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[EN] The ability of virtual reality (VR) to recreate controlled, immersive, and interactive environments that provide intensive and customized exercises has motivated its therapeutic use after stroke. Interaction and bodily presence in VR-based interventions is usually mediated through virtual selves, which synchronously represent body movements or responses to events on external input devices. Embodied self-representations in the virtual world not only provide an anchor for visuomotor tasks, but their morphologies can have behavioral implications. While research has focused on the underlying subjective mechanisms of exposure to VR on healthy individuals, the transference of these findings to individuals with stroke is not evident and remains unexplored, which could affect the experience and, ultimately, the clinical effectiveness of neurorehabilitation interventions. This study determined and compared the sense of embodiment and presence elicited by a virtual environment under different perspectives and levels of immersion in healthy subjects and individuals with stroke. Forty-six healthy subjects and 32 individuals with stroke embodied a gender-matched neutral avatar in a virtual environment that was displayed in a first-person perspective with a head-mounted display and in a third-person perspective with a screen, and the participants were asked to interact in a virtual task for 10 min under each condition in counterbalanced order, and to complete two questionnaires about the sense of embodiment and presence experienced during the interaction. The sense of body-ownership, self-location, and presence were more vividly experienced in a first-person than in a third-person perspective by both healthy subjects (p < 0.001, eta(2)(p) = 0.212; p = 0.005, eta(2)(p) = 0.101; p = 0.001, eta(2)(p) = 0.401, respectively) and individuals with stroke (p = 0.019, eta(2)(p) = 0.070; p = 0.001, eta(2)(p) = 0.135; p = 0.014, eta(2)(p) = 0.077, respectively). In contrast, no agency perspective-related differences were found in any group. All measures were consistently higher for healthy controls than for individuals with stroke, but differences between groups only reached statistical significance in presence under the first-person condition (p < 0.010, eta(2)(p) = 0.084). In spite of these differences, the participants experienced a vivid sense of embodiment and presence in almost all conditions. These results provide first evidence that, although less intensively, embodiment and presence are similarly experienced by individuals who have suffered a stroke and by healthy individuals, which could support the vividness of their experience and, consequently, the effectiveness of VR-based interventions., This study was funded by Ministerio de Economía y Competitividad of Spain (Project RTC-2017-6051-7 and Grant BES-2014-068218), Fundació la Marató de la TV3 (Grant 201701-10), and Universitat Politècnica de València (Grant PAID-10-18). We acknowledge the support of NVIDIA Corporation with the donation of the Titan Xp GPU used for this research.

Self-consciousness consists of several dissociable experiences, including the sense of ownership of one's body and the sense of agency over one's action consequences. The relationship between body-ownership and the sense of agency has been described by different neurocognitive models, each providing specific neurofunctional predictions. According to an “additive” model, the sense of agency entails body-ownership, while an alternative “independence” hypothesis suggests that they represent two qualitatively different processes, underpinned by distinct brain systems. We propose a third “interactive” model, arguing the interdependence between body-ownership and the sense of agency: these constructs might represent different experiences with specific and exclusive brain correlates, but they also could partly overlap at the neurofunctional level. Here we test these three neurocognitive models by reviewing the available neurofunctional literature of body-ownership and the sense of agency, with a quantitative meta-analytical approach that allowed us to compare their neural correlates statistically. We identified (i) a body-ownership-specific network including the left inferior parietal lobule and the left extra-striate body area, (ii) a sense-of-agency-specific network including the left SMA, the left posterior insula, the right postcentral gyrus, and the right superior temporal lobe and (iii) a shared network in the left middle insula. These results provide support for the interactive neurocognitive model of body-ownership and the sense of agency. Body-ownership involves a sensory network in which multisensory inputs are integrated to be self-attributed. On the other hand, the sense of agency is specifically associated with premotor and sensory-motor areas, typically involved in generating motor predictions and in action monitoring. Finally, body-ownership and the sense of agency interact at the level of the left middle insula, a high-level multisensory hub engaged in body and action awareness in general.

International audience; While the Sense of Agency (SoA) has so far been predominantly characterised in VR as a component of the Sense of Embodiment, other communities (e.g., in psychology or neurosciences) have investigated the SoA from a different perspective proposing complementary theories. Yet, despite the acknowledged potential benefits of catching up with these theories a gap remains. This paper first aims to contribute to fill this gap by introducing a theory according to which the SoA can be divided into two components, the feeling and the judgment of agency, and relies on three principles, namely the principles of priority, exclusivity and consistency. We argue that this theory could provide insights on the factors influencing the SoA in VR systems. Second, we propose novel approaches to manipulate the SoA in controlled VR experiments (based on these three principles) as well as to measure the SoA, and more specifically its two components based on neurophysiological markers, using ElectroEncephaloGraphy (EEG). We claim that these approaches would enable us to deepen our understanding of the SoA in VR contexts. Finally, we validate these approaches in an experiment. Our results (N=24) suggest that our approach was successful in manipulating the SoA as the modulation of each of the three principles induced significant decreases of the SoA (measured using questionnaires). In addition, we recorded participants' EEG signals during the VR experiment, and neurophysiological markers of the SoA, potentially reflecting the feeling and judgment of agency specifically, were revealed. Our results also suggest that users' profile, more precisely their Locus of Control (LoC), influences their level of immersion and SoA.

[EN] Stroke can lead to motor impairments that can affect the body structure and restraint mobility. We hypothesize that brain lesions and their motor sequelae can distort the body schema, a sensorimotor map of body parts and elements in the peripersonal space through which human beings embody the reachable space and ready the body for forthcoming movements. Two main constructs have been identified in the embodiment mechanism: body-ownership, the sense that the body that one inhabits is his/her own, and agency, the sense that one can move and control his/her body. To test this, the present study simultaneously investigated different embodiment subcomponents (body-ownership, localization, and agency) and different neurophysiological measures (galvanic skin response, skin temperature, and surface electromyographic activity), and the interaction between them, in clinically-controlled hemiparetic individuals with stroke and in healthy subjects after the rubber hand illusion. Individuals with stroke reported significantly stronger body-ownership and agency and reduced increase of galvanic skin response, skin temperature, and muscular activity in the stimulated hand. We suggest that differences in embodiment could have been motivated by increased plasticity of the body schema and pathological predominance of the visual input over proprioception. We also suggest that differences in neurophysiological responses could have been promoted by a suppression of the reflex activity of the sympathetic nervous system and by the involvement of the premotor cortex in the reconfiguration of the body schema. These results could evidence a body schema plasticity promoted by the brain lesion and a main role of the premotor cortex in this mechanism., This work was supported by Ministerio de Economia y Competitividad of Spain (Project NeuroVR, TIN2013-44741-R, Project REACT, TIN2014-61975-EXP, and Grant BES-2014-068218), and by Universitat Politecnica de Valencia (Grant PAID-10-14).

[EN] The Rubber Hand Illusion (RHI) is a perceptual illusion that enables integration of artificial limbs into the body representation through combined multisensory integration. Most previous studies investigating the RHI have involved young healthy adults within a very narrow age range (typically 20-30 years old). The purpose of this paper was to determine the influence of age on the RHI. The RHI was performed on 93 healthy adults classified into three groups of age (20-35 years old, N = 41; 36-60 years old, N = 28; and 61-80 years old, N = 24), and its effects were measured with subjective (Embodiment of Rubber Hand Questionnaire), behavioral (proprioceptive drift), and physiological (changes in skin temperature and conductance) measures. There were neither significant differences among groups in any response, nor significant covariability or correlation between age and other measures (but for skin temperature), which suggests that the RHI elicits similar responses across different age groups in the adult phase., This study was funded, in part, by Ministerio de Economia y Competitividad of Spain (Grant BES-2014-068218) and Universitat Politecnica de Valencia (Grant PAID-10-14 and Grant PAID-10-16).

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., Frontiers in Behavioral Neuroscience, 7, ISSN:1662-5153

International audience; Brain correlates of the sense of agency have recently received increased attention. However, the explorations remain largely restricted to the study of brains in isolation. The prototypical paradigm used so far consists of manipulating visual perception of own action while asking the subject to draw a distinction between self- versus externally caused action. However, the recent definition of agency as a multifactorial phenomenon combining bottom-up and top-down processes suggests the exploration of more complex situations. Notably there is a need of accounting for the dynamics of agency in a two-body context where we often experience the double faceted question of who is at the origin of what in an ongoing interaction. In a dyadic context of role switching indeed, each partner can feel body ownership, share a sense of agency and altogether alternate an ascription of the primacy of action to self and to other. To explore the brain correlates of these different aspects of agency, we recorded with dual EEG and video set-ups 22 subjects interacting via spontaneous versus induced imitation (II) of hand movements. The differences between the two conditions lie in the fact that the roles are either externally attributed (induced condition) or result from a negotiation between subjects (spontaneous condition). Results demonstrate dissociations between self- and other-ascription of action primacy in delta, alpha and beta frequency bands during the condition of II. By contrast a similar increase in the low gamma frequency band (38-47 Hz) was observed over the centro-parietal regions for the two roles in spontaneous imitation (SI). Taken together, the results highlight the different brain correlates of agency at play during live interactions.

Introduction. Aberrant experience of agency is characteristic of schizophrenia. An understanding of the neurobiological basis of such experience is therefore of considerable importance for developing successful models of the disease. We aimed to characterise the effects of ketamine, a drug model for psychosis, on sense of agency (SoA). SoA is associated with a subjective compression of the temporal interval between an action and its effects: This is known as "intentional binding". This action-effect binding provides an indirect measure of SoA. Previous research has found that the magnitude of binding is exaggerated in patients with schizophrenia. We therefore investigated whether ketamine administration to otherwise healthy adults induced a similar pattern of binding.Methods. 14 right-handed healthy participants (8 female; mean age 22.4 years) were given low-dose ketamine (100 ng/mL plasma) and completed the binding task. They also underwent structured clinical interviews.Results. Ketamine mimicked the performance of schizophrenia patients on the intentional binding task, significantly increasing binding relative to placebo. The size of this effect also correlated with aberrant bodily experiences engendered by the drug.Conclusions. These data suggest that ketamine may be able to mimic certain aberrant agency experiences that characterise schizophrenia. The link to individual changes in bodily experience suggests that the fundamental change produced by the drug has wider consequences in terms of individuals' experiences of their bodies and movements.

Artificial 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.