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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.

In two experiments, we compared the dynamics of corticospinal excitability when processing visually or linguistically presented tool-oriented hand actions in native speakers and sequential bilinguals. In a third experiment we used the same procedure to test non-motor, low-level stimuli, i.e. scrambled images and pseudo-words. Stimuli were presented in sequence: pictures (tool + tool-oriented hand action or their scrambled counterpart) and words (tool noun + tool-action verb or pseudo-words). Experiment 1 presented German linguistic stimuli to native speakers, while Experiment 2 presented English stimuli to non-natives. Experiment 3 tested Italian native speakers. Single-pulse trascranial magnetic stimulation (spTMS) was applied to the left motor cortex at five different timings: baseline, 200 ms after tool/noun onset, 150, 350 and 500 ms after hand/verb onset with motor-evoked potentials (MEPs) recorded from the first dorsal interosseous (FDI) and abductor digiti minimi (ADM) muscles. We report strong similarities in the dynamics of corticospinal excitability across the visual and linguistic modalities. MEPs’ suppression started as early as 150 ms and lasted for the duration of stimulus presentation (500 ms). Moreover, we show that this modulation is absent for stimuli with no motor content. Overall, our study supports the notion of a core, overarching system of action semantics shared by different modalities.

According to embodied cognition, processing language with motor content involves a simulation of this content by the brain motor system. Patients with brain lesions involving the motor system are characterized by deficits in action verbs processing in the absence of dementia. We sought to assess whether action verbs interfere with the motor behavior of patients with Parkinson's disease (PD) having tremor dominant symptoms. PD tremor is considered to result from dysfunction of cortical-subcortical motor circuits driven by dopamine depletion. In addition, PD tremor is reduced during active movement execution. Therefore, likewise movement execution, the motor simulation of bodily actions predicted by the embodiment may show to be effective in modifying tremor by interfering with a dysfunctional motor system. Here, we asked to simply read and repeat words expressing a hand-related bodily action. Abstract verbs served as control. Changes in tremor kinematics were evaluated using a monoaxial accelerometer. Seventeen PD patients with rest tremor of the upper limbs were enrolled. Tremor amplitude was significantly smaller when reading action verbs as compared to abstract verbs. We provide empirical evidence supporting the embodied cognition theory by showing that circuits mediating tremor of PD patients are distinctively affected by processing action language.

In two experiments, we compared the dynamics of corticospinal excitability when processing visually or linguistically presented tool-oriented hand actions in native speakers and sequential bilinguals. In a third experiment we used the same procedure to test non-motor, low-level stimuli, i.e. scrambled images and pseudo-words. Stimuli were presented in sequence: pictures (tool + tool-oriented hand action or their scrambled counterpart) and words (tool noun + tool-action verb or pseudo-words). Experiment 1 presented German linguistic stimuli to native speakers, while Experiment 2 presented English stimuli to non-natives. Experiment 3 tested Italian native speakers. Single-pulse trascranial brain stimulation (spTMS) was applied to the left motor cortex at five different timings: baseline, 200ms after tool/noun onset, 150, 350 and 500ms after hand/verb onset with motor-evoked potentials (MEPs) recorded from the first dorsal interosseous (FDI) and abductor digiti minimi (ADM) muscles.We report strong similarities in the dynamics of corticospinal excitability across the visual and linguistic modalities. MEPs’ suppression started as early as 150ms and lasted for the duration of stimulus presentation (500ms). Moreover, we show that this modulation is absent for stimuli with no motor content. Overall, our study supports the notion of a core, overarching system of action semantics shared by different modalities.

According to embodiment, the recruitment of the motor system is necessary to process language material expressing a motor content. Coherently, an impairment of the motor system should affect the capacity to process language items with a motor content. The aim of the present study was to assess the capacity to process graspable objects and their nouns in Parkinson's disease (PD) patients and healthy controls. Participants saw photos and nouns depicting graspable and non-graspable objects. Scrambled images and pseudo-words served as control stimuli. At 150 msec after stimulus presentation, they had to respond when the stimulus referred to a real object, and refrain from responding when it was meaningless (go-no go paradigm). In the control group, participants gave slower motor responses for stimuli (both photos and nouns) related to graspable objects as compared to non-graspable ones. This in keeping with data obtained in a previous study with young healthy participants. In the PD group, motor responses were similar for both graspable and non-graspable items. Moreover, error number was significantly greater than in controls. These findings support the notion that when the motor circuits are lesioned, like in PD, patients do not show the typical modulation of motor responses and have troubles in processing graspable objects and their nouns.

Different accounts have been proposed to explain the nature of concept representations. Embodied accounts claim a key involvement of sensory-motor systems during semantic processing while more traditional accounts posit that concepts are abstract mental entities independent of perceptual and motor brain systems. While the involvement of sensory-motor areas in concrete language processing is supported by a large number of studies, this involvement is far from being established when considering abstract language. The present study addressed abstract and concrete verb processing, by investigating the spatiotemporal dynamics of evoked responses by means of high density EEG while participants performed a semantic decision task. In addition, RTs to the same set of stimuli were collected. In both early and late time intervals, ERP scalp topography significantly differed according to word categories. Concrete verbs showed involvement of parieto-frontal networks for action, according to the implied body effector. In contrast, abstract verbs recruited mostly frontal regions outside the motor system, suggesting a non-motor semantic processing for this category. In addition, differently from what has been reported during action observation, the parietal recruitment related to concrete verbs presentation followed the frontal one. The present findings suggest that action word semantic is grounded in sensory-motor systems, provided a bodily effector is specified, while abstract concepts׳ representation cannot be easily explained by a motor embodiment.

Previous repetitive Transcranial Magnetic Stimulation and neuroimaging studies showed that Broca's area is involved in the interaction between gestures and words. However, in these studies the nature of this interaction was not fully investigated; consequently, we addressed this issue in three behavioral experiments. When compared to the expression of one signal at a time, arm kinematics slowed down and voice parameters were amplified when congruent words plus gestures were simultaneously produced (experiment 1). When word and gesture were incongruent, arm kinematics did not change regardless of word category, whereas the gesture induced variation in vocal parameters of communicative and action words only (experiments 2 and 3). Data are discussed according to the hypothesis that integration between gesture and word occurs by transferring the social intention to interact directly with the interlocutor from the gesture to the word.

Arm movements can influence language comprehension much as semantics can influence arm movement planning. Arm movement itself can be used as a linguistic signal. We reviewed neurophysiological and behavioural evidence that manual gestures and vocal language share the same control system. Studies of primate premotor cortex and, in particular, of the so-called “mirror system”, including humans, suggest the existence of a dual hand/mouth motor command system involved in ingestion activities. This may be the platform on which a combined manual and vocal communication system was constructed. In humans, speech is typically accompanied by manual gesture, speech production itself is influenced by executing or observing transitive hand actions, and manual actions play an important role in the development of speech, from the babbling stage onwards. Behavioural data also show reciprocal influence between word and symbolic gestures. Neuroimaging and repetitive transcranial magnetic stimulation (rTMS) data suggest that the system governing both speech and gesture is located in Broca's area. In general, the presented data support the hypothesis that the hand motor-control system is involved in higher order cognition.

Studies of primate premotor cortex, and, in particular, of the so-called “mirror system” suggest a double hand/mouth motor command system that may have evolved initially in the context of ingestion, and later formed a platform for combined manual and vocal communication. In humans, manual gesture, when it accompanies speech, is tightly integrated with speech production. Speech production itself is influenced by executing or observing transitive actions, and manual actions also play an important role in language development in children, from the babbling stage onwards. Behavioural data reported here even show a reciprocal influence between word and symbolic gestures. Studies employing neuroimaging and repetitive Transcranial Magnetic Stimulation (rTMS) techniques suggest that the system governing both speech and gesture is located in Broca’s area.

The present study aimed to determine whether the representation of object affordances requires specification of the effector potentially interacting with the object: specifically, in this study, vision of the interacting hand. In Experiment 1 we used an apparatus by which a fruit to be reached and grasped was identified by word reading, whereas another (interfering) fruit was visually perceived at the same location as the target. The apparatus allowed visual presentation of the agent's interacting hand or prevented it. When visually presented, the hand was perceived as still at the start position even when it moved to grasp the fruit. An interference effect on the grasp congruent with the distractor size was observed only when the hand was visible. In Experiment 2, interference was observed also when a hand different from the agent's own was visually presented. In both Experiments 1 and 2 the visible fruit interfered with the arm's reach, but the effect was independent of its size and less dependent on the visually-presented hand. A control experiment (Experiment 3) enabled comparison of the interference of visual stimuli on targets identified by word reading (Experiments 1 and 2) with that of objects identified by word reading on visually-presented targets (Experiment 3). The interference induced by visual stimuli was stronger than the interference induced by objects identified by words (i.e. affordances evoked by visual stimuli were stronger than affordances evoked by semantics). Taken together, the results of the present study suggest that the specification of the agent's effector is necessary for the elicitation of affordances. However, the elicitation of these affordances was observed for interactions between object and hand (grasp), rather than for interactions between object and arm (reach). Finally, our data confirm the influence of semantics on the control of arm movements, though less strong than that due to visual input.

BACKGROUND: The observation of action done by others determines a desynchronization of the rhythms recorded from cortical central regions. Here, we examined whether the observation of different types of hand movements (target directed, non-target directed, cyclic and non-cyclic) elicits different EEG cortical temporal patterns. METHODOLOGY: Video-clips of four types of hand movements were shown to right-handed healthy participants. Two were target directed (grasping and pointing) motor acts; two were non-target directed (supinating and clenching) movements. Grasping and supinating were performed once, while pointing and clenching twice (cyclic movements). High-density EEG was recorded and analyzed by means of wavelet transform, subdividing the time course in time bins of 200 ms. The observation of all presented movements produced a desynchronization of alpha and beta rhythms in central and parietal regions. The rhythms desynchronized as soon as the hand movement started, the nadir being reached around 700 ms after movement onset. At the end of the movement, a large power rebound occurred for all bands. Target and non-target directed movements produced an alpha band desynchronization in the central electrodes at the same time, but with a stronger desynchronization and a prolonged rebound for target directed motor acts. Most interestingly, there was a clear correlation between the velocity profile of the observed movements and beta band modulation. SIGNIFICANCE: Our data show that the observation of motor acts determines a modulation of cortical rhythm analogous to that occurring during motor act execution. In particular, the cortical motor system closely follows the velocity of the observed movements. This finding provides strong evidence for the presence in humans of a mechanism (mirror mechanism) mapping action observation on action execution motor programs.

The present study aimed at determining whether and what factors affect the control of motor sequences related to interactions between conspecifics. Experiment 1 demonstrated that during interactions between conspecifics guided by the social intention of feeding, a social affordance was activated, which modified the kinematics of sequences constituted by reaching-grasping and placing. This was relative to the same sequence directed to an inanimate target. Experiments 2 and 4 suggested that the related-to-feeding social request emitted by the receiver (i.e. the request gesture of mouth opening) is prerequisite in order to activate a social affordance. Specifically, the two experiments showed that the social request to be fed activated a social affordance even when the sequences directed towards a conspecific were not finalized to feed. Experiment 3 showed that moving inside the peripersonal space of a conspecific, who did not produce any social request, marginally affected the sequence. Finally, experiments 5 and 6 indicated that the gaze of a conspecific is necessary to make a social request effective at activating a social affordance. Summing up, the results of the present study suggest that the control of motor sequences can be changed by the interaction between giver and receiver: the interaction is characterized by a social affordance that the giver activates on the basis of social requests produced by the receiver. The gaze of the receiver is a prerequisite to make a social request effective.

The present study aimed at verifying whether and why sequences of actions directed to oneself are facilitated when compared to action sequences directed to conspecifics. In experiment 1, participants reached to grasp and brought a piece of food either to their own mouth for self-feeding or to the mouth of a conspecific for feeding. In control conditions, they executed the same sequence to place the piece of food into a mouth-like aperture in a flat container placed upon either their own mouth or the mouth of a conspecific. Kinematic analysis showed that the actions of reaching and bringing were faster when directed to the participant's own body, especially for self-feeding. The data support the hypothesis that reaching to grasp and bringing to one's own body and, in particular, one's own mouth for self-feeding, form an automatic sequence, because this is the result of more frequent execution and coordination between different effectors of one's own body, such as arm and mouth. In contrast, the same sequence directed toward a conspecific is not automatic and requires more accuracy probably because it is guided by social intentions. This hypothesis was supported by the results of control experiment 2 in which we compared the kinematics of reaching to grasp and placing the piece of food into the mouth of a conspecific (i.e. feeding) with those of reaching to grasp and placing the same piece of food into a mouth-like aperture in a human body shape (i.e. placing). Indeed, the entire sequence was slowed down during feeding when compared to placing.

A brush of 64 microwires was chronically implanted in the ventral premotor cortex of a macaque monkey. Contrary to common approaches, the wires were inserted from the white matter side. This approach, by avoiding mechanical pressure on the dura and pia mater during penetration, disturbed only minimally the cortical recording site. With this approach isolated potentials and multiunit activity were recorded for more than seven years in about one third of electrodes. The indirect insertion method also provided an excellent stability within each recording session, and in some cases even allowed recording from the same neurons for several years. Histological examination of the implanted brain region shows only a very marginal damage the recording area. Advantages and problems related to long-term recording are discussed.

Is the motor system involved in language processing? In order to clarify this issue, we carried out three behavioral experiments, using go-no-go and choice paradigms. In all the experiments, we used a semantic decision task with an early delivery of the go signal (during processing language material). Italian verbs expressing hand actions, foot actions or an abstract content served as stimuli. Participants executed intransitive (Experiment 1) or transitive (Experiment 2) actions with their right hand in response to the acoustic presentation of action-related verbs and refrained from responding to abstract verbs. The kinematics of the actions was slowed down by hand action-related verbs when compared with foot action-related verbs. In Experiment 3, hand-related and foot-related verbs were presented. Participants responded to hand-related and foot-related verbs with their hand and their foot (compatible condition) and in another block of trials they responded to hand-related and foot-related verbs with their foot and their hand (incompatible condition), respectively. In the compatible condition, the beginning of the action was faster, whereas the kinematics of the action was slower. The present findings suggest complete activation of verb-related motor programs during language processing. The data are discussed in support of the hypothesis that this complete activation is necessary requisite to understand the exact meaning of action words because goal and consequence of the actions are represented.

Does the mirror system affect the control of speech? This issue was addressed in behavioral and Transcra- nial Magnetic Stimulation (TMS) experiments. In behavioral experiment 1, participants pronounced the syllable /da/ while observing (1) a hand grasping large and small objects with power and precision grasps, respectively, (2) a foot interacting with large and small objects and (3) differently sized objects presented alone. Voice formant 1 was higher when observing power as compared to precision grasp, whereas it remained unaffected by observation of the different types of foot interaction and objects alone. In TMS experiment 2, we stimulated hand motor cortex, while participants observed the two types of grasp. Motor Evoked Potentials (MEPs) of hand muscles active during the two types of grasp were greater when observing power than precision grasp. In experiments 3–5, TMS was applied to tongue motor cortex of participants silently pronouncing the syllable /da/ and simultaneously observing power and precision grasps, pantomimes of the two types of grasps, and differently sized objects presented alone. Tongue MEPs were greater when observing power than precision grasp either executed or pantomimed. Finally, in TMS experiment 6, the observation of foot interaction with large and small objects did not modulate tongue MEPs. We hypothesized that grasp observation activated motor commands to the mouth as well as to the hand that were congruent with the hand kinematics implemented in the observed type of grasp. The commands to the mouth selectively affected postures of phonation organs and consequently basic features of phonological units.

The present study aimed to determine whether the observation of different grasps of the same object elicits automatic imitation of the kinematics of those grasps and this process influences the estimation of intrinsic target properties. In experiments 1 and 2, participants reached and grasped differently sized spheres after observation of the same objects grasped using two different types of grasp (power and precision grasp) and hand kinematics. The observed grasp kinematics were imitated especially when the vision of the target and the acting hand were precluded. In experiments 3, 4 and 5 participants matched the diameter of the spheres, either perceived or imagined, by opening their thumb and index finger (i.e. the fingers used to grasp the objects) after observation of the two types of grasp. Finger opening was larger after observation of power grasp than precision grasp, consistently with the notion that power grasp and precision grasp are preferentially used to grasp large and small objects, respectively. However, the effect was poorly observed for the small object, this depending on the fact that the participants imitated also the final position of the thumb and index finger, which were closer to each other in the power grasp. Finally, those participants, for whom the effect was stronger, reported to have perceived more differently sized objects than those really presented. The results suggest that imitation evoked by a mirror system is involved in planning how to interact with an object and in the estimation of the properties extracted for sensory-motor integration.

The aim of the present study was to determine whether Broca's area is involved in translating some aspects of arm gesture representations into mouth articulation gestures. In Experiment 1, we applied low-frequency repetitive transcranial magnetic stimulation over Broca's area and over the symmetrical loci of the right hemisphere of participants responding verbally to communicative spoken words, to gestures, or to the simultaneous presentation of the two signals. We performed also sham stimulation over the left stimulation loci. In Experiment 2, we performed the same stimulations as in Experiment 1 to participants responding with words congruent and incongruent with gestures. After sham stimulation voicing parameters were enhanced when responding to communicative spoken words or to gestures as compared to a control condition of word reading. This effect increased when participants responded to the simultaneous presentation of both communicative signals. In contrast, voicing was interfered when the verbal responses were incongruent with gestures. The left stimulation neither induced enhancement on voicing parameters of words congruent with gestures nor interference on words incongruent with gestures. We interpreted the enhancement of the verbal response to gesturing in terms of intention to interact directly. Consequently, we proposed that Broca's area is involved in the process of translating into speech aspects concerning the social intention coded by the gesture. Moreover, we discussed the results in terms of evolution to support the theory [Corballis, M. C. (2002). From hand to mouth: The origins of language. Princeton, NJ: Princeton University Press] proposing spoken language as evolved from an ancient communication system using arm gestures.

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