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.

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.

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.

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.

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.

Integration of predictions and afferent signals in body ownership.

We aimed at investigating the contribution of sensory predictions triggered by the sight of an object moving towards the body for the sense of body ownership. We used a recently developed psychophysical discrimination task to assess body ownership in the rubber hand illusion. In this task, the participants had to choose which of the two right rubber hands in view felt most like their own, and the ownership discriminations were fitted to psychometric curves. In the current study, we occluded the visual impressions of the object moving towards one of the rubber hands (during the first two-thirds of the path) and only revealed the final third of the object's movement trajectory when it touched the rubber hand (approach-occluded condition). Alternatively, we occluded only the final part so that the main part of the movement towards the model hand was visible (touch-occluded). We compared these two conditions to an illusion baseline condition where the object was visible during the entire trajectory and contact (no-occlusion). The touch-occluded condition produced equally strong hand ownership as the baseline condition with no occlusion, while ownership perception was significantly reduced when vision of the object approaching the rubber hand was occluded (approach-occluded). Our results show that tactile predictions generated from seeing an object moving towards the body are temporally exact, and they contribute to the rubber hand illusion by integrating with temporally congruent afferent sensory signals. This finding highlights the importance of multisensory predictions in peripersonal space, object permanence, and the interplay between bottom-up sensory signals and top-down predictions in body ownership.

Which hand is mine? Discriminating body ownership perception in a two-alternative forced-choice task.

The experience of one's body as one's own is referred to as the sense of body ownership. This central part of human conscious experience determines the boundary between the self and the external environment, a crucial distinction in perception, action, and cognition. Although body ownership is known to involve the integration of signals from multiple sensory modalities, including vision, touch, and proprioception, little is known about the principles that determine this integration process, and the relationship between body ownership and perception is unclear. These uncertainties stem from the lack of a sensitive and rigorous method to quantify body ownership. Here, we describe a two-alternative forced-choice discrimination task that allows precise and direct measurement of body ownership as participants decide which of two rubber hands feels more like their own in a version of the rubber hand illusion. In two experiments, we show that the temporal and spatial congruence principles of multisensory stimulation, which determine ownership discrimination, impose tighter constraints than previously thought and that texture congruence constitutes an additional principle; these findings are compatible with theoretical models of multisensory integration. Taken together, our results suggest that body ownership constitutes a genuine perceptual multisensory phenomenon that can be quantified with psychophysics in discrimination experiments.

Seeing Your Foot Move Changes Muscle Proprioceptive Feedback.

Multisensory effects are found when the input from single senses combines, and this has been well researched in the brain. Presently, we examined in humans the potential impact of visuo-proprioceptive interactions at the peripheral level, using microneurography, and compared it with a similar behavioral task. We used a paradigm where participants had either proprioceptive information only (no vision) or combined visual and proprioceptive signals (vision). We moved the foot to measure changes in the sensitivity of single muscle afferents, which can be altered by the descending fusimotor drive. Visual information interacted with proprioceptive information, where we found that for the same passive movement, the response of muscle afferents increased when the proprioceptive channel was the only source of information, as compared with when visual cues were added, regardless of the attentional level. Behaviorally, when participants looked at their foot moving, they more accurately judged differences between movement amplitudes, than in the absence of visual cues. These results impact our understanding of multisensory interactions throughout the nervous system, where the information from different senses can modify the sensitivity of peripheral receptors. This has clinical implications, where future strategies may modulate such visual signals during sensorimotor rehabilitation.

What's left of the mirror illusion when the mirror can no longer be seen? Bilateral integration of proprioceptive afferents!

Recent data suggest that manipulating the muscle afferents of one arm affects both ipsilateral and contralateral perceptual estimates. Here, we used the mirror paradigm to study the bimanual integration of kinesthetic muscle afferents. The reflection of a moving hand in a mirror positioned in the sagittal plane creates an illusion of symmetrical bimanual movement. Although vision clearly has a role in kinesthesia, its role in the mirror illusion might have been overestimated. Conversely, the role of bimanual integration of muscle afferents might have been underestimated. We hypothesized that muscle-proprioceptive afferents of the passively displaced arm (the image of which was reflected in the mirror) are involved in this illusion. We evoked in 19 healthy adult participants the mirror illusion by displacing passively their left arm, the image of which was reflected in the mirror. Once participants experienced the illusion that their hidden right arm was moving, we then either occluded their view of the mirror (using occlusive glasses) and/or prevent the passive left arm displacement. Participants' illusion characteristics (duration and kinematic) under these conditions were compared with classical mirror illusion (without visual occlusion). We found that as long as the arm was still moving, the kinesthetic illusion decayed slowly after visual occlusion. These findings suggest that the mirror illusion results from the combination of visuo-proprioceptive signals from the two arms and is not purely visual in origin. Our findings also support the more general concept whereby proprioceptive afferents are integrated bilaterally for the purpose of kinesthesia during bimanual tasks.

The kinaesthetic mirror illusion: How much does the mirror matter?

The reflection of a moving hand in a mirror positioned in the sagittal plane can create an illusion of symmetrical, bimanual movement. This illusion is implicitly presumed to be of visual origin. However, muscle proprioceptive afferents of the arm reflected in the mirror might also affect the perceived position and movement of the other arm. We characterized the relative contributions of visual and proprioceptive cues by performing two experiments. In Experiment 1, we sought to establish whether kinaesthetic illusions induced using the mirror paradigm would survive marked visual impoverishment (obtained by covering between 0 and 100 % of the mirror in 16 % steps). We found that the mirror illusion was only significantly influenced when the visual degradation was 84 % or more. In Experiment 2, we masked the muscle proprioceptive afferents of the arm reflected in the mirror by co-vibrating antagonistic muscles. We found that masking the proprioceptive afferents reduced the velocity of the illusory displacement of the other arm. These results confirm that the mirror illusion is not a purely visual illusion but emerges from a combination of congruent signals from the two arms, i.e. visual afferents from the virtually moving arm and proprioceptive afferents from the contralateral, moving arm.

Kinaesthetic mirror illusion and spatial congruence.

Position sense and kinaesthesia are mainly derived from the integration of somaesthetic and visual afferents to form a single, coherent percept. However, visual information related to the body can play a dominant role in these perceptual processes in some circumstances, and notably in the mirror paradigm. The objective of the present study was to determine whether or not the kinaesthetic illusions experienced in the mirror paradigm obey one of the key rules of multisensory integration: spatial congruence. In the experiment, the participant's left arm (the image of which was reflected in a mirror) was either passively flexed/extended with a motorized manipulandum (to induce a kinaesthetic illusion in the right arm) or remained static. The right (unseen) arm remained static but was positioned parallel to the left arm's starting position or placed in extension (from 15° to 90°, in steps of 15°), relative to the left arm's flexed starting position. The results revealed that the frequency of the illusion decreased only slightly as the incongruence prior to movement onset between the reflected left arm and the hidden right arm grew and remained quite high even in the most incongruent settings. However, the greater the incongruence between the visually and somaesthetically specified positions of the right forearm (from 15° to 90°), the later the onset and the lower the perceived speed of the kinaesthetic illusion. Although vision dominates perception in a context of visuoproprioceptive conflict (as in the mirror paradigm), our results show that the relative weightings allocated to proprioceptive and visual signals vary according to the degree of spatial incongruence prior to movement onset.