Humans normally experience the conscious self as localized within their bodily borders. This spatial unity may break down in certain neurological conditions such as out-of-body experiences, leading to a striking disturbance of bodily self-consciousness. On the basis of these clinical data, we designed an experiment that uses conflicting visual-somatosensory input in virtual reality to disrupt the spatial unity between the self and the body. We found that during multisensory conflict, participants felt as if a virtual body seen in front of them was their own body and mislocalized themselves toward the virtual body, to a position outside their bodily borders. Our results indicate that spatial unity and bodily selfconsciousness can be studied experimentally and are based on multisensory and cognitive processing of bodily information.

Self-consciousness has mostly been approached by philosophical enquiry and not by empirical neuroscientific study, leading to an overabundance of diverging theories and an absence of data-driven theories. Using robotic technology, we achieved specific bodily conflicts and induced predictable changes in a fundamental aspect of self-consciousness by altering where healthy subjects experienced themselves to be (self-location). Functional magnetic resonance imaging revealed that temporo-parietal junction (TPJ) activity reflected experimental changes in self-location that also depended on the first-person perspective due to visuo-tactile and visuo-vestibular conflicts. Moreover, in a large lesion analysis study of neurological patients with a well-defined state of abnormal self-location, brain damage was also localized at TPJ, providing causal evidence that TPJ encodes self-location. Our findings reveal that multisensory integration at the TPJ reflects one of the most fundamental subjective feelings of humans: the feeling of being an entity localized at a position in space and perceiving the world from this position and perspective.

BackgroundThe spatial unity between self and body can be disrupted by employing conflicting visual-somatosensory bodily input, thereby bringing neurological observations on bodily self-consciousness under scientific scrutiny. Here we designed a novel paradigm linking the study of bodily self-consciousness to the spatial representation of visuo-tactile stimuli by measuring crossmodal congruency effects (CCEs) for the full body.Methodology/Principal FindingsWe measured full body CCEs by attaching four vibrator-light pairs to the trunks (backs) of subjects who viewed their bodies from behind via a camera and a head mounted display (HMD). Subjects made speeded elevation (up/down) judgments of the tactile stimuli while ignoring light stimuli. To modulate self-identification for the seen body subjects were stroked on their backs with a stick and the felt stroking was either synchronous or asynchronous with the stroking that could be seen via the HMD.We found that (1) tactile stimuli were mislocalized towards the seen body (2) CCEs were modulated systematically during visual-somatosensory conflict when subjects viewed their body but not when they viewed a body-sized object, i.e. CCEs were larger during synchronous than during asynchronous stroking of the body and (3) these changes in the mapping of tactile stimuli were induced in the same experimental condition in which predictable changes in bodily self-consciousness occurred.Conclusions/SignificanceThese data reveal that systematic alterations in the mapping of tactile stimuli occur in a full body illusion and thus establish CCE magnitude as an online performance proxy for subjective changes in global bodily self-consciousness.

a b s t r a c tVisual, somatosensory, and perspectival cues normally provide congruent information about where the self is experienced. Separating those cues by virtual reality techniques, recent studies found that self-location was systematically biased to where a visual-tactile event was seen. Here we developed a novel, repeatable and implicit measure of self-location to compare and extend previous protocols. We investigated illusory self-location and associated phenomenological aspects in a lying body position that facilitates clinically observed abnormal self-location (as on out-of-body experiences). The results confirm that the self is located to where touch is seen. This leads to either predictable lowering or elevation of self-localization, and the latter was accompanied by sensations of floating, as during out-of-body experiences. Using a novel measurement we show that the unitary and localized character of the self can be experimentally separated from both the origin of the visual perspective and the location of the seen body, which is compatible with clinical data.

a b s t r a c tArtificial stimulation of the peripheral vestibular system has been shown to improve ownership of body parts in neurological patients, suggesting vestibular contributions to bodily self-consciousness. Here, we investigated whether galvanic vestibular stimulation (GVS) interferes with the mechanisms underlying ownership, touch, and the localization of one's own hand in healthy participants by using the ''rubber hand illusion" paradigm. Our results show that left anodal GVS increases illusory ownership of the fake hand and illusory location of touch. We propose that these changes are due to vestibular interference with spatial and/or temporal mechanisms of visual-tactile integration leading to an enhancement of visual capture. As only left anodal GVS lead to such changes, and based on neurological data on body part ownership, we suggest that this vestibular interference is mediated by the right temporo-parietal junction and the posterior insula.

The vestibular system analyses angular and linear accelerations of the head that are important information for perceiving the location of one's own body in space. Vestibular stimulation and in particular galvanic vestibular stimulation (GVS) that allow a systematic modiWcation of vestibular signals has so far mainly been used to investigate vestibular inXuence on sensori-motor integration in eye movements and postural control. Comparatively, only a few behavioural and imaging studies have investigated how cognition of space and body may depend on vestibular processing. This study was designed to diVerentiate the inXuence of left versus right anodal GVS compared to sham stimulation on object-based versus egocentric mental transformations. While GVS was applied, subjects made leftright judgments about pictures of a plant or a human body presented at diVerent orientations in the roll plane. All subjects reported illusory sensations of body self-motion and/ or visual Weld motion during GVS. Response times in the mental transformation task were increased during right but not left anodal GVS for the more diYcult stimuli and the larger angles of rotation. Post-hoc analyses suggested that the interfering eVect of right anodal GVS was only present in subjects who reported having imagined turning themselves to solve the mental transformation task (egocentric transformation) as compared to those subjects having imagined turning the picture in space (object-based mental transformation). We suggest that this eVect relies on shared functional and cortical mechanisms in the posterior parietal cortex associated with both right anodal GVS and mental imagery.

a b s t r a c tPain and the conscious mind (or the self) are experienced in our body. Both are intimately linked to the subjective quality of conscious experience. Here, we used virtual reality technology and visuo-tactile conflicts in healthy subjects to test whether experimentally induced changes of bodily self-consciousness (self-location; self-identification) lead to changes in pain perception. We found that visuo-tactile stroking of a virtual body but not of a control object led to increased pressure pain thresholds and self-location. This increase was not modulated by the synchrony of stroking as predicted based on earlier work. This differed for self-identification where we found as predicted that synchrony of stroking increased self-identification with the virtual body (but not a control object), and positively correlated with an increase in pain thresholds. We discuss the functional mechanisms of self-identification, self-location, and the visual perception of human bodies with respect to pain perception.

Increasing evidence suggests that the basic foundations of the self lie in the brain systems that represent the body. Specific sensorimotor stimulation has been shown to alter the bodily self. However, little is known about how disconnection of the brain from the body affects the phenomenological sense of the body and the self. Spinal cord injury (SCI) patients who exhibit massively reduced somatomotor processes below the lesion in the absence of brain damage are suitable for testing the influence of body signals on two important components of the self–the sense of disembodiment and body ownership. We recruited 30 SCI patients and 16 healthy participants, and evaluated the following parameters: (i) depersonalization symptoms, using the Cambridge Depersonalization Scale (CDS), and (ii) measures of body ownership, as quantified by the rubber hand illusion (RHI) paradigm. We found higher CDS scores in SCI patients, which show increased detachment from their body and internal bodily sensations and decreasing global body ownership with higher lesion level. The RHI paradigm reveals no alterations in the illusory ownership of the hand between SCI patients and controls. Yet, there was no typical proprioceptive drift in SCI patients with intact tactile sensation on the hand, which might be related to cortical reorganization in these patients. These results suggest that disconnection of somatomotor inputs to the brain due to spinal cord lesions resulted in a disturbed sense of an embodied self. Furthermore, plasticity-related cortical changes might influence the dynamics of the bodily self.