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The mirror neuron system (MNS) has been theorized to play a neurobiological role in a number of social cognitive abilities and is commonly indexed putatively in humans via interpersonal motor resonance (IMR) and mu suppression. Although both indices are thought to measure similar neuronal populations (i.e., "mirror neurons"), it has been suggested that these methods are unrelated, and therefore, incompatible. However, prior studies reporting no relationships were typically conducted in small and underpowered samples. Thus, we aimed to investigate this potential association in a large sample of neurotypical adults (N = 116; 72 females). Participants underwent transcranial magnetic stimulation (TMS), electromyography (EMG), and electroencephalography (EEG) during the observation of videos of actors performing grasping actions in order to index IMR and mu suppression (in beta, lower alpha, and upper alpha bandwidths). A series of linear regressions revealed no associations between IMR and each of the mu suppression bandwidths. Supplementary Bayesian analyses provided further evidence in favor of the null (B01 = 8.85-8.93), providing further support for no association between the two indices of MNS activity. Our findings suggest that these two measures may indeed be unrelated indices that perhaps assess different neurophysiological aspects of the MNS. These results have important implications for future studies examining the MNS.

Given the rising incidence of stroke, several technology-driven methods for rehabilitation have recently been developed. Virtual reality (VR) is a promising therapeutic technology among them. We recently developed a neuroscientifically grounded VR system to aid recovery of motor function poststroke. The developed system provides unilateral and bilateral upper extremity (UE) training in a fully immersive virtual environment that may stimulate and activate mirror neurons (MNs) in the brain necessary for UE rehabilitation. Twenty-three participants were randomized to a VR group (n = 12) to receive VR intervention (8 h within 2 weeks) plus 8-h occupational therapy (OT) or a control group (n = 11) to receive time-matched OT alone. Treatment effects on motor recovery and cortical reorganization were investigated using the Barthel Index (BI), Fugl-Meyer Upper Extremity (FM-UE), and resting-state fMRI. Both groups significantly improved BI (P < 0.05), reflecting the recovery of UE motor function. The VR group revealed significant improvements on FM-UE scores (P < 0.05) than the control group. Neural activity increased after the intervention, particularly in the brain areas implicating MNs, such as in the primary motor cortex. Overall, results suggested that using a neuroscientifically grounded VR system might offer additional benefits for UE rehabilitation in patients receiving OT.

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 compara-ble 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.

Dysfunctional frontal cortical areas associated with clinical features are observed in children with autism spectrum disorder (ASD). This study attempted to identify any potential therapeutic effects of bilateral anodal transcranial direct current stimulation (tDCS) applied over the left and right prefrontal and motor areas on the clinical characteristics of children with ASD. Fifty children with confirmed ASD medical diagnoses were divided equally and randomly into a tDCS treatment group and a control group. The tDCS treatment group underwent 10 sessions (20-min durations, five per week) of bilateral anodal tDCS stimulation applied simultaneously over the left and right prefrontal and motor areas, whereas the control group underwent the same procedures but with the use of sham tDCS stimulation. Total scores and sub-scores of autism treatment evaluation checklist (ATEC) (language and communication; sociability; sensory awareness; and behavioral, health, and physical conditions) were measured before and after the tDCS treatment sessions of both groups. There were significant decreases in total ATEC scores (P = 0.014), sociability sub-scores (P = 0.021), and behavioral, health, and physical condition sub-scores (P = 0.011) in the tDCS treatment group. No significant changes were observed in total ATEC scores and sub-scores in the control group. In conclusion, compared to the control group, bilateral anodal tDCS showed potential therapeutic effects on children with ASD in terms of improvements in sociability, behavior, health, and physical conditions with no reported side effects. Autism Res 2020, 13: 828-836. © 2020 International Society for Autism Research, Wiley Periodicals, Inc. LAY SUMMARY: Dysfunctional frontal cortical areas are associated with clinical features in children with autism spectrum disorder (ASD). Transcranial direct current stimulation (tDCS) is found to be a safe, noninvasive method to stimulate cortical regions and thus have therapeutic effects on children with ASD. © 2020 International Society for Autism Research, Wiley Periodicals, Inc.

The finding of mirror neurons (MNs) has provided a biological substrate to a new concept of cognition, relating data on actions and perceptions not only to integrate perception in action planning and execution but also as a neural mechanism supporting a wide range of cognitive functions. Here we first summarize data on MN localization and role in primates, then we report findings in normal human subjects: functional magnetic resonance imaging and neurophysiological studies sustain that MNs have a role in motor learning and recognizing actions and intentions of others, and they also support an embodied view of language, empathy, and memory. Then, we detail the results of literature searching on MNs and embodied cognition in Parkinson's disease (PD), frontotemporal dementia (FTD)/amyotrophic lateral sclerosis (ALS), and in mild cognitive impairment (MCI)/Alzheimer's disease (AD). In PD the network of MN could be altered, but its hyperactivation might support motor and cognitive performances at least in early stages. In the ALS/FTD continuum, preliminary evidence points out to an involvement of the MN network, which could explain language and inter-subjectivity deficits shown in patients affected by these clinical entities. In the MCI/AD spectrum, a few recent studies suggest a possible progressive involvement from posterior to anterior areas of the MN network, with the brain putting in place compensatory mechanisms in early stages. Reinterpreting neurodegenerative diseases at the light of the new views about brain organization stemming from the discovery of MN could help to better comprehend clinical manifestations and open new pathways to rehabilitation.

Action Observation Treatment (AOT) has been shown to be effective in the functional recovery of several clinical populations. However, little is known about the neural underpinnings of the clinical efficacy of AOT in children with Cerebral Palsy (CP). Using electroencephalography (EEG), we recorded µ rhythm desynchronization as an index of sensorimotor cortex modulation during a passive action observation task before and after AOT. The relationship between sensorimotor modulation and clinical outcomes was also assessed. Eight children with CP entered the present randomized controlled crossover pilot study in which the experimental AOT preceded or followed a control Videogame Observation Treatment (VOT). Results provide further evidence of the clinical efficacy of AOT for improving hand motor function in CP, as assessed with the Assisting Hand Assessment (AHA) and Melbourne Assessment of Unilateral Upper Limb Function Scale (MUUL). The novel finding is that AOT increases µ rhythm desynchronization at scalp locations corresponding to the hand representation areas. This effect is associated to functional improvement assessed with the MUUL. These preliminary findings, although referred to as a small sample, suggest that AOT may affect upper limb motor recovery in children with CP and modulate the activation of sensorimotor areas, offering a potential neurophysiological correlate to support the clinical utility of AOT.

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 (f MRI ) 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 f MRI 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 MN s across the human brain consider the use of MVPA.

Summary The Mirror Neuron System (MNS) plays a crucial role in action perception and imitative behavior, which is suggested to be impaired in Autism Spectrum Disorders (ASDs). In this review, we discuss the plausibility and empirical evidence of a neural interaction between the MNS, action perception, empathy, imitative behavior, and their impact on social decision making in ASDs. To date, there is no consensus regarding a particular theory in ASDs and its underlying mechanisms. Some theories have completely focused on social difficulties, others have emphasized sensory aspects. Based on the current studies, we suggest a multilayer neural network model including the MNS on a first layer and transforming this information to a higher layer network responsible for reasoning. Future studies with ASD participants combining behavioral tasks with neuroimaging methods and transcranial brain stimulation as well as computational modeling can help validate and complement this suggested model. Moreover, we propose applying the behavioral paradigms, and the neurophysiological markers mentioned in this review article for evaluating psychiatric treatment approaches in ASDs. The investigation of modulating effects of different treatment approaches on the neurophysiological markers of the MNS can help find specific subgroups of ASDs patients and support tailored psychiatric interventions.

When watching performing arts, a wide and complex network of brain processes emerge. These processes can be shaped by professional expertise. When compared to laymen, dancers have enhanced processes in observation of short dance movement and listening to music. But how do the cortical processes differ in musicians and dancers when watching an audio-visual dance performance? In our study, we presented the participants long excerpts from the contemporary dance choreography of Carmen. During multimodal movement of a dancer, theta phase synchrony over the fronto-central electrodes was stronger in dancers when compared to musicians and laymen. In addition, alpha synchrony was decreased in all groups during large rapid movement when compared to nearly motionless parts of the choreography. Our results suggest an enhanced cortical communication in dancers when watching dance and, further, that this enhancement is rather related to multimodal, cognitive and emotional processes than to simple observation of dance movement. peerReviewed

The growth of online shopping increases consumers' dependence on vicarious sensory experiences, such as observing others touching products in commercials. However, empirical evidence on whether observing others' sensory experiences increases purchasing intention is still scarce. In the present study, participants observed others interacting with products in the first- or third-person perspective in video clips, and their neural responses were measured with functional magnetic resonance imaging (fMRI). We investigated (1) whether and how vicariously touching certain products affected purchasing intention, and the neural correlates of this process; and (2) how visual perspective interacts with vicarious tactility. Vicarious tactile experiences were manipulated by hand actions touching or not touching the products, while the visual perspective was manipulated by showing the hand actions either in first- or third-person perspective. During the fMRI scanning, participants watched the video clips and rated their purchasing intention for each product. The results showed that, observing others touching (vs. not touching) the products increased purchasing intention, with vicarious neural responses found in mirror neuron systems (MNS) and lateral occipital complex (LOC). Moreover, the stronger neural activities in MNS was associated with higher purchasing intention. The effects of visual perspectives were found in left superior parietal lobule (SPL), while the interaction of tactility and visual perspective was shown in precuneus and precuneus-LOC connectivity. The present study provides the first evidence that vicariously touching a given product increased purchasing intention and the neural activities in bilateral MNS, LOC, left SPL and precuneus are involved in this process. Hum Brain Mapp 00:000–000, 2017. © 2017 Wiley Periodicals, Inc.

Imitative stimuli presented from a first-person perspective (FPP) produce stronger visuomotor effects than those presented from a third-person perspective (TPP) due to the relatively greater response of the mirror neuron system (MNS) to FPP stimuli. Some previous studies utilizing TPP stimuli have reported no differences in MNS activity between moving and static bodies’ stimuli. However, few studies have compared visuomotor effects of such stimuli when presented in the FPP. To clarify this issue, we measured cortical activation in 17 participants during a functional magnetic resonance imaging (MRI) imitation task involving three conditions: moving (a lifting finger was presented), static (an “X” appeared on a static finger), and control (an “X” appeared on a button). All stimuli were presented from the FPP or TPP. Participants were asked to lift the finger corresponding to the imitative stimulus. In the FPP condition, moving stimuli elicited greater MNS activation than static stimuli. Furthermore, such movement effects were stronger in the MNS and insula (a region associated with body-ownership) for FPP stimuli than for TPP stimuli. Psychophysiological interaction analysis revealed increased connectivity between the MNS and insula for moving stimuli in the FPP condition. These findings suggest that bodily movements presented in the FPP elicit a greater visuomotor response than static body presented in the FPP, and that the visuomotor effects of bodily movements were greater in the FPP condition than in the TPP condition. Our analyses further indicated that such responses are processed via the neural system underlying body-ownership. Hum Brain Mapp, 2017. © 2017 Wiley Periodicals, Inc.

Musical expertise is visible both in the morphology and functionality of the brain. Recent research indicates that functional integration between multi-sensory, somato-motor, default-mode (DMN), and salience (SN) networks of the brain differentiates musicians from non-musicians during resting state. Here, we aimed at determining whether brain networks differentially exchange information in musicians as opposed to non-musicians during naturalistic music listening. Whole-brain graph-theory analyses were performed on participants' fMRI responses. Group-level differences revealed that musicians' primary hubs comprised cerebral and cerebellar sensorimotor regions whereas non-musicians' dominant hubs encompassed DMN-related regions. Community structure analyses of the key hubs revealed greater integration of motor and somatosensory homunculi representing the upper limbs and torso in musicians. Furthermore, musicians who started training at an earlier age exhibited greater centrality in the auditory cortex, and areas related to top-down processes, attention, emotion, somatosensory processing, and non-verbal processing of speech. We here reveal how brain networks organize themselves in a naturalistic music listening situation wherein musicians automatically engage neural networks that are action-based while non-musicians use those that are perception-based to process an incoming auditory stream. Hum Brain Mapp 38:2955-2970, 2017. © 2017 Wiley Periodicals, Inc.

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.

Considering the properties of mirror neurons (MNs) in terms of development and phylogeny, we offer a novel, unifying, and testable account of their evolution according to the available data and try to unify apparently discordant research, including the plasticity of MNs during development, their adaptive value and their phylogenetic relationships and continuity. We hypothesize that the MN system reflects a set of interrelated traits, each with an independent natural history due to unique selective pressures, and propose that there are at least three evolutionarily significant trends that gave raise to three subtypes: hand visuomotor, mouth visuomotor, and audio–vocal. Specifically, we put forward a mosaic evolution hypothesis, which posits that different types of MNs may have evolved at different rates within and among species. This evolutionary hypothesis represents an alternative to both adaptationist and associative models. Finally, the review offers a strong heuristic potential in predicting the circumstances under which specific variations and properties of MNs are expected. Such predictive value is critical to test new hypotheses about MN activity and its plastic changes, depending on the species, the neuroanatomical substrates, and the ecological niche.

There is much disagreement about how extensive a role theoretical mindreading, behavior-reading, and simulation each have and need to have in our knowing and understanding other minds, and how each method is implemented in the brain, but less discussion of the epistemological question what it is about the products of these methods that makes them count as knowledge or understanding. This question has become especially salient recently as some have the intuition that mirror neurons can bring understanding of another’s action despite involving no higher-order processing, whereas most epistemologists writing about understanding think that it requires reflective access to one’s grounds, which is closer to the intuitions of other commenters on mirror neurons. I offer a definition of what it is that makes something understanding that is compelling independently of the context of cognition of other minds, and use it to show two things: 1) that theoretical mind-reading and simulation bring understanding in virtue of the same epistemic feature, and 2) why the kind of motor representation without propositional attitudes that is done by mirror neurons is sufficient for action understanding. I further suggest that more attention should be paid to the potential disadvantages of a simulative method of knowing. Though it can be more efficient in some cases, it can also bring vulnerability, wear and tear on one’s personal equipment, and unintended mimicry.

It has been suggested that an early deficit in the human mirror neuron system (MNS) is an important feature of autism. Recent findings related to simple hand and finger movements do not support a general dysfunction of the MNS in autism. Studies investigating facial actions (e.g., emotional expressions) have been more consistent, however, mostly relied on passive observation tasks. We used a new variant of a compatibility task for the assessment of automatic facial mimicry responses that allowed for simultaneous control of attention to facial stimuli. We used facial electromyography in 18 children and adolescents with Autism spectrum disorder (ASD) and 18 typically developing controls (TDCs). We observed a robust compatibility effect in ASD, that is, the execution of a facial expression was facilitated if a congruent facial expression was observed. Time course analysis of RT distributions and comparison to a classic compatibility task (symbolic Simon task) revealed that the facial compatibility effect appeared early and increased with time, suggesting fast and sustained activation of motor codes during observation of facial expressions. We observed a negative correlation of the compatibility effect with age across participants and in ASD, and a positive correlation between self-rated empathy and congruency for smiling faces in TDC but not in ASD. This pattern of results suggests that basic motor mimicry is intact in ASD, but is not associated with complex social cognitive abilities such as emotion understanding and empathy. Autism Res 2017, 10: 298-310. © 2016 International Society for Autism Research, Wiley Periodicals, Inc.

Very little is known about the action observation network and the mirror neuron system (AON/MNS) in children and its age-related properties compared with those observed in adults. In the present fMRI study we explored the activation of areas belonging to the AON/MNS in children and adults during observation of complex hand-grasping actions, as compared to observation of simple grasping acts executed with the left and the right hand, seen from a first person perspective. The results indicate that during the action observation tasks in children there was activation of a cortical network similar to that found in adults, including the premotor cortex, the posterior part of the inferior frontal gyrus and the posterior parietal lobe. However, the activation in children was more widespread and showed a higher inter-subject variability compared with adults. Furthermore, the activated network seems more lateralized to the left hemisphere in adults and more bilateral in children, with a linear growth of lateralization index as a function of age. Finally, in children the activation in the anterior intraparietal cortex (AIP) of each hemisphere was higher during observation of the contralateral hand (hand identity effect) and during the observation of complex actions relative to simple grasping acts, confirming the role of AIP for action-related hand identity previously described in adults. These results support the assumption that structure and size of action representations are sensitive to mechanisms of development and show physiological plasticity. These properties of the AON/MNS could constitute a powerful tool for spontaneous reorganization and recovery of motor deficits after brain injury in children and in adults, as well as for specific rehabilitation programmes.

Recent research in cognitive science has provided broad empirical support for the model of cognition and self-development formulated by G.H. Mead. This is demonstrated specifically through three distinct cognitive mechanisms: basic-level categories, mirror neurons, and joint-attention schemes. These mechanisms illustrate a form of embodied sociality that underscores the intersubjective foundation of mind and self articulated by Mead. Research on basic-level categories, mirror neurons, and joint-attention schemes further extends Mead's relevance outside standard sociological and social psychological circles to the field of cognitive science and its emerging focus on sociality and social interaction.

Resonance with the inner states of another social actor is regarded as a hallmark of emotional closeness. Nevertheless, sensitivity to potential incongruities between one's own and an intimate partner's subjective experience is reportedly also important for close relationship quality. Here, we tested whether perceivers show greater neurobehavioral responsiveness to a spouse's positive (rather than negative) context-incongruent emotions, and whether this effect is influenced by the perceiver's satisfaction with the relationship. Thus, we used fMRI to scan older long-term married female perceivers while they judged either their spouse's or a stranger's affect, based on incongruent nonverbal and verbal cues. The verbal cues were selected to evoke strongly polarized affective responses. Higher perceiver marital satisfaction predicted greater neural processing of the spouse's (rather than the strangers) nonverbal cues. Nevertheless, across all perceivers, greater neural processing of a spouse's (rather than a stranger's) nonverbal behavior was reliably observed only when the behavior was positive and the context was negative. The spouse's positive (rather than negative) nonverbal behavior evoked greater activity in putative mirror neuron areas, such as the bilateral inferior parietal lobule (IPL). This effect was related to a stronger inhibitory influence of cognitive control areas on mirror system activity in response to a spouse's negative nonverbal cues, an effect that strengthened with increasing perceiver marital satisfaction. Our valence-asymmetric findings imply that neurobehavioral responsiveness to a close other's emotions may depend, at least partly, on cognitive control resources, which are used to support the perceiver's interpersonal goals (here, goals that are relevant to relationship stability).

A lack of imitative behavior is frequently described as a core feature of Autism Spectrum Disorder (ASD), and is consistent with claims of mirror neuron system dysfunction in these individuals. Previous research has questioned this characterization of ASD however, arguing that when tests of automatic imitation are used--which do not require higher-level cognitive processing--imitative behavior is intact or even enhanced in individuals with ASD. In Experiment 1, 60 adult individuals with ASD and a matched Control group completed an automatic imitation task in which they were required to perform an index or a middle finger lift while observing a hand making either the same, or the alternate, finger movement. Both groups demonstrated a significant imitation effect whereby actions were executed faster when preceded by observation of the same action, than when preceded by the alternate action. The magnitude of this "imitation effect" was statistically indistinguishable in the ASD and Control groups. Experiment 2 utilized an improved automatic imitation paradigm to demonstrate that, when automatic imitation effects are isolated from those due to spatial compatibility, increasing autism symptom severity is associated with an increased tendency to imitate. Notably, there was no association between autism symptom severity and spatial compatibility, demonstrating the specificity of the link between ASD symptoms and increased imitation. These results provide evidence against claims of a lack of imitative behavior in ASD, and challenge the "Broken Mirror Theory of Autism."

Previous studies using neural activity recording and neuroimaging techniques have reported functional deficits in the mirror neuron system (MNS) for individuals with autism spectrum disorder (ASD). However, a few studies focusing on gray and white matter structures of the MNS have yielded inconsistent results. The current study recruited adolescents and young adults with ASD (aged 15-26 years) and age-matched typically developing (TD) controls (aged 14-25 years). The cortical thickness (CT) and microstructural integrity of the tracts connecting the regions forming the classical MNS were investigated. High-resolution T1-weighted imaging and diffusion spectrum imaging were performed to quantify the CT and tract integrity, respectively. The structural covariance of the CT of the MNS regions revealed a weaker coordination of the MNS network in ASD. A strong correlation was found between the integrity of the right frontoparietal tracts and the social communication subscores measured by the Chinese version of the Social Communication Questionnaire. The results showed that there were no significant mean differences in the CTs and tract integrity between the ASD and TD groups, but revealed a moderate or even reverse age effect on the frontal MNS structures in ASD. In conclusion, aberrant structural coordination may be an underlying factor affecting the function of the MNS in ASD patients. The association between the right frontoparietal tracts and social communication performance implies a neural correlate of communication processing in the autistic brain. This study provides evidence of abnormal MNS structures and their influence on social communication in individuals with ASD.

Social impairments in individuals with autism spectrum disorders (ASD) may be in part due to difficulty perceiving and recognizing the actions of others. Evidence from imitation studies, which involves both observation and execution of an action, suggests differences, in individuals with ASD, between the ability to imitate goal-directed actions involving objects (transitive actions) and the ability to imitate actions that do not involve objects (intransitive actions). In the present study, we examined whether there were differences in how ASD adolescents encoded transitive and intransitive actions compared to typically developing (TD) adolescents, by having participants view videos of a hand reaching across a screen toward an object or to where an object would be while functional magnetic resonance images were collected. Analyses focused on areas within the action observation network (AON), which is activated during the observation of actions performed by others. We hypothesized that the AON would differentiate transitive from intransitive actions only in the ASD group. However, results revealed that object presence modulated activity in the right inferior frontal gyrus and supramarginal gyrus of the TD group, a differentiation that was not seen in the ASD group. Furthermore, there were no significant group differences between the TD and ASD groups in any of the conditions. This suggests that there is not a global deficit of the AON in individuals with ASD while observing transitive and intransitive actions.

The discovery of mirror neurons compellingly shows that the monkey premotor area F5 is active not only during the execution but also during the observation of goal-directed motor acts. Previous studies have addressed the functioning of the mirror-neuron system at the single-unit level. Here, we tackled this research question at the network level by analysing local field potentials in area F5 while the monkey was presented with goal-directed actions executed by a human or monkey actor and observed either from a first-person or third-person perspective. Our analysis showed that rhythmic responses are not only present in area F5 during action observation, but are also modulated by the point of view. Observing an action from a subjective point of view produced significantly higher power in the low-frequency band (2-10 Hz) than observing the same action from a frontal view. Interestingly, an increase in power in the 2-10 Hz band was also produced by the execution of goal-directed motor acts. Independently of the point of view, action observation also produced a significant decrease in power in the 15-40 Hz band and an increase in the 60-100 Hz band. These results suggest that, depending on the point of view, action observation might activate different processes in area F5. Furthermore, they may provide information about the functional architecture of action perception in primates.

Mirror neurons, originally described in the monkey premotor area F5, are embedded in a frontoparietal network for action execution and observation. A similar Mirror Neuron System (MNS) exists in humans, including precentral gyrus, inferior parietal lobule, and superior temporal sulcus. Controversial is the inclusion of Broca's area, as homologous to F5, a relevant issue in light of the mirror hypothesis of language evolution, which postulates a key role of Broca's area in action/speech perception/production. We assess “mirror” properties of this area by combining neuroimaging and intraoperative neurophysiological techniques. Our results show that Broca's area is minimally involved in action observation and has no motor output on hand or phonoarticulatory muscles, challenging its inclusion in the MNS. The presence of these functions in premotor BA6 makes this area the likely homologue of F5 suggesting that the MNS may be involved in the representation of articulatory rather than semantic components of speech. Hum Brain Mapp 36:1010–1027, 2015. © 2014 Wiley Periodicals, Inc.

A number of psychoanalysts have become excited about mirror neurons, as they are called by neuroscientists. Mirror neurons have the remarkable property of responding identically to an action I intend as well as an action you intend. The argument of some psychoanalysts is that mirror neurons open a new pathway to understanding the intentions of other. They make possible a new type of empathy, more direct and less mediated by the typical defenses. One result of such a perspective on psychoanalysis is the virtual death of the countertransference. If one has direct empathic contact with another mind, then countertransferential experience is only a barrier, not a guide. The essay not only looks at the evidence for mirror neurons, which is ambiguous, but also at what need they might be filling in our contemporary culture. Copyright © 2014 John Wiley & Sons, Ltd.

Action perception and recognition are core abilities fundamental for human social interaction. A parieto-frontal network (the mirror neuron system) matches visually presented biological motion information onto observers' motor representations. This process of matching the actions of others onto our own sensorimotor repertoire is thought to be important for action recognition, providing a non-mediated "motor perception" based on a bidirectional flow of information along the mirror parieto-frontal circuits. State-of-the-art machine learning strategies for hand action identification have shown better performances when sensorimotor data, as opposed to visual information only, are available during learning. As speech is a particular type of action (with acoustic targets), it is expected to activate a mirror neuron mechanism. Indeed, in speech perception, motor centers have been shown to be causally involved in the discrimination of speech sounds. In this paper, we review recent neurophysiological and machine learning-based studies showing (a) the specific contribution of the motor system to speech perception and (b) that automatic phone recognition is significantly improved when motor data are used during training of classifiers (as opposed to learning from purely auditory data).

Birdsong is a form of sensorimotor learning that involves a mirror-like system that activates with both song hearing and production. Early models of song learning, based on behavioral measures, identified key features of vocal plasticity, such as the requirements for memorization of a tutor song and auditory feedback during song practice. The concept of a comparator, which compares the memory of the tutor song to auditory feedback, featured prominently. Later models focused on linking anatomically-defined neural modules to behavioral concepts, such as the comparator. Exploiting the anatomical modularity of the songbird brain, localized lesions illuminated mechanisms of the neural song system. More recent models have integrated neuronal mechanisms identified in other systems with observations in songbirds. While these models explain multiple aspects of song learning, they must incorporate computational elements based on unknown biological mechanisms to bridge the motor-to-sensory delay and/or transform motor signals into the sensory domain. Here, I introduce the stabilizing critic hypothesis, which enables sensorimotor learning by (1) placing a purely sensory comparator afferent of the song system and (2) endowing song system disinhibitory interneuron networks with the capacity both to bridge the motor-sensory delay through prolonged bursting and to stabilize song segments selectively based on the comparator signal. These proposed networks stabilize an otherwise variable signal generated by both putative mirror neurons and a cortical-basal ganglia-thalamic loop. This stabilized signal then temporally converges with a matched premotor signal in the efferent song motor cortex, promoting spike-timing-dependent plasticity in the premotor circuitry and behavioral song learning.

Investigating the underlying neural mechanisms of autism spectrum disorder (ASD) has recently been influenced by the discovery of mirror neurons. These neurons, active during both observation and execution of actions, are thought to play a crucial role in imitation and other social-communicative skills that are often impaired in ASD. In the current electroencephalographic study, we investigated mu suppression, indicating neural mirroring in children with ASD between the ages of 24 and 48 months and age-matched typically developing children, during observation of goal-directed actions and non-goal-directed mimicked hand movements, as well as during action execution. Results revealed no significant group differences with significant central mu suppression in the ASD children and control children during both execution and observation of goal-directed actions and during observation of hand movements. Furthermore, no significant correlations between mu suppression on one hand and quality of imitation, age, and social communication questionnaire scores on the other hand were found. These findings challenge the "broken mirror" hypothesis of ASD, suggesting that impaired neural mirroring is not a distinctive feature of ASD.

Sobell Department of Motor Neuroscience and Movement Disorders, Institute of Neurology, Queen Square, London,United KingdomAccepted 21 November 2013ABSTRACT: We propose that amyotrophic lateral sclerosis(ALS), and frontotemporal dementia may be viewed as a failureof interlinked functional complexes having their origins in keyevolutionary adaptations. We discuss how hand-arm function,locomotion, brainstem function (involving vocalization/speech,swallowing and breathing), and cognitive impairment share com-plex, interdependent evolutionary adaptations that can be tracedback several million years. Fine movements of the hand facili-tated tool-making and use enhanced by development of bipedal-ism. Development of the larynx and integration of respiratorycontrol were central to vocalization, which when combined withgesture are intermediary to human language. These adaptationswere accompanied by progressive encephalization, with develop-ment of Theory of Mind to facilitate socialization. The varied clini-cal phenotypes of ALS can thus be understood in the context ofinter-related functional complexes that subserve “Tools and Talk”;they have a long evolutionary history and are related to specificdevelopmental neural and gene networks.

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.

Motor learning has mostly been studied in laboratory settings where participants execute the task alone, unlike in real life, where people perform activities together with others. Recent discovery of mirror neuron system (MNS)—linking perception of others' actions and execution of our own—makes the scarcity of studies on social aspects of motor learning even more peculiar. Here, we investigated motor learning during interaction with a partner. Participants engaged in 2 consecutive sessions executing the task alone or in pairs. We found that social interaction improved motor performance only when participants had prior individual experience with the task. This effect was amplified if subjects could first observe their partners' actions, suggesting that it may be mediated by prior MNS configuration.

Previous studies suggest that about 50% of rhesus macaque infants engage in neonatal imitation of facial gestures. Here we measured whether individual differences in newborn macaques' (n = 49) visual attention may explain why some infants imitate lipsmacking (LPS) and tongue protrusion (TP) gestures. LPS imitators, but not TP imitators, looked more to a human experimenter's face and to a control stimulus compared to nonimitators (p = .017). LPS imitation was equally accurate when infants were looking at faces and when they were looking away (p = .221); TP imitation was more accurate when infants were looking at faces (p = .001). Potentially, less attention is necessary for LPS imitation compared to TP imitation, as LPS is part of macaques' natural communicative repertoire. These findings suggest that facial gestures may differentially engage imitators and nonimitators, and infants' visual attention during neonatal assessments may uncover the conditions that support this skill.

The goal of the current study was to determine whether activation of the mirror neuron system, as measured by mu rhythm desynchronization, varied as a function of image content in smokers compared with nonsmokers. EEG activity was recorded while participants passively viewed images depicting smoking-related and nonsmoking-related stimuli. In half of the images, cues were depicted alone (inactive), while for the remaining images, cues were depicted with humans interacting with them (active). For the nonsmoking stimuli, smokers and nonsmokers showed greater mu suppression to the active cues compared to the inactive cues. However, for the smoking-related stimuli, smokers showed greater perception-action coupling for the active cues as reflected in their enhanced mu suppression, compared to nonsmokers. The results of the current study support the involvement of the perceptual-motor system in the activation of motivated drug use behaviors.

Mirror neurons are widely regarded as an important key to social cognition. Despite such wide agreement, there is very little consensus on how or why they are important. The goal of this article is to explicate clearly the exact role mirror neurons play in social cognition. I aim to answer two questions about the relationship between mirroring and social cognition: What kind of social understanding is involved with mirroring? How is mirroring related to that understanding? I argue that philosophical and empirical considerations lead us to accord a fairly minimal role for mirror neurons in social cognition.

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.

Since their discovery in the early 1990s, mirror neurons have been proposed to be related to many social-communicative abilities, such as imitation. However, research into the early manifestations of the putative neural mirroring system and its role in early social development is still inconclusive. In the current EEG study, mu suppression, generally thought to reflect activity in neural mirroring systems was investigated in 18- to 30-month-olds during the observation of object manipulations as well as mimicked actions. EEG power data recorded from frontal, central, and parietal electrodes were analysed. As predicted, based on previous research, mu wave suppression was found over central electrodes during action observation and execution. In addition, a similar suppression was found during the observation of intransitive, mimicked hand movements. To a lesser extent, the results also showed mu suppression at parietal electrode sites, over all three conditions. Mu wave suppression during the observation of hand movements and during the execution of actions was significantly correlated with quality of imitation, but not with age or language level.

Since the beginnings of speech research, the issue of variability has been a central topic in the field. Until recently, solutions to this problem have focused mainly on resolving linguistic forms from variable acoustic realizations, with little systematic consideration of talker variability or social settings. However, acoustic-phonetic variability within and between talkers does not arise by chance – such variation is attributable to a variety of causes including talker physiology, dialect, affect, and social/situational factors. Studies of conversational interaction demonstrate the influence of situational factors on many aspects of spoken communication. In particular, interacting talkers converge and diverge in acoustic-phonetic form over the course of a single conversational setting. Although phonetic convergence likely reflects a close connection between speech perception and production, it is not an automatic consequence of priming mechanisms or a mirror neuron system. In studies of the neural activity associated with speech perception, it is increasingly clear that utterances are processed in separate dorsal and ventral cortical streams, one that is part of the self-regulation of spoken utterances, and another that is part of conceptual understanding. Links between perception and production appear to be part of the self-regulatory system that participates in speech production control monitoring, which makes this system a likely candidate for talker control of acoustic-phonetic variability. Whether talkers converge or diverge depends on multiple aspects of the social setting, which have a much greater influence on the form of phonetic expression than automatic links or mirror mechanisms.

Several studies have shown activation of the mirror neuron system (MNS), comprising the temporal, posterior parietal, and sensorimotor areas when observing plausible actions, but far less is known on how these cortical areas interact during the recognition of a plausible action. Here, we recorded neural activity with magnetoencephalography while subjects viewed point-light displays of biologically plausible and scrambled versions of actions. We were interested in modulations of oscillatory activity and, specifically, in coupling of oscillatory activity between visual and motor areas. Both plausible and scrambled actions elicited modulations of θ (5-7 Hz), α (7-13 Hz), β (13-35 Hz), and γ (55-100 Hz) power within visual and motor areas. When comparing between the two actions, we observed sequential and spatially distinct increases of γ (∼65 Hz), β (∼25 Hz), and α (∼11 Hz) power between 0.5 and 1.3 s in parieto-occipital, sensorimotor, and left temporal areas. In addition, significant clusters of γ (∼65 Hz) and α/β (∼15 Hz) power decrease were observed in right temporal and parieto-occipital areas between 1.3 and 2.0 s. We found β-power in sensorimotor areas to be positively correlated on a trial-by-trial basis with parieto-occipital γ and left temporal α-power for the plausible but not for the scrambled condition. These results provide new insights in the neuronal oscillatory activity of the areas involved in the recognition of plausible action movements and their interaction. The power correlations between specific areas underscore the importance of interactions between visual and motor areas of the MNS during the recognition of a plausible action.

There is still no clear consensus as to which of the many functional and structural changes in the brain in schizophrenia are of most importance, although the main focus to date has been on those in the frontal and cingulate cortices. In the present study, we have used a novel holistic approach to identify brain-wide functional connectivity changes in medicated schizophrenia patients, and functional connectivity changes were analyzed using resting-state fMRI data from 69 medicated schizophrenia patients and 62 healthy controls. As far as we are aware, this is the largest population reported in the literature for a resting-state study. Voxel-based morphometry was also used to investigate gray and white matter volume changes. Changes were correlated with illness duration/symptom severity and a support vector machine analysis assessed predictive validity. A network involving the inferior parietal lobule, superior parietal gyrus, precuneus, superior marginal, and angular gyri was by far the most affected (68% predictive validity compared with 82% using all connections) and different components correlated with illness duration and positive and negative symptom severity. Smaller changes occurred in emotional memory and sensory and motor processing networks along with weakened interhemispheric connections. Our findings identify the key functional circuitry altered in schizophrenia involving the default network midline cortical system and the cortical mirror neuron system, both playing important roles in sensory and cognitive processing and particularly self-processing, all of which are affected in this disorder. Interestingly, the functional connectivity changes with the strongest links to schizophrenia involved parietal rather than frontal regions. Hum Brain Mapp, 2012.