The "embreathment" illusion highlights the role of breathing in corporeal awareness.

Recent theories posit that physiological signals contribute to corporeal awareness, the basic feeling that one has a body (body ownership) that acts according to one's will (body agency) and occupies a specific position (body location). Combining physiological recordings with immersive virtual reality, we found that an ecological mapping of real respiratory patterns onto a virtual body illusorily changes corporeal awareness. This new way of inducing a respiratory bodily illusion, called "embreathment," revealed that breathing is almost as important as visual appearance for inducing body ownership and more important than any other cue for body agency. These effects were moderated by individual levels of interoception, as assessed through a standard heartbeat-counting task and a new "pneumoception" task. By showing that respiratory, visual, and spatial signals exert a specific and weighted influence on the fundamental feeling that one is an embodied agent, we pave the way for a comprehensive hierarchical model of corporeal awareness.NEW & NOTEWORTHY Our body is the only object we sense from the inside; however, it is unclear how much inner physiology contributes to the global sensation of having a body and controlling it. We combine respiration recordings with immersive virtual reality and find that making a virtual body breathe like the real body gives an illusory sense of ownership and agency over the avatar, elucidating the role of a key physiological process like breathing in corporeal awareness.

Transitory Inhibition of the Left Anterior Intraparietal Sulcus Impairs Joint Actions: A Continuous Theta-Burst Stimulation Study.

Although temporal coordination is a hallmark of motor interactions, joint action (JA) partners do not simply synchronize; rather, they dynamically adapt to each other to achieve a joint goal. We created a novel paradigm to tease apart the processes underlying synchronization and JA and tested the causal contribution of the left anterior intraparietal sulcus (aIPS) in these behaviors. Participants had to synchronize their congruent or incongruent movements with a virtual partner in two conditions: (i) being instructed on what specific action to perform, independently from what action the partner performed (synchronization), and (ii) being instructed to adapt online to the partner's action (JA). Offline noninvasive inhibitory brain stimulation (continuous theta-burst stimulation) over the left aIPS selectively modulated interpersonal synchrony in JA by boosting synchrony during congruent interactions and impairing it during incongruent ones, while leaving performance in the synchronization condition unaffected. These results suggest that the left aIPS plays a causal role in supporting online adaptation to a partner's action goal, whereas it is not necessarily engaged in social situations where the goal of the partner is irrelevant. This indicates that, during JAs, the integration of one's own and the partner's action goal is supported by aIPS.

Theta synchronization over occipito-temporal cortices during visual perception of body parts.

Categorical clustering in the visual system is thought to have evolved as a function of intrinsic (intra-areal) and extrinsic (interareal) connectivity and experience. In the visual system, the extrastriate body area (EBA), an occipito-temporal region, responds to full body and body part images under the organizational principle of their functional/semantic meaning. Although frequency-specific modulations of neural activity associated with perceptive and cognitive functions are increasingly attracting the interest of neurophysiologists and cognitive neuroscientists, perceiving single body parts with different functional meaning and full body images induces time-frequency modulations over occipito-temporal electrodes are yet to be described. Here, we studied this issue by measuring EEG in participants who passively observed fingers, hands, arms and faceless full body images with four control plant stimuli, each bearing hierarchical analogy with the body stimuli. We confirmed that occipito-temporal electrodes (compatible with the location of EBA) show a larger event-related potential (ERP, N190) for body-related images. Furthermore, we identified a body part-specific (i.e. selective for hands and arms) theta event-related synchronization increase under the same electrodes. This frequency modulation associated with the perception of body effectors over occipito-temporal cortices is in line with recent findings of categorical organization of neural responses to human effectors in the visual system.

Midfrontal theta transcranial alternating current stimulation modulates behavioural adjustment after error execution.

Cognitive control during conflict monitoring, error processing, and post-error adjustment appear to be associated with the occurrence of midfrontal theta (MFÏ´). While this association is supported by correlational EEG studies, much less is known about the possible causal link between MFÏ´ and error and conflict processing. In the present study, we aimed to explore the role of band-specific effects in modulating the error system during a conflict resolution. In turn, we delivered transcranial alternating current stimulation (tACS) at different frequency bands (delta δ, theta θ, alpha α, beta ß, gamma γ) and sham stimulation over the medial frontal cortex (MFC) in 36 healthy participants performing a modified version of the Flanker task. Task performance and reports about the sensations (e.g. visual flickering, cutaneous burning) induced by the different frequency bands, were also recorded. We found that online θ-tACS increased the response speed to congruent stimuli after error execution with respect to sham stimulation. Importantly, the accuracy following the errors did not decrease because of speed-accuracy trade off. Moreover, tACS evoked visual and somatosensory sensations were significantly stronger at α-tACS and ß-tACS compared to other frequencies. Our findings suggest that theta activity plays a causative role in modulating behavioural adjustments during perceptual choices in a stimulus-response conflict task.

Visual appearance of a virtual upper limb modulates the temperature of the real hand: a thermal imaging study in Immersive Virtual Reality.

To explore the link between Sense of Embodiment (SoE) over a virtual hand and physiological regulation of skin temperature, 24 healthy participants were immersed in virtual reality through a Head Mounted Display and had their real limb temperature recorded by means of a high-sensitivity infrared camera. Participants observed a virtual right upper limb (appearing either normally, or with the hand detached from the forearm) or limb-shaped non-corporeal control objects (continuous or discontinuous wooden blocks) from a first-person perspective. Subjective ratings of SoE were collected in each observation condition, as well as temperatures of the right and left hand, wrist and forearm. The observation of these complex, body and body-related virtual scenes resulted in increased real hand temperature when compared to a baseline condition in which a 3d virtual ball was presented. Crucially, observation of non-natural appearances of the virtual limb (discontinuous limb) and limb-shaped non-corporeal objects elicited high increase in real hand temperature and low SoE. In contrast, observation of the full virtual limb caused high SoE and low temperature changes in the real hand with respect to the other conditions. Interestingly, the temperature difference across the different conditions occurred according to a topographic rule that included both hands. Our study sheds new light on the role of an external hand's visual appearance and suggests a tight link between higher-order bodily self-representations and topographic regulation of skin temperature.

Susceptibility to the rubber hand illusion does not tell the whole body-awareness story.

The rubber hand illusion (RHI) is an enigmatic illusion that creates a feeling of owning an artificial limb. Enthusiasts of this paradigm assert that it operationalizes bodily self-awareness, but there are reasons to doubt such a clear link. Because little is known about other functional contributions to the RHI, including effects of context-dependent visual processing and cognitive control or the ability to resolve intermodal conflict, we carried out two complementary experiments. In the first, we examined the relationships between the RHI and (1) body awareness, as assessed by the Body Perception Questionnaire (BPQ); (2) context-dependent visual processing, as assessed by the rod-and-frame test (RFT); and (3) conflict resolution, as assessed by the Stroop test. We found a significant positive correlation between the RHI-associated proprioceptive drift and context-dependent visual processing on the RFT, but not between the RHI and body awareness on the BPQ. In the second experiment, we examined the RHI in advanced yoga practitioners with an embodied lifestyle and a heightened sense of their own body in space. They succumbed to the illusion just as much as did yoga-naïve control participants, despite significantly greater body awareness on the BPQ. These findings suggest that susceptibility to the RHI and awareness of one's own body are at least partially independent processes.

Fooling the kickers but not the goalkeepers: behavioral and neurophysiological correlates of fake action detection in soccer.

Studies demonstrate that elite athletes are able to extract kinematic information of observed domain-specific actions to predict their future course. Little is known, however, on the perceptuo-motor processes and neural correlates of the athletes' ability to predict fooling actions. Combining psychophysics and transcranial magnetic stimulation, we explored the impact of motor and perceptual expertise on the ability to predict the fate of observed actual or fake soccer penalty kicks. We manipulated the congruence between the model's body kinematics and the subsequent ball trajectory and investigated the prediction performance and cortico-spinal reactivity of expert kickers, goalkeepers, and novices. Kickers and goalkeepers outperformed novices by anticipating the actual kick direction from the model's initial body movements. However, kickers were more often fooled than goalkeepers and novices in cases of incongruent actions. Congruent and incongruent actions engendered a comparable facilitation of kickers' lower limb motor representation, but their neurophysiological response was correlated with their greater susceptibility to be fooled. Moreover, when compared with actual actions, motor facilitation for incongruent actions was lower among goalkeepers and higher among novices. Thus, responding to fooling actions requires updation of simulative motor representations of others' actions and is facilitated by visual rather than by motor expertise.

Compensatory plasticity in the action observation network: virtual lesions of STS enhance anticipatory simulation of seen actions.

Observation of snapshots depicting ongoing motor acts increases corticospinal motor excitability. Such motor facilitation indexes the anticipatory simulation of observed (implied) actions and likely reflects computations occurring in the parietofrontal nodes of a cortical network subserving action perception (action observation network, AON). However, direct evidence for the active role of AON in simulating the future of seen actions is lacking. Using a perturb-and-measure transcranial magnetic stimulation (TMS) approach, we show that off-line TMS disruption of regions within (inferior frontal cortex, IFC) and upstream (superior temporal sulcus, STS) the parietofrontal AON transiently abolishes and enhances the motor facilitation to observed implied actions, respectively. Our findings highlight the critical role of IFC in anticipatory motor simulation. More importantly, they show that disruption of STS calls into play compensatory motor simulation activity, fundamental for counteracting the noisy visual processing induced by TMS. Thus, short-term plastic changes in the AON allow motor simulation to deal with any gap or ambiguity of ever-changing perceptual worlds. These findings support the active, compensatory, and predictive role of frontoparietal nodes of the AON in the perception and anticipatory simulation of implied actions.

The pattern theory of compassion.

Concepts of empathy, sympathy and compassion are often confused in a variety of literatures. This article proposes a pattern-theoretic approach to distinguishing compassion from empathy and sympathy. Drawing on psychology, Western philosophy, affective neuroscience, and contemplative science, we clarify the nature of compassion as a specific pattern of dynamically related factors that include physiological, cognitive, and affective processes, relational/intersubjective processes, and motivational/action tendencies. We also show that the dynamic nature of the compassion pattern is reflected in neuroscientific findings, as well as in compassion practice. The pattern theory of compassion allows us to make several clear distinctions between compassion, empathy, and sympathy.

Putting yourself in the skin of a black avatar reduces implicit racial bias.

Although it has been shown that immersive virtual reality (IVR) can be used to induce illusions of ownership over a virtual body (VB), information on whether this changes implicit interpersonal attitudes is meager. Here we demonstrate that embodiment of light-skinned participants in a dark-skinned VB significantly reduced implicit racial bias against dark-skinned people, in contrast to embodiment in light-skinned, purple-skinned or with no VB. 60 females participated in this between-groups experiment, with a VB substituting their own, with full-body visuomotor synchrony, reflected also in a virtual mirror. A racial Implicit Association Test (IAT) was administered at least three days prior to the experiment, and immediately after the IVR exposure. The change from pre- to post-experience IAT scores suggests that the dark-skinned embodied condition decreased implicit racial bias more than the other conditions. Thus, embodiment may change negative interpersonal attitudes and thus represent a powerful tool for exploring such fundamental psychological and societal phenomena.

Long- and short-term plastic modeling of action prediction abilities in volleyball.

Athletes show superior abilities not only in executing complex actions, but also in anticipating others' moves. Here, we explored how visual and motor experiences contribute to forge elite action prediction abilities in volleyball players. Both adult athletes and supporters were more accurate than novices in predicting the fate of volleyball floating services by viewing the initial ball trajectory, while only athletes could base their predictions on body kinematics. Importantly, adolescents assigned to physical practice training improved their ability to predict the fate of the actions by reading body kinematics, while those assigned to the observational practice training improved only in understanding the ball trajectory. The results suggest that physical and observational practice might provide complementary and mutually reinforcing contributions to the superior perceptual abilities of elite athletes. Moreover, direct motor experience is required to establish novel perceptuo-motor representations that are used to predict others' actions ahead of realization.

Gut markers of bodily self-consciousness in men.

Bodily self-consciousness, the state of mind that allows humans to be aware of their own body, forms the backdrop for almost every human experience, yet its underpinnings remain elusive. Here we combine an ingestible, minimally invasive capsule with surface electrogastrography to probe if gut physiology correlates with bodily self-consciousness in a sample of healthy men during a virtual bodily illusion. We discover that specific patterns of stomach and bowel activity (temperature, pressure, and pH) covary with specific facets of bodily self-consciousness (feelings of body location, agency, and disembodiment). These results uncover the hitherto untapped potential of minimally invasive probes to study the link between mental and gut states and show the significance of deep visceral organs in the self-conscious perception of ourselves as embodied beings.

Gesture discrimination in primary progressive aphasia: the intersection between gesture and language processing pathways.

The issue of the relationship between language and gesture processing and the partial overlap of their neural representations is of fundamental importance to neurology, psychology, and social sciences. Patients suffering from primary progressive aphasia, a clinical syndrome characterized by comparatively isolated language deficits, may provide direct evidence for anatomical and functional association between specific language deficits and gesture discrimination deficits. A consecutive series of 16 patients with primary progressive aphasia and 16 matched control subjects participated. Our nonverbal gesture discrimination task consisted of 19 trials. In each trial, participants observed three video clips showing the same gesture performed correctly in one clip and incorrectly in the other two. Subjects had to indicate which of the three versions was correct. Language and gesture production were evaluated by means of conventional tasks. All participants underwent high-resolution structural and diffusion tensor magnetic resonance imaging. Ten of the primary progressive aphasia patients showed a significant deficit on the nonverbal gesture discrimination task. A factor analysis revealed that this deficit clustered with gesture imitation, word and pseudoword repetition, and writing-to-dictation. Individual scores on this cluster correlated with volume in the left anterior inferior parietal cortex extending into the posterior superior temporal gyrus. Probabilistic tractography indicated this region comprised the cortical relay station of the indirect pathway connecting the inferior frontal gyrus and the superior temporal cortex. Thus, the left perisylvian temporoparietal area may underpin verbal imitative behavior, gesture imitation, and gesture discrimination indicative of a partly shared neural substrate for language and gesture resonance.

Representation of body identity and body actions in extrastriate body area and ventral premotor cortex.

Although inherently linked, body form and body action may be represented in separate neural substrates. Using repetitive transcranial magnetic stimulation in healthy individuals, we show that interference with the extrastriate body area impairs the discrimination of bodily forms, and interference with the ventral premotor cortex impairs the discrimination of bodily actions. This double dissociation suggests that whereas extrastriate body area mainly processes actors' body identity, premotor cortex is crucial for visual discriminations of actions.

Anosognosia for limb and bucco-facial apraxia as inferred from the recognition of gestural errors.

Anosognosia is a multifaceted syndrome characterized by a lack of awareness of motor, cognitive, or emotional deficits. While most studies have focused on basic motor disorders such as hemiplegia, only recently, the issue of whether anosognosia also concerns higher-order motor disorders like apraxia has been addressed. Here, we explore the existence of a specific form of anosognosia for apraxia in forty patients with uni-hemispheric vascular lesions. The patients were requested to imitate actions involving upper limb or bucco-facial body parts and then judge their performance. Successively, they were also asked to observe video recordings of the same actions performed by themselves or by other patients and judge the accuracy of the displayed actions. The comparison of participants versus examiner judgement and between error recognition of others' versus self's actions was considered as an index of awareness deficit for the online and offline conditions, respectively. Evidence was found that awareness deficits occurred both immediately after action execution (online anosognosia) and in the video recording task (offline anosognosia). Moreover, bucco-facial and limb apraxic patients were specifically unaware of their errors in bucco-facial and limb actions, respectively, indicating for the first time a topographical organization of the syndrome. Our approach allowed us to distinguish awareness deficits from more general disorders in error recognition; indeed, anosognosic patients were able to identify errors when the same action was executed by another patient but not when the video showed their own actions. Finally, we provide evidence that anosognosia for apraxia might be associated with frontal cortical and subcortical networks.

Visually induced analgesia: seeing the body reduces pain.

Given previous reports of strong interactions between vision and somatic senses, we investigated whether vision of the body modulates pain perception. Participants looked into a mirror aligned with their body midline at either the reflection of their own left hand (creating the illusion that they were looking directly at their own right hand) or the reflection of a neutral object. We induced pain using an infrared laser and recorded nociceptive laser-evoked potentials (LEPs). We also collected subjective ratings of pain intensity and unpleasantness. Vision of the body produced clear analgesic effects on both subjective ratings of pain and the N2/P2 complex of LEPs. Similar results were found during direct vision of the hand, without the mirror. Furthermore, these effects were specific to vision of one's own hand and were absent when viewing another person's hand. These results demonstrate a novel analgesic effect of non-informative vision of the body.

The body in the brain revisited.

Corporeal awareness is a difficult concept which refers to perception, knowledge and evaluation of one's own body as well as of other bodies. We discuss here some controversies regarding the significance of the concepts of body schema and body image, as variously entertained by different authors, for the understanding of corporeal awareness, and consider some newly proposed alternatives. We describe some recent discoveries of cortical areas specialized for the processing of bodily forms and bodily actions, as revealed by neuroimaging, neurophysiological, and lesion studies. We further describe new empirical and theoretical evidence for the importance of interoception, in addition to exteroception and proprioception, for corporeal awareness, and discuss how itch, a typical interoceptive input, has been wrongly excluded from the classic concept of the proprioceptive-tactile body schema. Finally, we consider the role of the insular cortex as the terminal cortical station of interoception and other bodily signals, along with Craig's proposal that the human insular cortex sets our species apart from other species by supporting consciousness of the body and the self. We conclude that corporeal awareness depends on the spatiotemporally distributed activity of many bodies in the brain, none of which is isomorphic with the actual body.

Representing actions through their sound.

Since the discovery of 'mirror neurons' in the monkey premotor and parietal cortex, an increasing body of evidence in animals and humans alike has supported the notion of the inextricable link between action execution and action perception. Although research originally focused on the relationship between performed and viewed actions, more recent studies highlight the importance of representing the actions of others through audition. In the first part of this article, we discuss animal studies, which provide direct evidence that action is inherently linked to multi-sensory cues, as well as the studies carried out on healthy subjects by using state-of-the-art cognitive neuroscience techniques such as functional magnetic resonance imaging (fMRI), event-related potentials (ERP), magnetoencephalography (MEG), and transcranial magnetic stimulation (TMS). In the second section, we review the lesion analysis studies in brain-damaged patients demonstrating the link between 'resonant' fronto-parieto-temporal networks and the ability to represent an action by hearing its sound. Moreover, we examine the evidence in favour of somatotopy as a possible representational rule underlying the auditory mapping of actions and consider the links between language and audio-motor action mapping. We conclude with a discussion of some outstanding questions for future research on the link between actions and the sounds they produce.

Visual similarity and psychological closeness are neurally dissociable in the brain response to vicarious pain.

Personal and vicarious experience of pain activate partially overlapping brain networks. This brain activity is further modulated by low- and high-order factors, e.g., the perceived intensity of the model's pain and the model's similarity with the onlooker, respectively. We investigated which specific aspect of similarity modulates such empathic reactivity, focusing on the potential differentiation between visual similarity and psychological closeness between the onlooker and different types of models. To this aim, we recorded fMRI data in neurotypical participants who observed painful and tactile stimuli delivered to an adult human hand, a baby human hand, a puppy dog paw, and an anthropomorphic robotic hand. The interaction between type of vicarious experience (pain, touch) and nature of model (adult, baby, dog, robot) showed that the right supramarginal gyrus (rSMG) was selectively active for visual similarity (more active during vicarious pain for the adult and baby models), while the anterior cingulate cortex (ACC) was more sensitive to psychological closeness (specifically linked to vicarious pain for the baby model). These findings indicate that visual similarity and psychological closeness between onlooker and model differentially affect the activity of brain regions specifically implied in encoding interindividual sharing of sensorimotor and affective aspects of vicarious pain, respectively.

Midline frontal and occipito-temporal activity during error monitoring in dyadic motor interactions.

Discrepancies between sensory predictions and action outcome are at the base of error coding. However, these phenomena have mainly been studied focussing on individual performance. Here, we explored EEG responses to motor prediction errors during a human-avatar interaction and show that Theta/Alpha activity of the frontal error-monitoring system works in phase with activity of the occipito-temporal node of the action observation network. Our motor interaction paradigm required healthy individuals to synchronize their reach-to-grasp movements with those of a virtual partner in conditions that did (Interactive) or did not require (Cued) movement prediction and adaptation to the partner's actions. Crucially, in 30% of the trials the virtual partner suddenly and unpredictably changed its movement trajectory thereby violating the human participant's expectation. These changes elicited error-related neuromarkers (ERN/Pe - Theta/Alpha modulations) over fronto-central electrodes during the Interactive condition. Source localization and connectivity analyses showed that the frontal Theta/Alpha activity induced by violations of the expected interactive movements was in phase with occipito-temporal Theta/Alpha activity. These results expand current knowledge about the neural correlates of on-line interpersonal motor interactions linking the frontal error-monitoring system to visual, body motion-related, responses.