Primate Amygdala Neurons Simulate Decision Processes of Social Partners.

By observing their social partners, primates learn about reward values of objects. Here, we show that monkeys' amygdala neurons derive object values from observation and use these values to simulate a partner monkey's decision process. While monkeys alternated making reward-based choices, amygdala neurons encoded object-specific values learned from observation. Dynamic activities converted these values to representations of the recorded monkey's own choices. Surprisingly, the same activity patterns unfolded spontaneously before partner's choices in separate neurons, as if these neurons simulated the partner's decision-making. These "simulation neurons" encoded signatures of mutual-inhibitory decision computation, including value comparisons and value-to-choice conversions, resulting in accurate predictions of partner's choices. Population decoding identified differential contributions of amygdala subnuclei. Biophysical modeling of amygdala circuits showed that simulation neurons emerge naturally from convergence between object-value neurons and self-other neurons. By simulating decision computations during observation, these neurons could allow primates to reconstruct their social partners' mental states.

Live vs video interaction: sensorimotor and visual cortical oscillations during action observation.

Increasingly, in the field of communication, education, and business, people are switching to video interaction, and interlocutors frequently complain that the perception of nonverbal information and concentration suffer. We investigated this issue by analyzing electroencephalogram (EEG) oscillations of the sensorimotor (mu rhythm) and visual (alpha rhythm) cortex of the brain in an experiment with action observation live and on video. The mu rhythm reflects the activity of the mirror neuron system, and the occipital alpha rhythm shows the level of visual attention. We used 32-channel EEG recorded during live and video action observation in 83 healthy volunteers. The ICA method was used for selecting the mu- and alpha-components; the Fourier Transform was used to calculate the suppression index relative to the baseline (stationary demonstrator) of the rhythms. The main range of the mu rhythm was indeed sensitive to social movement and was highly dependent on the conditions of interaction-live or video. The upper mu-range appeared to be less sensitive to the conditions, but more sensitive to different movements. The alpha rhythm did not depend on the type of movement; however, a live performance initially caused a stronger concentration of visual attention. Thus, subtle social and nonverbal perceptions may suffer in remote video interactions.

Increased sensorimotor activity during categorisation of emotionally ambiguous faces.

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.

Three-stage Dynamic Brain-cognitive Model of Understanding Action Intention Displayed by Human Body Movements.

The ability to comprehend the intention conveyed through human body movements is crucial for effective interpersonal interactions. If people can't understand the intention behind other individuals' isolated or interactive actions, their actions will become meaningless. Psychologists have investigated the cognitive processes and neural representations involved in understanding action intention, yet a cohesive theoretical explanation remains elusive. Hence, we mainly review existing literature related to neural correlates of action intention, and primarily propose a putative Three-stage Dynamic Brain-cognitive Model of understanding action intention, which involves body perception, action identification and intention understanding. Specifically, at the first stage, body parts/shapes are processed by those brain regions such as extrastriate and fusiform body areas; During the second stage, differentiating observed actions relies on configuring relationships between body parts, facilitated by the activation of the Mirror Neuron System; The last stage involves identifying various intention categories, utilizing the Mentalizing System for recruitment, and different activation patterns concerning the nature of the intentions participants dealing with. Finally, we delves into the clinical practice, like intervention training based on a theoretical model for individuals with autism spectrum disorders who encounter difficulties in interpersonal communication.

Neural basis of lower-limb visual feedback therapy: an EEG study in healthy subjects.

BACKGROUND: Video-feedback observational therapy (VOT) is an intensive rehabilitation technique based on movement repetition and visualization that has shown benefits for motor rehabilitation of the upper and lower limbs. Despite an increase in recent literature on the neurophysiological effects of VOT in the upper limb, there is little knowledge about the cortical effects of visual feedback therapies when applied to the lower limbs. The aim of our study was to better understand the neurophysiological effects of VOT. Thus, we identified and compared the EEG biomarkers of healthy subjects undergoing lower limb VOT during three tasks: passive observation, observation and motor imagery, observation and motor execution. METHODS: We recruited 38 healthy volunteers and monitored their EEG activity while they performed a right ankle dorsiflexion task in the VOT. Three graded motor tasks associated with action observation were tested: action observation alone (O), motor imagery with action observation (OI), and motor execution synchronized with action observation (OM). The alpha and beta event-related desynchronization (ERD) and event-related synchronization (or beta rebound, ERS) rhythms were used as biomarkers of cortical activation and compared between conditions with a permutation test. Changes in connectivity during the task were computed with phase locking value (PLV). RESULTS: During the task, in the alpha band, the ERD was comparable between O and OI activities across the precentral, central and parietal electrodes. OM involved the same regions but had greater ERD over the central electrodes. In the beta band, there was a gradation of ERD intensity in O, OI and OM over central electrodes. After the task, the ERS changes were weak during the O task but were strong during the OI and OM (Cz) tasks, with no differences between OI and OM. CONCLUSION: Alpha band ERD results demonstrated the recruitment of mirror neurons during lower limb VOT due to visual feedback. Beta band ERD reflects strong recruitment of the sensorimotor cortex evoked by motor imagery and action execution. These results also emphasize the need for an active motor task, either motor imagery or motor execution task during VOT, to elicit a post-task ERS, which is absent during passive observation. Trial Registration NCT05743647.

Mu-Suppression Neurofeedback Training Targeting the Mirror Neuron System: A Pilot Study.

Neurofeedback training (NFT) is a promising adjuvant intervention method. The desynchronization of mu rhythm (8-13 Hz) in the electroencephalogram (EEG) over centro-parietal areas is known as a valid indicator of mirror neuron system (MNS) activation, which has been associated with social skills. Still, the effect of neurofeedback training on the MNS requires to be well investigated. The present study examined the possible impact of NFT with a mu suppression training protocol encompassing 15 NFT sessions (45 min each) on 16 healthy neurotypical participants. In separate pre- and post-training sessions, 64-channel EEG was recorded while participants (1) observed videos with various types of movements (including complex goal-directed hand movements and social interaction scenes) and (2) performed the "Reading the Mind in the Eyes Test" (RMET). EEG source reconstruction analysis revealed statistically significant mu suppression during hand movement observation across MNS-attributed fronto-parietal areas after NFT. The frequency analysis showed no significant mu suppression after NFT, despite the fact that numerical mu suppression appeared to be visible in a majority of participants during goal-directed hand movement observation. At the behavioral level, RMET accuracy scores did not suggest an effect of NFT on the ability to interpret subtle emotional expressions, although RMET response times were reduced after NFT. In conclusion, the present study exhibited preliminary and partial evidence that mu suppression NFT can induce mu suppression in MNS-attributed areas. More powerful experimental designs and longer training may be necessary to induce substantial and consistent mu suppression, particularly while observing social scenarios.

Effects of mirror neuron activation therapies on functionality in older adults: Systematic review and meta-analysis.

PURPOSE: To identify the effects of mirror neuron activation (MNAT) combined or not with physical exercise (PE) in healthy older adults, on functionality, balance, gait velocity and risk of falls. METHODS: A systematic electronic search was performed in PubMed/MEDLINE, Cochrane, and Embase databases. RESULTS: Thirteen randomized controlled trials were included in the qualitative analysis, and eleven in the quantitative analysis. All studies showed fair to high quality and the most frequent high-risk bias was "Blinding of participants and personnel". Compared to the control condition, higher improvement was shown in older people who received MNAT, on functionality (1.57 [0.57, 2.62], balance (1.95 [1.32, 2.572]), and gait velocity (1.20 [0.30, 2.11]). Compared to PE, MNAT combined with PE does not improve functionality. More studies are needed to assess MNAT effectiveness in the rest of the outcomes. CONCLUSIONS: Neuron system activation through MNAT improves relevant abilities in older adults, with better results when including functional activities. However, the beneficial effects on these variables of adding MNAT to a PE program are controversial.

Increased interbrain synchronization and neural efficiency of the frontal cortex to enhance human coordinative behavior: A combined hyper-tES and fNIRS study.

Coordination is crucial for individuals to achieve common goals; however, the causal relationship between coordination behavior and neural activity has not yet been explored. Interbrain synchronization (IBS) and neural efficiency in cortical areas associated with the mirror neuron system (MNS) are considered two potential brain mechanisms. In the present study, we attempted to clarify how the two mechanisms facilitate coordination using hypertranscranial electrical stimulation (hyper-tES). A total of 124 healthy young adults were randomly divided into three groups (the hyper-tACS, hyper-tDCS and sham groups) and underwent modulation of the right inferior frontal gyrus (IFG) during functional near-infrared spectroscopy (fNIRS). Increased IBS of the PFC or neural efficiency of the right IFG (related to the MNS) was accompanied by greater coordination behavior; IBS had longer-lasting effects on behavior. Our findings highlight the importance of IBS and neural efficiency of the frontal cortex for coordination and suggest potential interventions to improve coordination in different temporal windows.

The value of corticospinal excitability and intracortical inhibition in predicting motor skill improvement driven by action observation.

The observation of other's actions represents an essential element for the acquisition of motor skills. While action observation is known to induce changes in the excitability of the motor cortices, whether such modulations may explain the amount of motor improvement driven by action observation training (AOT) remains to be addressed. Using transcranial magnetic stimulation (TMS), we first assessed in 41 volunteers the effect of action observation on corticospinal excitability, intracortical inhibition, and transcallosal inhibition. Subsequently, half of the participants (AOT-group) were asked to observe and then execute a right-hand dexterity task, while the controls had to observe a no-action video before practicing the same task. AOT participants showed greater performance improvement relative to controls. More importantly, the amount of improvement in the AOT group was predicted by the amplitude of corticospinal modulation during action observation and, even more, by the amount of intracortical inhibition induced by action observation. These relations were specific for the AOT group, while the same patterns were not found in controls. Taken together, our findings demonstrate that the efficacy of AOT in promoting motor learning is rooted in the capacity of action observation to modulate the trainee's motor system excitability, especially its intracortical inhibition. Our study not only enriches the picture of the neurophysiological effects induced by action observation onto the observer's motor excitability, but linking them to the efficacy of AOT, it also paves the way for the development of models predicting the outcome of training procedures based on the observation of other's actions.

Differential responses in the mirror neuron system during imitation of individual emotional facial expressions and association with autistic traits.

The mirror neuron system (MNS), including the inferior frontal gyrus (IFG), inferior parietal lobule (IPL) and superior temporal sulcus (STS) plays an important role in action representation and imitation and may be dysfunctional in autism spectrum disorder (ASD). However, it's not clear how these three regions respond and interact during the imitation of different basic facial expressions and whether the pattern of responses is influenced by autistic traits. Thus, we conducted a natural facial expression (happiness, angry, sadness and fear) imitation task in 100 healthy male subjects where expression intensity was measured using facial emotion recognition software (FaceReader) and MNS responses were recorded using functional near-infrared spectroscopy (fNIRS). Autistic traits were measured using the Autism Spectrum Quotient questionnaire. Results showed that imitation of happy expressions produced the highest expression intensity but a small deactivation in MNS responses, suggesting a lower processing requirement compared to other expressions. A cosine similarity analysis indicated a distinct pattern of MNS responses during imitation of each facial expression with functional intra-hemispheric connectivity between the left IPL and left STS being significantly higher during happy compared to other expressions, while inter-hemispheric connectivity between the left and right IPL differed between imitation of fearful and sad expressions. Furthermore, functional connectivity changes during imitation of each different expression could reliably predict autistic trait scores. Overall, the results provide evidence for distinct patterns of functional connectivity changes between MNS regions during imitation of different emotions which are also associated with autistic traits.

The widespread action observation/execution matching system for facial expression processing.

Observing and understanding others' emotional facial expressions, possibly through motor synchronization, plays a primary role in face-to-face communication. To understand the underlying neural mechanisms, previous functional magnetic resonance imaging (fMRI) studies investigated brain regions that are involved in both the observation/execution of emotional facial expressions and found that the neocortical motor regions constituting the action observation/execution matching system or mirror neuron system were active. However, it remains unclear (1) whether other brain regions in the limbic, cerebellum, and brainstem regions could be also involved in the observation/execution matching system for processing facial expressions, and (2) if so, whether these regions could constitute a functional network. To investigate these issues, we performed fMRI while participants observed dynamic facial expressions of anger and happiness and while they executed facial muscle activity associated with angry and happy facial expressions. Conjunction analyses revealed that, in addition to neocortical regions (i.e., the right ventral premotor cortex and right supplementary motor area), bilateral amygdala, right basal ganglia, bilateral cerebellum, and right facial nerve nucleus were activated during both the observation/execution tasks. Group independent component analysis revealed that a functional network component involving the aforementioned regions were activated during both observation/execution tasks. The data suggest that the motor synchronization of emotional facial expressions involves a widespread observation/execution matching network encompassing the neocortex, limbic system, basal ganglia, cerebellum, and brainstem.

Action viewing and language in adolescents with autism spectrum disorder.

The mirror neuron system consists of fronto-parietal regions and responds to both goal-directed action execution and observation. The broader action observation network is specifically involved in observation of actions and is thought to play a role in understanding the goals of the motor act, the intention of others, empathy, and language. Many, but not all, studies have found mirror neuron system or action observation network dysfunction in autism spectrum disorder. The objective of this study was to use observation of a goal-directed action fMRI paradigm to examine the action observation network in autism spectrum disorder and to determine whether fronto-parietal activation is associated with language ability. Adolescents with autism spectrum disorder (n = 23) were compared to typically developing adolescents (n = 20), 11-17 years. Overall, there were no group differences in activation, however, the autism spectrum group with impaired expressive language (n = 13) had significantly reduced inferior frontal and inferior parietal activation during action viewing. In controls, right supramarginal gyrus activation was associated with higher expressive language; bilateral supramarginal and left pars opercularis activation was associated with better verbal-gesture integration. Results suggest that action-observation network dysfunction may characterize a subgroup of individuals with autism spectrum disorder with expressive language deficits. Therefore, interventions that target this dysfunctional network may improve expressive language in this autism spectrum subgroup. Future treatment studies should individualize therapeutic approaches based on brain-behavior relationships.

Activation of Cerebellum, Basal Ganglia and Thalamus During Observation and Execution of Mouth, hand, and foot Actions.

Humans and monkey studies showed that specific sectors of cerebellum and basal ganglia activate not only during execution but also during observation of hand actions. However, it is unknown whether, and how, these structures are engaged during the observation of actions performed by effectors different from the hand. To address this issue, in the present fMRI study, healthy human participants were required to execute or to observe grasping acts performed with different effectors, namely mouth, hand, and foot. As control, participants executed and observed simple movements performed with the same effectors. The results show that: (1) execution of goal-directed actions elicited somatotopically organized activations not only in the cerebral cortex but also in the cerebellum, basal ganglia, and thalamus; (2) action observation evoked cortical, cerebellar and subcortical activations, lacking a clear somatotopic organization; (3) in the territories displaying shared activations between execution and observation, a rough somatotopy could be revealed in both cortical, cerebellar and subcortical structures. The present study confirms previous findings that action observation, beyond the cerebral cortex, also activates specific sectors of cerebellum and subcortical structures and it shows, for the first time, that these latter are engaged not only during hand actions observation but also during the observation of mouth and foot actions. We suggest that each of the activated structures processes specific aspects of the observed action, such as performing internal simulation (cerebellum) or recruiting/inhibiting the overt execution of the observed action (basal ganglia and sensory-motor thalamus).

The transition from invariant to action-dependent visual object representation in human dorsal pathway.

The human brain can efficiently process action-related visual information, which supports our ability to quickly understand and learn others' actions. The visual information of goal-directed action is extensively represented in the parietal and frontal cortex, but how actions and goal-objects are represented within this neural network is not fully understood. Specifically, which part of this dorsal network represents the identity of goal-objects? Is such goal-object information encoded at an abstract level or highly interactive with action representations? Here, we used functional magnetic resonance imaging with a large number of participants (n = 94) to investigate the neural representation of goal-objects and actions when participants viewed goal-directed action videos. Our results showed that the goal-directed action information could be decoded across much of the dorsal pathway, but in contrast, the invariant goal-object information independent of action was mainly localized in the early stage of dorsal pathway in parietal cortex rather than the down-stream areas of the parieto-frontal cortex. These results help us to understand the relationship between action and goal-object representations in the dorsal pathway, and the evolution of interactive representation of goal-objects and actions along the dorsal pathway during goal-directed action observation.

Altered activation in the action observation system during synchronization in high loneliness individuals.

Lonely people tend to evaluate social exchanges negatively and to display difficulties in interactions. Interpersonal synchronization is crucial for achieving positive interactions, promoting affinity, closeness, and satisfaction. However, little is known about lonely individuals' ability to synchronize and about their brain activity while synchronizing. Following the screening of 303 participants, we recruited 32 low and 32 high loneliness participants. They were scanned while engaged in movement synchronization, using a novel dyadic interaction paradigm. Results showed that high loneliness individuals exhibited a reduced ability to adapt their movement to their partner's movement. Intriguingly, during movement adaptation periods, high loneliness individuals showed increased activation in the action observation (AO) system, specifically in the inferior frontal gyrus and the inferior parietal lobule. They did not show increased activation in the dorsomedial prefrontal cortex, which in the context of synchronization was suggested to be related to gap-monitoring. Based on these findings, we propose a model according to which lonely people may require stronger activation of their AO system for alignment, to compensate for some deficiency in their synchronization ability. Despite this hyperactivation, they still suffer from reduced synchronization capacity. Consequently, synchronization may be a relevant intervention area for the amelioration of loneliness.

Mirror neuron activity depending on the content and stage of the observed action: a TMS study.

Background/aim: The firing rate of the mirror neuron system in monkeys decreases systematically with more repetitions. The aim of this study is to investigate whether the activity of the mirror neuron system varies based on the observed movement and the contents of the action, as well as whether there is inhibition in the mirror neuron system when humans observe repeated actions. If inhibition is present, the second question of the study is whether it is related to the organization of the observed action. Materials and methods: Fourteen healthy volunteers participated in the study. Transcranial magnetic stimulation was applied to the left primary motor cortex and motor evoked potentials (MEPs) were recorded from the right first dorsal interosseous and abductor pollicis brevis muscles while the participants were watching videos specially prepared for the study. Results: There were no significant changes in MEP amplitudes compared to baseline MEPs while observing aimless action. However, while participants watched the repeated action video, the mean MEP amplitude increased at the beginning of the movement, but neither facilitation nor inhibition was detected when the participants watched the phase of grasping the object of the action compared to the baseline MEP amplitude. On the other hand, while participants were watching different activities, an increased MEP amplitude was observed at the beginning of the movement and in the grasping of the object of the action. Additionally, there was no significant reduction in MEP amplitude during any movement stages while observing the repeated action video. Conclusion: The findings of this study suggest that the activation of the mirror neuron system in humans depends on the content and stages of the observed movement. Additionally, there was no inhibition or systematic reduction in MEP amplitudes while watching a repeated action.

The mirror neuron system compensates for amygdala dysfunction - associated social deficits in individuals with higher autistic traits.

The amygdala is a core node in the social brain which exhibits structural and functional abnormalities in Autism spectrum disorder and there is evidence that the mirror neuron system (MNS) can functionally compensate for impaired emotion processing following amygdala lesions. In the current study, we employed an fMRI paradigm in 241 subjects investigating MNS and amygdala responses to observation, imagination and imitation of dynamic facial expressions and whether these differed in individuals with higher (n = 77) as opposed to lower (n = 79) autistic traits. Results indicated that individuals with higher compared to lower autistic traits showed worse recognition memory for fearful faces, smaller real-life social networks, and decreased left basolateral amygdala (BLA) responses to imitation. Additionally, functional connectivity between the left BLA and the left inferior frontal gyrus (IFG) as well as some other MNS regions was increased in individuals with higher autistic traits, especially during imitation of fearful expressions. The left BLA-IFG connectivity significantly moderated the autistic group differences on recognition memory for fearful faces, indicating that increased amygdala-MNS connectivity could diminish the social behavioral differences between higher and lower autistic trait groups. Overall, findings demonstrate decreased imitation-related amygdala activity in individuals with higher autistic traits in the context of increased amygdala-MNS connectivity which may functionally compensate for amygdala dysfunction and social deficits. Training targeting the MNS may capitalize on this compensatory mechanism for therapeutic benefits in Autism spectrum disorder.

Enhanced mirror neuron network activity and effective connectivity during live interaction among female subjects.

Facial expressions are indispensable in daily human communication. Previous neuroimaging studies investigating facial expression processing have presented pre-recorded stimuli and lacked live face-to-face interaction. Our paradigm alternated between presentations of real-time model performance and pre-recorded videos of dynamic facial expressions to participants. Simultaneous functional magnetic resonance imaging (fMRI) and facial electromyography activity recordings, as well as post-scan valence and arousal ratings were acquired from 44 female participants. Live facial expressions enhanced the subjective valence and arousal ratings as well as facial muscular responses. Live performances showed greater engagement of the right posterior superior temporal sulcus (pSTS), right inferior frontal gyrus (IFG), right amygdala and right fusiform gyrus, and modulated the effective connectivity within the right mirror neuron system (IFG, pSTS, and right inferior parietal lobule). A support vector machine algorithm could classify multivoxel activation patterns in brain regions involved in dynamic facial expression processing in the mentalizing networks (anterior and posterior cingulate cortex). These results indicate that live social interaction modulates the activity and connectivity of the right mirror neuron system and enhances spontaneous mimicry, further facilitating emotional contagion.

Mirror neuron brain regions contribute to identifying actions, but not intentions.

Previous studies have struggled to determine the relationship between mirror neuron brain regions and two distinct "action understanding" processes: identifying actions and identifying the intentions underlying those actions. This may be because the identification of intentions from others' actions requires an initial action identification process. Disruptive transcranial magnetic stimulation was administered to left inferior frontal gyrus (lIFG) during a novel cognitive task to determine which of these "action understanding" processes is subserved by mirror neuron brain regions. Participants identified either the actions performed by observed hand actions or the intentions underlying those actions. The extent to which intention identification was disrupted by lIFG (vs. control site) stimulation was dependent on the level of disruption to action identification. We subsequently performed functional magnetic resonance imaging during the same task. During action identification, responses were widespread within mirror neuron areas including lIFG and inferior parietal lobule. However, no independent responses were found in mirror neuron brain regions during intention identification. Instead, responses occurred in brain regions associated with two distinct mentalizing localizer tasks. This supports an account in which mirror neuron brain regions are involved in an initial action identification process, but the subsequent identification of intentions requires additional processing in mentalizing brain regions.

Decoding point-light displays and fully visible hand grasping actions within the action observation network.

Action observation typically recruits visual areas and dorsal and ventral sectors of the parietal and premotor cortex. This network has been collectively termed as extended action observation network (eAON). Within this network, the elaboration of kinematic aspects of biological motion is crucial. Previous studies investigated these aspects by presenting subjects with point-light displays (PLDs) videos of whole-body movements, showing the recruitment of some of the eAON areas. However, studies focused on cortical activation during observation of PLDs grasping actions are lacking. In the present functional magnetic resonance imaging (fMRI) study, we assessed the activation of eAON in healthy participants during the observation of both PLDs and fully visible hand grasping actions, excluding confounding effects due to low-level visual features, motion, and context. Results showed that the observation of PLDs grasping stimuli elicited a bilateral activation of the eAON. Region of interest analyses performed on visual and sensorimotor areas showed no significant differences in signal intensity between PLDs and fully visible experimental conditions, indicating that both conditions evoked a similar motor resonance mechanism. Multivoxel pattern analysis (MVPA) revealed significant decoding of PLDs and fully visible grasping observation conditions in occipital, parietal, and premotor areas belonging to eAON. Data show that kinematic features conveyed by PLDs stimuli are sufficient to elicit a complete action representation, suggesting that these features can be disentangled within the eAON from the features usually characterizing fully visible actions. PLDs stimuli could be useful in assessing which areas are recruited, when only kinematic cues are available, for action recognition, imitation, and motor learning.