Multisensory conflict affects body schema and reaching space.

Multisensory integration plays a crucial role in building the sense of body ownership, i.e., the perceptual status of one's body for which the body is perceived as belonging to oneself. Temporal and spatial mismatching of visual and tactile signals coming from one's body can reduce ownership feelings towards the body and its parts, i.e., produce disownership feelings. Here, we investigated whether visuo-tactile conflict also affects the sensorimotor representation of the body in space (i.e., body schema) and the perception of the space around the body in terms of action potentiality (i.e., reaching space). In two experiments, body schema (Experiment 1) and reaching space (Experiment 2) were assessed before and after either synchronous or asynchronous visuo-tactile stimulation. Results showed that the asynchronous condition, provoking multisensory conflict, caused disownership over one's hand and concurrently affected the body schema and the reaching space. These findings indicate that body schema and reaching space could be dynamically shaped by the multisensory regularities that build up the sense of body ownership.

Malleability of the sense of bodily self in early childhood: 5- and 6-year-old children show the enfacement illusion.

The mechanisms underlying the developing sense of bodily self are debated. Whereas some scholars stress the role of sensory factors, others propose the importance of contextual factors. By manipulating multisensory stimulation and social familiarity with the other person, we explored two factors that are proposed to relate to young children's developing sense of bodily self. Including an adult sample allowed us to investigate age-related differences of the malleability of the bodily self. To this end, the study implemented an enfacement illusion with children (N = 64) and adults (N = 33). Participants were exposed to one trial with synchronous interpersonal multisensory stimulation and one trial with asynchronous interpersonal multisensory stimulation-either with a stranger or with the mother as the other person. A self-recognition task using morph videos of self and other and an enfacement questionnaire were implemented as dependent measures. Results revealed evidence for the presence of the enfacement effect in children in both measures. The identity of the other person had a significant effect on the self-recognition task. Contrary to our hypothesis, the effect was significantly smaller in the caregiver condition. No significant differences between children and adults emerged. Our results demonstrate the role of both multisensory stimulation and contextual-here social familiarity-factors for the construction and development of a bodily self. The study provides developmental science with a novel approach to the bodily self by showing the validity of the self-recognition task in a child sample. Overall, the study supports proposals that the sense of bodily self is malleable early in development.

Does Embodying a Divine Avatar Influence Moral Decisions? An Immersive Virtual Reality Study.

The term Proteus effect refers to the changes in attitudes and behavior induced by the characteristics of an embodied virtual agent. Whether the effect can extend to the moral sphere is currently unknown. To deal with this issue, we investigated if embodying virtual agents (i.e., avatars) with different characteristics modulate people's moral standards differentially. Participants were requested to embody an avatar resembling the Christian God in His anthropomorphic appearance or a control human avatar and to perform a text-based version of incidental and instrumental dilemmas in a virtual environment. For each participant, we recorded (1) chosen options (deontological vs. utilitarian), (2) decision times, (3) postdecision feelings, and (4) physiological reactions (skin conductance response and heart rate). We found that embodying God vs. a control avatar did not change the performance in the moral dilemma task, indicating that no strong Proteus effect was at play in our experimental conditions. We interpreted this result by examining the constraints and limitations of our task, reasoning about the necessary conditions for eliciting the Proteus effect, and discussing future developments and advances in the field. Moreover, we presented compelling effects concerning dilemma type, chosen option, personality traits, and religion affiliation, thus supporting and extending literature on decision making in moral dilemmas.

Dissociative Symptoms and Interoceptive Integration.

Dissociative symptoms and disorders of dissociation are characterised by disturbances in the experience of the self and the surrounding world, manifesting as a breakdown in the normal integration of consciousness, memory, identity, emotion, and perception. This paper aims to provide insights into dissociative symptoms from the perspective of interoception, the sense of the body's internal physiological state, adopting a transdiagnostic framework.Dissociative symptoms are associated with a blunting of autonomic reactivity and a reduction in interoceptive precision. In addition to the central function of interoception in homeostasis, afferent visceral signals and their neural and mental representation have been shown to shape emotional feeling states, support memory encoding, and contribute to self-representation. Changes in interoceptive processing and disrupted integration of interoceptive signals into wider cognition may contribute to detachment from the body and the world, blunted emotional experience, and altered subjective recall, as experienced by individuals who suffer from dissociation.A better understanding of the role of altered interoceptive integration across the symptom areas of dissociation could thus provide insights into the neurophysiological mechanisms underlying dissociative disorders. As new therapeutic approaches targeting interoceptive processing emerge, recognising the significance of interoceptive mechanisms in dissociation holds potential implications for future treatment targets.

Development and validation of a pictographic assessment embodiment scale.

Embodiment is a complex concept related to the subjective perception of an object as it belongs to its own body. In general, this construct has been evaluated by means of questionnaires, but validation studies in other cultures and limitations related with barriers of language received little attention. The purpose of the present investigation was twofold: to validate the factorial structure of embodiment questionnaire (EQ) and to construct a pictographic scale (PAE) to measure embodiment without relapse verbal representations. In the first experiment, 136 participants underwent a Rubber Hand Illusion (RHI) procedure following both congruent and incongruent (control) visuo-tactile stimulations. Then, they evaluated embodiment illusion in EQ using a Likert-type scale to rate their agreement or disagreement with 27 statements and with a pictographic scale designed to assess their subjective experience of the illusion. Principal components analysis in EQ scores identified four components that emerged in both conditions: Embodiment, Disembodiment, Affect and Deafference. PAE scale was highly correlated with embodiment factor and can differentiate between conditions. In a second experiment, 30 participants underwent the RHI procedure, and they were assessed using PAE and proprioceptive drift. Results indicate a high positive correlation between PAE and post-illusion drift score. These results provide evidence about the consistency of the factorial structure of EQ across cultures, and we also provide a new pictographic tool that allows quick measurement of embodiment overcoming language barriers.

Is self-awareness necessary to have a theory of mind?

Forty years ago, Gallup proposed that theory of mind presupposes self-awareness. Following Humphrey, his hypothesis was that individuals can infer the mental states of others thanks to the ability to monitor their own mental states in similar circumstances. Since then, advances in several disciplines, such as comparative and developmental psychology, have provided empirical evidence to test Gallup's hypothesis. Herein, we review and discuss this evidence.

Visual perspective and body ownership modulate vicarious pain and touch: A systematic review.

We conducted a systematic review investigating the influence of visual perspective and body ownership (BO) on vicarious brain resonance and vicarious sensations during the observation of pain and touch. Indeed, the way in which brain reactivity and the phenomenological experience can be modulated by blurring the bodily boundaries of self-other distinction is still unclear. We screened Scopus and WebOfScience, and identified 31 articles, published from 2000 to 2022. Results show that assuming an egocentric perspective enhances vicarious resonance and vicarious sensations. Studies on synaesthetes suggest that vicarious conscious experiences are associated with an increased tendency to embody fake body parts, even in the absence of congruent multisensory stimulation. Moreover, immersive virtual reality studies show that the type of embodied virtual body can affect high-order sensations such as appropriateness, unpleasantness, and erogeneity, associated with the touched body part and the toucher's social identity. We conclude that perspective plays a key role in the resonance with others' pain and touch, and full-BO over virtual avatars allows investigation of complex aspects of pain and touch perception which would not be possible in reality.

Hierarchical and dynamic relationships between body part ownership and full-body ownership.

What is the relationship between experiencing individual body parts and the whole body as one's own? We theorised that body part ownership is driven primarily by the perceptual binding of visual and somatosensory signals from specific body parts, whereas full-body ownership depends on a more global binding process based on multisensory information from several body segments. To examine this hypothesis, we used a bodily illusion and asked participants to rate illusory changes in ownership over five different parts of a mannequin's body and the mannequin as a whole, while we manipulated the synchrony or asynchrony of visual and tactile stimuli delivered to three different body parts. We found that body part ownership was driven primarily by local visuotactile synchrony and could be experienced relatively independently of full-body ownership. Full-body ownership depended on the number of synchronously stimulated parts in a nonlinear manner, with the strongest full-body ownership illusion occurring when all parts received synchronous stimulation. Additionally, full-body ownership influenced body part ownership for nonstimulated body parts, and skin conductance responses provided physiological evidence supporting an interaction between body part and full-body ownership. We conclude that body part and full-body ownership correspond to different processes and propose a hierarchical probabilistic model to explain the relationship between part and whole in the context of multisensory awareness of one's own body.

Body ownership and kinaesthetic illusions: Dissociated bodily experiences for distinct levels of body consciousness?

Seeing an embodied humanoid avatar move its arms can induce in the observer the illusion that its own (static) arms are moving accordingly, the kinematic signals emanating from this avatar thus being considered like those from the biological body. Here, we investigated the causal relationship between these kinaesthetic illusions and the illusion of body ownership, manipulated through visuomotor synchronisation. The results of two experiments revealed that the sense of body ownership over an avatar seen from a first-person perspective was intimately linked to visuomotor synchrony. This was not the case for kinaesthetic illusions indicating that when superimposed on the biological body, the avatar is inevitably treated at the sensorimotor level as one's own body, whether consciously considered as such or not. The question of whether these two bodily experiences (body ownership and kinaesthetic illusion) are underpinned by distinct representations, the body image, and the body schema, is discussed.

Texture congruence modulates perceptual bias but not sensitivity to visuotactile stimulation during the rubber hand illusion.

The sense of body ownership is the feeling that one's body belongs to oneself. To study body ownership, researchers use bodily illusions, such as the rubber hand illusion (RHI), which involves experiencing a visible rubber hand as part of one's body when the rubber hand is stroked simultaneously with the hidden real hand. The RHI is based on a combination of vision, touch, and proprioceptive information following the principles of multisensory integration. It has been posited that texture incongruence between rubber hand and real hand weakens the RHI, but the underlying mechanisms remain poorly understood. To investigate this, we recently developed a novel psychophysical RHI paradigm. Based on fitting psychometric functions, we discovered the RHI resulted in shifts in the point of subjective equality when the rubber hand and the real hand were stroked with matching materials. We analysed these datasets further by using signal detection theory analysis, which distinguishes between the participants' sensitivity to visuotactile stimulation and the associated perceptual bias. We found that texture incongruence influences the RHI's perceptual bias but not its sensitivity to visuotactile stimulation. We observed that the texture congruence bias effect was the strongest in shorter visuotactile asynchronies (50-100 ms) and weaker in longer asynchronies (200 ms). These results suggest texture-related perceptual bias is most prominent when the illusion's sensitivity is at its lowest. Our findings shed light on the intricate interactions between top-down and bottom-up processes in body ownership, the links between body ownership and multisensory integration, and the impact of texture congruence on the RHI.

With hand on heart: A cardiac Rubber Hand Illusion.

Body illusions such as the Rubber Hand Illusion (RHI) have highlighted how multisensory integration underpins the sense of one's own body. Much of this research has focused on senses arising from outside the body (e.g. vision and touch), but sensations from within the body may also play a role. In a pre-registered study, participants completed a cardiac variation of the RHI, where taps to the finger occurred in or out of time with the heartbeat. We replicated the RHI effect, showing that synchronous but not asynchronous taps to the real and rubber hand increased sensations of embodiment over the rubber hand and caused a shift in the perceived hand location. However, there were no significant influences of cardiac timing on embodiment, nor did it interact with visuo-tactile synchrony. An exploratory analysis found a three-way interaction between synchrony, cardiac timing and interoceptive accuracy as measured by a heartbeat counting task, such that greater interoceptive accuracy was associated with lower embodiment ratings in the systole condition compared to diastole, but only during synchronous stimulation. Although our novel methodology successfully replicated the RHI, our findings suggest that the cooccurence of vision and touch with cardiac signals may make little contribution to the sense of one's body.

Active self-touch restores bodily proprioceptive spatial awareness following disruption by 'rubber hand illusion'.

Bodily self-awareness relies on a constant integration of visual, tactile, proprioceptive, and motor signals. In the 'rubber hand illusion' (RHI), conflicting visuo-tactile stimuli lead to changes in self-awareness. It remains unclear whether other, somatic signals could compensate for the alterations in self-awareness caused by visual information about the body. Here, we used the RHI in combination with robot-mediated self-touch to systematically investigate the role of tactile, proprioceptive and motor signals in maintaining and restoring bodily self-awareness. Participants moved the handle of a leader robot with their right hand and simultaneously received corresponding tactile feedback on their left hand from a follower robot. This self-touch stimulation was performed either before or after the induction of a classical RHI. Across three experiments, active self-touch delivered after-but not before-the RHI, significantly reduced the proprioceptive drift caused by RHI, supporting a restorative role of active self-touch on bodily self-awareness. The effect was not present during involuntary self-touch. Unimodal control conditions confirmed that both tactile and motor components of self-touch were necessary to restore bodily self-awareness. We hypothesize that active self-touch transiently boosts the precision of proprioceptive representation of the touched body part, thus counteracting the visual capture effects that underlie the RHI.

Subcortical contributions to the sense of body ownership.

The sense of body ownership (i.e. the feeling that our body or its parts belong to us) plays a key role in bodily self-consciousness and is believed to stem from multisensory integration. Experimental paradigms such as the rubber hand illusion have been developed to allow the controlled manipulation of body ownership in laboratory settings, providing effective tools for investigating malleability in the sense of body ownership and the boundaries that distinguish self from other. Neuroimaging studies of body ownership converge on the involvement of several cortical regions, including the premotor cortex and posterior parietal cortex. However, relatively less attention has been paid to subcortical structures that may also contribute to body ownership perception, such as the cerebellum and putamen. Here, on the basis of neuroimaging and neuropsychological observations, we provide an overview of relevant subcortical regions and consider their potential role in generating and maintaining a sense of ownership over the body. We also suggest novel avenues for future research targeting the role of subcortical regions in making sense of the body as our own.

Sense of own body shapes neural processes of memory encoding and reinstatement.

How is the fundamental sense of one's body, a basic aspect of selfhood, incorporated into memories for events? Disrupting bodily self-awareness during encoding impairs functioning of the left posterior hippocampus during retrieval, which implies weakened encoding. However, how changes in bodily self-awareness influence neural encoding is unknown. We investigated how the sense of body ownership, a core aspect of the bodily self, impacts encoding in the left posterior hippocampus and additional core memory regions including the angular gyrus. Furthermore, we assessed the degree to which memories are reinstated according to body ownership during encoding and vividness during retrieval as a measure of memory strength. We immersed participants in naturalistic scenes where events unfolded while we manipulated feelings of body ownership with a full-body-illusion during functional magnetic resonance imaging scanning. One week later, participants retrieved memories for the videos during functional magnetic resonance imaging scanning. A whole brain analysis revealed that patterns of activity in regions including the right hippocampus and angular gyrus distinguished between events encoded with strong versus weak body ownership. A planned region-of-interest analysis showed that patterns of activity in the left posterior hippocampus specifically could predict body ownership during memory encoding. Using the wider network of regions sensitive to body ownership during encoding and the left posterior hippocampus as separate regions-of-interest, we observed that patterns of activity present at encoding were reinstated more during the retrieval of events encoded with strong body ownership and high memory vividness. Our results demonstrate how the sense of physical self is bound within an event during encoding, which facilitates reactivation of a memory trace during retrieval.

The self and the Bayesian brain: Testing probabilistic models of body ownership through a self-localization task.

Simple multisensory manipulations can induce the illusory misattribution of external objects to one's own body, allowing to experimentally investigate body ownership. In this context, body ownership has been conceptualized as the result of the online Bayesian optimal estimation of the probability that one object belongs to the body from the congruence of multisensory inputs. This idea has been highly influential, as it provided a quantitative basis to bottom-up accounts of self-consciousness. However, empirical evidence fully supporting this view is scarce, as the optimality of the putative inference process has not been assessed rigorously. This pre-registered study aimed at filling this gap by testing a Bayesian model of hand ownership based on spatial and temporal visuo-proprioceptive congruences. Model predictions were compared to data from a virtual-reality reaching task, whereby reaching errors induced by a spatio-temporally mismatching virtual hand have been used as an implicit proxy of hand ownership. To rigorously test optimality, we compared the Bayesian model versus alternative non-Bayesian models of multisensory integration, and independently assess unisensory components and compare them to model estimates. We found that individually measured values of proprioceptive precision correlated with those fitted from our reaching task, providing compelling evidence that the underlying visuo-proprioceptive integration process approximates Bayesian optimality. Furthermore, reaching errors correlated with explicit ownership ratings at the single individual and trial level. Taken together, these results provide novel evidence that body ownership, a key component of self-consciousness, can be truly described as the bottom-up, behaviourally optimal processing of multisensory inputs.

No reduction in motor-evoked potential amplitude during the rubber hand illusion.

INTRODUCTION: In the rubber hand illusion (RHI), touches are applied to a fake hand at the same time as touches are applied to a participant's real hand that is hidden in a congruent position. Synchronous (but not asynchronous) tactile stimulation of the two hands may induce the sensation that the fake hand is the participant's own. As such, the illusion is commonly used to examine the sense of body ownership. Some studies indicate that in addition to the subjective experience of limb ownership reported by participants, the RHI can also reduce corticospinal excitability (e.g., as reflected in motor-evoked potential [MEP] amplitude) and alter parietal-motor cortical connectivity in passive participants. These findings have been taken to support a link between motor cortical processing and the subjective experience of body ownership. METHODS: In this study, we tried to replicate the reduction in MEP amplitude associated with the RHI and uncover the components of the illusion that might explain these changes. As such, we used single-pulse transcranial magnetic stimulation to probe the excitability of the corticospinal motor system as participants experienced the RHI. RESULTS: Despite participants reporting the presence of the illusion and showing shifts in perceived real hand position towards the fake limb supporting its elicitation, we did not observe any associated reduction in MEP amplitude. CONCLUSION: We conclude that a reduction in MEP amplitude is not a reliable outcome of the RHI and argue that if such effects do occur, they are unlikely to be large or functionally relevant.

Disentangling the neural correlates of agency, ownership and multisensory processing.

The experience of the self as an embodied agent in the world is an essential aspect of human consciousness. This experience arises from the feeling of control over one's bodily actions, termed the Sense of Agency, and the feeling that the body belongs to the self, Body Ownership. Despite longstanding philosophical and scientific interest in the relationship between the body and brain, the neural systems involved in Body Ownership and Sense of Agency, and especially their interactions, are not yet understood. In this preregistered study using the Moving Rubber Hand Illusion inside an MR-scanner, we aimed to uncover the relationship between Body Ownership and Sense of Agency in the human brain. Importantly, by using both visuomotor and visuotactile stimulations and measuring online trial-by-trial fluctuations in the illusion magnitude, we were able to disentangle brain systems related to objective sensory stimulation and subjective judgments of the bodily-self. Our results indicate that at both the behavioral and neural levels, Body Ownership and Sense of Agency are strongly interrelated. Multisensory regions in the occipital and fronto-parietal regions encoded convergence of sensory stimulation conditions. The subjective judgments of the bodily-self were related to BOLD fluctuations in the Somatosensory cortex and in regions not activated by the sensory conditions, such as the insular cortex and precuneus. Our results highlight the convergence of multisensory processing in specific neural systems for both Body Ownership and Sense of Agency with partially dissociable regions for subjective judgments in regions of the Default Mode Network.

Mu Rhythm Desynchronization while Observing Rubber Hand Movement in the Mirror: The Interaction of Body Representation with Visuo-Tactile Stimulation.

During rubber hand illusion (RHI), participants feel that a rubber (fake) hand is their own (i.e., embodiment of the rubber hand) if the unseen real hand and seen rubber hand are stroked synchronously (i.e., visuo-tactile stimuli). The RHI is also evoked if the real and rubber hands are placed in the same position (i.e., visual-proprioceptive congruency), which can be performed using a mirror setting. Using electroencephalography (EEG) and mirror settings, we compared µ rhythm (8-13 Hz) event-related desynchronization (ERD; an index of sensorimotor activation) while watching the movements of embodied or non-embodied rubber hands, which was preceded by an observation of the rubber hand with or without synchronous visuo-tactile stimuli. The illusory ownership of the fake hand was manipulated using visual continuity with (RHI) and without (non-RHI) a fake forearm. Resultantly, an ownership-dependent µ rhythm ERD was found when delivering visuo-tactile stimuli; a greater and more persistent µ rhythm ERD during the rubber hand movement was identified in the RHI in comparison to the non-RHI condition. However, no difference was observed between the two when observing the fake hand alone. These findings suggest the possibility that a self-related multisensory interaction between body representation (top-down processing) and visuo-tactile inputs (bottom-up processing) before a fake hand movement produces ownership-dependent sensorimotor activations during subsequent movement observations.

Quantifying body ownership information processing and perceptual bias in the rubber hand illusion.

Bodily illusions have fascinated humankind for centuries, and researchers have studied them to learn about the perceptual and neural processes that underpin multisensory channels of bodily awareness. The influential rubber hand illusion (RHI) has been used to study changes in the sense of body ownership - that is, how a limb is perceived to belong to one's body, which is a fundamental building block in many theories of bodily awareness, self-consciousness, embodiment, and self-representation. However, the methods used to quantify perceptual changes in bodily illusions, including the RHI, have mainly relied on subjective questionnaires and rating scales, and the degree to which such illusory sensations depend on sensory information processing has been difficult to test directly. Here, we introduce a signal detection theory (SDT) framework to study the sense of body ownership in the RHI. We provide evidence that the illusion is associated with changes in body ownership sensitivity that depend on the information carried in the degree of asynchrony of correlated visual and tactile signals, as well as with perceptual bias and sensitivity that reflect the distance between the rubber hand and the participant's body. We found that the illusion's sensitivity to asynchrony is remarkably precise; even a 50 ms visuotactile delay significantly affected body ownership information processing. Our findings conclusively link changes in a complex bodily experience such as body ownership to basic sensory information processing and provide a proof of concept that SDT can be used to study bodily illusions.