Interoception, network physiology and the emergence of bodily self-awareness.

The interplay between the brain and interoceptive signals is key in maintaining internal balance and orchestrating neural dynamics, encompassing influences on perceptual and self-awareness. Central to this interplay is the differentiation between the external world, others and the self, a cornerstone in the construction of bodily self-awareness. This review synthesizes physiological and behavioral evidence illustrating how interoceptive signals can mediate or influence bodily self-awareness, by encompassing interactions with various sensory modalities. To deepen our understanding of the basis of bodily self-awareness, we propose a network physiology perspective. This approach explores complex neural computations across multiple nodes, shifting the focus from localized areas to large-scale neural networks. It examines how these networks operate in parallel with and adapt to changes in visceral activities. Within this framework, we propose to investigate physiological factors that disrupt bodily self-awareness, emphasizing the impact of interoceptive pathway disruptions, offering insights across several clinical contexts. This integrative perspective not only can enhance the accuracy of mental health assessments but also paves the way for targeted interventions.

Individual Differences in Bodily Self-Consciousness and Its Neural Basis.

Bodily self-consciousness (BSC), a subject of interdisciplinary interest, refers to the awareness of one's bodily states. Previous studies have noted the existence of individual differences in BSC, while neglecting the underlying factors and neural basis of such individual differences. Considering that BSC relied on integration from both internal and external self-relevant information, we here review previous findings on individual differences in BSC through a three-level-self model, which includes interoceptive, exteroceptive, and mental self-processing. The data show that cross-level factors influenced individual differences in BSC, involving internal bodily signal perceptibility, multisensory processing principles, personal traits shaped by environment, and interaction modes that integrate multiple levels of self-processing. Furthermore, in interoceptive processing, regions like the anterior cingulate cortex and insula show correlations with different perceptions of internal sensations. For exteroception, the parietal lobe integrates sensory inputs, coordinating various BSC responses. Mental self-processing modulates differences in BSC through areas like the medial prefrontal cortex. For interactions between multiple levels of self-processing, regions like the intraparietal sulcus involve individual differences in BSC. We propose that diverse experiences of BSC can be attributed to different levels of self-processing, which moderates one's perception of their body. Overall, considering individual differences in BSC is worth amalgamating diverse methodologies for the diagnosis and treatment of some diseases.

The foundations of informed consent and bodily self-sovereignty: a positive suggestion.

In medical care, the obtaining of informed consent is taken to be required prior to treatment in order to ensure that patients sufficiently understand the potential risks and benefits of a given medical procedure. In this paper, I begin by looking at the history of informed consent and consider how the norms and laws in medicine have evolved away from benevolent paternalism and toward a blanket obligation to obtain informed consent. In so doing, I consider what values might be taken to underpin such a requirement. After dismissing some unsatisfactory answers, I offer a positive view as to the constellation of values I think informed consent ought to be protecting. I call these bodily self-sovereignty, which I take to be a coupling of two groups of values: autonomy and non-domination on the one hand, and self-ownership and personal integrity on the other. Given the connection between autonomy and responsibility, autonomy is both required for the act of consenting, and respected by allowing it. And, because of our special and inescapable relationship to our own bodies, this authorization is particularly important when our bodies are involved. So, I suggest that informed consent protects our self-sovereignty over our own bodies.

Embodiment of underweight and normal-weight avatars affects bodily self-representations in anorexia nervosa.

Body image distortion (BID) is a crucial aspect of anorexia nervosa (AN), leading to body overestimation, dissatisfaction, and low self-esteem. BID significantly influences the onset, maintenance, and relapse of the pathology. We assessed whether a Full Body Illusion (FBI) using under and normal-weight avatars' bodies affects perceptual body image and body schema estimations in both individuals with anorexia nervosa (AN) and healthy controls (HC). After each embodiment procedure, we asked participants to estimate the width of their hips (Perceptual Body Image Task) and the minimum aperture width of a virtual door necessary to pass through it (Body Schema Task). Additionally, we asked participants to rate the avatars in terms of self-similarity, attractiveness, and implicit disgust (i.e., pleasant/unpleasant body odour). Whereas participants with AN (N = 26) showed changes in body schema estimations after embodying the normal-weight avatar, no changes were found in HC (N = 25), highlighting increased bodily self-plasticity in AN. Notably, individuals with AN rated the normal weight avatar as the most similar to their real body, which was also considered the least attractive and the most repulsive. These ratings correlated with BID severity. Furthermore, at the explicit level, all participants reported feeling thinner than usual after embodying the underweight avatar. Overall, our findings suggest that BID in AN engages multiple sensory channels (from visual to olfactory) and components (from perceptual to affective), offering potential targets for innovative non-invasive treatments aimed at modifying flexible aspects of body representation.

A quantitative study on peripersonal space in anorexia nervosa and healthy subjects: Role of social variables and association with psychopathology.

Objectives: This study investigates peripersonal space (PPS) modulation in patients with anorexia nervosa (AN) versus healthy controls (HCs) and explores associations between PPS, eating-related, and general psychopathology. Method: Forty-six patients and 42 HCs completed a computer-based task observing videos of an approaching actor (male or female) displaying different facial expressions along with a non-social condition. Then, participants completed self-report questionnaires assessing eating-related and general psychopathology. Results: Mixed-models revealed that both groups adjusted PPS based on task conditions, with a gender effect favoring closer proximity to female actor. HCs reduced PPS amplitude progressively during the task, while patients did not show this effect. In patients, wider PPS correlated with lower self-esteem and facial expression identification accuracy, while in HCs, PPS was associated to body dissatisfaction and anxiety symptoms. Conclusion: These findings enhance understanding of bodily self-consciousness, suggesting PPS consideration in therapeutic interactions with patients with AN and as a potential target in treatments addressing social impairment.

Malleability of the sense of bodily self in early childhood: 5- and 6-year-old children show the enfacement illusion.

The mechanisms underlying the developing sense of bodily self are debated. Whereas some scholars stress the role of sensory factors, others propose the importance of contextual factors. By manipulating multisensory stimulation and social familiarity with the other person, we explored two factors that are proposed to relate to young children's developing sense of bodily self. Including an adult sample allowed us to investigate age-related differences of the malleability of the bodily self. To this end, the study implemented an enfacement illusion with children (N = 64) and adults (N = 33). Participants were exposed to one trial with synchronous interpersonal multisensory stimulation and one trial with asynchronous interpersonal multisensory stimulation-either with a stranger or with the mother as the other person. A self-recognition task using morph videos of self and other and an enfacement questionnaire were implemented as dependent measures. Results revealed evidence for the presence of the enfacement effect in children in both measures. The identity of the other person had a significant effect on the self-recognition task. Contrary to our hypothesis, the effect was significantly smaller in the caregiver condition. No significant differences between children and adults emerged. Our results demonstrate the role of both multisensory stimulation and contextual-here social familiarity-factors for the construction and development of a bodily self. The study provides developmental science with a novel approach to the bodily self by showing the validity of the self-recognition task in a child sample. Overall, the study supports proposals that the sense of bodily self is malleable early in development.

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.

Predicting the bodily self in space and time.

To understand how the human brain distinguishes itself from external stimulation, it was examined if motor predictions enable healthy adult volunteers to infer self-location and to distinguish their body from the environment (and other agents). By uniquely combining a VR-setup with full-body motion capture, a full-body illusion paradigm (FBI) was developed with different levels of motion control: (A) a standard, passive FBI in which they had no motion control; (B) an active FBI in which they made simple, voluntary movements; and (C) an immersive game in which they real-time controlled a human-sized avatar in third person. Systematic comparisons between measures revealed a causal relationship between (i) motion control (prospective agency), (ii) self-other identification, and (iii) the ability to locate oneself. Healthy adults could recognise their movements in a third-person avatar and psychologically align with it (action observation); but did not lose a sense of place (self-location), time (temporal binding), nor who they are (self/other). Instead, motor predictions enabled them to localise their body and to distinguish self from other. In the future, embodied games could target and strengthen the brain's control networks in psychosis and neurodegeneration; real-time motion simulations could help advance neurorehabilitation techniques by fine-tuning and personalising therapeutic settings.

Bodily experiences of trauma and psychosis risk.

Bodily self-disturbances including anomalous embodiment of emotions are observed in psychosis-spectrum conditions. Psychosis is also associated with trauma exposure but the relationship between altered bodily experiences and trauma has not been extensively investigated in individuals at risk for psychosis (HR). We implemented a mapping task to localize felt sensations associated with trauma. Results show that trauma experiences were always localized in the body. HR reported increased rates of traumatic experiences than low-risk group (LR). HR reported sensations associated with trauma across widespread body areas. Further research is needed to elucidate how trauma might lead to psychotic-like experiences via bodily self-disturbances.

Schizophrenia and the bodily self.

Despite the historically consolidated psychopathological perspective, on the one hand, contemporary organicistic psychiatry often highlights abnormalities in neurotransmitter systems like dysregulation of dopamine transmission, neural circuitry, and genetic factors as key contributors to schizophrenia. Neuroscience, on the other, has so far almost entirely neglected the first-person experiential dimension of this syndrome, mainly focusing on high-order cognitive functions, such as executive function, working memory, theory of mind, and the like. An alternative view posits that schizophrenia is a self-disorder characterized by anomalous self-experience and awareness. This view may not only shed new light on the psychopathological features of psychosis but also inspire empirical research targeting the bodily and neurobiological changes underpinning this disorder. Cognitive neuroscience can today address classic topics of phenomenological psychopathology by adding a new level of description, finally enabling the correlation between the first-person experiential aspects of psychiatric diseases and their neurobiological roots. Recent empirical evidence on the neurobiological basis of a minimal notion of the self, the bodily self, is presented. The relationship between the body, its motor potentialities and the notion of minimal self is illustrated. Evidence on the neural mechanisms underpinning the bodily self, its plasticity, and the blurring of self-other distinction in schizophrenic patients is introduced and discussed. It is concluded that brain-body function anomalies of multisensory integration, differential processing of self- and other-related bodily information mediating self-experience, might be at the basis of the disruption of the self disorders characterizing schizophrenia.

Network Structure of Childhood Trauma, Bodily Disturbances, and Schizotypy in Schizophrenia and Nonclinical Controls.

Background and Hypothesis: Exposure to childhood trauma has been linked to the development of psychosis and bodily self-disturbances, 2 hallmarks of schizophrenia (SZ). Prior work demonstrated that bodily disturbances serve as a bridge between childhood trauma and SZ symptomatology, but the diagnostic specificity of these connections remains unknown. This study uses network analysis to bridge this gap by comparing the interplays between childhood trauma, bodily self-disturbances, and schizotypy in clinical and general populations. Study Design: Networks were constructed to examine the relationships between schizotypy (Schizotypal Personality Questionnaire; SPQ), bodily self-disturbances (Perceptual Aberration Scale; PAS), and childhood trauma (Childhood Trauma Questionnaire, CTQ) in 152 people with SZ and 162 healthy comparison participants (HC). The Fused Graphical Lasso was used to jointly estimate the networks in the 2 groups and the structure and strength of the networks were compared. Node centrality and shortest paths between CTQ, PAS, and schizotypy were examined. Study Results: When comparing SZ and HC, the network of bodily self-disturbances, childhood trauma, and schizotypy were similarly structured, but the network was significantly stronger in SZ than HC. In both groups, bodily self-disturbances were on one of the shortest paths between childhood trauma to schizotypal experiences. Conclusions: Our findings revealed reliable associations between childhood trauma, bodily self-disturbance, and schizotypy, with bodily disturbances acting as a bridge from childhood trauma to schizotypy. The elevated strength of the SZ network indicates a more highly interconnected, and therefore reactive network in which exposure to childhood trauma can more easily activate bodily disturbances and schizotypy.

Mindfulness Affects the Boundaries of Bodily Self-Representation: The Effect of Focused-Attention Meditation in Fading the Boundary of Peripersonal Space.

Peripersonal space (PPS) is a dynamic multisensory representation of the space around the body, influenced by internal and external sensory information. The malleability of PPS boundaries, as evidenced by their expansion after tool use or modulation through social interactions, positions PPS as a crucial element in understanding the subjective experiences of self and otherness. Building on the existing literature highlighting both the cognitive and bodily effects of mindfulness meditation, this study proposes a novel approach by employing focused-attention meditation (FAM) and a multisensory audio-tactile task to assess PPS in both the extension and sharpness of its boundaries. The research hypothesis posits that FAM, which emphasizes heightened attention to bodily sensations and interoception, may reduce the extension of PPS and make its boundaries less sharp. We enrolled 26 non-meditators who underwent a repeated measure design in which they completed the PPS task before and after a 15-min FAM induction. We found a significant reduction in the sharpness of PPS boundaries but no significant reduction in PPS extension. These results provide novel insights into the immediate effects of FAM on PPS, potentially shedding light on the modulation of self-other representations in both cognitive and bodily domains. Indeed, our findings could have implications for understanding the intricate relationship between mindfulness practices and the subjective experience of self within spatial contexts.

Interplay between childhood trauma, bodily self-disturbances, and clinical phenomena in schizophrenia spectrum disorders: A network analysis.

BACKGROUND: Bodily self-disturbances have long been considered central to schizophrenia. Exposure to childhood trauma has been linked to the development of both psychosis and bodily self-disturbances, yet little work has examined the role of bodily self-disturbances in the relationship between childhood trauma and schizophrenia symptomatology. This study uses network analysis to bridge this gap. METHODS: Networks were constructed to examine relationships between schizophrenia symptoms (Positive and Negative Symptom Scale; PANSS), bodily self-disturbances (Perceptual Aberration Scale; PAS), and self-reported exposure to childhood trauma (Childhood Trauma Questionnaire, Short-Form; CTQ-SF) in 152 people with a schizophrenia-spectrum disorder. Shortest path and bridge analyses were conducted to assess the role of bodily self-disturbances in linking childhood trauma to schizophrenia symptomatology. Three networks were constructed: 1) PAS, childhood trauma, and PANSS sub-scale composites (positive, negative, general); 2) PAS, childhood trauma, and positive symptoms, 3) PAS, childhood trauma, and distress symptoms. RESULTS: Shortest path analysis revealed that bodily self-disturbances were on the shortest path between childhood trauma and positive and general symptoms (Network 1), between trauma and hallucinations (Network 2), and between trauma and depression (Network 3). Bodily self-disturbances were also found to serve as a bridge between childhood trauma and positive symptoms of schizophrenia, particularly delusions and hallucinations. CONCLUSIONS: Using a novel, data-driven approach, we showed that bodily self-disturbances play a key role in linking childhood trauma to positive and co-morbid affective symptoms of schizophrenia. Threat experiences (i.e., abuse) specifically relate to bodily self-disturbances and psychotic symptoms.

Active self-touch restores bodily proprioceptive spatial awareness following disruption by 'rubber hand illusion'.

Bodily self-awareness relies on a constant integration of visual, tactile, proprioceptive, and motor signals. In the 'rubber hand illusion' (RHI), conflicting visuo-tactile stimuli lead to changes in self-awareness. It remains unclear whether other, somatic signals could compensate for the alterations in self-awareness caused by visual information about the body. Here, we used the RHI in combination with robot-mediated self-touch to systematically investigate the role of tactile, proprioceptive and motor signals in maintaining and restoring bodily self-awareness. Participants moved the handle of a leader robot with their right hand and simultaneously received corresponding tactile feedback on their left hand from a follower robot. This self-touch stimulation was performed either before or after the induction of a classical RHI. Across three experiments, active self-touch delivered after-but not before-the RHI, significantly reduced the proprioceptive drift caused by RHI, supporting a restorative role of active self-touch on bodily self-awareness. The effect was not present during involuntary self-touch. Unimodal control conditions confirmed that both tactile and motor components of self-touch were necessary to restore bodily self-awareness. We hypothesize that active self-touch transiently boosts the precision of proprioceptive representation of the touched body part, thus counteracting the visual capture effects that underlie the RHI.

Beyond peripersonal boundaries: insights from crossmodal interactions.

We experience our self as a body located in space. However, how information about self-location is integrated into multisensory processes underlying the representation of the peripersonal space (PPS), is still unclear. Prior studies showed that the presence of visual information related to oneself modulates the multisensory processes underlying PPS. Here, we used the crossmodal congruency effect (CCE) to test whether this top-down modulation depends on the spatial location of the body-related visual information. Participants responded to tactile events on their bodies while trying to ignore a visual distractor presented on the mirror reflection of their body (Self) either in the peripersonal space (Near) or in the extrapersonal space (Far). We found larger CCE when visual events were presented on the mirror reflection in the peripersonal space, as compared to the extrapersonal space. These results suggest that top-down modulation of the multisensory bodily self is only possible within the PPS.

Brain-inspired bodily self-perception model for robot rubber hand illusion.

The core of bodily self-consciousness involves perceiving ownership of one's body. A central question is how body illusions like the rubber hand illusion (RHI) occur. Existing theoretical models still lack satisfying computational explanations from connectionist perspectives, especially for how the brain encodes body perception and generates illusions from neuronal interactions. Moreover, the integration of disability experiments is also neglected. Here, we integrate biological findings of bodily self-consciousness to propose a brain-inspired bodily self-perception model by which perceptions of bodily self are autonomously constructed without any supervision signals. We successfully validated the model with six RHI experiments and a disability experiment on an iCub humanoid robot and simulated environments. The results show that our model can not only well-replicate the behavioral and neural data of monkeys in biological experiments but also reasonably explain the causes and results of RHI at the neuronal level, thus contributing to the revelation of mechanisms underlying RHI.

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.

Development and validation of the Exteroceptive Body Awareness (EBA-q) questionnaire.

The conscious processing of body signals influences higher-order psychological and cognitive functions, including self-awareness. Dysfunctions in the processing of these signals has been connected to neurological and psychiatric disorders characterized by altered states of self-consciousness. Studies indicate that perceiving the body through interoceptive signals (e.g., from internal organs such as heartbeat and breathing) is distinct from perceiving the body through exteroceptive signals (e.g., by relying on visual, tactile and olfactory cues). While questionnaires are available for assessing interoception, there are no validated self-report instruments for measuring bodily exterception. To fill this gap, we performed three studies to develop and validate a novel scale designed to assess bodily self-consciousness based on the processing of exteroceptive bodily signals. Exploratory factor analysis (Study 1, N = 302) led to an 18-item questionnaire comprised of four factors. We called this instrument Exteroceptive Body Awareness questionnaire (EBA-q). Confirmatory factor analysis (Study 2, N = 184) run on a second sample showed an acceptable fit for a bifactor model, suggesting researchers may use the questionnaire as a unidimensional scale reflecting exteroceptive bodily self-consciousness, or use each of its four sub-scales, reflecting "visuo-tactile body awareness", "spatial coordination", "awareness of body changes" and "awareness of clothing fit". Overall EBA-q showed good internal consistency. Convergent and divergent validity were assessed via cross-validation with existing body awareness questionnaires (Study 3, N = 366) and behavioral measures (Study 3, N = 64) of exteroceptive and interoceptive bodily self-consciousness. Research applications are discussed within a multi-faceted model of exteroception and interoception as distinct, but at the same time interconnected, dimensions of bodily self-consciousness.

Bodily self-recognition and body size overestimation in restrictive anorexia nervosa: implicit and explicit mechanisms.

It is widely known that among others, a pervasive symptom characterizing anorexia nervosa (AN) concerns body image overestimation, which largely contributes to the onset and maintenance of eating disorders. In the present study, we investigated the nature of the body image distortion by recording accuracy and reaction times in both a group of healthy controls and AN patients during two validated tasks requiring an implicit or explicit recognition of self/other hand stimuli, in which the perceived size of the stimuli was manipulated. Our results showed that (1) the perceived size of hand stimuli modulated both the implicit and explicit processing of body parts in both groups; (2) the implicit self-advantage emerged in both groups, but the bodily self, at an explicit level (perceptual, psycho-affective, cognitive) together with the integration and the distinction between self and other, was altered only in restrictive anorexia patients. Although further investigations will be necessary, these findings shed new light on the relationship between the different layers of self-experience and bodily self-disorders.