than during real play. The modulation of responses in medial prefrontal cortex by game events was strong during playing Hide & Seek, but weak during observing Hide & Seek. We conclude the rat prefrontal cortex does not mirror play events under our experimental conditions. [ABSTRACT FROM AUTHOR]

Mirror neurons (MN) have been proposed as the neural substrate for a wide range of clinical, social and cognitive phenomena. Over the last decade, a commonly used tool for investigating MN activity in the human brain has been functional magnetic resonance (fMRI) repetition suppression (RS) paradigms. However, the available evidence is mixed, largely owing to inconsistent application of the methodological criteria necessary to infer MN properties. This raises concerns about the degree to which one can infer the presence (or absence) of MN activity from earlier accounts that adopted RS paradigms. We aimed to clarify this issue using a well‐validated fMRI RS paradigm and tested for mirror properties by rigorously applying the widely accepted criteria necessary to demonstrate MN activity using traditional univariate techniques and Multivariate Pattern Analysis (MVPA). While univariate whole brain analysis in healthy adults showed uni‐modal RS effects within the supplementary motor area, no evidence for cross‐modal RS effects consistent with mirror neuron activity was found. MVPA on the other hand revealed a region along the anterior intraparietal sulcus that met the criteria for MN activity. Taken together, these results clarify disparate evidence from earlier RS studies, highlighting that traditional univariate analysis of RS data may not be sensitive for detecting MN activity when rigorously applying the requisite criteria. In light of these findings, we recommend that short of increasing sample sizes substantially, future studies using RS paradigms to investigate MNs across the human brain consider the use of MVPA. [ABSTRACT FROM AUTHOR]

Abstract: The extent to which affective empathy is impaired in Autism Spectrum Disorder (ASD) remains unclear, as some—but not all—previous neuroimaging studies investigating empathy for pain in ASD have shown similar activation levels to those of neurotypicals individuals. These inconsistent results could be due to the use of different empathy‐eliciting stimuli. While some studies used pictures of faces exhibiting a painful expression, others used pictures of limbs in painful situations. In this study, we used fMRI to compare activation in areas associated with empathy processing (empathy network) for these two types of stimuli in 31 participants (16 with ASD, 15 controls). We found a group difference in the inferior frontal gyrus (IFG) and the thalamus when participants viewed stimuli of limbs in painful situations, but not when they viewed face stimuli with a painful expression. Both groups of participants activated their empathy network more when viewing pictures of limbs in painful situations than when viewing pictures of faces with a painful expression; this increased activation for limbs versus faces was significantly enhanced in controls relative to ASD participants, especially in the secondary somatosensory cortex (SII). Our findings suggest that empathy defect of people with ASD is contingent upon the type of stimuli used, and may be related to the level of Mirror Neuron System involvement, as brain regions showing group differences (IFG, SII) underlie embodiment. We discuss the potential clinical implications of our findings in terms of developing interventions boosting the empathetic abilities of people with ASD. [ABSTRACT FROM AUTHOR]

Questions regarding the malleability of the mirror neuron system ( MNS) continue to be debated. MNS activation has been reported when people observe another person performing biological goal-directed behaviors, such as grasping a cup. These findings support the importance of mapping goal-directed biological behavior onto one's motor repertoire as a means of understanding the actions of others. Still, other evidence supports the Associative Sequence Learning ( ASL) model which predicts that the MNS responds to a variety of stimuli after sensorimotor learning, not simply biological behavior. MNS activity develops as a consequence of developing stimulus-response associations between a stimulus and its motor outcome. Findings from the ideomotor literature indicate that stimuli that are more ideomotor compatible with a response are accompanied by an increase in response activation compared to less compatible stimuli; however, non-compatible stimuli robustly activate a constituent response after sensorimotor learning. Here, we measured changes in the mu-rhythm, an EEG marker thought to index MNS activity, predicting that stimuli that differ along dimensions of ideomotor compatibility should show changes in mirror neuron activation as participants learn the respective stimulus-response associations. We observed robust mu-suppression for ideomotor-compatible hand actions and partially compatible dot animations prior to learning; however, compatible stimuli showed greater mu-suppression than partially or non-compatible stimuli after explicit learning. Additionally, non-compatible abstract stimuli exceeded baseline only after participants explicitly learned the motor responses associated with the stimuli. We conclude that the empirical differences between the biological and ASL accounts of the MNS can be explained by Ideomotor Theory. [ABSTRACT FROM AUTHOR]

Our interactions with other people - and our processing of their actions - are shaped by their reputation. Research has identified an Action Observation Network ( AON) which is engaged when observing other people's actions. Yet, little is known about how the processing of others' actions is influenced by another's reputation. Is the response of the AON modulated by the reputation of the actor? We developed a variant of the ultimatum game in which participants watched either the visible or occluded actions of two 'proposers'. These actions were tied to decisions of how to split a pot of money although the proposers' decisions on each trial were not known to participants when observing the actions. One proposer made fair offers on the majority of trials, establishing a positive reputation, whereas the other made predominantly, unfair offers resulting in a negative reputation. We found significant activations in two regions of the left dorsal premotor cortex ( dPMC). The first of these showed a main effect of reputation with greater activation for the negative reputation proposer than the positive reputation proposer. Furthermore individual differences in trust ratings of the two proposers covaried with activation in the right primary motor cortex (M1). The second showed an interaction between visibility and reputation driven by a greater effect of reputation when participants were observing an occluded action. Our findings show that the processing of others' actions in the AON is modulated by an actor's reputation, and suggest a predictive role for the PMC during action observation. [ABSTRACT FROM AUTHOR]

The ability to understand competitive games is closely connected to the mirror neuron system ( MNS). This network is activated not only when an action is performed, but also when it is observed. Apart from allowing the understanding of actions performed by others, the MNS has been implicated in predicting subsequent actions. However, the results concerning the modulation of this network by the final outcome of these predictions are contradictory. These contradictions may be related to the use of complex experimental conditions. The aim of this research is to identify changes in the activity of the MNS when the predictions are or are not satisfied in a simple intransitive action-based game. An event-related functional magnetic resonance imaging study was conducted. It consisted of the observation of videos with two actors playing the well-known rock-paper-scissors game. The participants were asked to predict the response of the second actor when the first actor performed one of the three possible actions. In some videos (congruents) the prediction was satisfied, but in the rest of the videos (incongruents) the prediction was not satisfied. When the result was shown, higher activity in the MNS was observed in the congruent videos than in the incongruent ones. Therefore, the observation of a simple manual game leads to a significant activation of the MNS, and this activity seems to be modulated by the final outcome of a prediction, and when predictions are satisfied the activity is higher. [ABSTRACT FROM AUTHOR]

Mirror neurons ( MNs) of the monkey ventral premotor cortex (area F5) are a class of cells that match the visual descriptions of others' actions with correspondent motor representations in the observer's brain. Several human studies suggest that one's own motor representations activated during action observation play a role in directing proactive eye movements to the site of the upcoming hand-target interaction. However, there are no data on the possible relationship between gaze behaviour and MN activity. Here we addressed this issue by simultaneously recording eye position and F5 MN activity in two macaques during free observation of a grasping action. More than half of the recorded neurons discharged stronger when the monkey looked at the action than when it did not look at it, but their firing rate was better predicted by 'when' rather than by 'how long' the monkey gazed at the location of the upcoming hand-target interaction. Interestingly, the onset of MN response was linked to the onset of the experimenter's movement, thus making motor representations potentially exploitable to drive eye movements. Furthermore, MNs discharged stronger and earlier when the gaze was 'proactive' compared with 'reactive', indicating that gaze behaviour influences MN activity. We propose that feedforward, automatic representations of other's actions could lead eye movements that, in turn, would provide the motor system with feedback information that enhances the neural representations of the ongoing action. [ABSTRACT FROM AUTHOR]

The Rehabilitation Gaming System ( RGS) has been designed as a flexible, virtual-reality ( VR)-based device for rehabilitation of neurological patients. Recently, training of visuomotor processing with the RGS was shown to effectively improve arm function in acute and chronic stroke patients. It is assumed that the VR-based training protocol related to RGS creates conditions that aid recovery by virtue of the human mirror neuron system. Here, we provide evidence for this assumption by identifying the brain areas involved in controlling the catching of approaching colored balls in the virtual environment of the RGS. We used functional magnetic resonance imaging of 18 right-handed healthy subjects (24 ± 3 years) in both active and imagination conditions. We observed that the imagery of target catching was related to activation of frontal, parietal, temporal, cingulate and cerebellar regions. We interpret these activations in relation to object processing, attention, mirror mechanisms, and motor intention. Active catching followed an anticipatory mode, and resulted in significantly less activity in the motor control areas. Our results provide preliminary support for the hypothesis underlying RGS that this novel neurorehabilitation approach engages human mirror mechanisms that can be employed for visuomotor training. [ABSTRACT FROM AUTHOR]

The ventral agranular frontal cortex of the macaque monkey is formed by a mosaic of anatomically distinct areas. Although each area has been explored by several neurophysiological studies, most of them focused on small sectors of single areas, thus leaving to be clarified which is the general anatomo-functional organization of this wide region. To fill this gap, we studied the ventral convexity of the frontal cortex in two macaque monkeys ( Macaca nemestrina) using intracortical microstimulation and extracellular recording. Functional data were then matched with the cytoarchitectonic parcellation of the recorded region. The results demonstrated the existence of a dorso-ventral functional border, encompassing the anatomical boundary between areas F4 and F1, and a rostro-caudal anatomo-functional border between areas F5 and F4. The ventral subdivision of areas F4 and F1 was highly electrically excitable, represented simple mouth movements and lacked visual properties; in contrast, their dorsal counterpart showed a higher stimulation threshold, represented forelimb and mouth motor acts and hosted different types of visual properties. The data also showed that area F5 was scarcely excitable, and displayed various motor specificity (e.g. for the type of grip) and complex visual (i.e. mirror responses) properties. Overall, the posterior areas F4 and F1 appear to be involved in organizing and controlling goal-directed mouth motor acts and simple movements within different parts of the external (dorsal sector) and internal (ventral sector) space, whereas area F5 code motor acts at a more abstract level, thus enabling the emergence of higher order socio-cognitive functions. [ABSTRACT FROM AUTHOR]

In the last decade there has been a great amount of research investigating the role of simulation in our ability to infer the underlying intentions of any observed action. The majority of studies have focussed on the role of mirror neurons and the network of cortical areas active during action observation (AON) in inferring the goal of an observed action. However, it remains unclear what precisely is simulated when we observe an action and how such simulations can enable the observer to infer the underlying intention of that action. In particular it is not known how simulation in the AON enables the inference of the same goal when the kinematics observed to achieve that goal differ, such as when reaching to grasp an object with the left or right hands. Here we performed a behavioural study with healthy human subjects to address this question. We show that the subjects were able to detect very subtle changes in the kinematics of an observed action. In addition, we fitted the behavioural responses with a model based on the predictive coding account of mirror neurons. This is a Bayesian account of action observation that can be explained by the free-energy principle. Here we show that we can model all the effects observed when the action observation system is considered within a predictive coding framework. [ABSTRACT FROM AUTHOR]

Activity of the primary motor cortex (M1) during action observation is thought to reflect motor resonance. Here, we conducted three studies using transcranial magnetic stimulation (TMS)-induced motor-evoked potentials (MEPs) of the first dorsal interosseus muscle (FDI) during action observation to determine: (i) the time course of M1 corticospinal excitability during the observation of a simple finger movement; (ii) the specificity of M1 modulation in terms of type of movement and muscle; and (iii) the relationship between M1 activity and measures of empathy and autistic traits. In a first study, we administered single-pulse TMS at 30-ms intervals during the observation of simple finger movements. Results showed enhanced corticospinal excitability occurring between 60 and 90 ms after movement onset. In a second experiment, TMS-induced MEPs were recorded from the FDI and abductor digiti minimi muscles while pulses were delivered 90 ms after movement onset during observation of simple finger movement and dot movement. Increased corticospinal excitability was restricted to finger movement and was present in both muscles. Finally, in an exploratory experiment, single-pulse TMS was administered at 30, 90 and 150 ms after movement onset, and participants were asked to complete the Empathy Quotient (EQ) and the Autism Spectrum Quotient (AQ). Correlational analysis revealed a significant link between motor facilitation at 90 ms and the EQ and AQ scores. These results suggest that corticospinal excitability modulation seen at M1 during action observation is the result of a rapid and crude automatic process, which may be related to social functioning. [ABSTRACT FROM AUTHOR]

The general view on the functional role of the monkey inferior parietal lobule (IPL) convexity mainly derives from studies carried out more than two decades ago and does not account for the functional complexity suggested by more recent neuroanatomical findings. We investigated this issue by recording multi- and single units in the IPL convexity of two monkeys and characterizing their somatosensory, visual and motor responses, using a naturalistic (ethologically relevant) approach. These properties were then matched with IPL cytoarchitectonic parcellation. A further aim of this study was to describe the general properties and the localization of IPL mirror neurons, until now not investigated in detail. Results showed that each studied cytoarchitectonic subdivision of the IPL (PF, PFG, PG) is characterized by specific sensory and motor properties. A key feature of the recorded motor neurons is that of coding goal-directed motor acts. Motor responses are somatotopically organized in a rostro-caudal fashion, with mouth, hand and arm represented in PF, PFG and PG, respectively, with a certain degree of overlap between adjacent representations. In each subdivision the motor activity is associated with specific somatosensory and visual responses, suggesting that each area organizes motor acts in different space sectors. Mirror neurons have been found mainly in area PFG and their general features appear to be very similar to those of ventral premotor mirror neurons. The present data suggest that the IPL plays an important role in both action organization and action understanding and should be considered part of the motor system. [ABSTRACT FROM AUTHOR]

The mirror system, comprising cortical areas that allow the actions of others to be represented in the observer’s own motor system, is thought to be crucial for the development of social cognition in humans. Despite the importance of the human mirror system, little is known about its origins. We investigated the role of sensorimotor experience in the development of the mirror system. Functional magnetic resonance imaging was used to measure neural responses to observed hand and foot actions following one of two types of training. During training, participants in the Compatible (control) group made mirror responses to observed actions (hand responses were made to hand stimuli and foot responses to foot stimuli), whereas the Incompatible group made counter-mirror responses (hand to foot and foot to hand). Comparison of these groups revealed that, after training to respond in a counter-mirror fashion, the relative action observation properties of the mirror system were reversed; areas that showed greater responses to observation of hand actions in the Compatible group responded more strongly to observation of foot actions in the Incompatible group. These results suggest that, rather than being innate or the product of unimodal visual or motor experience, the mirror properties of the mirror system are acquired through sensorimotor learning. [ABSTRACT FROM AUTHOR]

We presented professional dancers and non-dancers with videos of two movement styles, dance movements and everyday movements. Participants were asked to indicate by a button press to which category a movement belonged. We computed event-related desynchronization (ERD) in alpha and beta frequency bands between 7.5 and 25 Hz relative to a visual baseline condition. Power in alpha and lower beta frequency bands was significantly reduced if dancers watched dance movements but not if non-dancers watched dance movements, in particular between 1 and 2 s after movement onset. During observation of everyday movements no such group difference was evident. Thus, ERD in alpha and beta frequency bands was modulated by a participant’s expertise with a certain movement style. The results are discussed in light of a human observation–execution matching system similar to the macaque mirror neuron system and strengthen the idea of a functional relationship between such a system and rhythmical activity in the alpha and beta frequency bands. [ABSTRACT FROM AUTHOR]

In the adult human brain, passive observation of actions performed by others activates some of the same cortical areas that are involved in the execution of actions, thereby contributing to action recognition. This mechanism appears to occur through activation of a population of action-coding cells known as mirror neurons (MN). In the adult motor cortex, performing actions and observing human movement reduces the magnitude of the mu (8–13 Hz) rhythm, possibly reflecting MN system activity. Despite the wealth of information available regarding the adult MN system, little is known about its existence in children. Here, we used EEG to probe mu rhythm modulation in 15 children during observation and execution of hand actions. Our data show that mu rhythm attenuation occurs in children under 11 years old during observation of hand movements. Similarly to what has been reported in adults, observation of goal/object-orientated movement produces greater modulation of the mu rhythm than intransitive movement. These data confirm the existence of an observation–execution matching system in the immature human brain and may be of clinical value in the understanding of neurodevelopmental disorders associated with a faulty MN system, such as autism spectrum disorder. [ABSTRACT FROM AUTHOR]

Transcranial magnetic stimulation (TMS) studies report that viewing a given action performed by a model activates the neural representation of the onlooker's muscles that are activated during the actual execution of the observed action. Here we sought to determine whether this mirror observation-execution facilitation reflects only muscular specificity or whether it is also influenced by postural congruency between onlooker/model body parts. We recorded motor potentials evoked by single-pulse TMS from the first dorsal interosseous (FDI) and abductor digiti minimi (ADM) muscles during observation of the right index and little finger abduction/adduction movements of models who kept their hands in a palm-down or palm-up position. Moreover, in different experiments observers kept their right hand palm down or palm up. Selective motor facilitation was observed during observation of movements that map the motor function of the targeted muscles, regardless of the posture of the observed hand. Modulation of FDI, however, was obtained only when participants kept their hand palm down; by contrast, modulation of ADM was obtained only when participants kept their hand palm up. Interestingly, electromyographic recordings showed that FDI is mostly active when index abduction/adduction movements are performed in the palm-down position, whereas ADM is mostly active when little finger abduction/adduction movements are performed in the palm-up position. Results show that the influence of the onlooker's hand posture is comparable in action execution and observation, thus indicating a fine-grain functional correspondence between these two processes. [ABSTRACT FROM AUTHOR]

In order to explore the activation dynamics of the human action recognition system, we investigated electrophysiological distinctions between the brain responses to sounds produced by human finger and tongue movements. Of special interest were the questions of how early these differences may occur, and whether the neural activation at the early stages of processing involves cortical motor representations related to the generation of these sounds. For this purpose we employed a high-density EEG set-up and recorded mismatch negativity (MMN) using a recently developed novel multideviant paradigm which allows acquisition of a high number of trials within a given time period. Deviant stimuli were naturally recorded finger and tongue clicks, as well as control stimuli with similar physical features but without the clear action associations (this was tested in a separate behavioural experiment). Both natural stimuli produced larger MMNs than their respective control stimuli at ∼ 100 ms, indicating activation of memory traces for familiar action-related sounds. Furthermore, MMN topography at this latency differed between the brain responses to the natural finger and natural tongue sounds. Source estimation revealed the strongest sources for finger sounds in centrolateral areas of the left hemisphere, suggesting that hearing a sound related to finger actions evokes activity in motor areas associated with the dominant hand. Furthermore, tongue sounds produced activation in more inferior brain areas. Our data suggest that motor areas in the human brain are part of neural systems subserving the early automatic recognition of action-related sounds. [ABSTRACT FROM AUTHOR]

Transcranial magnetic stimulation (TMS) was applied to motor areas in the left language-dominant hemisphere while right-handed human subjects made lexical decisions on words related to actions. Response times to words referring to leg actions (e.g.kick) were compared with those to words referring to movements involving the arms and hands (e.g.pick). TMS of hand and leg areas influenced the processing of arm and leg words differentially, as documented by a significant interaction of the factors Stimulation site and Word category. Arm area TMS led to faster arm than leg word responses and the reverse effect, faster lexical decisions on leg than arm words, was present when TMS was applied to leg areas. TMS-related differences between word categories were not seen in control conditions, when TMS was applied to hand and leg areas in the right hemisphere and during sham stimulation. Our results show that the left hemispheric cortical systems for language and action are linked to each other in a category-specific manner and that activation in motor and premotor areas can influence the processing of specific kinds of words semantically related to arm or leg actions. By demonstrating specific functional links between action and language systems during lexical processing, these results call into question modular theories of language and motor functions and provide evidence that the two systems interact in the processing of meaningful information about language and action. [ABSTRACT FROM AUTHOR]

Previous studies indicate that the motor areas of both hemispheres are active when observing actions. Here we explored how the motor areas of each hemisphere respond to the sounds associated with actions. We used transcranial magnetic stimulation (TMS) to measure motor corticospinal excitability of hand muscles while listening to sounds. Sounds associated with bimanual actions produced greater motor corticospinal excitability than sounds associated with leg movements or control sounds. This facilitation was exclusively lateralized to the left hemisphere, the dominant hemisphere for language. These results are consistent with the hypothesis that action coding may be a precursor of language. [ABSTRACT FROM AUTHOR]

Abstract The precise neural mechanisms underlying speech perception are still to a large extent unknown. The most accepted view is that speech perception depends on auditory-cognitive mechanisms specifically devoted to the analysis of speech sounds. An alternative view is that, crucial for speech perception, it is the activation of the articulatory (motor) gestures that generate these sounds. The listener understands the speaker when his/her articulatory gestures are activated (motor theory of speech perception). Here, by using transcranial magnetic stimulation (TMS), we demonstrate that, during speech listening, there is an increase of motor-evoked potentials recorded from the listeners' tongue muscles when the presented words strongly involve, when pronounced, tongue movements. Although these data do not prove the motor theory of speech perception, they demonstrate for the first time that word listening produces a phoneme specific activation of speech motor centres. [ABSTRACT FROM AUTHOR]

Abstract The precise mechanisms of how speech may have developed are still unknown to a large extent. Gestures have proven a powerful concept for explaining how planning and analysing of motor acts could have evolved into verbal communication. According to this concept, development of an action-perception network allowed for coding and decoding of communicative gestures. These were manual or manual/articulatory in the beginning and then became increasingly elaborate in the articulatory mode. The theory predicts that listening to the ‘gestures’ that compose spoken language should activate an extended articulatory and manual action-perception network. To examine this hypothesis, we assessed the effects of language on cortical excitability of the hand muscle representation by transcranial magnetic stimulation. We found the hand motor system to be activated by linguistic tasks, most notably pure linguistic perception, but not by auditory or visuospatial processing. The amount of motor system activation was comparable in both hemispheres. Our data support the theory that language may have evolved within a general and bilateral action-perception network. [ABSTRACT FROM AUTHOR]

Abstract The posterior sector of Broca's area (Brodmann area 44), a brain region critical for language, may have evolved from neurons active during observation and execution of manual movements. Imaging studies showing increased Broca's activity during execution, imagination, imitation and observation of hand movements support this hypothesis. Increased Broca's activity in motor task, however, may simply be due to inner speech. To test whether Broca's area is essential to imitation, we used repetitive transcranial magnetic stimulation (rTMS), which is known to transiently disrupt functions in stimulated areas. Subjects imitated finger key presses (imitation) or executed finger key presses in response to spatial cues (control task). While performing the tasks, subjects received rTMS over the left and right pars opercularis of the inferior frontal gyrus (where Brodmann area 44 is probabilistically located) and over the occipital cortex. There was significant impairment in imitation, but not in the control task, during rTMS over left and right pars opercularis compared to rTMS over the occipital cortex. This suggests that Broca's area is a premotor region essential to finger movement imitation. [ABSTRACT FROM AUTHOR]