The question of how we experience ownership of an entire body distinct from the external world is a fundamental problem in psychology and neuroscience [1-6]. Earlier studies suggest that integration of visual, tactile, and proprioceptive information in multisensory areas [7-11] mediates self-attribution of single limbs. However, it is still unknown how ownership of individual body parts translates into the unitary experience of owning a whole body. Here, we used a "body-swap" illusion [12], in which people experienced an artificial body to be their own, in combination with functional magnetic resonance imaging to reveal a coupling between the experience of full-body ownership and neural responses in bilateral ventral premotor and left intraparietal cortices, and left putamen. Importantly, activity in the ventral premotor cortex reflected the construction of ownership of a whole body from the parts, because it was stronger when the stimulated body part was attached to a body, was present irrespective of whether the illusion was triggered by stimulation of the hand or the abdomen, and displayed multivoxel patterns carrying information about full-body ownership. These findings suggest that the unitary experience of owning an entire body is produced by neuronal populations that integrate multisensory information across body segments.

The ability to identify and localize our own limbs is crucial for survival. Indeed, the majority of our interactions with objects occur within the space surrounding the hands. In non-human primates, neurons in the posterior parietal and premotor cortices dynamically represent the space near the upper limbs in hand-centered coordinates. Neuronal populations selective for the space near the hand also exist in humans. It is unclear whether these remap the peri-hand representation as the arm is moved in space. Furthermore, no combined neuronal and behavioral data are available about the possible involvement of peri-hand neurons in the perception of the upper limbs in any species. We used fMRI adaptation to demonstrate dynamic hand-centered encoding of space by reporting response suppression in human premotor and posterior parietal cortices to repeated presentations of an object near the hand for different arm postures. Furthermore, we show that such spatial representation is related to changes in body perception, being remapped onto a prosthetic hand if perceived as one's own during an illusion. Interestingly, our results further suggest that peri-hand space remapping in the premotor cortex is most tightly linked to the subjective feeling of ownership of the seen limb, whereas remapping in the posterior parietal cortex closely reflects changes in the position sense of the arm. These findings identify the neural bases for dynamic hand-centered encoding of peripersonal space in humans and provide hitherto missing evidence for the link between the peri-hand representation of space and the perceived self-attribution and position of the upper limb.

The dynamic integration of signals from different sensory modalities plays a key role in bodily self-perception. When visual information is used in the multisensory process of localizing and identifying one's own limbs, the sight of a body part often plays a dominant role. For example, it has repeatedly been shown that a viewed object must resemble a humanoid body part to permit illusory self-attribution of that object. Here, we report a perceptual illusion that challenges these assumptions by demonstrating that healthy (nonamputated) individuals can refer somatic sensations to a discrete volume of empty space and experience having an invisible hand. In 10 behavioral and one fMRI experiment, we characterized the perceptual rules and multisensory brain mechanisms that produced this "invisible hand illusion." Our behavioral results showed that the illusion depends on visuotactile-proprioceptive integration that obeys key spatial and temporal multisensory rules confined to near-personal space. The fMRI results associate the illusion experience with increased activity in regions related to the integration of multisensory body-related signals, most notably the bilateral ventral premotor, intraparietal, and cerebellar cortices. We further showed that a stronger feeling of having an invisible hand is associated with a higher degree of effective connectivity between the intraparietal and ventral premotor cortices. These findings demonstrate that the integration of temporally and spatially congruent multisensory signals in a premotor-intraparietal circuit is sufficient to redefine the spatial boundaries of the bodily self, even when visual information directly contradicts the presence of a physical limb at the location of the perceived illusory hand.

The perception of our limbs in space is built upon the integration of visual, tactile, and proprioceptive signals. Accumulating evidence suggests that these signals are combined in areas of premotor, parietal, and cerebellar cortices. However, it remains to be determined whether neuronal populations in these areas integrate hand signals according to basic temporal and spatial congruence principles of multisensory integration. Here, we developed a setup based on advanced 3D video technology that allowed us to manipulate the spatiotemporal relationships of visuotactile (VT) stimuli delivered on a healthy human participant's real hand during fMRI and investigate the ensuing neural and perceptual correlates. Our experiments revealed two novel findings. First, we found responses in premotor, parietal, and cerebellar regions that were dependent upon the spatial and temporal congruence of VT stimuli. This multisensory integration effect required a simultaneous match between the seen and felt postures of the hand, which suggests that congruent visuoproprioceptive signals from the upper limb are essential for successful VT integration. Second, we observed that multisensory conflicts significantly disrupted the default feeling of ownership of the seen real limb, as indexed by complementary subjective, psychophysiological, and BOLD measures. The degree to which self-attribution was impaired could be predicted from the attenuation of neural responses in key multisensory areas. These results elucidate the neural bases of the integration of multisensory hand signals according to basic spatiotemporal principles and demonstrate that the disintegration of these signals leads to "disownership" of the seen real hand.

The senses of owning a body and being localized somewhere in space are two key components of human self-consciousness. Despite a wealth of neurophysiological and neuroimaging research on the representations of the spatial environment in the parietal and medial temporal cortices, the relationship between body ownership and self-location remains unexplored. To investigate this relationship, we used a multisensory out-of-body illusion to manipulate healthy participants' perceived self-location, head direction, and sense of body ownership during high-resolution fMRI. Activity patterns in the hippocampus and the posterior cingulate, retrosplenial, and intraparietal cortices reflected the sense of self-location, whereas the sense of body ownership was associated with premotor-intraparietal activity. The functional interplay between these two sets of areas was mediated by the posterior cingulate cortex. These results extend our understanding of the role of the posterior parietal and medial temporal cortices in spatial cognition by demonstrating that these areas not only are important for ecological behaviors, such as navigation and perspective taking, but also support the perceptual representation of the bodily self in space. Our results further suggest that the posterior cingulate cortex has a key role in integrating the neural representations of self-location and body ownership.