Occlusion of LTP-like plasticity in human primary motor cortex by action observation

Passive observation of motor actions induces cortical activity in the primary motor cortex (M1) of the onlooker, which could potentially contribute to motor learning. While recent studies report modulation of motor performance following action observation, the neurophysiological mechanism supporting these behavioral changes remains to be specifically defined. Here, we assessed whether the observation of a repetitive thumb movement--similarly to active motor practice--would inhibit subsequent long-term potentiation-like (LTP) plasticity induced by paired-associative stimulation (PAS). Before undergoing PAS, participants were asked to either 1) perform abductions of the right thumb as fast as possible; 2) passively observe someone else perform thumb abductions; or 3) passively observe a moving dot mimicking thumb movements. Motor evoked potentials (MEP) were used to assess cortical excitability before and after motor practice (or observation) and at two time points following PAS. Results show that, similarly to participants in the motor practice group, individuals observing repeated motor actions showed marked inhibition of PAS-induced LTP, while the "moving dot" group displayed the expected increase in MEP amplitude, despite differences in baseline excitability. Interestingly, LTP occlusion in the action-observation group was present even if no increase in cortical excitability or movement speed was observed following observation. These results suggest that mere observation of repeated hand actions is sufficient to induce LTP, despite the absence of motor learning.     

Impaired motor facilitation during action observation in individuals with autism spectrum disorder

It has been suggested that social impairments observed in individuals with autism spectrum disorder (ASD) can be partly explained by an abnormal mirror neuron system (MNS) 1., 2.. Studies on monkeys have shown that mirror neurons are cells in premotor area F5 that discharge when a monkey executes or sees a specific action or when it hears the corresponding action-related sound 3., 4., 5.. Evidence for the presence of a MNS in humans comes in part from studies using transcranial magnetic stimulation (TMS), where a change in the amplitude of the TMS-induced motor-evoked potentials (MEPs) during action observation has been demonstrated 6., 7., 8., 9.. These data suggest that actions are understood when the representation of that action is mapped onto the observer's own motor structures [10]. To determine if the neural mechanism matching action observation and execution is anomalous in individuals with ASD, TMS was applied over the primary motor cortex (M1) during observation of intransitive, meaningless finger movements. We show that overall modulation of M1 excitability during action observation is significantly lower in individuals with ASD compared with matched controls. In addition, we find that basic motor cortex abnormalities do not underlie this impairment.     

Aplasics born without hands mirror the goal of hand actions with their feet

The premotor and parietal mirror neuron system (MNS) is thought to contribute to the understanding of observed actions by mapping them onto "corresponding" motor programs of the observer [1-24], but how would the MNS respond to the observation of hand actions if the observer never had hands? Would it not show changes of blood-oxygen-level dependent (BOLD) signal, because the observer lacks motor programs that can resonate [12, 25, 26], or would it show significant changes because the observer has motor programs for the foot or mouth with corresponding goals [15, 17, 19, 27, 28]? We scanned two aplasic subjects, born without arms or hands, while they watched hand actions and compared their brain activity with that of 16 control subjects. All subjects additionally executed actions with different effectors (feet, mouth, and, for controls, hands). The BOLD signal of aplasic individuals within the putative MNS was augmented when they watched hand actions, demonstrating the brain's capacity to mirror actions that deviate from the embodiment of the observer by recruiting voxels involved in the execution of actions that achieve corresponding goals by different effectors. This sheds light on the functional organization of the MNS and predominance of goals in imitation.     

How frontoparietal brain regions mediate imitative and complementary actions: an FMRI study

The 'mirror neuron system' (MNS), located within inferior parietal lobe (IPL) and inferior frontal gyrus (IFG), creates an internal motor representation of the actions we see and has been implicated in imitation. Recently, the MNS has been implicated in non-identical responses: when the actions we must execute do not match those that we observe. However, in such conflicting situations non action-specific cognitive control networks also located in frontoparietal regions may be involved. In the present functional magnetic resonance imaging (fMRI) study participants made both similar and dissimilar actions within two action contexts: imitative and complementary. We aimed to determine whether activity within IPL/IFG depends on (i) responding under an imitative versus complementary context (ii) responding with similar versus dissimilar responses, and (iii) observing hand actions versus symbolic arrow cue stimuli. Activity within rIPL/rIFG regions was largest during observation of hand actions compared with arrow cues. Specifically, rIPL/rIFG were recruited only during the imitative context, when participants responded with similar actions. When responding to symbolic arrow cues, rIPL/rIFG activity increased during dissimilar responses, reflecting increased demands placed on general cognitive control mechanisms. These results suggest a specific role of rIPL/rIFG during imitation of hand actions, and also a general role of frontoparietal areas in mediating dissimilar responses to both hand actions and symbolic stimuli. We discuss our findings in relation to recent work that has examined the role of frontoparietal brain structures in joint-actions and inter-actor cooperation. We conclude that the specific brain regions identified here to show increased activation during action observation conditions are likely to form part of a mechanism specifically involved in matching observed actions directly with internal motor plans. Conversely, observation of arrow cues recruited part of a wider cognitive control network involved in the rapid remapping of stimulus-response associations.     

Motor simulation without motor expertise: enhanced corticospinal excitability in visually experienced dance spectators

The human "mirror-system" is suggested to play a crucial role in action observation and execution, and is characterized by activity in the premotor and parietal cortices during the passive observation of movements. The previous motor experience of the observer has been shown to enhance the activity in this network. Yet visual experience could also have a determinant influence when watching more complex actions, as in dance performances. Here we tested the impact visual experience has on motor simulation when watching dance, by measuring changes in corticospinal excitability. We also tested the effects of empathic abilities. To fully match the participants' long-term visual experience with the present experimental setting, we used three live solo dance performances: ballet, Indian dance, and non-dance. Participants were either frequent dance spectators of ballet or Indian dance, or "novices" who never watched dance. None of the spectators had been physically trained in these dance styles. Transcranial magnetic stimulation was used to measure corticospinal excitability by means of motor-evoked potentials (MEPs) in both the hand and the arm, because the hand is specifically used in Indian dance and the arm is frequently engaged in ballet dance movements. We observed that frequent ballet spectators showed larger MEP amplitudes in the arm muscles when watching ballet compared to when they watched other performances. We also found that the higher Indian dance spectators scored on the fantasy subscale of the Interpersonal Reactivity Index, the larger their MEPs were in the arms when watching Indian dance. Our results show that even without physical training, corticospinal excitability can be enhanced as a function of either visual experience or the tendency to imaginatively transpose oneself into fictional characters. We suggest that spectators covertly simulate the movements for which they have acquired visual experience, and that empathic abilities heighten motor resonance during dance observation.     

Viewing lip forms: cortical dynamics

Viewing other persons' actions automatically activates brain areas belonging to the mirror-neuron system (MNS) assumed to link action execution and observation. We followed, by magnetoencephalographic cortical dynamics, subjects who observed still pictures of lip forms, on-line imitated them, or made similar forms in a self-paced manner. In all conditions and in both hemispheres, cortical activation progressed in 20-70 ms steps from the occipital cortex to the superior temporal region (where the strongest activation took place), the inferior parietal lobule, and the inferior frontal lobe (Broca's area), and finally, 50-140 ms later, to the primary motor cortex. The signals of Broca's area and motor cortex were significantly stronger during imitation than other conditions. These results demonstrate that still pictures, only implying motion, activate the human MNS in a well-defined temporal order.     

Observing a Movement Correction during Walking Affects Evoked Responses but Not Unperturbed Walking

Seeing an action activates neurons in the premotor, motor, and somatosensory cortex. Since a significant fraction of these pyramidal neurons project to the spinal motor circuits, a central question is why we do not automatically perform the actions that we see. Indeed, seeing an action increases both cortical and spinal excitability of consistent motor patterns that correspond to the observed ones. Thus, it is believed that such imitative motor patterns are either suppressed or remain at a sub-threshold level. This would predict, however, that seeing someone make a corrective movement while one is actively involved in the same action should either suppress evoked responses or suppress or modulate the action itself. Here we tested this prediction, and found that seeing someone occasionally stepping over an obstacle while walking on a treadmill did not affect the normal walking pattern at all. However, cutaneously evoked reflexes in the anterior tibial and soleus muscles were modulated as if the subject was stepping over an obstacle. This result thus indicates that spinal activation was not suppressed and was neither at sub-threshold motor resonance. Rather, the spinal modulation from observed stepping reflects an adaptive mechanism for regulating predictive control mechanisms. We conclude that spinal excitability during action observation is not an adverse side-effect of action understanding but reflects adaptive and predictive motor control.     

Young children with autism spectrum disorder use predictive eye movements in action observation

Does a dysfunction in the mirror neuron system (MNS) underlie the social symptoms defining autism spectrum disorder (ASD)? Research suggests that the MNS matches observed actions to motor plans for similar actions, and that these motor plans include directions for predictive eye movements when observing goal-directed actions. Thus, one important question is whether children with ASD use predictive eye movements in action observation. Young children with ASD as well as typically developing children and adults were shown videos in which an actor performed object-directed actions (human agent condition). Children with ASD were also shown control videos showing objects moving by themselves (self-propelled condition). Gaze was measured using a corneal reflection technique. Children with ASD and typically developing individuals used strikingly similar goal-directed eye movements when observing others’ actions in the human agent condition. Gaze was reactive in the self-propelled condition, suggesting that prediction is linked to seeing a hand–object interaction. This study does not support the view that ASD is characterized by a global dysfunction in the MNS.     

Interpersonal Liking Modulates Motor-Related Neural Regions

Observing someone perform an action engages brain regions involved in motor planning, such as the inferior frontal, premotor, and inferior parietal cortices. Recent research suggests that during action observation, activity in these neural regions can be modulated by membership in an ethnic group defined by physical differences. In this study we expanded upon previous research by matching physical similarity of two different social groups and investigating whether likability of an outgroup member modulates activity in neural regions involved in action observation. Seventeen Jewish subjects were familiarized with biographies of eight individuals, half of the individuals belonged to Neo-Nazi groups (dislikable) and half of which did not (likable). All subjects and actors in the stimuli were Caucasian and physically similar. The subjects then viewed videos of actors portraying the characters performing simple motor actions (e.g. grasping a water bottle and raising it to the lips), while undergoing fMRI. Using multivariate pattern analysis (MVPA), we found that a classifier trained on brain activation patterns successfully discriminated between the likable and dislikable action observation conditions within the right ventral premotor cortex. These data indicate that the spatial pattern of activity in action observation related neural regions is modulated by likability even when watching a simple action such as reaching for a cup. These findings lend further support for the notion that social factors such as interpersonal liking modulate perceptual processing in motor-related cortices.     

Brain Activation in Motor Sequence Learning Is Related to the Level of Native Cortical Excitability

Cortical excitability may be subject to changes through training and learning. Motor training can increase cortical excitability in motor cortex, and facilitation of motor cortical excitability has been shown to be positively correlated with improvements in performance in simple motor tasks. Thus cortical excitability may tentatively be considered as a marker of learning and use-dependent plasticity. Previous studies focused on changes in cortical excitability brought about by learning processes, however, the relation between native levels of cortical excitability on the one hand and brain activation and behavioral parameters on the other is as yet unknown. In the present study we investigated the role of differential native motor cortical excitability for learning a motor sequencing task with regard to post-training changes in excitability, behavioral performance and involvement of brain regions. Our motor task required our participants to reproduce and improvise over a pre-learned motor sequence. Over both task conditions, participants with low cortical excitability (CElo) showed significantly higher BOLD activation in task-relevant brain regions than participants with high cortical excitability (CEhi). In contrast, CElo and CEhi groups did not exhibit differences in percentage of correct responses and improvisation level. Moreover, cortical excitability did not change significantly after learning and training in either group, with the exception of a significant decrease in facilitatory excitability in the CEhi group. The present data suggest that the native, unmanipulated level of cortical excitability is related to brain activation intensity, but not to performance quality. The higher BOLD mean signal intensity during the motor task might reflect a compensatory mechanism in CElo participants.     

Male human motor cortex stimulus-response characteristics are not altered by aging

Evidence suggests that there are aging-related changes in corticospinal stimulus-response curve characteristics in later life. However, there is also limited evidence that these changes may only be evident in postmenopausal women and not in men. This study compared corticospinal stimulus-response curves from a group of young men [19.8 ± 1.6 yr (range 17-23 yr)] and a group of old men [n = 18, aged 64.1 ± 5.0 yr (range 55-73 yr)]. Transcranial magnetic stimulation (TMS) over the contralateral motor cortex was used to evoke motor potentials at a range of stimulus intensities in the first dorsal interosseous muscle of each hand separately. There was no effect of age group or hemisphere (i.e., left vs. right motor cortex) on motor evoked potential (MEP) amplitude or any other stimulus-response characteristic. MEP variability was strongly modulated by resting motor threshold but not by age. M-wave (but not F-wave) amplitude was reduced in old men, but expressing MEP amplitude as a ratio of M-wave amplitude did not reveal any age-related differences in cortically evoked stimulus-response characteristics. We conclude that male corticospinal stimulus-response characteristics are not altered by advancing age and that previously reported age-related changes in motor cortical excitability assessed with TMS are likely due to changes inherent in the female participants only. Future studies are warranted to fully elucidate the relationship between, and functional significance of, changes in circulating neuroactive sex hormones and motor function in later life.     

Gender differences in changes of motor cortex excitability during elevated blood lactate levels

Gender differences in cortical excitability have been detected by using transcranial magnetic stimulation (TMS). The present study was carried out to compare the effects of high blood lactate levels, induced by performing a maximal exhausting exercise, on the excitability of the primary motor cortex in young male and female athletes. The study was carried out on 21 young males and 20 females from the Middle Distance Track Team of our university. Before the exercise, at the end, as well as 5 and 10 min after the conclusion, venous blood lactate and glucose were measured and excitability of the motor cortex was evaluated by using TMS. We observed a similar enhancement of excitability of primary motor cortex, concomitantly with an increase of blood lactate, in both young male and female athletes. However, the improvement was significantly higher (p < 0.05) in women (37.4% ± 3.97) than in men (42.0% ± 6.43), suggesting a greater sensitiveness of female cerebral cortex to blood lactate.     

Dissociation of motor task-induced cortical excitability and pain perception changes in healthy volunteers

Background

There is evidence that interventions aiming at modulation of the motor cortex activity lead to pain reduction. In order to understand further the role of the motor cortex on pain modulation, we aimed to compare the behavioral (pressure pain threshold) and neurophysiological effects (transcranial magnetic stimulation (TMS) induced cortical excitability) across three different motor tasks.

Methodology/Principal Findings

Fifteen healthy male subjects were enrolled in this randomized, controlled, blinded, cross-over designed study. Three different tasks were tested including motor learning with and without visual feedback, and simple hand movements. Cortical excitability was assessed using single and paired-pulse TMS measures such as resting motor threshold (RMT), motor-evoked potential (MEP), intracortical facilitation (ICF), short intracortical inhibition (SICI), and cortical silent period (CSP). All tasks showed significant reduction in pain perception represented by an increase in pressure pain threshold compared to the control condition (untrained hand). ANOVA indicated a difference among the three tasks regarding motor cortex excitability change. There was a significant increase in motor cortex excitability (as indexed by MEP increase and CSP shortening) for the simple hand movements.

Conclusions/Significance

Although different motor tasks involving motor learning with and without visual feedback and simple hand movements appear to change pain perception similarly, it is likely that the neural mechanisms might not be the same as evidenced by differential effects in motor cortex excitability induced by these tasks. In addition, TMS-indexed motor excitability measures are not likely good markers to index the effects of motor-based tasks on pain perception in healthy subjects as other neural networks besides primary motor cortex might be involved with pain modulation during motor training.     

Human motor cortex excitability during the perception of others' action

Neuroscience research during the past ten years has fundamentally changed the traditional view of the motor system. In monkeys, the finding that premotor neurons also discharge during visual stimulation (visuomotor neurons) raises new hypotheses about the putative role played by motor representations in perceptual functions. Among visuomotor neurons, mirror neurons might be involved in understanding the actions of others and might, therefore, be crucial in interindividual communication. Functional brain imaging studies enabled us to localize the human mirror system, but the demonstration that the motor cortex dynamically replicates the observed actions, as if they were executed by the observer, can only be given by fast and focal measurements of cortical activity. Transcranial magnetic stimulation enables us to instantaneously estimate corticospinal excitability, and has been used to study the human mirror system at work during the perception of actions performed by other individuals. In the past ten years several TMS experiments have been performed investigating the involvement of motor system during others' action observation. Results suggest that when we observe another individual acting we strongly 'resonate' with his or her action. In other words, our motor system simulates underthreshold the observed action in a strictly congruent fashion. The involved muscles are the same as those used in the observed action and their activation is temporally strictly coupled with the dynamics of the observed action.     

Modulation of corticospinal excitability during acquisition of action sequences by observation

Excitability of the corticospinal pathway increases during observation of an action. However, how corticospinal excitability changes during observation of sequential actions in the course of acquiring novel skills (observational learning) remains unexplored. To investigate this, we used a previously unpracticed sequence of ten hand postures. Participants were asked to repeat observation and replication of the sequence. This block of observation and replication was repeated 5 times. During observation of a given hand posture (OK sign), motor-evoked potentials (MEPs) elicited by transcranial magnetic stimulation were recorded from hand muscles. In experiment 1, the OK sign appeared in the 9th position of the sequence. Almost all participants could replicate the OK sign only at the 5th block of the experiment. MEP amplitude was greater than that in the control, and decreased with the stages. This suggested that during observational learning of sequential hand postures MEP changed with the progress of the learning. To evaluate this idea, we performed two additional experiments. In experiment 2, the OK sign appeared in the 2nd position. Almost all participants replicated the OK sign even in the 1st block. The MEP amplitude did not change across stages. In experiment 3, the OK sign appeared in the 9th position, but the order of other signs was randomized in every stage. Many participants were not able to replicate the OK sign even during the 5th block of the experiment. The MEP amplitude did not change across stages. These results suggest that: (1) During observational learning modulation of corticospinal excitability is associated with the learning process. (2) Corticospinal excitability decreases as learning progresses.     

The role of ventral premotor cortex in action execution and action understanding

The human ventral premotor cortex overlaps, at least in part, with Broca's region in the dominant cerebral hemisphere, that is known to mediate the production of language and contributes to language comprehension. This region is constituted of Brodmann's areas 44 and 45 in the inferior frontal gyrus. We summarize the evidence that the motor related part of Broca's region is localized in the opercular portion of the inferior frontal cortex, mainly in area 44 of Brodmann. According to our own data, there seems to be a homology between Brodmann area 44 in humans and the monkey area F5. The non-language related motor functions of Broca's region comprise complex hand movements, associative sensorimotor learning and sensorimotor integration. Brodmann's area 44 is also a part of a specialized parieto-premotor network and interacts significantly with the neighbouring premotor areas. In the ventral premotor area F5 of monkeys, the so called mirror neurons have been found which discharge both when the animal performs a goal-directed hand action and when it observes another individual performing the same or a similar action. More recently, in the same area mirror neurons responding not only to the observation of mouth actions, but also to sounds characteristic to actions have been found. In humans, through an fMRI study, it has been shown that the observation of actions performed with the hand, the mouth and the foot leads to the activation of different sectors of Broca's area and premotor cortex, according to the effector involved in the observed action, following a somatotopic pattern which resembles the classical motor cortex homunculus. On the other hand the evidence is growing that human ventral premotor cortex, especially Brodmann's area 44, is involved in polymodal action processing. These results strongly support the existence of an execution-observation matching system (mirror neuron system). It has been proposed that this system is involved in polymodal action recognition and might represent a precursor of language processing. Experimental evidence in favour of this hypothesis both in the monkey and humans is shortly reviewed.     

Action Processing and Mirror Neuron Function in Patients with Amyotrophic Lateral Sclerosis: An fMRI Study

Amyotrophic lateral sclerosis (ALS) is a highly debilitating and rapidly fatal neurodegenerative disease. It has been suggested that social cognition may be affected, such as impairment in theory of mind (ToM) ability. Despite these findings, research in this area is scarce and the investigation of neural mechanisms behind such impairment is absent. Nineteen patients with ALS and eighteen healthy controls participated in this study. Because the mirror neuron system (MNS) is thought to be involved in theory of mind, we first implemented a straightforward action-execution and observation task to assess basic MNS function. Second, we examined the social-cognitive ability to understand actions of others, which is a component of ToM. We used fMRI to assess BOLD activity differences between groups during both experiments. Theory of mind was also measured behaviorally using the Reading the Mind in the Eyes test (RME). ALS patients displayed greater BOLD activity during the action-execution and observation task, especially throughout right anterior cortical regions. These areas included the right inferior operculum, premotor and primary motor regions, and left inferior parietal lobe. A conjunction analysis showed significantly more co-activated voxels during both the observation and action-execution conditions in the patient group throughout MNS regions. These results support a compensatory response in the MNS during action processing. In the action understanding experiment, healthy controls performed better behaviorally and subsequently recruited greater regions of activity throughout the prefrontal cortex and middle temporal gyrus. Lastly, action understanding performance was able to cluster patients with ALS into high and lower performing groups, which then differentiated RME performance. Collectively, these data suggest that social cognition, particularly theory of mind, may be affected in a subset of patients with ALS. This impairment may be related to functioning of the MNS and other regions related to action processing and understanding. Implications for future research are discussed.     

Gender differences in the mu rhythm of the human mirror-neuron system

Background

Psychologically, females are usually thought to be superior in interpersonal sensitivity than males. The human mirror-neuron system is considered to provide the basic mechanism for social cognition. However, whether the human mirror-neuron system exhibits gender differences is not yet clear.

Methodology/Principal Findings

We measured the electroencephalographic mu rhythm, as a reliable indicator of the human mirror-neuron system activity, when female (N = 20) and male (N = 20) participants watched either hand actions or a moving dot. The display of the hand actions included androgynous, male, and female characteristics. The results demonstrate that females displayed significantly stronger mu suppression than males when watching hand actions. Instead, mu suppression was similar across genders when participants observed the moving dot and between the perceived sex differences (same-sex vs. opposite-sex). In addition, the mu suppressions during the observation of hand actions positively correlated with the personal distress subscale of the interpersonal reactivity index and negatively correlated with the systemizing quotient.

Conclusions/Significance

The present findings indirectly lend support to the extreme male brain theory put forward by Baron-Cohen (2005), and may cast some light on the mirror-neuron dysfunction in autism spectrum disorders. The mu rhythm in the human mirror-neuron system can be a potential biomarker of empathic mimicry.     

Distinct olfactory cross-modal effects on the human motor system

Background

Converging evidence indicates that action observation and action-related sounds activate cross-modally the human motor system. Since olfaction, the most ancestral sense, may have behavioural consequences on human activities, we causally investigated by transcranial magnetic stimulation (TMS) whether food odour could additionally facilitate the human motor system during the observation of grasping objects with alimentary valence, and the degree of specificity of these effects.

Methodology/Principal Findings

In a repeated-measure block design, carried out on 24 healthy individuals participating to three different experiments, we show that sniffing alimentary odorants immediately increases the motor potentials evoked in hand muscles by TMS of the motor cortex. This effect was odorant-specific and was absent when subjects were presented with odorants including a potentially noxious trigeminal component.The smell-induced corticospinal facilitation of hand muscles during observation of grasping was an additive effect which superimposed to that induced by the mere observation of grasping actions for food or non-food objects. The odour-induced motor facilitation took place only in case of congruence between the sniffed odour and the observed grasped food, and specifically involved the muscle acting as prime mover for hand/fingers shaping in the observed action.

Conclusions/Significance

Complex olfactory cross-modal effects on the human corticospinal system are physiologically demonstrable. They are odorant-specific and, depending on the experimental context, muscle- and action-specific as well. This finding implies potential new diagnostic and rehabilitative applications.     

Observational learning of new movement sequences is reflected in fronto-parietal coherence

Mankind is unique in her ability for observational learning, i.e. the transmission of acquired knowledge and behavioral repertoire through observation of others' actions. In the present study we used electrophysiological measures to investigate brain mechanisms of observational learning. Analysis investigated the possible functional coupling between occipital (alpha) and motor (mu) rhythms operating in the 10 Hz frequency range for translating "seeing" into "doing". Subjects observed movement sequences consisting of six consecutive left or right hand button presses directed at one of two target-buttons for subsequent imitation. Each movement sequence was presented four times, intervened by short pause intervals for sequence rehearsal. During a control task subjects observed the same movement sequences without a requirement for subsequent reproduction. Although both alpha and mu rhythms desynchronized during the imitation task relative to the control task, modulations in alpha and mu power were found to be largely independent from each other over time, arguing against a functional coupling of alpha and mu generators during observational learning. This independence was furthermore reflected in the absence of coherence between occipital and motor electrodes overlaying alpha and mu generators. Instead, coherence analysis revealed a pair of symmetric fronto-parietal networks, one over the left and one over the right hemisphere, reflecting stronger coherence during observation of movements than during pauses. Individual differences in fronto-parietal coherence were furthermore found to predict imitation accuracy. The properties of these networks, i.e. their fronto-parietal distribution, their ipsilateral organization and their sensitivity to the observation of movements, match closely with the known properties of the mirror neuron system (MNS) as studied in the macaque brain. These results indicate a functional dissociation between higher order areas for observational learning (i.e. parts of the MNS as reflected in 10 Hz coherence measures) and peripheral structures (i.e. lateral occipital gyrus for alpha; central sulcus for mu) that provide low-level support for observation and motor imagery of action sequences.