Balancing the Senses: Electrophysiological Responses Reveal the Interplay between Somatosensory and Visual Processing During Body-Related Multisensory Conflict.

In the study of bodily awareness, the predictive coding theory has revealed that our brain continuously modulates sensory experiences to integrate them into a unitary body representation. Indeed, during multisensory illusions (e.g., the rubber hand illusion, RHI), the synchronous stroking of the participant's concealed hand and a fake visible one creates a visuotactile conflict, generating a prediction error. Within the predictive coding framework, through sensory processing modulation, prediction errors are solved, inducing participants to feel as if touches originated from the fake hand, thus ascribing the fake hand to their own body. Here, we aimed to address sensory processing modulation under multisensory conflict, by disentangling somatosensory and visual stimuli processing that are intrinsically associated during the illusion induction. To this aim, we designed two EEG experiments, in which somatosensory- (SEPs; Experiment 1; N = 18; F = 10) and visual-evoked potentials (VEPs; Experiment 2; N = 18; F = 9) were recorded in human males and females following the RHI. Our results show that, in both experiments, ERP amplitude is significantly modulated in the illusion as compared with both control and baseline conditions, with a modality-dependent diametrical pattern showing decreased SEP amplitude and increased VEP amplitude. Importantly, both somatosensory and visual modulations occur in long-latency time windows previously associated with tactile and visual awareness, thus explaining the illusion of perceiving touch at the sight location. In conclusion, we describe a diametrical modulation of somatosensory and visual processing as the neural mechanism that allows maintaining a stable body representation, by restoring visuotactile congruency under the occurrence of multisensory conflicts.

Neural Substrates of Body Ownership and Agency during Voluntary Movement.

Body ownership and the sense of agency are two central aspects of bodily self-consciousness. While multiple neuroimaging studies have investigated the neural correlates of body ownership and agency separately, few studies have investigated the relationship between these two aspects during voluntary movement when such experiences naturally combine. By eliciting the moving rubber hand illusion with active or passive finger movements during functional magnetic resonance imaging, we isolated activations reflecting the sense of body ownership and agency, respectively, as well as their interaction, and assessed their overlap and anatomic segregation. We found that perceived hand ownership was associated with activity in premotor, posterior parietal, and cerebellar regions, whereas the sense of agency over the movements of the hand was related to activity in the dorsal premotor cortex and superior temporal cortex. Moreover, one section of the dorsal premotor cortex showed overlapping activity for ownership and agency, and somatosensory cortical activity reflected the interaction of ownership and agency with higher activity when both agency and ownership were experienced. We further found that activations previously attributed to agency in the left insular cortex and right temporoparietal junction reflected the synchrony or asynchrony of visuoproprioceptive stimuli rather than agency. Collectively, these results reveal the neural bases of agency and ownership during voluntary movement. Although the neural representations of these two experiences are largely distinct, there are interactions and functional neuroanatomical overlap during their combination, which has bearing on theories on bodily self-consciousness.SIGNIFICANCE STATEMENT How does the brain generate the sense of being in control of bodily movement (agency) and the sense that body parts belong to one's body (body ownership)? Using fMRI and a bodily illusion triggered by movement, we found that agency is associated with activity in premotor cortex and temporal cortex, and body ownership with activity in premotor, posterior parietal, and cerebellar regions. The activations reflecting the two sensations were largely distinct, but there was overlap in premotor cortex and an interaction in somatosensory cortex. These findings advance our understanding of the neural bases of and interplay between agency and body ownership during voluntary movement, which has implications for the development of advanced controllable prosthetic limbs that feel like real limbs.

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.

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.

Bi-directional and multi-modal effects of dexamphetamine on spatial binding windows in healthy individuals.

OBJECTIVES: Stimuli that are separated by a short window of space or time, known as spatial and temporal binding windows (SBW/TBWs), may be perceived as separate. Widened TBWs are evidenced in schizophrenia, although it is unclear if the SBW is similarly affected. The current study aimed to assess if dexamphetamine (DEX) may increase SBWs in a multimodal visuo-tactile illusion, potentially validating usefulness as an experimental model for multimodal visuo-tactile hallucinations in schizophrenia, and to examine a possible association between altered binding windows (BWs) and working memory (WM) suggested by previous research. METHODS: A placebo-controlled, double-blinded, and counter-balanced crossover design was employed. Permuted block randomisation was used for drug order. Healthy participants received DEX (0.45 mg/kg, PO, b.i.d.) or placebo (glucose powder) in capsules. The Rubber Hand Illusion (RHI) and Wechsler Adult Intelligence Scale Spatial Span was employed to determine whether DEX would alter SBWs and WM, respectively. Schizotypy was assessed with a variety of psychological scales. RESULTS: Most participants did not experience the RHI even under normal circumstances. Bi-directional and multimodal effects of DEX on individual SBWs and schizotypy were observed, but not on WM. CONCLUSIONS: Bidirectional multimodal effects of DEX on the RHI and SBWs were observed in individuals, although not associated with alterations in WM.

Causal Inference of Body Ownership in the Posterior Parietal Cortex.

How do we come to sense that a hand in view belongs to our own body or not? Previous studies have suggested that the integration of vision and somatosensation in the frontoparietal areas plays a critical role in the sense of body ownership (i.e., the multisensory perception of limbs and body parts as our own). However, little is known about how these areas implement the multisensory integration process at the computational level and whether activity predicts illusion elicitation in individual participants on a trial-by-trial basis. To address these questions, we used functional magnetic resonance imaging and a rubber hand illusion-detection task and fitted the registered neural responses to a Bayesian causal inference model of body ownership. Thirty healthy human participants (male and female) performed 12 s trials with varying degrees of asynchronously delivered visual and tactile stimuli of a rubber hand (in view) and a (hidden) real hand. After the 12 s period, participants had to judge whether the rubber hand felt like their own. As hypothesized, activity in the premotor and posterior parietal cortices was related to illusion elicitation at the level of individual participants and trials. Importantly, activity in the posterior parietal cortex fit the predicted probability of illusion emergence of the Bayesian causal inference model based on each participant's behavioral response profile. Our findings suggest an important role for the posterior parietal cortex in implementing Bayesian causal inference of body ownership and reveal how trial-by-trial variations in neural signatures of multisensory integration relate to the elicitation of the rubber hand illusion.SIGNIFICANCE STATEMENT How does the brain create a coherent perceptual experience of one's own body based on information from the different senses? We examined how the likelihood of eliciting a classical bodily illusion that depends on vision and touch-the rubber hand illusion-is related to neural activity measured by functional magnetic resonance imaging. We found that trial-by-trial variations in the neural signal in the posterior parietal cortex, a well known center for sensory integration, fitted a statistical function that describes how likely it is that the brain infers that a rubber hand is one's own given the available visual and tactile evidence. Thus, probabilistic analysis of sensory information in the parietal lobe underlies our unitary sense of bodily self.

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.

Sixty Years After Orne's American Psychologist Article: A Conceptual Framework for Subjective Experiences Elicited by Demand Characteristics.

Study participants form beliefs based on cues present in a testing situation (demand characteristics). These beliefs can alter study outcomes (demand effects). Neglecting demand effects can threaten the internal and external validity of studies (including their replication). While demand characteristics garnered much attention following Orne's introduction of this notion, consideration of their effects has become sparse in experimental reports. Moreover, the concept remains confusing. Here, we introduce a conceptual framework for subjective experiences elicited by demand characteristics. The model distinguishes between participants' awareness of the hypothesis, their motivation to comply with it, and the strategy they use to meet situational requirements. We stress that demand characteristics can give rise to genuine experiences. To illustrate, we apply the model to Evaluative Conditioning and the Rubber Hand Illusion. In the General Discussion, we discuss risks and opportunities associated with demand characteristics, and we explain that they remain highly relevant to current research.

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.

The role of visual expectations in acupuncture analgesia: A quantitative electroencephalography study.

Acupuncture is a complex treatment comprising multisensory stimulation, including visual and tactile sensations and experiences of body ownership. The purpose of this study was to investigate the role of these three components of acupuncture stimulation in acupuncture analgesia. 40 healthy volunteers participated in the study and received acupuncture treatment under three different conditions (real-hand, rubber-hand synchronous, and rubber-hand asynchronous). The tolerance for heat pain stimuli was measured before and after treatment. Brain oscillation changes were also measured using electroencephalography (EEG). The pain tolerance was significantly increased after acupuncture treatment under all three conditions. Noticeable deqi (needle) sensations in response to acupuncture stimulation of the rubber hand were found under both rubber-hand synchronous and rubber-hand asynchronous conditions. Deqi sensations were significantly correlated with acupuncture analgesia only under the rubber-hand synchronous condition. Increased delta and decreased theta, alpha, beta, and gamma waves were observed after acupuncture treatment under all three conditions. Our findings clarified the role of cognitive components of acupuncture treatment in acupuncture analgesia through the rubber-hand illusion. This study is a first step toward separating various components of acupuncture analgesia, i.e. visual, tactile, and body ownership, and utilizing those components to maximize analgesic effects.

Effects of body ownership illusion during exposure to disgusting stimuli.

BACKGROUND: The Rubber Hand Illusion (RHI) refers to the feeling of ownership of a rubber hand by synchronously stroking the rubber hand and the own hand of a person. Previous research has shown that RHI can be used to simulate skin contact with a disgust-eliciting stimulus. We used a primary disgust elicitor (a living maggot) to replicate this finding and to gather data on disgust habituation during RHI, and effects on in-vivo exposure. METHODS: A total of 82 healthy participants (25 males, 57 females) were randomly assigned to one of two conditions; synchronous stroking or asynchronous stroking (control condition) of the rubber/own hand. Subsequently, a maggot was placed on the rubber hand for five minutes. Participants rated experienced disgust at the beginning and end of the exposure. They were also asked if the maggot could be placed on their own hand. RESULTS: Synchronous stroking successfully elicited RHI, which was associated with higher disgust ratings for the maggot at the beginning of exposure compared to asynchronous stroking. The two conditions did not differ in disgust habituation and the willingness to expose the own hand to the maggot. CONCLUSION: RHI successfully simulated skin contact with a disgust stimulus. Future studies should apply longer exposure intervals and test individuals with higher disgust propensity to detect possible RHI effects on disgust habituation.

Understanding components of embodiment: Evidence from the mirror box illusion.

Past studies have examined embodiment in the rubber hand illusion, using principal components analysis (PCA) to identify factors from questionnaire responses during synchronous and asynchronous stroking. To better understand the phenomenology of embodiment, we used PCA in the mirror box illusion to examine performance across conditions that varied in movement synchrony to examine multisensory integration and movement type to vary the amount of multisensory congruence. We found three dissociable components in all conditions: embodiment, deafference and attentiveness. We also examined how these embodiment ratings varied across the four conditions. As hypothesized, embodiment ratings were highest for synchronous movement, with feelings of deafference highest for asynchronous movement. Furthermore, there was a movement by timing interaction, such that sliding resulted in greater differences in synchronous versus asynchronous ratings than tapping. These results suggest that embodiment or deafference can be changed as a function of the amount of multisensory congruence.

Tactile temporal order judgment during rubber hand illusion: Distinct modulation of the point of subjective simultaneity and temporal resolution.

During the rubber hand illusion (RHI), individuals feel a fake hand as their own (ownership) and a perceived position of their real hand shifts toward the fake hand (proprioceptive drift; PD), which represents updating of multisensory hand representations. Bimanual tactile temporal order judgment (TOJ) includes processes of localizing tactile stimuli in space, for which multisensory hand representations are essential. According to the common processes, we examined tactile TOJ performance during the RHI and non-RHI. Temporal resolution (TR) as TOJ accuracy worsened during the non-RHI compared to the RHI. Additionally, a significant correlation between TR and PD was observed only in the non-RHI condition. However, the point of subjective simultaneity (PSS), which offers relative weighting of tactile inputs from the right and left hands, was correlated with illusory hand ownership. These results suggest that PSS and TR from tactile TOJ during RHI relate to self-attribution and localization of the hand, respectively.

Exploring the development of high-level contributions to body representation using the rubber hand illusion and the monkey hand illusion.

During development our body undergoes significant changes, yet we are able to maintain a coherent experience of our body and sense of self. Bodily experience is thought to comprise integration of multisensory signals (vision, touch, and proprioception) constrained by top-down knowledge of body appearance. Evidence from developmental studies suggests that low-level multisensory integration develops throughout childhood, reaching adult levels by 10 years of age. However, how high-level cognitive knowledge changes during childhood to constrain our multisensory body experience is unknown. This study describes four experiments examining high-level contributions to the bodily experience in children compared with adults using the rubber hand illusion and a monkey hand illusion. We found that children (5-17 years of age) exhibited more flexible body representations, showing stronger illusions for small and fantastical (monkey) fake hands compared with adults. Conversely, using a task indirectly capturing changes in hand size, we found that children and adults demonstrated statistically equivalent increases and decreases in hand size following illusions over large and small hands, respectively. Interestingly, at baseline children showed a bias in reporting larger hand size judgments that decreased with age. Finally, we did not find a relationship between individual differences in fantasy proneness and illusion strength for a fantastical (monkey) hand for children or adults, suggesting that developmental changes of top-down constraints are not purely driven by more diffuse boundaries between imagination and reality. These data suggest that high-level constraints acting on our multisensory body experience change during development, allowing children a more flexible bodily experience compared with adults.

Multisensory and Sensorimotor Integration in the Embodied Self: Relationship between Self-Body Recognition and the Mirror Neuron System.

The embodied self is rooted in the self-body in the "here and now". The senses of self-ownership and self-agency have been proposed as the basis of the sense of embodied self, and many experimental studies have been conducted on this subject. This review summarizes the experimental research on the embodied self that has been conducted over the past 20 years, mainly from the perspective of multisensory integration and sensorimotor integration regarding the self-body. Furthermore, the phenomenon of back projection, in which changes in an external object (e.g., a rubber hand) with which one has a sense of ownership have an inverse influence on the sensation and movement of one's own body, is discussed. This postulates that the self-body illusion is not merely an illusion caused by multisensory and/or sensorimotor integration, but is the incorporation of an external object into the self-body representation in the brain. As an extension of this fact, we will also review research on the mirror neuron system, which is considered to be the neural basis of recognition of others, and discuss how the neural basis of self-body recognition and the mirror neuron system can be regarded as essentially the same.

Sustained rubber hand illusion after the end of visuotactile stimulation with a similar time course for the reduction of subjective ownership and proprioceptive drift.

The rubber hand illusion is a perceptual illusion in which participants experience an inanimate rubber hand as their own when they observe this model hand being stroked in synchrony with strokes applied to the person's real hand, which is hidden. Earlier studies have focused on the factors that determine the elicitation of this illusion, the relative contribution of vision, touch and other sensory modalities involved and the best ways to quantify this perceptual phenomenon. Questionnaires serve to assess the subjective feeling of ownership, whereas proprioceptive drift is a measure of the recalibration of hand position sense towards the rubber hand when the illusion is induced. Proprioceptive drift has been widely used and thought of as an objective measure of the illusion, although the relationship between this measure and the subjective illusion is not fully understood. Here, we examined how long the illusion is maintained after the synchronous visuotactile stimulation stops with the specific aim of clarifying the temporal relationship in the reduction of both subjective ownership and proprioceptive drift. Our results show that both the feeling of ownership and proprioceptive drift are sustained for tens of seconds after visuotactile stroking has ceased. Furthermore, our results indicate that the reduction of proprioceptive drift and the feeling of ownership follow similar time courses in their reduction, suggesting that the two phenomena are temporally correlated. Collectively, these findings help us better understand the relationships of multisensory stimulation, subjective ownership, and proprioceptive drift in the rubber hand illusion.

Self-consciousness impairments in schizophrenia with and without first rank symptoms using the moving rubber hand illusion.

INTRODUCTION: The moving rubber hand illusion allows the evaluation both the sense of body ownership and agency using visuo-motor stimulations. METHODS: We used the moving rubber hand illusion in anatomic congruence with explicit measures to compare active asynchronous and passive synchronous movements in patients diagnosed with schizophrenia with first rank symptoms (FRS) (n = 31) versus without FRS (n = 25). RESULTS: Patients with FRS are characterized by a lack of agency in active asynchronous condition. The two groups had no sense of ownership in synchronous passive condition. Using a multivariate regression model, we found an association between agency and body ownership measures in the active asynchronous condition in two groups (OR: 1.825, p < 0.001). In the passive condition, this association was only present in the group with first rank symptoms (OR: 2.04, p < 0.001). CONCLUSION: Temporal proximity and sensorimotor information are essential in the understanding of self-consciousness disorders in schizophrenia.

The Impact of Experimentally Induced Limb Ischemia on the Rubber Hand Illusion.

The rubber hand illusion (RHI) is a perceptual illusion in which one experiences an object as part of their body when synchronous visuo-tactile stimulation is applied to one's hand and the object. There are a variety of factors that can impact the occurrence of the RHI. In the present study, we demonstrate that experimentally induced peripheral ischemia can impact the RHI, namely it can result in larger alterations to the perceived position of one's hand. This study suggests that alterations to the cardiovascular system may be a source of individual differences in the RHI literature. Future studies with samples of individuals with cardiovascular pathology are recommended.

Awe liberates the feeling that "my body is mine".

Awe is an emotional response to perceptually vast stimuli that transcend one's current frames of reference. Previous research indicated that awe promotes a smaller self, which led to the creation of a small-self hypothesis. Thus, we shed new light on this hypothesis in terms of sense of body ownership using a rubber hand illusion experiment; through it, we showed that awe evokes an increased sense of body ownership over the rubber hand and this effect was prominent among participants who experienced small self. Our findings suggest that awe might provoke a "liberation of the self" in terms of a sense of body ownership as awe has been thought to liberate existing schemas, hence informing the demonstrable implications of the psychological mechanisms of awe.

The rubber hand illusion in children: What are we measuring?

The rubber hand illusion (RHI) is a much-studied bodily illusion that has been used in a wide number of populations to investigate the plasticity of the mental body representation. In adult participants, the wide adoption of the illusion has led to a proliferation of experimental variations of the illusion, and with that, considerable apparent inconsistencies in both empirical results and conceptual interpretations. In turn, this makes it challenging to integrate empirical findings and to identify what those findings together can tell us about the representation of the body in the brain. More recently, scientists have started applying the illusion to populations of children, in order to better understand how body representations develop in both typically developing children and in clinical populations. With this field now starting to expand, we believe it is both urgent and important to prevent unintended methodological variability from hindering the consistency of the paediatric literature as it has the adult literature. With this aim in mind, we review the 12 currently available paediatric RHI studies, and summarise their key methodological choices and conceptual definitions. We highlight a number of important discrepancies, particularly where seemingly equivalent analysis choices might significantly affect the interpretation of results, and make recommendations for future studies. We hope this will allow this important and emerging field to benefit from the synergy that results from multiple studies using convergent and consistent empirical methods.