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.

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.

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.

Thermosensation and emotion: Thermosensory accuracy in a dynamic thermal matching task is linked to depression and anxiety symptomatology.

Interoception is related to the generation of bodily feelings and the awareness of ourselves as 'sentient beings', informing the organism about its bodily needs to guarantee survival. Previous studies have reported links among interoception, emotion processing, and mental health. For example, the alignment of interoceptive dimensions (i.e., accuracy, sensibility, awareness) can predict emotional symptoms, such as anxiety. Here, we aimed to investigate the relationship between the perception of a certain type of skin-mediated interoceptive signal, i.e., thermosensation, and self-reported depression, anxiety, and stress. One hundred seventy participants completed the Depression Anxiety Stress Scale (DASS-21) and a dynamic thermal matching task, a static temperature detection task, and a heartbeat counting task. Our results revealed that self-reported anxiety and depression were related to the perception of temperature on hairy and non-hairy skin, respectively: higher anxiety was related to better performance on the thermal matching task on the forearm, while higher depression was related to poorer performance on dynamic and static temperature tasks on the palm. Discrepancies between thermosensory accuracy and sensibility measures ('trait prediction error') were related to heightened anxiety, in line with previous studies. No significant correlations were found between DASS-21 scores and heartbeat counting accuracy. In conclusion, this study suggests that individual differences in thermosensory perception in different areas of the body are associated with self-reported anxiety and depression.

Are blind individuals immune to bodily illusions? Somatic rubber hand illusion in the blind revisited.

Multisensory awareness of one's own body relies on the integration of signals from various sensory modalities such as vision, touch, and proprioception. But how do blind individuals perceive their bodies without visual cues, and does the brain of a blind person integrate bodily senses differently from a sighted person? To address this question, we aimed to replicate the only two previous studies on this topic, which claimed that blind individuals do not experience the somatic rubber hand illusion, a bodily illusion triggered by the integration of correlated tactile and proprioceptive signals from the two hands. We used a larger sample size than the previous studies and added Bayesian analyses to examine statistical evidence in favor of the lack of an illusion effect. Moreover, we employed tests to investigate whether enhanced tactile acuity and cardiac interoceptive accuracy in blind individuals could also explain the weaker illusion. We tested 36 blind individuals and 36 age- and sex-matched sighted volunteers. The results show that blind individuals do not experience the somatic rubber hand illusion based on questionnaire ratings and behavioral measures that assessed changes in hand position sense toward the location of the rubber hand. This conclusion is supported by Bayesian evidence in favor of the null hypothesis. The findings confirm that blind individuals do not experience the somatic rubber hand illusion, indicating that lack of visual experience leads to permanent changes in multisensory bodily perception. In summary, our study suggests that changes in multisensory integration of tactile and proprioceptive signals may explain why blind individuals are "immune" to the nonvisual version of the rubber hand illusion.

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.

Proprioceptive uncertainty promotes the rubber hand illusion.

Body ownership is the multisensory perception of a body as one's own. Recently, the emergence of body ownership illusions like the visuotactile rubber hand illusion has been described by Bayesian causal inference models in which the observer computes the probability that visual and tactile signals come from a common source. Given the importance of proprioception for the perception of one's body, proprioceptive information and its relative reliability should impact this inferential process. We used a detection task based on the rubber hand illusion where participants had to report whether the rubber hand felt like their own or not. We manipulated the degree of asynchrony of visual and tactile stimuli delivered to the rubber hand and the real hand under two levels of proprioceptive noise using tendon vibration applied to the lower arm's antagonist extensor and flexor muscles. As hypothesized, the probability of the emergence of the rubber hand illusion increased with proprioceptive noise. Moreover, this result, well fitted by a Bayesian causal inference model, was best described by a change in the a priori probability of a common cause for vision and touch. These results offer new insights into how proprioceptive uncertainty shapes the multisensory perception of one's own body.

Brief Temporal Perturbations in Somatosensory Reafference Disrupt Perceptual and Neural Attenuation and Increase Supplementary Motor Area-Cerebellar Connectivity.

Intrinsic delays in sensory feedback can be detrimental for motor control. As a compensation strategy, the brain predicts the sensory consequences of movement via a forward model on the basis of a copy of the motor command. Using these predictions, the brain attenuates somatosensory reafference to facilitate the processing of exafferent information. Theoretically, this predictive attenuation is disrupted by (even minimal) temporal errors between the predicted and actual reafference; however, direct evidence of such disruption is lacking as previous neuroimaging studies contrasted nondelayed reafferent input with exafferent input. Here, we combined psychophysics with functional magnetic resonance imaging to test whether subtle perturbations in the timing of somatosensory reafference disrupt its predictive processing. Twenty-eight participants (14 women) generated touches on their left index finger by tapping a sensor with their right index finger. The touches on the left index finger were delivered close to the time of contact of the two fingers or with a temporal perturbation (i.e., 153 ms delay). We found that such a brief temporal perturbation disrupted the attenuation of the somatosensory reafference at both the perceptual and neural levels, leading to greater somatosensory and cerebellar responses and weaker somatosensory connectivity with the cerebellum, proportional to the perceptual changes. We interpret these effects as the failure of the forward model to predictively attenuate the perturbed somatosensory reafference. Moreover, we observed increased connectivity of the supplementary motor area with the cerebellum during the perturbations, which could indicate the communication of the temporal prediction error back to the motor centers.SIGNIFICANCE STATEMENT Our brain receives somatosensory feedback from our movements with a delay. To counteract these delays, motor control theories postulate that the brain predicts the timing of somatosensory consequences of our movements and attenuates sensations received at that time. Thus, a self-generated touch feels weaker than an identical external touch. However, how subtle temporal errors between the predicted and actual somatosensory feedback perturb this predictive attenuation remains unknown. We show that such errors make the otherwise attenuated touch feel stronger, elicit stronger somatosensory responses, weaken cerebellar connectivity with somatosensory areas, and increase this connectivity with motor areas. These findings show that motor and cerebellar areas are fundamental in forming temporal predictions about the sensory consequences of our movements.

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.

Heartbeat counting accuracy is enhanced in blind individuals.

Blind individuals have superior abilities to perform perceptual tasks that rely on exteroceptive information, since visual deprivation is associated with heightened cross-modal plasticity. However, it is unknown whether neuroplasticity after visual loss also affects interoception, that is, the sensations arising from one's inner organs that convey information about the physiological state of the body. Herein, we examine the influence of blindness on cardiac interoception, which is an interoceptive submodality that has important links to emotional processing and bodily self-awareness. We tested 36 blind and 36 age- and sex-matched sighted volunteers and examined their cardiac interoceptive ability using the heartbeat counting task. The results showed that blind individuals had significantly higher accuracy in perceiving their heartbeat than did individuals in a matched sighted control group. In contrast, there were no significant differences between the groups in the metacognitive dimensions of cardiac interoception or the purely physiological measurement of heart rate, thereby underscoring that the improved accuracy likely reflects a superior perceptual sensitivity to cardiac interoceptive signals in blind individuals. We conclude that visual deprivation leads to an enhanced ability to count one's own heartbeats, which has important implications for the study of the extent of cross-modal plasticity after visual loss, understanding emotional processing in blind individuals, and learning how bodily self-awareness can develop and be sustained in the absence of visual experience. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Synchronous motor imagery and visual feedback of finger movement elicit the moving rubber hand illusion, at least in illusion-susceptible individuals.

Recent evidence suggests that imagined auditory and visual sensory stimuli can be integrated with real sensory information from a different sensory modality to change the perception of external events via cross-modal multisensory integration mechanisms. Here, we explored whether imagined voluntary movements can integrate visual and proprioceptive cues to change how we perceive our own limbs in space. Participants viewed a robotic hand wearing a glove repetitively moving its right index finger up and down at a frequency of 1 Hz, while they imagined executing the corresponding movements synchronously or asynchronously (kinesthetic-motor imagery); electromyography (EMG) from the participants' right index flexor muscle confirmed that the participants kept their hand relaxed while imagining the movements. The questionnaire results revealed that the synchronously imagined movements elicited illusory ownership and a sense of agency over the moving robotic hand-the moving rubber hand illusion-compared with asynchronously imagined movements; individuals who affirmed experiencing the illusion with real synchronous movement also did so with synchronous imagined movements. The results from a proprioceptive drift task further demonstrated a shift in the perceived location of the participants' real hand toward the robotic hand in the synchronous versus the asynchronous motor imagery condition. These results suggest that kinesthetic motor imagery can be used to replace veridical congruent somatosensory feedback from a moving finger in the moving rubber hand illusion to trigger illusory body ownership and agency, but only if the temporal congruence rule of the illusion is obeyed. This observation extends previous studies on the integration of mental imagery and sensory perception to the case of multisensory bodily awareness, which has potentially important implications for research into embodiment of brain-computer interface controlled robotic prostheses and computer-generated limbs in virtual reality.

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.

The perception of affective and discriminative touch in blind individuals.

Enhanced tactile acuity in blindness is among the most widely reported results of neuroplasticity following prolonged visual deprivation. However, tactile submodalities other than discriminative touch are profoundly understudied in blind individuals. Here, we examined the influence of blindness on two tactile submodalities, affective and discriminative touch, the former being vital for social functioning and emotional processing. We tested 36 blind individuals and 36 age- and sex-matched sighted volunteers. In Experiment 1, we measured the perception of affective tactile signals by asking participants to rate the pleasantness of touch delivered on the palm (nonhairy skin, sparsely innervated with C tactile [CT] fibers) or the forearm (hairy skin, densely innervated with CT fibers) in a CT-optimal versus a CT-nonoptimal manner using a paradigm grounded in studies on tactile sensory neurophysiology. In Experiment 2, we implemented a classic task assessing discriminative touch abilities, the grating orientation task. We found that blind individuals rated the touch as more pleasant when delivered on the palm than on the forearm, while the opposite pattern was observed for sighted participants, who rated stimulation on the forearm as more pleasant than stimulation on the palm. We also replicated the previous findings showing enhanced discriminative tactile acuity in blind individuals. Altogether, our results suggest that blind individuals might experience affective touch differently than sighted individuals, with relatively greater pleasantness perceived on the palm. These results provide a broader insight into somatosensory perception in blind individuals, for the first time taking into consideration the socioemotional aspect of touch.

The Role of the Skin in Interoception: A Neglected Organ?

In the past 2 decades, interoception has received increasing attention in the fields of psychology and cognitive science, as well as neuroscience and physiology. A plethora of studies adopted the perception of cardiac signals as a proxy for interoception. However, recent findings have cast doubt on the methodological and intrinsic validity of the tasks used thus far. Therefore, there is an ongoing effort to improve the existing cardiac interoceptive tasks and to identify novel channels to target the perception of the physiological state of the body. Amid such scientific abundancy, one could question whether the field has been partially neglecting one of our widest organs in terms of dimensions and functions: the skin. According to some views grounded on anatomical and physiological evidence, skin-mediated signals such as affective touch, pain, and temperature have been redefined as interoceptive. However, there is no agreement in this regard. Here, we discuss some of the anatomical, physiological, and experimental arguments supporting the scientific study of interoception by means of skin-mediated signals. We argue that more attention should be paid to the skin as a sensory organ that monitors the bodily physiological state and further propose thermosensation as a particularly attractive model of skin-mediated interoception.

Combination of visuo-tactile and visuo-vestibular correlations in illusory body ownership and self-motion sensations.

Previous studies have shown that illusory ownership over a mannequin's body can be induced through synchronous visuo-tactile stimulation as well as through synchronous visuo-vestibular stimulation. The current study aimed to elucidate how three-way combinations of correlated visual, tactile and vestibular signals contribute to the senses of body ownership and self-motion. Visuo-tactile temporal congruence was manipulated by touching the mannequin's body and the participant's unseen real body on the trunk with a small object either synchronously or asynchronously. Visuo-vestibular temporal congruence was manipulated by synchronous or asynchronous presentation of a visual motion cue (the background rotating around the mannequin in one direction) and galvanic stimulation of the vestibular nerve generating a rotation sensation (in the same direction). The illusory experiences were quantified using a questionnaire; threat-evoked skin-conductance responses (SCRs) provided complementary indirect physiological evidence for the illusion. Ratings on the illusion questionnaire statement showed significant main effects of synchronous visuo-vestibular and synchronous visuo-tactile stimulations, suggesting that both of these pairs of bimodal correlations contribute to the ownership illusion. Interestingly, visuo-tactile synchrony dominated because synchronous visuo-tactile stimulation combined with asynchronous visuo-vestibular stimulation elicited a body ownership illusion of similar strength as when both bimodal combinations were synchronous. Moreover, both visuo-tactile and visuo-vestibular synchrony were associated with enhanced self-motion perception; self-motion sensations were even triggered when visuo-tactile synchrony was combined with visuo-vestibular asynchrony, suggesting that ownership enhanced the relevance of visual information as a self-motion cue. Finally, the SCR results suggest that synchronous stimulation of either modality pair led to a stronger illusion compared to the asynchronous conditions. Collectively, the results suggest that visuo-tactile temporal correlations have a stronger influence on body ownership than visuo-vestibular correlations and that ownership boosts self-motion perception. We present a Bayesian causal inference model that can explain how visuo-vestibular and visuo-tactile information are combined in multisensory own-body perception.

Modeling affective touch pleasantness across skin types at the individual level reveals a reliable and stable basic function.

Touch is perceived most pleasant when delivered at velocities known to optimally activate the C-tactile afferent system. At the group level, pleasantness ratings of touch delivered at velocities in the range between 0.3 and 30 cm/s follow an inverted-U shape curve, with maximum pleasantness between 1 and 10 cm/s. However, the prevalence, reliability, and stability of this function at the individual level and across skin types based on hair density remains unknown. Here, we tested a range of seven velocities (0.3, 1, 3, 6, 9, 18, and 27 cm/s) delivered with a soft brush, on both hairy (forearm and dorsal hand) and nonhairy skin (palm) in 123 participants. Our results suggest that the relationship between pleasantness and velocity of touch is significantly best described by a negative quadratic model at the individual level in the majority of participants both on hairy (67.1%) and nonhairy (62.6%) skin, a larger extent than previously reported. Higher interoceptive accuracy and self-reported depression were related to a better fit of the quadratic model and the steepness of the curve, respectively. The prevalence of the quadratic model at the individual level was stable across body sites (62.6%, experiment 1), across two experimental sessions (73%-78%, experiment 2), and regardless of the number of repetitions of each velocity (experiment 3). Thus, the individual perception of tactile pleasantness follows a characteristic velocity-dependent function across skin types and shows trait characteristics. Future studies can investigate further the possibility to use affective touch as a behavioral biomarker for mental health disorders.NEW & NOTEWORTHY Touch is perceived as most pleasant when delivered at slow, caress-like velocities, known to activate C-tactile afferents. At the group level, tactile pleasantness and velocity of touch show a reliable pattern of relationship on hairy skin. Here, we found that the perception of tactile pleasantness follows a consistent pattern also at the individual level, across skin types and testing sessions. However, individual differences in interoceptive abilities and self-reported depression do play a role.

Uncertainty-based inference of a common cause for body ownership.

Many studies have investigated the contributions of vision, touch, and proprioception to body ownership, i.e., the multisensory perception of limbs and body parts as our own. However, the computational processes and principles that determine subjectively experienced body ownership remain unclear. To address this issue, we developed a detection-like psychophysics task based on the classic rubber hand illusion paradigm, where participants were asked to report whether the rubber hand felt like their own (the illusion) or not. We manipulated the asynchrony of visual and tactile stimuli delivered to the rubber hand and the hidden real hand under different levels of visual noise. We found that: (1) the probability of the emergence of the rubber hand illusion increased with visual noise and was well predicted by a causal inference model involving the observer computing the probability of the visual and tactile signals coming from a common source; (2) the causal inference model outperformed a non-Bayesian model involving the observer not taking into account sensory uncertainty; (3) by comparing body ownership and visuotactile synchrony detection, we found that the prior probability of inferring a common cause for the two types of multisensory percept was correlated but greater for ownership, which suggests that individual differences in rubber hand illusion can be explained at the computational level as differences in how priors are used in the multisensory integration process. These results imply that the same statistical principles determine the perception of the bodily self and the external world.

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.

Visuo-thermal congruency modulates the sense of body ownership.

Thermosensation has been redefined as an interoceptive modality that provides information about the homeostatic state of the body. However, the contribution of thermosensory signals to the sense of body ownership remains unclear. Across two rubber hand illusion (RHI) experiments (N = 73), we manipulated the visuo-thermal congruency between the felt and seen temperature, on the real and rubber hand respectively. We measured the subjectively experienced RHI, the perceived hand location and temperature of touch, and monitored skin temperature. We found that visuo-thermal incongruencies between the seen and felt touch reduced the subjective and behavioural RHI experience (Experiment 1). Visuo-thermal incongruencies also gave rise to a visuo-thermal illusion effect, but only when the rubber hand was placed in a plausible position (Experiment 2) and when considering individual differences in interoceptive sensibility. Thus, thermosensation contributes to the sense of body ownership by a mechanism of dynamic integration of visual and thermosensory signals.

Multisensory Integration Dominates Hypnotisability and Expectations in the Rubber Hand Illusion.

Some recent papers by P. Lush and colleagues have argued that the rubber hand illusion (RHI), where participants can feel a rubber hand as their own under appropriate multisensory stimulation, may be caused mainly by hypnotic suggestibility and expectations (demand characteristics). These papers rely primarily on a study with 353 participants who took part in a RHI experiment carried out in a classical way with brush stroking. Participants experienced a synchronous condition where the rubber hand was seen to be touched in synchrony with touch felt on their corresponding hidden real hand, or the touches were applied asynchronously as a control. Each participant had a related measure of their hypnotisability on a scale known as the Sussex-Waterloo Scale of Hypnotisability (SWASH). The authors found a correlation between the questionnaire ratings of the RHI in the synchronous condition and the SWASH score. From this, they concluded that the RHI is largely driven by suggestibility and further proposed that suggestibility and expectations may even entirely explain the RHI. Here we examine their claims in a series of extensive new analyses of their data. We find that at every level of SWASH, the synchronous stimulation results in greater levels of the illusion than the asynchronous condition; moreover, proprioceptive drift is greater in the synchronous case at every level of SWASH. Thus, while the level of hypnotisability does modestly influence the subjective reports (higher SWASH is associated with somewhat higher illusion ratings), the major difference between the synchronous and asynchronous stimulation is always present. Furthermore, by including in the model the participants' expectancy ratings of how strongly they initially believed they would experience the RHI in the two conditions, we show that expectations had a very small effect on the illusion ratings; model comparisons further demonstrate that the multisensory condition is two-to-three-times as dominant as the other factors, with hypnotisability contributing modestly and expectations negligibly. Thus, although the results indicate that trait suggestibility may modulate the RHI, presumably through intersubject variations in top-down factors, the findings also suggest that the primary explanation for the RHI is as a multisensory bodily illusion.