The role of motor inhibition in implicit negation processing: two Go/No-Go behavioral studies.

Several studies demonstrated that explicit forms of negation processing (e.g., "I don't know") recruits motor inhibitory mechanisms. However, whether this is also true for implicit negation, in which the negative meaning is implicated but not explicitly lexicalized in the sentence (e.g., "I ignore"), has never been studied before. Two Go/No-Go studies, which differed only for the time-windows to respond to the Go stimulus, were carried out. In each, participants (N = 86 in experiment 1; N = 87 in experiment 2) respond to coloured circle while reading task-irrelevant affirmative, explicit negative and implicit negative sentences. We aimed to investigate whether: (i) the processing of implicit negations recruits inhibitory mechanisms; (ii) these inhibitory resources are differently modulated by implicit and explicit negations. Results show that implicit negative sentences recruit the inhibitory resources more strongly when compared to explicit ones, probably due to their inferential nature, likely requiring deeper processing of the negative meaning. Implicit and inferential meaning (i.e., pragmatic information) are grounded too in the same mechanisms that integrate action with perception. Such findings provide further evidence to the embodied account of language, showing that even abstract aspects, like implicit negation, are grounded in the sensory-motor system, by means of functional link between language and motor activity.

Is motor inhibition involved in the processing of sentential negation? An assessment via the Stop-Signal Task.

In the last decades, the embodied approach to cognition and language gained momentum in the scientific debate, leading to evidence in different aspects of language processing. However, while the bodily grounding of concrete concepts seems to be relatively not controversial, abstract aspects, like the negation logical operator, are still today one of the main challenges for this research paradigm. In this framework, the present study has a twofold aim: (1) to assess whether mechanisms for motor inhibition underpin the processing of sentential negation, thus, providing evidence for a bodily grounding of this logic operator, (2) to determine whether the Stop-Signal Task, which has been used to investigate motor inhibition, could represent a good tool to explore this issue. Twenty-three participants were recruited in this experiment. Ten hand-action-related sentences, both in affirmative and negative polarity, were presented on a screen. Participants were instructed to respond as quickly and accurately as possible to the direction of the Go Stimulus (an arrow) and to withhold their response when they heard a sound following the arrow. This paradigm allows estimating the Stop Signal Reaction Time (SSRT), a covert reaction time underlying the inhibitory process. Our results show that the SSRT measured after reading negative sentences are longer than after reading affirmative ones, highlighting the recruitment of inhibitory mechanisms while processing negative sentences. Furthermore, our methodological considerations suggest that the Stop-Signal Task is a good paradigm to assess motor inhibition's role in the processing of sentence negation.

Mirroring the self: testing neurophysiological correlates of disturbed self-experience in schizophrenia spectrum.

BACKGROUND: Self-disorders (SDs) have been described as a core schizophrenia spectrum vulnerability phenotype, both in classic and contemporary psychopathological literature. However, such a core phenotype has not yet been investigated adopting a trans-domain approach that combines the phenomenological and the neurophysiological levels of analysis. The aim of this study is to investigate the relation between SDs and subtle, schizophrenia-specific impairments of emotional resonance that are supposed to reflect abnormalities in the mirror neurons mechanism. Specifically, we tested whether electromyographic response to emotional stimuli (i.e. a proxy for subtle changes in facial mimicry and related motor resonance mechanisms) would predict the occurrence of anomalous subjective experiences (i.e. SDs). SAMPLING AND METHODS: Eighteen schizophrenia spectrum (SzSp) patients underwent a comprehensive psychopathological examination and were contextually tested with a multimodal paradigm, recording facial electromyographic activity of muscles in response to positive and negative emotional stimuli. Experiential anomalies were explored with the Bonn Scale for the Assessment of Basic Symptoms (BSABS) and then condensed into rational subscales mapping SzSp anomalous self-experiences. RESULTS: SzSp patients showed an imbalance in emotional motor resonance with a selective bias toward negative stimuli, as well as a multisensory integration impairment. Multiple regression analysis showed that electromyographic facial reactions in response to negative stimuli presented in auditory modality specifically and strongly correlated with SD subscore. CONCLUSIONS: The study confirms the potential of SDs as target phenotype for neurobiological research and encourages research into disturbed motor/emotional resonance as possible body-level correlate of disturbed subjective experiences in SzSp.

Moving mirrors: a high-density EEG study investigating the effect of camera movements on motor cortex activation during action observation.

Action execution-perception links (mirror mechanism) have been repeatedly suggested to play crucial roles in social cognition. Remarkably, the designs of most studies exploring this topic so far excluded even the simplest traces of social interaction, such as a movement of the observer toward another individual. This study introduces a new design by investigating the effects of camera movements, possibly simulating the observer's own approaching movement toward the scene. We conducted a combined high-density EEG and behavioral study investigating motor cortex activation during action observation measured by event-related desynchronization and resynchronization (ERD/ERS) of the mu rhythm. Stimuli were videos showing a goal-related hand action filmed while using the camera in four different ways: filming from a fixed position, zooming in on the scene, approaching the scene by means of a dolly, and approaching the scene by means of a steadycam. Results demonstrated a consistently stronger ERD of the mu rhythm for videos that were filmed while approaching the scene with a steadycam. Furthermore, videos in which the zoom was applied reliably demonstrated a stronger rebound. A rating task showed that videos in which the camera approached the scene were felt as more involving and the steadycam was most able to produce a visual experience close to the one of a human approaching the scene. These results suggest that filming technique predicts time course specifics of ERD/ERS during action observation with only videos simulating the natural vision of a walking human observer eliciting a stronger ERD than videos filmed from a fixed position. This demonstrates the utility of ecologically designed studies for exploring social cognition.

The effect of Surround sound on embodiment and sense of presence in cinematic experience: a behavioral and HD-EEG study.

Although many studies have investigated spectators' cinematic experience, only a few of them explored the neurophysiological correlates of the sense of presence evoked by the spatial characteristics of audio delivery devices. Nevertheless, nowadays both the industrial and the consumer markets have been saturated by some forms of spatial audio format that enrich the audio-visual cinematic experience, reducing the gap between the real and the digitally mediated world. The increase in the immersive capabilities corresponds to the instauration of both the sense of presence and the psychological sense of being in the virtual environment and also embodied simulation mechanisms. While it is well-known that these mechanisms can be activated in the real world, it is hypothesized that they may be elicited even in a virtual acoustic spatial environment and could be modulated by the acoustic spatialization cues reproduced by sound systems. Hence, the present study aims to investigate the neural basis of the sense of presence evoked by different forms of mediation by testing different acoustic space sound delivery (Presentation modes: Monophonic, Stereo, and Surround). To these aims, a behavioral investigation and a high-density electroencephalographic (HD-EEG) study have been developed. A large set of ecological and heterogeneous stimuli extracted from feature films were used. Furthermore, participants were selected following the generalized listener selection procedure. We found a significantly higher event-related desynchronization (ERD) in the Surround Presentation mode when compared to the Monophonic Presentation mode both in Alpha and Low-Beta centro-parietal clusters. We discuss this result as an index of embodied simulation mechanisms that could be considered as a possible neurophysiological correlation of the instauration of the sense of presence.

The influence of sensorimotor experience on beauty evaluation of preschool children.

Nowadays there is a broad consensus on the role of multimodality in the construction of an embodied aesthetic experience in adults, whereas little is known about the relationship between sensorimotor and aesthetic experience during development. To fill this gap, the present study investigated whether sensorimotor experience with sculpting natural materials (i.e., clay or sand) influences beauty judgments offered to abstract artifacts made by the same materials. Five years old children (n.47) were asked to rate tactile (How smooth is it?), visual (How dark is it?) and beauty (How much do you like it?) proprieties of two artifacts using a visual-analog measurement-tool ad hoc developed to fit children's cognitive skills. Participants rated the artifacts before and after a free-hands manipulation with only one of the two sculpting materials, either sand or clay. Results showed that the greater the sensorimotor interaction experienced with the artifacts, the higher the increment of beauty rating offered to the artifacts made by the same material previously manipulated. No modulations were found for tactile and visual ratings. These results demonstrate that, even in pre-school children, aesthetic experience is specifically linked to its sensorimotor component, supporting, from a developmental perspective, the definition of aesthetic experience as intrinsically rooted on beholders' bodily experience.

Emotional body postures affect inhibitory control only when task-relevant.

A classical theoretical frame to interpret motor reactions to emotional stimuli is that such stimuli, particularly those threat-related, are processed preferentially, i.e., they are capable of capturing and grabbing attention automatically. Research has recently challenged this view, showing that the task relevance of emotional stimuli is crucial to having a reliable behavioral effect. Such evidence indicated that emotional facial expressions do not automatically influence motor responses in healthy young adults, but they do so only when intrinsically pertinent to the ongoing subject's goals. Given the theoretical relevance of these findings, it is essential to assess their generalizability to different, socially relevant emotional stimuli such as emotional body postures. To address this issue, we compared the performance of 36 right-handed participants in two different versions of a Go/No-go task. In the Emotional Discrimination task, participants were required to withhold their responses at the display of emotional body postures (fearful or happy) and to move at the presentation of neutral postures. Differently, in the control task, the same images were shown, but participants had to respond according to the color of the actor/actress' t-shirt, disregarding the emotional content. Results showed that participants made more commission errors (instances in which they moved even though the No-go signal was presented) for happy than fearful body postures in the Emotional Discrimination task. However, this difference disappeared in the control task. Such evidence indicates that, like facial emotion, emotional body expressions do not influence motor control automatically, but only when they are task-relevant.

The consequences of COVID-19 on social interactions: an online study on face covering.

The COVID-19 pandemic has dramatically changed the nature of our social interactions. In order to understand how protective equipment and distancing measures influence the ability to comprehend others' emotions and, thus, to effectively interact with others, we carried out an online study across the Italian population during the first pandemic peak. Participants were shown static facial expressions (Angry, Happy and Neutral) covered by a sanitary mask or by a scarf. They were asked to evaluate the expressed emotions as well as to assess the degree to which one would adopt physical and social distancing measures for each stimulus. Results demonstrate that, despite the covering of the lower-face, participants correctly recognized the facial expressions of emotions with a polarizing effect on emotional valence ratings found in females. Noticeably, while females' ratings for physical and social distancing were driven by the emotional content of the stimuli, males were influenced by the "covered" condition. The results also show the impact of the pandemic on anxiety and fear experienced by participants. Taken together, our results offer novel insights on the impact of the COVID-19 pandemic on social interactions, providing a deeper understanding of the way people react to different kinds of protective face covering.

Visceromotor roots of aesthetic evaluation of pain in art: an fMRI study.

Empathy for pain involves sensory and visceromotor brain regions relevant also in the first-person pain experience. Focusing on brain activations associated with vicarious experiences of pain triggered by artistic or non-artistic images, the present study aims to investigate common and distinct brain activation patterns associated with these two vicarious experiences of pain and to assess whether empathy for pain brain regions contributes to the formation of an aesthetic judgement (AJ) in non-art expert observers. Artistic and non-artistic facial expressions (painful and neutral) were shown to participants inside the scanner and then aesthetically rated in a subsequent behavioural session. Results showed that empathy for pain brain regions (i.e. bilateral insular cortex, posterior sector of the anterior cingulate cortex and the anterior portion of the middle cingulate cortex) and bilateral inferior frontal gyrus are commonly activated by artistic and non-artistic painful facial expressions. For the artistic representation of pain, the activity recorded in these regions directly correlated with participants' AJ. Results also showed the distinct activation of a large cluster located in the posterior cingulate cortex/precuneus for non-artistic stimuli. This study suggests that non-beauty-specific mechanisms such as empathy for pain are crucial components of the aesthetic experience of artworks.

Responses of mirror neurons in area F5 to hand and tool grasping observation.

Mirror neurons are a distinct class of neurons that discharge both during the execution of a motor act and during observation of the same or similar motor act performed by another individual. However, the extent to which mirror neurons coding a motor act with a specific goal (e.g., grasping) might also respond to the observation of a motor act having the same goal, but achieved with artificial effectors, is not yet established. In the present study, we addressed this issue by recording mirror neurons from the ventral premotor cortex (area F5) of two monkeys trained to grasp objects with pliers. Neuron activity was recorded during the observation and execution of grasping performed with the hand, with pliers and during observation of an experimenter spearing food with a stick. The results showed that virtually all neurons responding to the observation of hand grasping also responded to the observation of grasping with pliers and, many of them to the observation of spearing with a stick. However, the intensity and pattern of the response differed among conditions. Hand grasping observation determined the earliest and the strongest discharge, while pliers grasping and spearing observation triggered weaker responses at longer latencies. We conclude that F5 grasping mirror neurons respond to the observation of a family of stimuli leading to the same goal. However, the response pattern depends upon the similarity between the observed motor act and the one executed by the hand, the natural motor template.

Time course and specificity of sensory-motor alpha modulation during the observation of hand motor acts and gestures: a high density EEG study.

The main aim of the present study was to explore, by means of high-density EEG, the intensity and the temporal pattern of event-related sensory-motor alpha desynchronization (ERD) during the observation of different types of hand motor acts and gestures. In particular, we aimed to investigate whether the sensory-motor ERD would show a specific modulation during the observation of hand behaviors differing for goal-relatedness (hand grasping of an object and meaningless hand movements) and social relevance (communicative hand gestures and grasping within a social context). Time course analysis of alpha suppression showed that all types of hand behaviors were effective in triggering sensory-motor alpha ERD, but to a different degree depending on the category of observed hand motor acts and gestures. Meaningless gestures and hand grasping were the most effective stimuli, resulting in the strongest ERD. The observation of social hand behaviors such as social grasping and communicative gestures, triggered a more dynamic time course of ERD compared to that driven by the observation of simple grasping and meaningless gestures. These findings indicate that the observation of hand motor acts and gestures evoke the activation of a motor resonance mechanism that differs on the basis of the goal-relatedness and the social relevance of the observed hand behavior.

Audience spontaneous entrainment during the collective enjoyment of live performances: physiological and behavioral measurements.

Cardiac synchrony is a crucial component of shared experiences, considered as an objective measure of emotional processes accompanying empathic interactions. No study has investigated whether cardiac synchrony among people engaged in collective situations links to the individual emotional evaluation of the shared experience. We investigated theatrical live performances as collective experiences evoking strong emotional engagement in the audience. Cross Recurrence Quantification Analysis was applied to obtain the cardiac synchrony of twelve spectators' quartets attending to two live acting performances. This physiological measure was then correlated with spectators' emotional intensity ratings. Results showed an expected increment in synchrony among people belonging to the same quartet during both performances attendance and rest periods. Furthermore, participants' cardiac synchrony was found to be correlated with audience's convergence in the explicit emotional evaluation of the performances they attended to. These findings demonstrate that the mere co-presence of other people sharing a common experience is enough for cardiac synchrony to occur spontaneously and that it increases in function of a shared and coherent explicit emotional experience.

Cortical control of grasp in non-human primates.

The skilled use of the hand for grasping and manipulation of objects is a fundamental feature of the primate motor system. Grasping movements involve transforming the visual information about an object into a motor command appropriate for the coordinated activation of hand and finger muscles. The cerebral cortex and its descending projections to the spinal cord are known to play a crucial role for the control of grasp. Recent studies in non-human primates have provided some striking new insights into the respective contribution of the parietal and frontal motor cortical areas to the control of grasp. Also, new approaches allowed investigating the coupling of grasp-related activity in different cortical areas for the control of the descending motor command.

Haptic Aesthetics and Bodily Properties of Ori Gersht's Digital Art: A Behavioral and Eye-Tracking Study.

Experimental aesthetics has shed light on the involvement of pre-motor areas in the perception of abstract art. However, the contribution of texture perception to aesthetic experience is still understudied. We hypothesized that digital screen-based art, despite its immateriality, might suggest potential sensorimotor stimulation. Original born-digital works of art were selected and manipulated by the artist himself. Five behavioral parameters: Beauty, Liking, Touch, Proximity, and Movement, were investigated under four experimental conditions: Resolution (high/low), and Magnitude (Entire image/detail). These were expected to modulate the quantity of material and textural information afforded by the image. While the Detail condition afforded less content-related information, our results show that it augmented the image's haptic appeal. High Resolution improved the haptic and aesthetic properties of the images. Furthermore, aesthetic ratings positively correlated with sensorimotor ratings. Our results demonstrate a strict relation between the aesthetic and sensorimotor/haptic qualities of the images, empirically establishing a relationship between beholders' bodily involvement and their aesthetic judgment of visual works of art. In addition, we found that beholders' oculomotor behavior is selectively modulated by the perceptual manipulations being performed. The eye-tracking results indicate that the observation of the Entire, original images is the only condition in which the latency of the first fixation is shorter when participants gaze to the left side of the images. These results thus demonstrate the existence of a left-side bias during the observation of digital works of art, in particular, while participants are observing their original version.

Embodying the camera: An EEG study on the effect of camera movements on film spectators´ sensorimotor cortex activation.

One key feature of film consists in its power to bodily engage the viewer. Previous research has suggested lens and camera movements to be among the most effective stylistic devices involved in such engagement. In an EEG experiment we assessed the role of such movements in modulating specific spectators´ neural and experiential responses, likely reflecting such engagement. We produced short video clips of an empty room with a still, a zooming and a moving camera (steadicam) that might simulate the movement of an observer in different ways. We found an event related desynchronization of the beta components of the rolandic mu rhythm that was stronger for the clips produced with steadicam than for those produced with a still or zooming camera. No equivalent modulation in the attention related occipital areas was found, thus confirming the sensorimotor nature of spectators´ neural responses to the film clips. The present study provides the first empirical evidence that filmic means such as camera movements alone can modulate spectators' bodily engagement with film.

How context influences the interpretation of facial expressions: a source localization high-density EEG study on the "Kuleshov effect".

Few studies have explored the specificities of contextual modulations of the processing of facial expressions at a neuronal level. This study fills this gap by employing an original paradigm, based on a version of the filmic "Kuleshov effect". High-density EEG was recorded while participants watched film sequences consisting of three shots: the close-up of a target person's neutral face (Face_1), the scene that the target person was looking at (happy, fearful, or neutral), and another close-up of the same target person's neutral face (Face_2). The participants' task was to rate both valence and arousal, and subsequently to categorize the target person's emotional state. The results indicate that despite a significant behavioural 'context' effect, the electrophysiological indexes still indicate that the face is evaluated as neutral. Specifically, Face_2 elicited a high amplitude N170 when preceded by neutral contexts, and a high amplitude Late Positive Potential (LPP) when preceded by emotional contexts, thus showing sensitivity to the evaluative congruence (N170) and incongruence (LPP) between context and Face_2. The LPP activity was mainly underpinned by brain regions involved in facial expressions and emotion recognition processing. Our results shed new light on temporal and neural correlates of context-sensitivity in the interpretation of facial expressions.

"Embodied Body Language": an electrical neuroimaging study with emotional faces and bodies.

To date, most investigations in the field of affective neuroscience mainly focused on the processing of facial expressions, overlooking the exploration of emotional body language (EBL), its capability to express our emotions notwithstanding. Few electrophysiological studies investigated the time course and the neural correlates of EBL and the integration of face and body emotion-related information. The aim of the present study was to investigate both the time course and the neural correlates underlying the integration of affective information conveyed by faces and bodies. We analysed EEG activities evoked during an expression matching task, requiring the judgment of emotional congruence between sequentially presented pairs of stimuli belonging to the same category (face-face or body-body), and between stimuli belonging to different categories (face-body or body-face). We focused on N400 time window and results showed that incongruent stimuli elicited a modulation of the N400 in all comparisons except for body-face condition. This modulation was mainly detected in the Middle Temporal Gyrus and within regions related to the mirror mechanism. More specifically, while the perception of incongruent facial expressions activates somatosensory-related representations, incongruent emotional body postures also require the activation of motor and premotor representations, suggesting a strict link between emotion and action.

Proactive Control Strategies for Overt and Covert Go/NoGo Tasks: An Electrical Neuroimaging Study.

Proactive and reactive inhibition are generally intended as mechanisms allowing the withholding or suppression of overt movements. Nonetheless, inhibition could also play a pivotal role during covert actions (i.e., potential motor acts not overtly performed, despite the activation of the motor system), such as Motor Imagery (MI). In a previous EEG study, we analyzed cerebral activities reactively triggered during two cued Go/NoGo tasks, requiring execution or withholding of overt or covert imagined actions, respectively. This study revealed activation of pre-supplementary motor area (pre-SMA) and right inferior frontal gyrus (rIFG), key nodes of the network underpinning reactive inhibition of overt responses in NoGo trials, also during MI enactment, enabling the covert nature of the imagined motor response. Taking into account possible proactive engagement of inhibitory mechanisms by cue signals, for an exhaustive interpretation of these previous findings in the present study we analyzed EEG activities elicited during the preparatory phase of our cued overt and covert Go/NoGo tasks. Our results demonstrate a substantial overlap of cerebral areas activated during proactive recruitment and subsequent reactive implementation of motor inhibition in both overt and covert actions; also, different involvement of pre-SMA and rIFG emerged, in accord with the intended type (covert or overt) of incoming motor responses. During preparation of the overt Go/NoGo task, the cue is encoded in a pragmatic mode, as it primes the possible overt motor response programs in motor and premotor cortex and, through preactivation of a pre-SMA-related decisional mechanism, it triggers a parallel preparation for successful response selection and/or inhibition during the response phase. Conversely, the preparatory strategy for the covert Go/NoGo task is centered on priming of an inhibitory mechanism in rIFG, tuned to the instructed covert modality of motor performance and instantiated during subsequent MI, which allows the imagined response to remain a potential motor act.

Correction: Proactive Control Strategies for Overt and Covert Go/NoGo Tasks: an Electrical Neuroimaging Study.

[This corrects the article DOI: 10.1371/journal.pone.0152188.].

Less Empathic and More Reactive: The Different Impact of Childhood Maltreatment on Facial Mimicry and Vagal Regulation.

Facial mimicry and vagal regulation represent two crucial physiological responses to others' facial expressions of emotions. Facial mimicry, defined as the automatic, rapid and congruent electromyographic activation to others' facial expressions, is implicated in empathy, emotional reciprocity and emotions recognition. Vagal regulation, quantified by the computation of Respiratory Sinus Arrhythmia (RSA), exemplifies the autonomic adaptation to contingent social cues. Although it has been demonstrated that childhood maltreatment induces alterations in the processing of the facial expression of emotions, both at an explicit and implicit level, the effects of maltreatment on children's facial mimicry and vagal regulation in response to facial expressions of emotions remain unknown. The purpose of the present study was to fill this gap, involving 24 street-children (maltreated group) and 20 age-matched controls (control group). We recorded their spontaneous facial electromyographic activations of corrugator and zygomaticus muscles and RSA responses during the visualization of the facial expressions of anger, fear, joy and sadness. Results demonstrated a different impact of childhood maltreatment on facial mimicry and vagal regulation. Maltreated children did not show the typical positive-negative modulation of corrugator mimicry. Furthermore, when only negative facial expressions were considered, maltreated children demonstrated lower corrugator mimicry than controls. With respect to vagal regulation, whereas maltreated children manifested the expected and functional inverse correlation between RSA value at rest and RSA response to angry facial expressions, controls did not. These results describe an early and divergent functional adaptation to hostile environment of the two investigated physiological mechanisms. On the one side, maltreatment leads to the suppression of the spontaneous facial mimicry normally concurring to empathic understanding of others' emotions. On the other side, maltreatment forces the precocious development of the functional synchronization between vagal regulation and threatening social cues facilitating the recruitment of fight-or-flight defensive behavioral strategies.