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KEY POINTS: Embodiment of a virtual body was induced and its movements were controlled by two different brain-computer interface (BCI) paradigms - one based on signals from sensorimotor versus one from visual cortical areas. BCI-control of movements engenders agency, but not equally for all paradigms. Cortical sensorimotor activation correlates with agency and responsibility. This has significant implications for neurological rehabilitation and neuroethics. ABSTRACT: Agency is the attribution of an action to the self and is a prerequisite for experiencing responsibility over its consequences. Here we investigated agency and responsibility by studying the control of movements of an embodied avatar, via brain-computer interface (BCI) technology, in immersive virtual reality. After induction of virtual body ownership by visuomotor correlations, healthy participants performed a motor task with their virtual body. We compared the passive observation of the subject's 'own' virtual arm performing the task with (1) the control of the movement through activation of sensorimotor areas (motor imagery) and (2) the control of the movement through activation of visual areas (steady-state visually evoked potentials). The latter two conditions were carried out using a BCI and both shared the intention and the resulting action. We found that BCI-control of movements engenders the sense of agency, which is strongest for sensorimotor area activation. Furthermore, increased activity of sensorimotor areas, as measured using EEG, correlates with levels of agency and responsibility. We discuss the implications of these results for the neural basis of agency.

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Changes in body representation may affect pain perception. The effect of a distorted body image, such as the telescoping effect in amputee patients, on pain perception, is unclear. This study aimed to investigate whether distorting an embodied virtual arm in virtual reality (simulating the telescoping effect in amputees) modulated pain perception and anticipatory responses to pain in healthy participants. Twenty-seven right-handed participants were immersed in virtual reality and the virtual arm was shown with three different levels of distortion with a virtual threatening stimulus either approaching or contacting the virtual hand. We evaluated pain/discomfort ratings, ownership, and skin conductance responses (SCRs) after each condition. Viewing a distorted virtual arm enhances the SCR to a threatening event with respect to viewing a normal control arm, but when viewing a reddened-distorted virtual arm, SCR was comparatively reduced in response to the threat. There was a positive relationship between the level of ownership over the distorted and reddened-distorted virtual arms with the level of pain/discomfort, but not in the normal control arm. Contact with the threatening stimulus significantly enhances SCR and pain/discomfort, while reduced SCR and pain/discomfort were seen in the simulated-contact condition. These results provide further evidence of a bi-directional link between body image and pain perception.

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The most robust and clear biological index differentiating persons with schizophrenia from healthy controls is the drastic reduction of the amplitude of their P300b event-related brain potential (ERP). However, the cause of that reduction remains obscure. Nevertheless, the P300b belongs to the family of the late posterior positivities (LPPs) which are closely related to the consciousness of the meaning of the stimulus in the task for the participants themselves (e.g., the: I am seeing the target stimulus for which I have to respond). The fragmentation of the self present in schizophrenia, could thus be the cause. If this were true, then P300bs should be somewhat reduced in healthy participants when their self representations are temporarily and minimally fragmented. We tested this hypothesis by using the innocuous fragmentation of the self that occurs in virtual reality (VR). There, participants can have a fragment of their self in an avatar they feel embodied in, within a VR room, while having another fragment of their self in their real body in the real room where they know they are. Our participants were thus equipped with a head mounted display in which they viewed a virtual room where a female humanoid avatar was facing them. She was lifting her right hand in synchrony with the participants, in order to induce in them a feeling of embodiment. Stimuli were a frequent green- and a rare red-disk, the oddball stimulus, occurring over the right hand of the avatar. Participants had to perform a Go/NoGo task, lifting their right hand to the frequent green disk and repressing this action for the oddball red disk. In the syncMove block of trials the avatar was lifting her right hand synchronously with the participant, disturbing her self representation as confirmed by the debriefing session. In the noMove block, the avatar remained immobile. In the classic block, only the red and the green disk were displayed on a monochrome background, neither the room nor the avatar were shown. As predicted, P300bs were found to be smaller in the syncMove block than in the noMove- and the classic-block in participants who had the classically large P300b oddball effect between ERPs to the frequent and those to the rare stimuli. Reduced P300bs of schizophrenia could thus be partly due to self fragmentation. Results may also open an avenue of research to the functional significance of LPPs and the content of the consciousness indexed by these potentials.

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Seeing one's own body has been reported to have analgesic properties. Analgesia has also been described when seeing an embodied virtual body colocated with the real one. However, there is controversy regarding whether this effect holds true when seeing an illusory-owned body part, such as during the rubber-hand illusion. A critical difference between these paradigms is the distance between the real and surrogate body part. Colocation of the real and surrogate arm is possible in an immersive virtual environment, but not during illusory ownership of a rubber arm. The present study aimed at testing whether the distance between a real and a virtual arm can explain such differences in terms of pain modulation. Using a paradigm of embodiment of a virtual body allowed us to evaluate heat pain thresholds at colocation and at a 30-cm distance between the real and the virtual arm. We observed a significantly higher heat pain threshold at colocation than at a 30-cm distance. The analgesic effects of seeing a virtual colocated arm were eliminated by increasing the distance between the real and the virtual arm, which explains why seeing an illusorily owned rubber arm does not consistently result in analgesia. These findings are relevant for the use of virtual reality in pain management. PERSPECTIVE: Looking at a virtual body has analgesic properties similar to looking at one's real body. We identify the importance of colocation between a real and a surrogate body for this to occur and thereby resolve a scientific controversy. This information is useful for exploiting immersive virtual reality in pain management.

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The somatotopic layout of the primary somatosensory cortex is known for its fine spatial structure as delineated in single cell recordings and macroscopic EEG evoked responses. While a gross somatotopic layout has been revealed also for neuronal oscillations responding to sensorimotor stimulation of distant body parts (e.g. hand vs. foot), it is still unclear whether these oscillatory dynamics exhibit fine spatial layout comparable to those found in evoked responses. In twelve healthy subjects we applied electric stimuli to the first (D1) and fifth finger (D5) of the same hand while performing high-density electroencephalography. We used Common Spatial Pattern analysis to optimally extract components showing the strongest Event-Related Desynchronization (ERD) in neuronal alpha oscillations. In agreement with the previous studies, dipole locations of Somatosensory Evoked Potentials (SEPs) confirmed the existence of spatially distinct representations of each finger. In contrast, dipole locations of alpha-ERD patterns did not yield spatially different source locations, indicating that the stimulation of different fingers most likely resulted in oscillatory activity of overlapping neuronal populations. When both fingers were stimulated simultaneously the SEP dipole strength was found increased in comparison to a stimulation of either finger alone, in agreement with spatially distinct SEP to finger stimulation. The strength of ERD, on the other hand, was the same regardless of whether either one or both fingers were stimulated. Our findings might reflect anatomical constraints on the sequential temporal activation of fingers' skin where almost simultaneous activation of many fingers usually occurs in everyday activities, such as grasping or holding objects. Such simultaneity is unlikely to benefit from slow amplitude modulation of alpha oscillations, which would rather be beneficial for contrasting somatosensory processing of distinct body parts.

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