Keeping up with ourselves: Multimodal processes underlying body ownership across the lifespan.

The sense of a bodily self is thought to depend on adaptive weighting and integration of bodily afferents and prior beliefs. While the physical body changes in shape, size, and functionality across the lifespan, the sense of body ownership remains relatively stable. Yet, little is known about how multimodal integration underlying such sense of ownership is altered in ontogenetic periods of substantial physical changes. We aimed to study this link for the motor and the tactile domain in a mixed-realty paradigm where participants ranging from 7 to 80 years old saw their own body with temporally mismatching multimodal signals. Participants were either stroked on their hand or moved it, while they saw it in multiple trials with different visual delays. For each trial, they judged the visuo-motor/tactile synchrony and rated the sense of ownership for the seen hand. Visual dependence and proprioceptive acuity were additionally assessed. The results show that across the lifespan body ownership decreases with increasing temporal multisensory mismatch, both in the tactile and the motor domain. We found an increased sense of ownership with increasing age independent of delay and modality. Delay sensitivity during multisensory conflicts was not consistently related to age. No effects of age were found on visual dependence or proprioceptive accuracy. The results are at least partly in line with an enhanced weighting of top-down and a reduced weighting of bottom-up signals for the momentary sense of bodily self with increasing age.

An interhemispheric frontoparietal network supports hypnotic states.

Understanding the neural substrate of altered conscious states is an important cultural, scientific, and clinical endeavour. Although hypnosis causes strong shifts in conscious perception and cognition, it remains largely unclear how hypnosis affects information processing in cortical networks. Here we manipulated the depth of hypnotic states to study information processing between cortical regions involved in attention and awareness. We used high-density Electroencephalography (EEG) to record resting-state cortical activity from 30 hypnosis experts during two hypnotic states with different depth. Each participant entered a light and a deep hypnotic state as well as two well-matched control states. Bridging top-down and lateralisation models of hypnosis, we found that interhemispheric frontoparietal connectivity distinguished hypnosis and control conditions, while no difference was found between the two hypnotic states. Using a graph-theoretic measure, we revealed that the amount of information passing through individual nodes (measured via betweenness centrality) is reduced during hypnosis relative to control states. Finally, we found that theta power was enhanced during hypnosis. Our result contributes to the current discussion around a role for theta power in bringing about hypnotic states, as well as other altered conscious states. Overall, our findings support the notion that altered top-down control in frontoparietal regions facilitates hypnosis by integrating information between cortical hemispheres.

Fluid face but not gender: Enfacement illusion through digital face filters does not affect gender identity.

It has been shown that observing a face being touched or moving in synchrony with our own face increases self-identification with the former which might alter both cognitive and affective processes. The induction of this phenomenon, termed enfacement illusion, has often relied on laboratory tools that are unavailable to a large audience. However, digital face filters applications are nowadays regularly used and might provide an interesting tool to study similar mechanisms in a wider population. Digital filters are able to render our faces in real time while changing important facial features, for example, rendering them more masculine or feminine according to normative standards. Recent literature using full-body illusions has shown that participants' own gender identity shifts when embodying a different gendered avatar. Here we studied whether participants' filtered faces, observed while moving in synchrony with their own face, may induce an enfacement illusion and if so, modulate their gender identity. We collected data from 35 female and 33 male participants who observed a stereotypically gender mismatched version of themselves either moving synchronously or asynchronously with their own face on a screen. Our findings showed a successful induction of the enfacement illusion in the synchronous condition according to a questionnaire addressing the feelings of ownership, agency and perceived similarity. However, we found no evidence of gender identity being modulated, neither in explicit nor in implicit measures of gender identification. We discuss the distinction between full-body and facial processing and the relevance of studying widely accessible devices that may impact the sense of a bodily self and our cognition, emotion and behaviour.

Examining the association between depersonalisation traits and the bodily self in waking and dreaming.

Depersonalisation (DP) is characterized by fundamental alterations to the sense of self that include feelings of detachment and estrangement from one's body. We conducted an online study in healthy participants (n = 514) with DP traits to investigate and quantify the subjective experience of body and self during waking and dreaming, as the vast majority of previous studies focussed on waking experience only. Investigating dreams in people experiencing DP symptoms may help us understand whether the dream state is a 'spared space' where people can temporarily 'retrieve' their sense of self and sense of bodily presence. We found that higher DP traits-i.e. higher scores on the Cambridge Depersonalisation Scale (CDS)-were associated with more frequent dream experiences from an outside observer perspective (r = 0.28) and more frequent dream experiences of distinct bodily sensations (r = 0.23). We also found that people with higher CDS scores had more frequent dream experiences of altered bodily perception (r = 0.24), more frequent nightmares (r = 0.33) and higher dream recall (r = 0.17). CDS scores were negatively correlated with body boundary scores (r = - 0.31) in waking states and there was a negative association between CDS scores and the degree of trust in interoceptive signals (r = - 0.52). Our study elucidates the complex phenomenology of DP in relation to bodily selfhood during waking and dreaming and suggests avenues for potential therapeutic interventions in people with chronic depersonalisation (depersonalisation -derealisation disorder).

Effects of environmental colours in virtual reality: Physiological arousal affected by lightness and hue.

It is a popular belief that colours impact one's psychological and affective functioning. However, clear-cut scientific evidence is still lacking, largely due to methodological challenges. Virtual reality (VR) enabled us to control and modify the environment. We exposed 60 participants to red or blue environments varying in lightness and saturation. We assessed participants' physiological responses (i.e. arousal) with heart rate and skin conductance measures, and their self-reported levels of valence and arousal in response to the coloured environments. The results revealed physiological effects of lightness and hue. When compared with the baseline measures, heart rate increased, and heart rate variability decreased more in the dark than the medium lightness rooms. Both measures signalled higher arousal in the darker room, irrespective of hue. Also, when compared with the baseline measures, skin conductance increased more in the red than the blue rooms, again signalling higher arousal in the red condition. The difference between the red and the blue conditions was detectable only on some saturation and lightness combinations. We conclude that being immersed in environments of different colours can change physiological arousal. However, not all changes are driven by hue and not all the effects are measurable on all physiological parameters.

Ownership of the affected leg is further reduced following deceptive behaviors in body integrity dysphoria.

Although predicted by the notion of embodied morality, it remains unknown whether a reduced sense of body ownership (SoO) is associated with increased or decreased dishonesty. To clarify this issue, we tested patients with body integrity dysphoria (BID), a clinical condition characterized by chronic reductions of SoO toward one leg that patients persistently desire to have amputated. Participants with BID played a card game in which they could voluntarily tell the truth or cheat an opponent, and thus either steal or give them money. To assess whether SoO toward the effector limb influences (im)moral decisions, responses were communicated with the affected or the unaffected leg. We found that a higher number of self-gain lies was followed by further reductions of SoO toward the affected leg. Our result supports the idea that reductions of SoO may follow immoral behaviors to distance from unwanted characteristics of the self, like one's own dishonesty.

From rubber hands to neuroprosthetics: Neural correlates of embodiment.

Our interaction with the world rests on the knowledge that we are a body in space and time, which can interact with the environment. This awareness is usually referred to as sense of embodiment. For the good part of the past 30 years, the rubber hand illusion (RHI) has been a prime tool to study embodiment in healthy and people with a variety of clinical conditions. In this paper, we provide a critical overview of this research with a focus on the RHI paradigm as a tool to study prothesis embodiment in individuals with amputation. The RHI relies on well-documented multisensory integration mechanisms based on sensory precision, where parietal areas are involved in resolving the visuo-tactile conflict, and premotor areas in updating the conscious bodily representation. This mechanism may be transferable to prosthesis ownership in amputees. We discuss how these results might transfer to technological development of sensorised prostheses, which in turn might progress the acceptability by users.

Cardiac interoception in infants: Behavioral and neurophysiological measures in various emotional and self-related contexts.

Interoception, the perception of internal bodily signals, is fundamental to our sense of self. Even though theoretical accounts suggest an important role for interoception in the development of the self, empirical investigations are limited, particularly in infancy. Previous studies used preferential-looking paradigms to assess the detection of sensorimotor and multisensory contingencies in infancy, usually related to proprioception and touch. So far, only one recent study reported that infants discriminated between audiovisual stimuli presented synchronously or asynchronously with their heartbeat. This discrimination was related to the amplitude of the infant's heartbeat evoked potentials (HEP), a neural correlate of interoception. In the current study, we measured looking preferences between synchronous and asynchronous visuocardiac (bimodal), and audiovisuocardiac (trimodal) stimuli as well as the HEP in conditions of different emotional contexts and with different degrees of self-relatedness in a mirror-like setup. While the infants preferred trimodal to bimodal stimuli, we did not observe the predicted differences between synchronous and asynchronous stimulation. Furthermore, the HEP was not modulated by emotional context or self-relatedness. These findings do not support previously published results and highlight the need for further studies on the early development of interoception in relation to the development of the self.

Visual perspective, distance, and felt presence of others in dreams.

The peripersonal space, that is, the limited space surrounding the body, involves multisensory coding and representation of the self in space. Previous studies have shown that peripersonal space representation and the visual perspective on the environment can be dramatically altered when neurotypical individuals self-identify with a distant avatar (i.e., in virtual reality) or during clinical conditions (i.e., out-of-body experience, heautoscopy, depersonalization). Despite its role in many cognitive/social functions, the perception of peripersonal space in dreams, and its relationship with the perception of other characters (interpersonal distance in dreams), remain largely uncharted. The present study aimed to explore the visuospatial properties of this space, which is likely to underlie self-location as well as self/other distinction in dreams. 530 healthy volunteers answered a web-based questionnaire to measure their dominant visuo-spatial perspective in dreams, the frequency of recall for felt distances between their dream self and other dream characters, and the dreamers' viewing angle of other dream characters. Most participants reported dream experiences from a first-person perspective (1PP) (82%) compared to a third-person perspective (3PP) (18%). Independent of their dream perspective, participants reported that they generally perceived other dream characters in their close space, that is, at distance of either between 0 and 90 cm, or 90-180 cm, than in further spaces (180-270 cm). Regardless of the perspective (1PP or 3PP), both groups also reported more frequently seeing other dream characters from eye level (0° angle of viewing) than from above (30° and 60°) or below eye level (-30° and -60°). Moreover, the intensity of sensory experiences in dreams, as measured by the Bodily Self-Consciousness in Dreams Questionnaire, was higher in individuals who habitually see other dream characters closer to their personal dream self (i.e., within 0-90 cm and 90-180 cm). These preliminary findings offer a new, phenomenological account of space representation in dreams with regards to the felt presence of others. They might provide insights not only to our understanding of how dreams are formed, but also to the type of neurocomputations involved in self/other distinction.

Neural bases of the bodily self as revealed by electrical brain stimulation: A systematic review.

An increasing amount of recent research has focused on the multisensory and neural bases of the bodily self. This pre-reflective form of self is considered as multifaceted, incorporating phenomenal components, such as self location, body ownership, first-person perspective, agency, and the perceptual body image. Direct electrical brain stimulation (EBS) during presurgical evaluation of epilepsy and brain tumor resection is a unique method to causally relate specific brain areas to the various phenomenal components of the bodily self. We conducted a systematic review of the literature describing altered phenomenal experience of the bodily self evoked by EBS. We included 42 articles and analyzed self reports from 221 patients. Three-dimensional density maps of EBS revealed that stimulation in the middle cingulum, inferior parietal lobule, supplementary motor area, posterior insula, hippocampal complex/amygdala, and precuneus most consistently altered one or several components of the bodily self. In addition, we found that only EBS in the parietal cortex induced disturbances of all five components of the bodily self considered in this review article. These findings inform current neuroscientific models of the bodily self.

Influence of bodily states on cognition: A web-based study in individuals with body integrity dysphoria.

There is a constant reciprocal flow of information between the malleable sensorimotor states of the body and cognitive functions, and some embodied cognition approaches argue that many cognitive-affective mechanisms depend on the physical characteristics of the body. To examine such influences of bodily state, the current study compared patients with body integrity dysphoria (BID) with an amputation desire of the lower limb to a healthy control group on an Implicit Association Test for self-identity and self-esteem, and a pain evaluation task. Patients with BID completed the tasks once while emulating their desired bodily state and once while simulating their undesired bodily state, while healthy controls were split into two groups: one control group completed the experiment once while either sitting on one leg and once while sitting in a normal position, whereas the other control group completed both experiments while sitting in a normal position. Results demonstrate that patients with BID implicitly identify more strongly with an amputated body, whereas healthy controls demonstrate stronger identification with a complete body, independent of bodily state. Furthermore, implicit self-esteem did not differ between the groups and was also not modulated by bodily state manipulation in any of the groups. Pain evaluation ratings were not influenced by bodily state manipulation, perspective, or consistency. Pain forced choice response times, however, revealed that individuals with BID were faster to judge whether the stimulus depicted was painful when simulating their desired bodily state. These results provide insightful information to how both the subjective sense of body, as well as more transient alterations of objective sensorimotor states of the physical body may exert selective pressure on certain cognitive tasks.

Happiness feels light and sadness feels heavy: introducing valence-related bodily sensation maps of emotions.

Bodily sensation mapping (BSM) is a recently developed self-report tool for the assessment of emotions in which people draw their sensations of activation in a body silhouette. Following the circumplex model of affect, activity and valence are the underling dimensions of every emotional experience. The aim of this study was to introduce the neglected valence dimension in BSM. We found that participants systematically report valence-related sensations of bodily lightness for positive emotions (happiness, love, pride), and sensations of bodily heaviness in response to negative emotions (e.g., anger, fear, sadness, depression) with specific body topography (Experiment 1). Further experiments showed that both computers (using a machine learning approach) and humans recognize emotions better when classification is based on the combined activity- and valence-related BSMs compared to either type of BSM alone (Experiments 2 and 3), suggesting that both types of bodily sensations reflect distinct parts of emotion knowledge. Importantly, participants found it clearer to indicate their bodily sensations induced by sadness and depression in terms of bodily weight than bodily activity (Experiment 2 and 4), suggesting that the added value of valence-related BSMs is particularly relevant for the assessment of emotions at the negative end of the valence spectrum.

Different armpits under my new nose: Olfactory sex but not gender affects implicit measures of embodiment.

Conflicting multisensory signals may alter embodiment to produce self-identification with a foreign body, but the role of olfaction in this process has been overlooked. We studied in healthy participants how sex (male and female sweat odors) and gender (male and female cosmetic scents) olfactory stimuli contribute to embodiment. Participants saw, on a head mounted display, the first-person perspective of a sex mismatching person. Synchronous visuotactile stimulation was applied to enhance illusory embodiment. Simultaneously, they smelled either sex- or gender- congruent or incongruent stimuli. We assessed implicit (skin conductance responses to visual threats) and explicit (questionnaire) measures of embodiment. Stronger responses to threat were found when participants smelled the sex-congruent compared to the sex-incongruent odor, while no such differences were found for the cosmetic scents. According to the questionnaire, embodiment did not differ between conditions. Post-experimental assessment of the presented cues, suggest that while both sweat odors were considered generally male, cosmetic scents were not. The presented scents were generally not associated with the embodied body. Our results suggest that sex-related body odors influence implicit but not explicit aspects of embodiment and are in line with unique characteristics of olfaction in other aspects of cognition.

The Scientific Study of Consciousness Cannot and Should Not Be Morally Neutral.

A target question for the scientific study of consciousness is how dimensions of consciousness, such as the ability to feel pain and pleasure or reflect on one's own experience, vary in different states and animal species. Considering the tight link between consciousness and moral status, answers to these questions have implications for law and ethics. Here we point out that given this link, the scientific community studying consciousness may face implicit pressure to carry out certain research programs or interpret results in ways that justify current norms rather than challenge them. We show that because consciousness largely determines moral status, the use of nonhuman animals in the scientific study of consciousness introduces a direct conflict between scientific relevance and ethics-the more scientifically valuable an animal model is for studying consciousness, the more difficult it becomes to ethically justify compromises to its well-being for consciousness research. Finally, in light of these considerations, we call for a discussion of the immediate ethical corollaries of the body of knowledge that has accumulated and for a more explicit consideration of the role of ideology and ethics in the scientific study of consciousness.

Is this my foot? Experimentally induced disownership in individuals with body integrity dysphoria.

In body integrity dysphoria (BID), otherwise healthy individuals feel like a part of their physical body does not belong to them despite normal sensorimotor functioning. Theoretical and empirical evidence suggested aweakened integration of the affected body part into higher-order multisensory cortical body networks. Here, we used a multisensory stimulation paradigm in mixed reality to modulate and investigate multisensory processing underlying body (dis)ownership in individuals with BID of the lower limb. In 20 participants with BID, delay perception and body ownership were measured after introducing delays between the visual and tactile information of viewed stroking applied to affected and unaffected body parts. Unlike predicted, delay perception did not differ between the two body parts. However, specifically for the affected limb, ownership was lower and more strongly modulated by delay. These findings might be following the idea of a stronger dependency on online bottom up sensory signals in BID.

Not my body, not my pain? Pain perception and placebo analgesia in individuals with body integrity dysphoria.

Acute and chronic states of physical pain are inherently linked to our bodily perception. Bodily illusion paradigms have demonstrated that an experimentally induced sense of body disownership can modulate both acute and chronic pain. Insight into the relationship between enduring clinical alterations in body perception and pain is much more limited. The current study examined both pain perception and placebo analgesia in Body Integrity Dysphoria (BID), a clinical model of long-term alterations of bodily disownership: in its most commonly studied variant, people feel like a part of their body does not belong to them, leading to a desire for amputation of a physically healthy limb. Heat stimulations were applied before and after a placebo intervention (sham analgesic cream) to the desired and the undesired leg of 19 patients with BID with a unilateral leg amputation desire. Pain perception was assessed using pain thresholds, and ratings for pain intensity and pain unpleasantness. Results show that pain perception and placebo efficacy were lower for the undesired than for the desired leg, demonstrating a potential link between a clinical disorder of body ownership, pain perception, and placebo analgesia.

White matter abnormalities in the amputation variant of body integrity dysphoria.

"Body integrity dysphoria" (BID) is a severe condition affecting nonpsychotic individuals. In the amputation variant of BID, a limb may be experienced as not being part of the body, despite normal anatomical development and intact sensorimotor functions. We previously demonstrated altered brain structural (gray matter) and functional connectivity in 16 men with BID with a long-lasting and exclusive desire for left leg amputation. Here, we aimed to identify, in the same sample, altered patterns of white matter structural connectivity. Fractional anisotropy (FA), derived from diffusion tensor imaging data, was considered as a measure of structural connectivity. Results showed reduced structural connectivity of: (i) the right superior parietal lobule (rSPL) with the right cuneus, with the superior occipital and with the posterior cingulate gyri, (ii) the pars orbitalis of the right middle frontal gyrus (rMFGOrb) with the putamen, and (iii) the left middle temporal gyrus (lMTG) with the pars triangularis of the left inferior frontal gyrus. Increased connectivity was found between the right paracentral lobule (rPLC) and the right caudate nucleus. By using a complementary method of investigation, we confirmed and extended previous results from the same sample of individuals with BID, showing structural alterations between areas tuned to the processing of the sensorimotor representations of the affected leg (rPCL), and to higher-order components of bodily representation such as the body image (rSPL) and visual processing. Alongside this network for bodily awareness, other networks such as the limbic (rMFGOrb) and the mirror (lMTG) systems showed alterations in structural connectivity. These findings consolidate current understanding of the neural correlates of the amputation variant of BID, which might in turn guide diagnostics and rehabilitative treatments.

Dissociation of motor control from motor awareness in awake sleepwalkers: An EEG study in virtual reality.

Recent behavioral evidence from a virtual reality (VR) study indicates that awake sleepwalkers show dissociation of motor control and motor awareness. This dissociation resembles the nocturnal disintegration of motor awareness and movement during episodes of sleepwalking. Here, we set out to examine the neural underpinnings of altered motor awareness in sleepwalkers by measuring EEG modulation during redirected walking in VR. To this end, we measured scalp EEG during ongoing motor behavior to provide information on motor processing and its modulation in VR. Using this approach, we discovered distinct EEG patterns associated to dual tasking and sub-threshold motor control in sleepwalkers compared to control subjects. These observations provide further electrophysiological evidence for the proposed brain-body dissociation in awake sleepwalkers. This study shows proof-of-principle that EEG biomarkers of movement in a VR setting add to the understanding of altered motor awareness in sleepwalkers. In a broader perspective, we confirm the feasibility of using the additional dimensionality in VR providing novel diagnostic biomarkers not accessible to conventional clinical investigations. In future studies, this approach could contribute to the diagnostic work-up of patients with a broad spectrum of neurological diseases.

Real Bodies Not Required? Placebo Analgesia and Pain Perception in Immersive Virtual and Augmented Reality.

Pain represents an embodied experience, wherein inferences are not only drawn from external sensory inputs, but also from bodily states. Previous research has demonstrated that a placebo administered to an embodied rubber hand can effectively induce analgesia, providing first evidence that placebos can work even when applied to temporarily embodied, artificial body parts. Using a heat pain paradigm, the present study investigates placebo analgesia and pain perception during virtual embodiment. We examined whether a virtual placebo (a sham heat protective glove) can successfully induce analgesia, even when administered to a virtual body. The analgesic efficacy of the virtual placebo to the real hand (augmented reality setting) or virtual hand (virtual reality setting) was compared to a physical placebo administered to the own, physical body (physical reality setting). Furthermore, pain perception and subjective embodiment were compared between settings. In this mixed design experiment, healthy participants (n = 48) were assigned to either an analgesia-expectation or control-expectation group, where subjective and objective pain was measured at pre- and post-intervention time points. Results demonstrate that pre-intervention pain intensity was lower in the virtual reality setting, and that participants in the analgesia-expectation group, after the intervention, exhibited significantly higher pain thresholds, and lower pain intensity and unpleasantness ratings than control-expectation participants, independent of the setting. Our findings show that a virtual placebo can elicit placebo analgesia comparable to that of a physical placebo, and that administration of a placebo does not necessitate physical bodily interaction to produce analgesic responses. PERSPECTIVE: This study demonstrates that a virtual placebo treatment, even when administered to a virtual body, can produce placebo analgesia. These findings indicate that the efficacy of a virtual placebo is comparable to that of a physical placebo, which could pave the way for effective new non-pharmacological approaches for pain management.