Orientation is a fundamental mental function that processes the relations between the behaving self to space (places), time (events), and person (people). Behavioral and neuroimaging studies have hinted at interrelations between processing of these three domains. To unravel the neurocognitive basis of orientation, we used highresolution 7T functional MRI as 16 subjects compared their subjective distance to different places, events, or people. Analysis at the individual-subject level revealed cortical activation related to orientation in space, time, and person in a precisely localized set of structures in the precuneus, inferior parietal, and medial frontal cortex. Comparison of orientation domains revealed a consistent order of cortical activity inside the precuneus and inferior parietal lobes, with space orientation activating posterior regions, followed anteriorly by person and then time. Core regions at the precuneus and inferior parietal lobe were activated for multiple orientation domains, suggesting also common processing for orientation across domains. The medial prefrontal cortex showed a posterior activation for time and anterior for person. Finally, the default-mode network, identified in a separate resting-state scan, was active for all orientation domains and overlapped mostly with person-orientation regions. These findings suggest that mental orientation in space, time, and person is managed by a specific brain system with a highly ordered internal organization, closely related to the default-mode network.cognitive map | disorientation | precuneus | default network | fMRI

Tales of ghosts, wraiths, and other apparitions have been reported in virtually all cultures. The strange sensation that somebody is nearby when no one is actually present and cannot be seen (feeling of a presence, FoP) is a fascinating feat of the human mind, and this apparition is often covered in the literature of divinity, occultism, and fiction. Although it is described by neurological and psychiatric patients and healthy individuals in different situations, it is not yet understood how the phenomenon is triggered by the brain. Here, we performed lesion analysis in neurological FoP patients, supported by an analysis of associated neurological deficits. Our data show that the FoP is an illusory own-body perception with well-defined characteristics that is associated with sensorimotor loss and caused by lesions in three distinct brain regions: temporoparietal, insular, and especially frontoparietal cortex. Based on these data and recent experimental advances of multisensory own-body illusions, we designed a master-slave robotic system that generated specific sensorimotor conflicts and enabled us to induce the FoP and related illusory own-body perceptions experimentally in normal participants. These data show that the illusion of feeling another person nearby is caused by misperceiving the source and identity of sensorimotor (tactile, proprioceptive, and motor) signals of one's own body. Our findings reveal the neural mechanisms of the FoP, highlight the subtle balance of brain mechanisms that generate the experience of "self" and "other," and advance the understanding of the brain mechanisms responsible for hallucinations in schizophrenia.

The processing of interoceptive signals in the insular cortex is thought to underlie self-awareness. However, the influence of interoception on visual awareness and the role of the insular cortex in this process remain unclear. Here, we show in a series of experiments that the relative timing of visual stimuli with respect to the heartbeat modulates visual awareness. We used two masking techniques and show that conscious access for visual stimuli synchronous to participants' heartbeat is suppressed compared with the same stimuli presented asynchronously to their heartbeat. Two independent brain imaging experiments using high-resolution fMRI revealed that the insular cortex was sensitive to both visible and invisible cardio-visual stimulation, showing reduced activation for visual stimuli presented synchronously to the heartbeat. Our results show that interoceptive insular processing affects visual awareness, demonstrating the role of the insula in integrating interoceptive and exteroceptive signals and in the processing of conscious signals beyond self-awareness.

Recent research has investigated self-consciousness associated with the multisensory processing of bodily signals (e.g., somatosensory, visual, vestibular signals), a notion referred to as bodily self-consciousness, and these studies have shown that the manipulation of bodily inputs induces changes in bodily self-consciousness such as self-identification. Another line of research has highlighted the importance of signals from the inside of the body (e.g., visceral signals) and proposed that neural representations of internal bodily signals underlie self-consciousness, which to date has been based on philosophical inquiry, clinical case studies, and behavioral studies. Here, we investigated the relationship of bodily self-consciousness with the neural processing of internal bodily signals. By combining electrical neuroimaging, analysis of peripheral physiological signals, and virtual reality technology in humans, we show that transient modulations of neural responses to heartbeats in the posterior cingulate cortex covary with changes in bodily self-consciousness induced by the full-body illusion. Additional analyses excluded that measured basic cardiorespiratory parameters or interoceptive sensitivity traits could account for this finding. These neurophysiological data link experimentally the cortical mapping of the internal body to selfconsciousness.

A central feature of our consciousness is the experience of the self as a unified entity residing in a physical body, termed bodily self-consciousness. This phenomenon includes aspects such as the sense of owning a body (also known as body ownership) and has been suggested to arise from the integration of sensory signals from the body. Several studies have shown that temporally synchronous tactile stimulation of the real body and visual stimulation of a fake or virtual body can induce changes in bodily self-consciousness, typically resulting in a sense of illusory ownership over the fake body. The present study assessed the effect of anatomical congruency of visuo-tactile stimulation on bodily self-consciousness. A virtual body was presented and temporally synchronous visuo-tactile stroking was applied simultaneously to the participants' body and to the virtual body. We manipulated the anatomical locations of the visuo-tactile stroking (i.e., on the back, on the leg), resulting in congruent stroking (stroking was felt and seen on the back or the leg) or incongruent stroking (i.e., stroking was felt on the leg and seen on the back). We measured self-identification with the virtual body and self-location as well as skin temperature. Illusory self-identification with the avatar as well as changes in self-location were experienced in the congruent stroking conditions. Participants showed a decrease in skin temperature across several body locations during congruent stimulation. These data establish that the full-body illusion (FBI) alters bodily self-consciousness and instigates widespread physiological changes in the participant's body.