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.

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.

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.

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.

Prosthesis embodiment and attenuation of prosthetic touch in upper limb amputees - A proof-of-concept study.

Sensory attenuation of self-touch, that is, the perceptual reduction of self-generated tactile stimuli, is considered a neurocognitive basis for self-other distinction. However, whether this effect can also be found in upper limb amputees using a prosthesis is unknown. Thirteen participants were asked to touch their foot sole with a) their intact hand (self-touch), b) their prosthesis (prosthesis-touch), or c) let it be touched by another person (other-touch). Intensity of touch was assessed with a questionnaire. In addition, prosthesis embodiment was assessed in nine participants. Self-touch as well as prosthesis-touch was characterized by significant perceptual attenuation compared to other-touch, while self- and prosthesis-touch did not differ. The more embodied the prosthesis was, the more similar was its elicited touch perception to actual self-touch. These findings - although preliminary - suggest that perceptually embodied prostheses can be represented as an actual limb by the users' sensorimotor system.

The monologue of the double: Allocentric reduplication of the own voice alters bodily self-perception.

During autoscopic phenomena, people perceive a double of themselves in extrapersonal space. Such clinical allocentric self-experiences sometimes co-occur with auditory hallucinations, yet experimental setups to induce similar illusions in healthy participants have generally neglected acoustic cues. We investigated whether feeling the presence of an auditory double could be provoked experimentally by recording healthy participants' own versus another person's voice and movements using binaural headphones from an egocentric (the participants' own) and an allocentric (a dummy head located elsewhere) perspective. When hearing themselves allocentrically, participants reported feeling a self-identified presence extracorporeally, an arguably distinct quality of autoscopy. Our results suggest that participants without hallucinatory experiences localized their own voice closer to themselves compared to that of another person. Explorative findings suggest that distinct patterns for hallucinators should be further investigated. This study suggests a successful induction of the feeling of an acoustic doppelganger, bridging clinical phenomena and experimental work.

Development of the bodily self: Effects of visuomotor synchrony and visual appearance on virtual embodiment in children and adults.

The sense of a bodily self is thought to depend on adaptive weighting and integration of bodily afferents and prior beliefs. Evidence from studies using paradigms such as the rubber hand illusion and full body illusion suggests changes in the integration of visuotactile bodily signals throughout childhood. Here, we extended this line of research by assessing how bottom-up visuomotor synchrony and expectancy, modulated by visual appearance of virtual avatars, contribute to embodiment in children. We compared responses to a first-person perspective virtual full body illusion from 8- to 12-year-old children and adults while manipulating synchrony of the avatar's movements (synchronous, 0.5-s delay, or 1-s delay compared with the participant's movements) and appearance of the avatar (human or skeleton). We measured embodiment with both subjective questionnaires and objective skin conductance responses to virtual threat. Results showed that children experienced ownership for the virtual avatar in a similar way as adults, which was reduced with increasing asynchrony, and for the skeleton avatar as compared with the human avatar. This modulation of ownership was not reflected in the skin conductance responses, which were equally high in all experimental conditions and only showed a modulation of repetition by age. In contrast, in children the subjective experience of agency was less affected by the dampening effects of visuomotor asynchrony or reduced human likeness and was overall higher. These findings suggest that children can easily embody a virtual avatar but that different aspects of embodiment develop at different rates, which could have important implications for applications of embodied virtual reality.

Uncertainty Promotes Neuroreductionism: A Behavioral Online Study on Folk Psychological Causal Inference from Neuroimaging Data.

INTRODUCTION: Increased efforts in neuroscience try to understand mental disorders as brain disorders. In the present study, we investigate how common a neuroreductionist inclination is among highly educated people. In particular, we shed light on implicit presuppositions of mental disorders little is known about in the public, exemplified here by the case of body integrity dysphoria (BID) that is considered a mental disorder for the first time in ICD-11. METHODS: Identically graphed, simulated data of mind-brain correlations were shown in 3 contexts with presumably different presumptions about causality. 738 highly educated lay people rated plausibility of causality attribution from the brain to mind and from mind to the brain for correlations between brain structural properties and mental phenomena. We contrasted participants' plausibility ratings of causality in the contexts of commonly perceived brain lesion-induced behavior (aphasia), behavior-induced training effects (piano playing), and a newly described mental disorder (BID). RESULTS: The findings reveal the expected context-dependent modulation of causality attributions in the contexts of aphasia and piano playing. Furthermore, we observed a significant tendency to more readily attribute causal inference from the brain to mind than vice versa with respect to BID. CONCLUSION: In some contexts, exemplified here by aphasia and piano playing, unidirectional causality attributions may be justified. However, with respect to BID, we critically discuss presumably unjustified neuroreductionist inclinations under causal uncertainty. Finally, we emphasize the need for a presupposition-free approach in psychiatry.

Being Short, Sweet, and Sour: Congruent Visuo-Olfactory Stimulation Enhances Illusory Embodiment.

Bodily self-identification has shown to be easily altered through spatiotemporally congruent multimodal signals. While such manipulations are mostly studied through visuo-tactile or visuo-motor stimulation, here we investigated whether congruent visuo-olfactory cues might enhance illusory self-identification with an arbitrary object. Using virtual reality, healthy individuals saw a grapefruit from its supposed first-person perspective that was touched in synchrony with their own body. The touch attempted to replicate what was seen as softly squeezing the grapefruit. Crucially, when we additionally presented the smell of a grapefruit in synchrony with the squeezing, they self-identified stronger with the fruit than when they smelled strawberry.

Tactile motion lacks momentum.

The displacement of the final position of a moving object in the direction of the observed motion path, i.e. an overestimation, is known as representational momentum. It has been described both in the visual and the auditory domain, and is suggested to be modality-independent. Here, we tested whether a representational momentum can also be demonstrated in the somatosensory domain. While the cognitive literature on representational momentum suggests that it can, previous work on the psychophysics of tactile motion perception would rather predict an underestimation of the perceived endpoint of a tactile stimulus. Tactile motion stimuli were applied on the left and the right dorsal forearms of 32 healthy participants, who were asked to indicate the subjectively perceived endpoint of the stimulation. Velocity, length and direction of the trajectory were varied. Contrary to the prediction based on the representational momentum literature, participants in our experiment significantly displaced the endpoint against the direction of movement (underestimation). The results are thus compatible with previous psychophysical findings on the perception of tactile motion. Further studies combining paradigms from classical psychophysics and cognitive psychology will be needed to resolve the apparently paradoxical predictions by the two literatures.

Minding gaps on the skin: Opposite bisection biases on forehead and back of one's head.

Humans perceive the world from an egocentric perspective, while being able to mentally take a third person's perspective. Graphesthesia tasks revealed that letters written on the back of one's own head are consistently perceived from an embodied perspective, while the perspective on one's front is less consistent and often disembodied. We developed a cutaneous gap bisection task as a more discrete measure of the perspective on the body. In analogy to a visual pseudoneglect, we expected bisections to deviate toward the left ear when perceived from an embodied perspective. While this hypothesis was confirmed for gap bisections on the back, the results on the front suggest overall a disembodied perspective. Contrary to our expectation, this pattern was not predicted by the spontaneous perspective participants took in a graphesthesia task, indicating different cognitive mechanisms. We discuss these findings in the frame of the current literature on spatial attention and perspective taking.

Binding body and self in visuo-vestibular conflicts.

Maintenance of the bodily self relies on the accurate integration of multisensory inputs in which visuo-vestibular cue integration is thought to play an essential role. Here, we tested in healthy volunteers how conflicting visuo-vestibular bodily input might impact on body self-coherence in a full body illusion set-up. Natural passive vestibular stimulation was provided on a motion platform, while visual input was manipulated using virtual reality equipment. Explicit (questionnaire) and implicit (skin temperature) measures were employed to assess illusory self-identification with either a mannequin or a control object. Questionnaire results pointed to a relatively small illusion, but hand skin temperature, plausibly an index of illusory body ownership, showed the predicted drop specifically in the condition when participants saw the mannequin moving in congruence with them. We argue that this implicit measure was accessible to visuo-vestibular modulation of the sense of self, possibly mediated by shared neural processes in the insula involved in vestibular and interoceptive signalling, thermoregulation and multisensory integration.

Don't look at my wheelchair! The plasticity of longlasting prejudice.

CONTEXT: Scientific research has consistently shown that prejudicial behaviour may contribute to discrimination and disparities in social groups. However, little is known about whether and how implicit assumptions and direct contact modulate the interaction and quality of professional interventions in education and health contexts. OBJECTIVES: This study was designed to examine implicit and explicit attitudes towards wheelchair users. METHODS: We investigated implicit and explicit attitudes towards wheelchair users in three different groups: patients with traumatic spinal cord injury (SCI); health professionals with intense contact with wheelchair users, and healthy participants without personal contact with wheelchair users. To assess the short-term plasticity of prejudices, we used a valid intervention that aims to change implicit attitudes through brief direct contact with a patient who uses a wheelchair in an ecologically valid real-life interaction. RESULTS: We found that: (i) wheelchair users with SCI held positive explicit but negative implicit attitudes towards their novel in-group; (ii) the amount of experience with wheelchair users affected implicit attitudes among health professionals, and (iii) interacting with a patient with SCI who contradicts prejudices modulated implicit negative bias towards wheelchair users in healthy participants. CONCLUSIONS: The use of a wheelchair immediately and profoundly affects how a person is perceived. However, our findings highlight the dynamic nature of perceptions of social identity, which are not only sensitive to personal beliefs, but also highly permeable to intergroup interactions. Having direct contact with people with disabilities might foster positive attitudes in multidisciplinary health care teams. Such interventions could be integrated into medical education programmes to successfully prevent or reduce hidden biases in a new generation of health professionals and to increase the general acceptance of disability in patients.

The bodily self and its disorders: neurological, psychological and social aspects.

PURPOSE OF REVIEW: The experience of ourselves as an embodied agent with a first-person perspective is referred to as 'bodily self'. We present a selective overview of relevant clinical and experimental studies. RECENT FINDINGS: Sharing multisensory body space with others can be observed in patients with structurally altered bodies (amputations, congenital absence of limbs), with altered functionality after hemiplegia, such as denial of limb ownership (somatoparaphrenia) and with alterations in bodily self-consciousness on the level of the entire body (e.g. in autoscopic phenomena). In healthy participants, the mechanisms underpinning body ownership and observer perspective are empirically investigated by multisensory stimulation paradigms to alter the bodily self. The resulting illusions have promoted the understanding of complex disturbances of the bodily self, such as out-of-body experiences. We discuss the role of interoception in differentiating between self and others and review current advances in the study of body integrity identity disorder, a condition shaped as much by neurological as by social-psychological factors. SUMMARY: We advocate a social neuroscience approach to the bodily self that takes into account the interactions between body, mind and society and might help close the divide between neurology and psychiatry.

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.

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).

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.

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.

Listening to your heart and feeling yourself: effects of exposure to interoceptive signals during the ultimatum game.

The ultimatum game (UG) is commonly used to study the tension between financial self-interest and social equity motives. Here, we investigated whether experimental exposure to interoceptive signals influences participants' behavior in the UG. Participants were presented with various bodily sounds--i.e., their own heart, another person's heart, or the sound of footsteps--while acting both in the role of responder and proposer. We found that listening to one's own heart sound, compared to the other bodily sounds: (1) increased subjective feelings of unfairness, but not rejection behavior, in response to unfair offers and (2) increased the unfair offers while playing in the proposer role. These findings suggest that heightened feedback of one's own visceral processes may increase a self-centered perspective and drive socioeconomic exchanges accordingly. In addition, this study introduces a valuable procedure to manipulate online the access to interoceptive signals and for exploring the interplay between viscero-sensory information and cognition.

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.