Beyond peripersonal boundaries: insights from crossmodal interactions.

We experience our self as a body located in space. However, how information about self-location is integrated into multisensory processes underlying the representation of the peripersonal space (PPS), is still unclear. Prior studies showed that the presence of visual information related to oneself modulates the multisensory processes underlying PPS. Here, we used the crossmodal congruency effect (CCE) to test whether this top-down modulation depends on the spatial location of the body-related visual information. Participants responded to tactile events on their bodies while trying to ignore a visual distractor presented on the mirror reflection of their body (Self) either in the peripersonal space (Near) or in the extrapersonal space (Far). We found larger CCE when visual events were presented on the mirror reflection in the peripersonal space, as compared to the extrapersonal space. These results suggest that top-down modulation of the multisensory bodily self is only possible within the PPS.

The Vestibulocerebellum and the Shattered Self: a Resting-State Functional Connectivity Study in Posttraumatic Stress Disorder and Its Dissociative Subtype.

The flocculus is a region of the vestibulocerebellum dedicated to the coordination of neck, head, and eye movements for optimal posture, balance, and orienting responses. Despite growing evidence of vestibular and oculomotor impairments in the aftermath of traumatic stress, little is known about the effects of chronic psychological trauma on vestibulocerebellar functioning. Here, we investigated alterations in functional connectivity of the flocculus at rest among individuals with post-traumatic stress disorder (PTSD) and its dissociative subtype (PTSD + DS) as compared to healthy controls. Forty-four healthy controls, 57 PTSD, and 32 PTSD + DS underwent 6-min resting-state MRI scans. Seed-based functional connectivity analyses using the right and left flocculi as seeds were performed. These analyses revealed that, as compared to controls, PTSD and PTSD + DS showed decreased resting-state functional connectivity of the left flocculus with cortical regions involved in bodily self-consciousness, including the temporo-parietal junction, the supramarginal and angular gyri, and the superior parietal lobule. Moreover, as compared to controls, the PTSD + DS group showed decreased functional connectivity of the left flocculus with the medial prefrontal cortex, the precuneus, and the mid/posterior cingulum, key regions of the default mode network. Critically, when comparing PTSD + DS to PTSD, we observed increased functional connectivity of the right flocculus with the right anterior hippocampus, a region affected frequently by early life trauma. Taken together, our findings point toward the crucial role of the flocculus in the neurocircuitry underlying a coherent and embodied self, which can be compromised in PTSD and PTSD + DS.

Visual perspective, distance, and felt presence of others in dreams.

The peripersonal space, that is, the limited space surrounding the body, involves multisensory coding and representation of the self in space. Previous studies have shown that peripersonal space representation and the visual perspective on the environment can be dramatically altered when neurotypical individuals self-identify with a distant avatar (i.e., in virtual reality) or during clinical conditions (i.e., out-of-body experience, heautoscopy, depersonalization). Despite its role in many cognitive/social functions, the perception of peripersonal space in dreams, and its relationship with the perception of other characters (interpersonal distance in dreams), remain largely uncharted. The present study aimed to explore the visuospatial properties of this space, which is likely to underlie self-location as well as self/other distinction in dreams. 530 healthy volunteers answered a web-based questionnaire to measure their dominant visuo-spatial perspective in dreams, the frequency of recall for felt distances between their dream self and other dream characters, and the dreamers' viewing angle of other dream characters. Most participants reported dream experiences from a first-person perspective (1PP) (82%) compared to a third-person perspective (3PP) (18%). Independent of their dream perspective, participants reported that they generally perceived other dream characters in their close space, that is, at distance of either between 0 and 90 cm, or 90-180 cm, than in further spaces (180-270 cm). Regardless of the perspective (1PP or 3PP), both groups also reported more frequently seeing other dream characters from eye level (0° angle of viewing) than from above (30° and 60°) or below eye level (-30° and -60°). Moreover, the intensity of sensory experiences in dreams, as measured by the Bodily Self-Consciousness in Dreams Questionnaire, was higher in individuals who habitually see other dream characters closer to their personal dream self (i.e., within 0-90 cm and 90-180 cm). These preliminary findings offer a new, phenomenological account of space representation in dreams with regards to the felt presence of others. They might provide insights not only to our understanding of how dreams are formed, but also to the type of neurocomputations involved in self/other distinction.

Did you hear your action? An ecological approach to the senses of ownership and agency.

The Sense of Ownership (SoO) and the Sense of Agency (SoA) are two key components of bodily self-consciousness. In this experiment, we investigated how they are affected by variations in the ecological validity of the moving Rubber Hand Illusion (mRHI) paradigm, which typically include three movement conditions: active congruent, passive congruent, and active incongruent. These conditions were either in a session in which no auditory feedback associated with finger-tapping was eliminated, or in a session in which such a feedback occurred. Since the presence of the auditory feedback more closely corresponds to what individuals experience in daily life when they tap their finger on a surface, sessions with feedback are more ecologically valid, and should thus result in a more marked SoO. Results indicated that in the active movement condition in which the illusion is typically found (congruent), the effect was enhanced when the feedback was present. This advantage emerged on both on objective and subjective measures of SoO. The SoA, on the other hand, is not affected by the auditory feedback.

From statistical regularities in multisensory inputs to peripersonal space representation and body ownership: Insights from a neural network model.

Peripersonal space (PPS), the interface between the self and the environment, is represented by a network of multisensory neurons with visual (or auditory) receptive fields anchored to specific body parts, and tactile receptive fields covering the same body parts. Neurophysiological and behavioural features of hand PPS representation have been previously modelled through a neural network constituted by one multisensory population integrating tactile inputs with visual/auditory external stimuli. Reference frame transformations were not explicitly modelled, as stimuli were encoded in pre-computed hand-centred coordinates. Here we present a novel model, aiming to overcome this limitation by including a proprioceptive population encoding hand position. We confirmed behaviourally the plausibility of the proposed architecture, showing that visuo-proprioceptive information is integrated to enhance tactile processing on the hand. Moreover, the network's connectivity was spontaneously tuned through a Hebbian-like mechanism, under two minimal assumptions. First, the plasticity rule was designed to learn the statistical regularities of visual, proprioceptive and tactile inputs. Second, such statistical regularities were simply those imposed by the body structure. The network learned to integrate proprioceptive and visual stimuli, and to compute their hand-centred coordinates to predict tactile stimulation. Through the same mechanism, the network reproduced behavioural correlates of manipulations implicated in subjective body ownership: the invisible and the rubber hand illusion. We thus propose that PPS representation and body ownership may emerge through a unified neurocomputational process; the integration of multisensory information consistently with a model of the body in the environment, learned from the natural statistics of sensory inputs.

Testosterone administration in women increases the size of their peripersonal space.

Peripersonal space (PPS) is the space immediately surrounding the body, conceptualised as a sensory-motor interface between body and environment. PPS size differs between individuals and contexts, with intrapersonal traits and states, as well as social factors having a determining role on the size of PPS. Testosterone plays an important role in regulating social-motivational behaviour and is known to enhance dominance motivation in an implicit and unconscious manner. We investigated whether the dominance-enhancing effects of testosterone reflect as changes in the representation of PPS in a within-subjects testosterone administration study in women (N = 19). Participants performed a visuo-tactile integration task in a mixed-reality setup. Results indicated that the administration of testosterone caused a significant enlargement of participants' PPS, suggesting that testosterone caused participants to implicitly appropriate a larger space as their own. These findings suggest that the dominance-enhancing effects of testosterone reflect at the level of sensory-motor processing in PPS.

The monologue of the double: Allocentric reduplication of the own voice alters bodily self-perception.

During autoscopic phenomena, people perceive a double of themselves in extrapersonal space. Such clinical allocentric self-experiences sometimes co-occur with auditory hallucinations, yet experimental setups to induce similar illusions in healthy participants have generally neglected acoustic cues. We investigated whether feeling the presence of an auditory double could be provoked experimentally by recording healthy participants' own versus another person's voice and movements using binaural headphones from an egocentric (the participants' own) and an allocentric (a dummy head located elsewhere) perspective. When hearing themselves allocentrically, participants reported feeling a self-identified presence extracorporeally, an arguably distinct quality of autoscopy. Our results suggest that participants without hallucinatory experiences localized their own voice closer to themselves compared to that of another person. Explorative findings suggest that distinct patterns for hallucinators should be further investigated. This study suggests a successful induction of the feeling of an acoustic doppelganger, bridging clinical phenomena and experimental work.

Influence of vestibular signals on bodily self-consciousness: Different sensory weighting strategies based on visual dependency.

Previous studies showed that the vestibular system is crucial for multisensory integration, however, its contribution to bodily self-consciousness more specifically on full-body illusions is not well understood. Thus, the current study examined the role of visuo-vestibular conflict on a full-body illusion (FBI) experiment that was induced during a supine body position. In a mixed design experiment, 56 participants underwent through a full-body illusion protocol. During the experiment, half of the participants received synchronous visuo-tactile stimulation, and the other half received asynchronous visuo-tactile stimulation, while their physical body was lying in a supine position, but the virtual body was standing. Additionally, the contribution of individual sensory weighting strategies was investigated via the Rod and Frame task (RFT), which was applied both before (pre-FBI standing and pre-FBI supine) and after the full-body illusion (post-FBI supine) protocol. Subjective reports of the participants confirmed previous findings suggesting that there was a significant increase in ownership over a virtual body during synchronous visuo-tactile stimulation. Additionally, further categorization of participants based on their visual dependency (by RFT) showed that those participants who rely more on visual information (visual field dependents) perceived the full-body illusion more strongly than non-visual field dependents during the synchronous visuo-tactile stimulation condition. Further analysis provided not only a quantitative demonstration of full-body illusion but also revealed changes in perceived self-orientation based on their field dependency. Altogether, findings of the current study make further contributions to our understanding of the vestibular system and brought new insight for individual sensory weighting strategies during a full-body illusion.

First-person body view modulates the neural substrates of episodic memory and autonoetic consciousness: A functional connectivity study.

Episodic memory (EM) is classically conceived as a memory for events, localized in space and time, and characterized by autonoetic consciousness (ANC) allowing to mentally travel back in time and subjectively relive an event. Building on recent evidence that the first-person visual co-perception of one's own body during encoding impacts EM, we used a scene recognition task in immersive virtual reality (VR) and measured how first-person body view would modulate peri-encoding resting-state fMRI, EM performance, and ANC. Specifically, we investigated the impact of body view on post-encoding functional connectivity in an a priori network of regions related either to EM or multisensory bodily processing and used these regions in a seed-to-whole brain analysis. Post-encoding connectivity between right hippocampus (rHC) and right parahippocampus (rPHC) was enhanced when participants encoded scenes while seeing their body. Moreover, the strength of connectivity between the rHC, rPHC and the neocortex displayed two main patterns with respect to body view. The connectivity with a sensorimotor fronto-parietal network, comprising primary somatosensory and primary motor cortices, correlated with ANC after - but not before - encoding, depending on body view. The opposite change of connectivity was found between rHC, rPHC and the medial parietal cortex (from being correlated with ANC before encoding to an absence of correlation after encoding), but irrespective of body view. Linking immersive VR and fMRI for the study of EM and ANC, these findings suggest that seeing one's own body during encoding impacts the brain activity related to EM formation by modulating the connectivity between the right hippocampal formation and the neocortical regions involved in the processing of multisensory bodily signals and self-consciousness.

Spontaneous expression of mirror self-recognition in monkeys after learning precise visual-proprioceptive association for mirror images.

Mirror self-recognition (MSR) is generally considered to be an intrinsic cognitive ability found only in humans and a few species of great apes. Rhesus monkeys do not spontaneously show MSR, but they have the ability to use a mirror as an instrument to find hidden objects. The mechanism underlying the transition from simple mirror use to MSR remains unclear. Here we show that rhesus monkeys could show MSR after learning precise visual-proprioceptive association for mirror images. We trained head-fixed monkeys on a chair in front of a mirror to touch with spatiotemporal precision a laser pointer light spot on an adjacent board that could only be seen in the mirror. After several weeks of training, when the same laser pointer light was projected to the monkey's face, a location not used in training, all three trained monkeys successfully touched the face area marked by the light spot in front of a mirror. All trained monkeys passed the standard face mark test for MSR both on the monkey chair and in their home cage. Importantly, distinct from untrained control monkeys, the trained monkeys showed typical mirror-induced self-directed behaviors in their home cage, such as using the mirror to explore normally unseen body parts. Thus, bodily self-consciousness may be a cognitive ability present in many more species than previously thought, and acquisition of precise visual-proprioceptive association for the images in the mirror is critical for revealing the MSR ability of the animal.

Illusory own body perceptions mapped in the cingulate cortex-An intracranial stimulation study.

Body awareness is the result of sensory integration in the posterior parietal cortex; however, other brain structures are part of this process. Our goal is to determine how the cingulate cortex is involved in the representation of our body. We retrospectively selected patients with drug-resistant epilepsy, explored by stereo-electroencephalography, that had the cingulate cortex sampled outside the epileptogenic zone. The clinical effects of high-frequency electrical stimulation were reviewed and only those sites that elicited changes related to body perception were included. Connectivity of the cingulate cortex and other cortical structures was assessed using the h2 coefficient, following a nonlinear regression analysis of the broadband EEG signal. Poststimulation changes in connectivity were compared between two sets of stimulations eliciting or not eliciting symptoms related to body awareness (interest and control groups). We included 17 stimulations from 12 patients that reported different types of body perception changes such as sensation of being pushed toward right/left/up, one limb becoming heavier/lighter, illusory sensation of movement, sensation of pressure, sensation of floating or detachment of one hemi-body. High-frequency stimulation in the cingulate cortex (1 anterior, 15 middle, 1 posterior part) elicits body perception changes, associated with a decreased connectivity of the dominant posterior insula and increased coupling between other structures, located particularly in the nondominant hemisphere.

Self-Grounded Vision: Hand Ownership Modulates Visual Location through Cortical ß and γ Oscillations.

Vision is known to be shaped by context, defined by environmental and bodily signals. In the Taylor illusion, the size of an afterimage projected on one's hand changes according to proprioceptive signals conveying hand position. Here, we assessed whether the Taylor illusion does not just depend on the physical hand position, but also on bodily self-consciousness as quantified through illusory hand ownership. Relying on the somatic rubber hand illusion, we manipulated hand ownership, such that participants embodied a rubber hand placed next to their own hand. We found that an afterimage projected on the participant's hand drifted depending on illusory ownership between the participants' two hands, showing an implication of self-representation during the Taylor illusion. Oscillatory power analysis of electroencephalographic signals showed that illusory hand ownership was stronger in participants with stronger α suppression over left sensorimotor cortex, whereas the Taylor illusion correlated with higher ß/γ power over frontotemporal regions. Higher γ connectivity between left sensorimotor and inferior parietal cortex was also found during illusory hand ownership. These data show that afterimage drifts in the Taylor illusion do not only depend on the physical hand position but also on subjective ownership, which itself is based on the synchrony of somatosensory signals from the two hands. The effect of ownership on afterimage drifts is associated with ß/γ power and γ connectivity between frontoparietal regions and the visual cortex. Together, our results suggest that visual percepts are not only influenced by bodily context but are self-grounded, mapped on a self-referential frame. SIGNIFICANCE STATEMENT: Vision is influenced by the body: in the Taylor illusion, the size of an afterimage projected on one's hand changes according to tactile and proprioceptive signals conveying hand position. Here, we report a new phenomenon revealing that the perception of afterimages depends not only on bodily signals, but also on the sense of self. Relying on the rubber hand illusion, we manipulated hand ownership, so that participants embodied a rubber hand placed next to their own hand. We found that visual afterimages projected on the participant's hand drifted laterally, only when the rubber hand was embodied. Electroencephalography revealed spectral dissociations between somatic and visual effects, and higher γ connectivity along the dorsal visual pathways when the rubber hand was embodied.

The cerebellum after trauma: Resting-state functional connectivity of the cerebellum in posttraumatic stress disorder and its dissociative subtype.

The cerebellum plays a key role not only in motor function but also in affect and cognition. Although several psychopathological disorders have been associated with overall cerebellar dysfunction, it remains unclear whether different regions of the cerebellum contribute uniquely to psychopathology. Accordingly, we compared seed-based resting-state functional connectivity of the anterior cerebellum (lobule IV-V), of the posterior cerebellum (Crus I), and of the anterior vermis across posttraumatic stress disorder (PTSD; n = 65), its dissociative subtype (PTSD + DS; n = 37), and non-trauma-exposed healthy controls (HC; n = 47). Here, we observed decreased functional connectivity of the anterior cerebellum and anterior vermis with brain regions involved in somatosensory processing, multisensory integration, and bodily self-consciousness (temporo-parietal junction, postcentral gyrus, and superior parietal lobule) in PTSD + DS as compared to PTSD and HC. Moreover, the PTSD + DS group showed increased functional connectivity of the posterior cerebellum with cortical areas related to emotion regulation (ventromedial prefrontal and orbito-frontal cortex, subgenual anterior cingulum) as compared to PTSD. By contrast, PTSD showed increased functional connectivity of the anterior cerebellum with cortical areas associated with visual processing (fusiform gyrus), interoceptive awareness (posterior insula), memory retrieval, and contextual processing (hippocampus) as compared to HC. Finally, we observed decreased functional connectivity between the posterior cerebellum and prefrontal regions involved in emotion regulation, in PTSD as compared to HC. These findings not only highlight the crucial role of each cerebellar region examined in the psychopathology of PTSD but also reveal unique alterations in functional connectivity distinguishing the dissociative subtype of PTSD versus PTSD.

Transient Modulations of Neural Responses to Heartbeats Covary with Bodily Self-Consciousness.

UNLABELLED: Recent research has investigated self-consciousness associated with the multisensory processing of bodily signals (e.g., somatosensory, visual, vestibular signals), a notion referred to as bodily self-consciousness, and these studies have shown that the manipulation of bodily inputs induces changes in bodily self-consciousness such as self-identification. Another line of research has highlighted the importance of signals from the inside of the body (e.g., visceral signals) and proposed that neural representations of internal bodily signals underlie self-consciousness, which to date has been based on philosophical inquiry, clinical case studies, and behavioral studies. Here, we investigated the relationship of bodily self-consciousness with the neural processing of internal bodily signals. By combining electrical neuroimaging, analysis of peripheral physiological signals, and virtual reality technology in humans, we show that transient modulations of neural responses to heartbeats in the posterior cingulate cortex covary with changes in bodily self-consciousness induced by the full-body illusion. Additional analyses excluded that measured basic cardiorespiratory parameters or interoceptive sensitivity traits could account for this finding. These neurophysiological data link experimentally the cortical mapping of the internal body to self-consciousness. SIGNIFICANCE STATEMENT: What are the brain mechanisms of self-consciousness? Prominent views propose that the neural processing associated with signals from the internal organs (such as the heart and the lung) plays a critical role in self-consciousness. Although this hypothesis dates back to influential views in philosophy and psychology (e.g., William James), definitive experimental evidence supporting this idea is lacking despite its recent impact in neuroscience. In the present study, we show that posterior cingulate activities responding to heartbeat signals covary with changes in participants' conscious self-identification with a body that were manipulated experimentally using virtual reality technology. Our finding provides important neural evidence about the long-standing proposal that self-consciousness is linked to the cortical processing of internal bodily signals.

Common and distinct brain regions processing multisensory bodily signals for peripersonal space and body ownership.

We take the feeling that our body belongs to us for granted. However, recent research has shown that it is possible to alter the subjective sensation of body ownership (BO) by manipulating multisensory bodily inputs. Several frontal and parietal regions are known to specifically process multisensory cues presented close to the body, i.e., within the peripersonal space (PPS). It has been proposed that these PPS fronto-parietal regions also underlie BO. However, most previous studies investigated the brain mechanisms of either BO or of PPS processing separately and by using a variety of paradigms. Here, we conducted an extensive meta-analysis of functional neuroimaging studies to investigate PPS and BO processing in humans in order to: a) assess quantitatively where each one of these functions was individually processed in the brain; b) identify whether and where these processes shared common or engaged distinct brain mechanisms; c) characterize these areas in terms of whole-brain co-activation networks and functions, respectively. We identified (i) a bilateral PPS network including superior parietal, temporo-parietal and ventral premotor regions and (ii) a BO network including posterior parietal cortex (right intraparietal sulcus, IPS; and left IPS and superior parietal lobule, SPL), right ventral premotor cortex, and the left anterior insula. Co-activation maps related to both PPS and BO encompassed largely overlapping fronto-parietal networks, but whereas the PPS network was more frequently associated with sensorimotor tasks, the BO network was rather associated with attention and awareness tasks. Finally, the conjunction analysis showed that (iii) PPS and BO tasks anatomically overlapped only in two clusters located in the left parietal cortex (dorsally at the intersection between the SPL, the IPS and area 2 and ventrally between areas 2 and IPS). Distinct activations were located for PPS at the temporo-parietal junction and for BO in the anterior insula. These results in PPS and BO and provide evidence-based insight about the overlap of the two processes in the IPS region and the extensive connectivity between the two associated co-activation networks. They also show significant dissociations, with PPS fronto-parietal areas located more proximal to the central sulcus than BO areas. Such anatomical distinction may also reflect the different functions of the two processes, whereby PPS areas underlie a multisensory-motor interface for body-objects interaction and BO areas being involved in bodily awareness and self-consciousness.

Increased functional connectivity between superior colliculus and brain regions implicated in bodily self-consciousness during the rubber hand illusion.

Bodily self-consciousness refers to bodily processes operating at personal, peripersonal, and extrapersonal spatial dimensions. Although the neural underpinnings of representations of personal and peripersonal space associated with bodily self-consciousness were thoroughly investigated, relatively few is known about the neural underpinnings of representations of extrapersonal space relevant for bodily self-consciousness. In the search to unravel brain structures generating a representation of the extrapersonal space relevant for bodily self-consciousness, we developed a functional magnetic resonance imaging (fMRI) study to investigate the implication of the superior colliculus (SC) in bodily illusions, and more specifically in the rubber hand illusion (RHi), which constitutes an established paradigm to study the neural underpinnings of bodily self-consciousness. We observed activation of the colliculus ipsilateral to the manipulated hand associated with eliciting of RHi. A generalized form of context-dependent psychophysiological interaction analysis unravelled increased illusion-dependent functional connectivity between the SC and some of the main brain areas previously involved in bodily self-consciousness: right temporoparietal junction (rTPJ), bilateral ventral premotor cortex (vPM), and bilateral postcentral gyrus. We hypothesize that the collicular map of the extrapersonal space interacts with maps of the peripersonal and personal space generated at rTPJ, vPM and the postcentral gyrus, producing a unified representation of space that is relevant for bodily self-consciousness. We suggest that processes of multisensory integration of bodily-related sensory inputs located in this unified representation of space constitute one main factor underpinning emergence of bodily self-consciousness.

A quantitative study on peripersonal space in anorexia nervosa and healthy subjects: Role of social variables and association with psychopathology.

Objectives: This study investigates peripersonal space (PPS) modulation in patients with anorexia nervosa (AN) versus healthy controls (HCs) and explores associations between PPS, eating-related, and general psychopathology. Method: Forty-six patients and 42 HCs completed a computer-based task observing videos of an approaching actor (male or female) displaying different facial expressions along with a non-social condition. Then, participants completed self-report questionnaires assessing eating-related and general psychopathology. Results: Mixed-models revealed that both groups adjusted PPS based on task conditions, with a gender effect favoring closer proximity to female actor. HCs reduced PPS amplitude progressively during the task, while patients did not show this effect. In patients, wider PPS correlated with lower self-esteem and facial expression identification accuracy, while in HCs, PPS was associated to body dissatisfaction and anxiety symptoms. Conclusion: These findings enhance understanding of bodily self-consciousness, suggesting PPS consideration in therapeutic interactions with patients with AN and as a potential target in treatments addressing social impairment.

Individual Differences in Bodily Self-Consciousness and Its Neural Basis.

Bodily self-consciousness (BSC), a subject of interdisciplinary interest, refers to the awareness of one's bodily states. Previous studies have noted the existence of individual differences in BSC, while neglecting the underlying factors and neural basis of such individual differences. Considering that BSC relied on integration from both internal and external self-relevant information, we here review previous findings on individual differences in BSC through a three-level-self model, which includes interoceptive, exteroceptive, and mental self-processing. The data show that cross-level factors influenced individual differences in BSC, involving internal bodily signal perceptibility, multisensory processing principles, personal traits shaped by environment, and interaction modes that integrate multiple levels of self-processing. Furthermore, in interoceptive processing, regions like the anterior cingulate cortex and insula show correlations with different perceptions of internal sensations. For exteroception, the parietal lobe integrates sensory inputs, coordinating various BSC responses. Mental self-processing modulates differences in BSC through areas like the medial prefrontal cortex. For interactions between multiple levels of self-processing, regions like the intraparietal sulcus involve individual differences in BSC. We propose that diverse experiences of BSC can be attributed to different levels of self-processing, which moderates one's perception of their body. Overall, considering individual differences in BSC is worth amalgamating diverse methodologies for the diagnosis and treatment of some diseases.

Predicting the bodily self in space and time.

To understand how the human brain distinguishes itself from external stimulation, it was examined if motor predictions enable healthy adult volunteers to infer self-location and to distinguish their body from the environment (and other agents). By uniquely combining a VR-setup with full-body motion capture, a full-body illusion paradigm (FBI) was developed with different levels of motion control: (A) a standard, passive FBI in which they had no motion control; (B) an active FBI in which they made simple, voluntary movements; and (C) an immersive game in which they real-time controlled a human-sized avatar in third person. Systematic comparisons between measures revealed a causal relationship between (i) motion control (prospective agency), (ii) self-other identification, and (iii) the ability to locate oneself. Healthy adults could recognise their movements in a third-person avatar and psychologically align with it (action observation); but did not lose a sense of place (self-location), time (temporal binding), nor who they are (self/other). Instead, motor predictions enabled them to localise their body and to distinguish self from other. In the future, embodied games could target and strengthen the brain's control networks in psychosis and neurodegeneration; real-time motion simulations could help advance neurorehabilitation techniques by fine-tuning and personalising therapeutic settings.

Bodily ownership and self-location: components of bodily self-consciousness.

Recent research on bodily self-consciousness has assumed that it consists of three distinct components: the experience of owning a body (body ownership); the experience of being a body with a given location within the environment (self-location); and the experience of taking a first-person, body-centered, perspective on that environment (perspective). Here we review recent neuroimaging studies suggesting that at least two of these components-body ownership and self-location-are implemented in rather distinct neural substrates, located, respectively, in the premotor cortex and in the temporo-parietal junction. We examine these results and consider them in relation to clinical evidence from patients with altered body perception and work on a variety of multisensory, body-related illusions, such as the rubber hand illusion, the full body illusion, the body swap illusion and the enfacement illusion. We conclude by providing a preliminary synthesis of the data on bodily self-consciousness and its neural correlates.