The capacity of action observation to drag the trainees' motor pattern toward the observed model.

Action Observation Training (AOT) promotes the acquisition of motor abilities. However, while the cortical modulations associated with the AOT efficacy are well known, few studies investigated the AOT peripheral neural correlates and whether their dynamics move towards the observed model during the training. We administered seventy-two participants (randomized into AOT and Control groups) with training for learning to grasp marbles with chopsticks. Execution practice was preceded by an observation session, in which AOT participants observed an expert performing the task, whereas controls observed landscape videos. Behavioral indices were measured, and three hand muscles' electromyographic (EMG) activity was recorded and compared with the expert. Behaviorally, both groups improved during the training, with AOT outperforming controls. The EMG trainee-model similarity also increased during the training, but only for the AOT group. When combining behavioral and EMG similarity findings, no global relationship emerged; however, behavioral improvements were "locally" predicted by the similarity gain in muscles and action phases more related to the specific motor act. These findings reveal that AOT plays a magnetic role in motor learning, attracting the trainee's motor pattern toward the observed model and paving the way for developing online monitoring tools and neurofeedback protocols.

Kicking in or kicking out? The role of the individual motor expertise in predicting the outcome of rugby actions.

In sports, understanding others' actions represents a fundamental skill that allows players to predict the outcome of teammates' and opponents' actions and counteract them properly. While it is well known that motor expertise sets better premises for predicting the result of an observed sports action, it remains untested whether this principle applies to a team where players cover different positions that imply different motor repertoires. To test this hypothesis, we selected rugby as a paradigmatic example in which only one or two players out of 22 train and perform placed kicks. We administered a placed kick outcome prediction task to three groups of participants, namely, rugby kickers, rugby non-kickers, and controls, thus spanning over different combinations of motor expertise and visual experience. Kickers outperformed both their non-kicking teammates and controls in overall prediction accuracy. We documented how the viewpoint of observation, the expertise of the observed kicker, and the position of the kick on the court influenced the prediction performance across the three groups. Finally, we revealed that within rugby players, the degree of motor expertise (but not the visual experience) causally affects accuracy, and such a result stands even after accounting for the level of visual experience. These findings extend the role of motor expertise in decoding and predicting others' behaviors to sports teammates, among which every member is equipped with a position-specific motor repertoire, advocating for new motor training procedures combining the gestures to-be-performed with those to-be-faced.

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.

Embodying Language through Gestures: Residuals of Motor Memories Modulate Motor Cortex Excitability during Abstract Words Comprehension.

There is a debate about whether abstract semantics could be represented in a motor domain as concrete language. A contextual association with a motor schema (action or gesture) seems crucial to highlighting the motor system involvement. The present study with transcranial magnetic stimulation aimed to assess motor cortex excitability changes during abstract word comprehension after conditioning word reading to a gesture execution with congruent or incongruent meaning. Twelve healthy volunteers were engaged in a lexical-decision task responding to abstract words or meaningless verbal stimuli. Motor cortex (M1) excitability was measured at different after-stimulus intervals (100, 250, or 500 ms) before and after an associative-learning training where the execution of the gesture followed word processing. Results showed a significant post-training decrease in hand motor evoked potentials at an early processing stage (100 ms) in correspondence to words congruent with the gestures presented during the training. We hypothesized that traces of individual semantic memory, combined with training effects, induced M1 inhibition due to the redundancy of evoked motor representation. No modulation of cortical excitability was found for meaningless or incongruent words. We discuss data considering the possible implications in research to understand the neural basis of language development and language rehabilitation protocols.

The vision of haptics tunes the somatosensory threshold.

The interaction between different sensory modalities represents a crucial issue in the neuroscience of consciousness: when the processing of one modality is deficient, the concomitant presentation of stimuli of other spared modalities may sustain the restoration of the damaged sensory functions. In this regard, visual enhancement of touch may represent a viable tool in rehabilitating tactile disorders, yet the specific visual features mostly modulating the somatosensory experience remain unsettled. In this study, healthy subjects underwent a tactile detection task during the observation of videos displaying different contents, including static gratings, meaningless motions and natural or point-lights reach-to-grasp-and-manipulate actions. Concurrently, near-threshold stimuli were delivered to the median nerve at different time-points. The subjective report was collected after each trial; the sensory detection rate was computed and compared across video conditions. Our results indicate that the specific presence of haptic contents (i.e., the vision of manipulation), either fully displayed or implied by point-lights, magnifies tactile sensitivity. The notion that such stimuli prompt a conscious tactile experience opens to novel rehabilitation approaches for tactile consciousness disorders.

The Proactive Synergy Between Action Observation and Execution in the Acquisition of New Motor Skills.

Motor learning can be defined as a process that leads to relatively permanent changes in motor behavior through repeated interactions with the environment. Different strategies can be adopted to achieve motor learning: movements can be overtly practiced leading to an amelioration of motor performance; alternatively, covert strategies (e.g., action observation) can promote neuroplastic changes in the motor system even in the absence of real movement execution. However, whether a training regularly alternating action observation and execution (i.e., Action Observation Training, AOT) may surpass the pure motor practice (MP) and observational learning (OL) remains to be established. To address this issue, we enrolled 54 subjects requiring them to learn tying nautical knots via one out of three types of training (AOT, MP, OL) with the scope to investigate which element mostly contributes to motor learning. We evaluated the overall improvement of each group, along with the predictive role that neuropsychological indexes exert on each treatment outcome. The AOT group exhibited the highest performance improvement (42%), indicating that the regular alternation between observation and execution biases participants toward a better performance. The reiteration of this sequence provides an incremental, adjunct value that super-adds onto the efficacy of motor practice or observational learning in isolation (42% > 25% + 10%, i.e., OL + MP). These findings extend the use of the AOT from clinical and rehabilitative contexts to daily routines requiring the learning and perfectioning of new motor skills such as sports training, music, and occupational activities requiring fine motor control.

Observation of others' actions during limb immobilization prevents the subsequent decay of motor performance.

There is rich clinical evidence that observing normally executed actions promotes the recovery of the corresponding action execution in patients with motor deficits. In this study, we assessed the ability of action observation to prevent the decay of healthy individuals' motor abilities following upper-limb immobilization. To this end, upper-limb kinematics was recorded in healthy participants while they performed three reach-to-grasp movements before immobilization and the same movements after 16 h of immobilization. The participants were subdivided into two groups; the experimental group observed, during the immobilization, the same reach-to-grasp movements they had performed before immobilization, whereas the control group observed natural scenarios. After bandage removal, motor impairment in performing reach-to-grasp movements was milder in the experimental group. These findings support the hypothesis that action observation, via the mirror mechanism, plays a protective role against the decline of motor performance induced by limb nonuse. From this perspective, action observation therapy is a promising tool for anticipating rehabilitation onset in clinical conditions involving limb nonuse, thus reducing the burden of further rehabilitation.

Catching the imposter in the brain: The case of Capgras delusion.

Here we describe a rare case of Capgras delusion - a misidentification syndrome characterized by the belief that a person has been replaced by an imposter - in a patient without evident neurological or psychiatric symptoms. Intriguingly, delusional belief was selective for both person and modality, as the patient believed that his son - not his daughter or other relatives - was substituted with an imposter only while being in presence of him and looking at his face, but not when merely listening to his voice. A neuroanatomical reconstruction obtained integrating morphological and functional patient's neuroimaging data highlighted two main peculiarities: a compression of the rostral portion of right temporal lobe due to a large arachnoid cyst, and a bilaterally reduced metabolism of frontal areas. Autonomic data obtained from thermal infra-red camera and skin conductance recordings showed that a higher sympathetic activation was evoked by the observation of daughter's face, relative to the observation of the son's face as well as of not-familiar faces; conversely, daughter and son voices elicited a similar sympathetic activation, higher relative to not-familiar voices, indicating a modality-dependent dissociation consistent with the delusional behavior. Our case supports the "two-hit hypothesis" about Capgras delusion etiopathogenesis: here, the first hit is represented by the right-temporal lesion impairing the association between familiar faces and emotional values, the second one is the frontal bilateral hypometabolism favoring delusional behavior. The selective occurrence of "imposter" delusion for a particular subject and for a specific perceptual modality suggests the involvement of modality-specific interactions in the retrieval of affective properties during familiar people recognition.

Observer-Agent Kinematic Similarity Facilitates Action Intention Decoding.

It is well known that the kinematics of an action is modulated by the underlying motor intention. In turn, kinematics serves as a cue also during action observation, providing hints about the intention of the observed action. However, an open question is whether decoding others' intentions on the basis of their kinematics depends solely on how much the kinematics varies across different actions, or rather it is also influenced by its similarity with the observer motor repertoire. The execution of reach-to-grasp and place actions, differing for target size and context, was recorded in terms of upper-limb kinematics in 21 volunteers and in an actor. Volunteers had later to observe the sole reach-to-grasp phase of the actor's actions, and predict the underlying intention. The potential benefit of the kinematic actor-participant similarity for recognition accuracy was evaluated. In execution, both target size and context modulated specific kinematic parameters. More importantly, although participants performed above chance in intention recognition, the similarity of motor patterns positively correlated with recognition accuracy. Overall, these data indicate that kinematic similarity exerts a facilitative role in intention recognition, providing further support to the view of action intention recognition as a visuo-motor process grounded in motor resonance.

Language, Gesture, and Emotional Communication: An Embodied View of Social Interaction.

Spoken language is an innate ability of the human being and represents the most widespread mode of social communication. The ability to share concepts, intentions and feelings, and also to respond to what others are feeling/saying is crucial during social interactions. A growing body of evidence suggests that language evolved from manual gestures, gradually incorporating motor acts with vocal elements. In this evolutionary context, the human mirror mechanism (MM) would permit the passage from "doing something" to "communicating it to someone else." In this perspective, the MM would mediate semantic processes being involved in both the execution and in the understanding of messages expressed by words or gestures. Thus, the recognition of action related words would activate somatosensory regions, reflecting the semantic grounding of these symbols in action information. Here, the role of the sensorimotor cortex and in general of the human MM on both language perception and understanding is addressed, focusing on recent studies on the integration between symbolic gestures and speech. We conclude documenting some evidence about MM in coding also the emotional aspects conveyed by manual, facial and body signals during communication, and how they act in concert with language to modulate other's message comprehension and behavior, in line with an "embodied" and integrated view of social interaction.

The effect of motor context on semantic processing: A TMS study.

BACKGROUND: Strong embodiment theories claimed that action language representation is grounded in the sensorimotor system, which would be crucially to semantic understanding. However, there is a large disagreement in literature about the neural mechanisms involved in abstract (symbolic) language comprehension. OBJECTIVE: In the present study, we investigated the role of motor context in the semantic processing of abstract language. We hypothesized that motor cortex excitability during abstract word comprehension could be modulated by previous presentation of a stimuli which associated a congruent motor content (i.e., a semantically related gesture) to the word. METHODS AND RESULTS: We administered a semantic priming paradigm where postures of gestures (primes) were followed by semantically congruent verbal stimuli (targets, meaningful or meaningless words). Transcranial Magnetic Stimulation was delivered to left motor cortex 100, 250 and 500 ms after the presentation of each target. Results showed that Motor evoked potentials of hand muscle significantly increased in correspondence to meaningful compared to meaningless words, but only in the earlier phase of semantic processing (100 and 250 ms from target onset). CONCLUSION: Results suggested that the gestural motor representation was integrated with corresponding word meaning in order to accomplish (and facilitate) the lexical task. We concluded that motor context resulted crucial to highlight motor system involvement during semantic processing of abstract language.

From meaning to categorization: The hierarchical recruitment of brain circuits selective for action verbs.

Sensorimotor and affective brain systems are known to be involved in language processing. However, to date it is still debated whether this involvement is a crucial step of semantic processing or, on the contrary, it is dependent on the specific context or strategy adopted to solve a task at hand. The present electroencephalographic (EEG) study is aimed at investigating which brain circuits are engaged when processing written verbs. By aligning event-related potentials (ERPs) both to the verb onset and to the motor response indexing the accomplishment of a semantic task of categorization, we were able to dissociate the relative stimulus-related and response-related cognitive components at play, respectively. EEG signal source reconstruction showed that while the recruitment of sensorimotor fronto-parietal circuits was time-locked with action verb onset, a left temporal-parietal circuit was time-locked to the task accomplishment. Crucially, by comparing the time course of both these bottom-up and top-down cognitive components, it appears that the frontal motor involvement precedes the task-related temporal-parietal activity. The present findings suggest that the recruitment of fronto-parietal sensorimotor circuits is independent of the specific strategy adopted to solve a semantic task and, given its temporal hierarchy, it may provide crucial information to brain circuits involved in the categorization task. Eventually, a discussion on how the present results may contribute to the clinical literature on patients affected by disorders specifically impairing the motor system is provided.

Vitality Forms Expressed by Others Modulate Our Own Motor Response: A Kinematic Study.

During social interaction, actions, and words may be expressed in different ways, for example, gently or rudely. A handshake can be gentle or vigorous and, similarly, tone of voice can be pleasant or rude. These aspects of social communication have been named vitality forms by Daniel Stern. Vitality forms represent how an action is performed and characterize all human interactions. In spite of their importance in social life, to date it is not clear whether the vitality forms expressed by the agent can influence the execution of a subsequent action performed by the receiver. To shed light on this matter, in the present study we carried out a kinematic study aiming to assess whether and how visual and auditory properties of vitality forms expressed by others influenced the motor response of participants. In particular, participants were presented with video-clips showing a male and a female actor performing a "giving request" (give me) or a "taking request" (take it) in visual, auditory, and mixed modalities (visual and auditory). Most importantly, requests were expressed with rude or gentle vitality forms. After the actor's request, participants performed a subsequent action. Results showed that vitality forms expressed by the actors influenced the kinematic parameters of the participants' actions regardless to the modality by which they are conveyed.

The Emotional Modulation of Facial Mimicry: A Kinematic Study.

It is well-established that the observation of emotional facial expression induces facial mimicry responses in the observers. However, how the interaction between emotional and motor components of facial expressions can modulate the motor behavior of the perceiver is still unknown. We have developed a kinematic experiment to evaluate the effect of different oro-facial expressions on perceiver's face movements. Participants were asked to perform two movements, i.e., lip stretching and lip protrusion, in response to the observation of four meaningful (i.e., smile, angry-mouth, kiss, and spit) and two meaningless mouth gestures. All the stimuli were characterized by different motor patterns (mouth aperture or mouth closure). Response Times and kinematics parameters of the movements (amplitude, duration, and mean velocity) were recorded and analyzed. Results evidenced a dissociated effect on reaction times and movement kinematics. We found shorter reaction time when a mouth movement was preceded by the observation of a meaningful and motorically congruent oro-facial gesture, in line with facial mimicry effect. On the contrary, during execution, the perception of smile was associated with the facilitation, in terms of shorter duration and higher velocity of the incongruent movement, i.e., lip protrusion. The same effect resulted in response to kiss and spit that significantly facilitated the execution of lip stretching. We called this phenomenon facial mimicry reversal effect, intended as the overturning of the effect normally observed during facial mimicry. In general, the findings show that both motor features and types of emotional oro-facial gestures (conveying positive or negative valence) affect the kinematics of subsequent mouth movements at different levels: while congruent motor features facilitate a general motor response, motor execution could be speeded by gestures that are motorically incongruent with the observed one. Moreover, valence effect depends on the specific movement required. Results are discussed in relation to the Basic Emotion Theory and embodied cognition framework.

The Effects of Meaning and Emotional Content of a Sentence on the Kinematics of a Successive Motor Sequence Mimiking the Feeding of a Conspecific.

AIM: Do the emotional content and meaning of sentences affect the kinematics of successive motor sequences? MATERIAL AND METHODS: Participants observed video-clips of an actor pronouncing sentences expressing positive or negative emotions and meanings (related to happiness or anger in Experiment 1 and food admiration or food disgust in Experiment 2). Then, they reached-to-grasp and placed a sugar lump on the actor's mouth. Participants acted in response to sentences whose content could convey (1) emotion (i.e., face expression and prosody) and meaning, (2) meaning alone, or (3) emotion alone. Within each condition, the kinematic effects of sentences expressing positive and negative emotions were compared. Stimuli (positive for food admiration and negative for food disgust), conveyed either by emotion or meaning affected similarly the kinematics of both grasp and reach. RESULTS: In Experiment 1, the kinematics did not vary between positive and negative sentences either when the content was expressed by both emotion and meaning, or meaning alone. In contrast, in the case of sole emotion, sentences with positive valence made faster the approach of the conspecific. In Experiment 2, the valence of emotions (positive for food admiration and negative for food disgust) affected the kinematics of both grasp and reach, independently of the modality. DISCUSSION: The lack of an effect of meaning in Experiment 1 could be due to the weak relevance of sentence meaning with respect to the motor sequence goal (feeding). Experiment 2 demonstrated that, indeed, this was the case, because when the meaning and the consequent emotion were related to the sequence goal, they affected the kinematics. In contrast, the sole emotion activated approach or avoidance toward the actor according to positive and negative valence. The data suggest a behavioral dissociation between effects of emotion and meaning.

Factors affecting athletes' motor behavior after the observation of scenes of cooperation and competition in competitive sport: the effect of sport attitude.

AIM: This study delineated how observing sports scenes of cooperation or competition modulated an action of interaction, in expert athletes, depending on their specific sport attitude. METHOD: In a kinematic study, athletes were divided into two groups depending on their attitude toward teammates (cooperative or competitive). Participants observed sport scenes of cooperation and competition (basketball, soccer, water polo, volleyball, and rugby) and then they reached for, picked up, and placed an object on the hand of a conspecific (giving action). Mixed-design ANOVAs were carried out on the mean values of grasping-reaching parameters. RESULTS: Data showed that the type of scene observed as well as the athletes' attitude affected reach-to-grasp actions to give. In particular, the cooperative athletes were speeded when they observed scenes of cooperation compared to when they observed scenes of competition. DISCUSSION: Participants were speeded when executing a giving action after observing actions of cooperation. This occurred only when they had a cooperative attitude. A match between attitude and intended action seems to be a necessary prerequisite for observing an effect of the observed type of scene on the performed action. It is possible that the observation of scenes of competition activated motor strategies which interfered with the strategies adopted by the cooperative participants to execute a cooperative (giving) sequence.

Gesture and word analysis: the same or different processes?

The present study aimed at determining whether elaboration of communicative signals (symbolic gestures and words) is always accompanied by integration with each other and, if present, this integration can be considered in support of the existence of a same control mechanism. Experiment 1 aimed at determining whether and how gesture is integrated with word. Participants were administered with a semantic priming paradigm with a lexical decision task and pronounced a target word, which was preceded by a meaningful or meaningless prime gesture. When meaningful, the gesture could be either congruent or incongruent with word meaning. Duration of prime presentation (100, 250, 400 ms) randomly varied. Voice spectra, lip kinematics, and time to response were recorded and analyzed. Formant 1 of voice spectra, and mean velocity in lip kinematics increased when the prime was meaningful and congruent with the word, as compared to meaningless gesture. In other words, parameters of voice and movement were magnified by congruence, but this occurred only when prime duration was 250 ms. Time to response to meaningful gesture was shorter in the condition of congruence compared to incongruence. Experiment 2 aimed at determining whether the mechanism of integration of a prime word with a target word is similar to that of a prime gesture with a target word. Formant 1 of the target word increased when word prime was meaningful and congruent, as compared to meaningless congruent prime. Increase was, however, present for whatever prime word duration. Experiment 3 aimed at determining whether symbolic prime gesture comprehension makes use of motor simulation. Transcranial Magnetic Stimulation was delivered to left primary motor cortex 100, 250, 500 ms after prime gesture presentation. Motor Evoked Potential of First Dorsal Interosseus increased when stimulation occurred 100 ms post-stimulus. Thus, gesture was understood within 100 ms and integrated with the target word within 250 ms. Experiment 4 excluded any hand motor simulation in order to comprehend prime word. Thus, the same type of integration with a word was present for both prime gesture and word. It was probably successive to understanding of the signal, which used motor simulation for gesture and direct access to semantics for words.

The spatial alignment effect in near and far space: a kinematic study.

The present experiment aimed at verifying whether the spatial alignment effect modifies kinematic parameters of pantomimed reaching-grasping of cups located at reachable and not reachable distance. The cup's handle could be oriented either to the right or to the left, thus inducing a grasp movement that could be either congruent or incongruent with the pantomime. The incongruence/congruence induced an increase/decrease in maximal finger aperture, which was observed when the cup was located near but not far from the body. This effect probably depended on influence of the size of the cup body on pantomime control when, in the incongruent condition, cup body was closer to the grasp hand as compared to the handle. Cup distance (near and far) influenced the pantomime even if it was actually executed in the same peripersonal space. Specifically, arm and hand temporal parameters were affected by actual cup distance as well as movement amplitudes. The results indicate that, when executing a reach-to-grasp pantomime, affordance related to the use of the object was instantiated (and in particular the spatial alignment effect became effective), but only when the object could be actually reached. Cup distance (extrinsic object property) influenced affordance, independently of the possibility to actually reach the target.

Does comprehension of symbolic gestures and corresponding-in-meaning words make use of motor simulation?

The present study aimed at determining whether or not the comprehension of symbolic gestures, and corresponding-in-meaning words, makes use of cortical circuits involved in movement execution control. Participants were presented with videos of an actress producing meaningful or meaningless gestures, pronouncing corresponding-in-meaning words or pseudo-words; they were required to judge whether the signal was meaningful or meaningless. Single pulse TMS was applied to forearm primary motor cortex area 150-200 ms after the point when the stimulus meaning could be understood. MEPs were significantly greater when processing meaningless signals as compared to a baseline condition presenting a still-and-silent actress. In contrast, this was not the case for meaningful signals whose motor activation did not differ from that for the baseline stimulus. MEPs were significantly greater for meaningless than meaningful signals and no significant difference was found between gesture and speech. On the basis of these results, we hypothesized that the observation-of/listening-to meaningless signals recruits motor areas. In contrast, this did not occur when the signals were meaningful. Overall, the data suggest that the processes related to comprehension of symbolic gestures and communicative words do not involve primary motor area and probably use brain areas involved in semantics.

Concatenation of observed grasp phases with observer's distal movements: a behavioural and TMS study.

The present study aimed at determining how actions executed by two conspecifics can be coordinated with each other, or more specifically, how the observation of different phases of a reaching-grasping action is temporary related to the execution of a movement of the observer. Participants observed postures of initial finger opening, maximal finger aperture, and final finger closing of grasp after observation of an initial hand posture. Then, they opened or closed their right thumb and index finger (experiments 1, 2 and 3). Response times decreased, whereas acceleration and velocity of actual finger movements increased when observing the two late phases of grasp. In addition, the results ruled out the possibility that this effect was due to salience of the visual stimulus when the hand was close to the target and confirmed an effect of even hand postures in addition to hand apparent motion due to the succession of initial hand posture and grasp phase. In experiments 4 and 5, the observation of grasp phases modulated even foot movements and pronunciation of syllables. Finally, in experiment 6, transcranial magnetic stimulation applied to primary motor cortex 300 ms post-stimulus induced an increase in hand motor evoked potentials of opponens pollicis muscle when observing the two late phases of grasp. These data suggest that the observation of grasp phases induced simulation which was stronger during observation of finger closing. This produced shorter response times, greater acceleration and velocity of the successive movement. In general, our data suggest best concatenation between two movements (one observed and the other executed) when the observed (and simulated) movement was to be accomplished. The mechanism joining the observation of a conspecific's action with our own movement may be precursor of social functions. It may be at the basis for interactions between conspecifics, and related to communication between individuals.