Subcortical contributions to the sense of body ownership.

The sense of body ownership (i.e. the feeling that our body or its parts belong to us) plays a key role in bodily self-consciousness and is believed to stem from multisensory integration. Experimental paradigms such as the rubber hand illusion have been developed to allow the controlled manipulation of body ownership in laboratory settings, providing effective tools for investigating malleability in the sense of body ownership and the boundaries that distinguish self from other. Neuroimaging studies of body ownership converge on the involvement of several cortical regions, including the premotor cortex and posterior parietal cortex. However, relatively less attention has been paid to subcortical structures that may also contribute to body ownership perception, such as the cerebellum and putamen. Here, on the basis of neuroimaging and neuropsychological observations, we provide an overview of relevant subcortical regions and consider their potential role in generating and maintaining a sense of ownership over the body. We also suggest novel avenues for future research targeting the role of subcortical regions in making sense of the body as our own.

Modeling affective touch pleasantness across skin types at the individual level reveals a reliable and stable basic function.

Touch is perceived most pleasant when delivered at velocities known to optimally activate the C-tactile afferent system. At the group level, pleasantness ratings of touch delivered at velocities in the range between 0.3 and 30 cm/s follow an inverted-U shape curve, with maximum pleasantness between 1 and 10 cm/s. However, the prevalence, reliability, and stability of this function at the individual level and across skin types based on hair density remains unknown. Here, we tested a range of seven velocities (0.3, 1, 3, 6, 9, 18, and 27 cm/s) delivered with a soft brush, on both hairy (forearm and dorsal hand) and nonhairy skin (palm) in 123 participants. Our results suggest that the relationship between pleasantness and velocity of touch is significantly best described by a negative quadratic model at the individual level in the majority of participants both on hairy (67.1%) and nonhairy (62.6%) skin, a larger extent than previously reported. Higher interoceptive accuracy and self-reported depression were related to a better fit of the quadratic model and the steepness of the curve, respectively. The prevalence of the quadratic model at the individual level was stable across body sites (62.6%, experiment 1), across two experimental sessions (73%-78%, experiment 2), and regardless of the number of repetitions of each velocity (experiment 3). Thus, the individual perception of tactile pleasantness follows a characteristic velocity-dependent function across skin types and shows trait characteristics. Future studies can investigate further the possibility to use affective touch as a behavioral biomarker for mental health disorders.NEW & NOTEWORTHY Touch is perceived as most pleasant when delivered at slow, caress-like velocities, known to activate C-tactile afferents. At the group level, tactile pleasantness and velocity of touch show a reliable pattern of relationship on hairy skin. Here, we found that the perception of tactile pleasantness follows a consistent pattern also at the individual level, across skin types and testing sessions. However, individual differences in interoceptive abilities and self-reported depression do play a role.

The role of affective touch in whole-body embodiment remains equivocal.

Previous studies have highlighted that affective touch delivered at slow velocities (1-10 cm/s) enhances body-part embodiment during multisensory illusions, yet its role towards whole-body embodiment is less established. Across two experiments, we investigated the role of affective touch towards subjective embodiment of a whole mannequin body within the full body illusion, amongst healthy females. Participants perceived affective touch to be more pleasant than non-affective touch, but this did not enhance subjective embodiment within the illusion and no interaction between synchrony (Experiment 1), or congruency (Experiment 2), and the velocity of touch was observed. Finally, the perceived pleasantness of touch was not modulated by subthreshold eating disorder psychopathology, as measured by means of a self-report questionnaire. Therefore, the present findings suggest that enhancement of embodiment due to affective touch may be body-part specific, and not generalise to greater ownership towards a whole body.

Embodied Precision: Intranasal Oxytocin Modulates Multisensory Integration.

Multisensory integration processes are fundamental to our sense of self as embodied beings. Bodily illusions, such as the rubber hand illusion (RHI) and the size-weight illusion (SWI), allow us to investigate how the brain resolves conflicting multisensory evidence during perceptual inference in relation to different facets of body representation. In the RHI, synchronous tactile stimulation of a participant's hidden hand and a visible rubber hand creates illusory body ownership; in the SWI, the perceived size of the body can modulate the estimated weight of external objects. According to Bayesian models, such illusions arise as an attempt to explain the causes of multisensory perception and may reflect the attenuation of somatosensory precision, which is required to resolve perceptual hypotheses about conflicting multisensory input. Recent hypotheses propose that the precision of sensorimotor representations is determined by modulators of synaptic gain, like dopamine, acetylcholine, and oxytocin. However, these neuromodulatory hypotheses have not been tested in the context of embodied multisensory integration. The present, double-blind, placebo-controlled, crossover study ( n = 41 healthy volunteers) aimed to investigate the effect of intranasal oxytocin (IN-OT) on multisensory integration processes, tested by means of the RHI and the SWI. Results showed that IN-OT enhanced the subjective feeling of ownership in the RHI, only when synchronous tactile stimulation was involved. Furthermore, IN-OT increased an embodied version of the SWI (quantified as estimation error during a weight estimation task). These findings suggest that oxytocin might modulate processes of visuotactile multisensory integration by increasing the precision of top-down signals against bottom-up sensory input.

If I were a grown-up: Children's response to the rubber hand illusion with different hand sizes.

During childhood, children's bodies undergo rapid physical growth, which may affect their ability to accurately perceive their own bodies as well as the external environment. Concurrently, multisensory processes underlying bodily self-consciousness gradually develop. However, little is known of the relation between changes in body size and corresponding update in bodily self-consciousness. In this study, 6- to 8-year-old children experienced the "rubber hand illusion" while watching a regular (child-like) or larger (adult-like) rubber hand being touched either synchronously or asynchronously with their own real hand. We measured proprioceptive drift and subjective ownership as well as changes in weight estimation of objects. Synchronous versus asynchronous visuotactile stroking with both the regular and larger hands caused a change in subjective body ownership but did not make a difference in perceived hand location and weight estimation of objects having the same size but different weight. Intriguingly, simply viewing the regular hand led to greater proprioceptive drift and to overestimation of the objects' weight relative to the larger hand. In addition, weight estimation was also influenced by hand size after visuotactile stroking. These results demonstrate that visuotactile information modulates the intrinsic properties of body representation in terms of size in children and further corroborate previous findings suggesting late developing optimal calibration of visual and proprioceptive signals.

Sense of body ownership in patients affected by functional motor symptoms (conversion disorder).

BACKGROUND AND AIM: Patients with functional neurological symptoms are commonly seen in neurological practice. Nevertheless their aetiopathology remains unclear. We have recently shown that patients affected by functional motor symptoms (FMS) present lower interoceptive awareness and higher alexithymia levels than healthy controls. Nevertheless sense of body ownership has never been studied in FMS patients. The aim of the present study was to systematically investigate the sense of body ownership, with the rubber hand illusion (RHI) paradigm, in patients with FMS and healthy controls. MATERIALS AND METHODS: We included in the study 16 patients with FMS and 18 healthy controls (HC). Patients and HC were asked to complete the Toronto Alexithymia Scale (TAS-20) and the self-consciousness scale (self-objectification questionnaire). All participants underwent the RHI paradigm: illusionary experience was measured by self-report and by proprioceptive alteration. RESULTS: A Mann-Whitney U test performed revealed that FMS (median=2.11) participants embodied the rubber hand to the same extent than HC participants (median=2.0, Z = -0.86, p>0.05, r = -0.15). The same test revealed no significant difference in the Proprioceptive Drift experience between FMS (median=0.0) and HC participants (median = -0.5, Z = -0.96, p>0.05, r = -0.16). CONCLUSIONS: Our study revealed that sense of body ownership is not impaired in patients affected by FMS. This, together with the results from our previous experiment (studying the interoceptive awareness), supports the hypothesis that interoceptive awareness and sense of body ownership may be dissociated in patients with FMS.

Thermosensation and emotion: Thermosensory accuracy in a dynamic thermal matching task is linked to depression and anxiety symptomatology.

Interoception is related to the generation of bodily feelings and the awareness of ourselves as 'sentient beings', informing the organism about its bodily needs to guarantee survival. Previous studies have reported links among interoception, emotion processing, and mental health. For example, the alignment of interoceptive dimensions (i.e., accuracy, sensibility, awareness) can predict emotional symptoms, such as anxiety. Here, we aimed to investigate the relationship between the perception of a certain type of skin-mediated interoceptive signal, i.e., thermosensation, and self-reported depression, anxiety, and stress. One hundred seventy participants completed the Depression Anxiety Stress Scale (DASS-21) and a dynamic thermal matching task, a static temperature detection task, and a heartbeat counting task. Our results revealed that self-reported anxiety and depression were related to the perception of temperature on hairy and non-hairy skin, respectively: higher anxiety was related to better performance on the thermal matching task on the forearm, while higher depression was related to poorer performance on dynamic and static temperature tasks on the palm. Discrepancies between thermosensory accuracy and sensibility measures ('trait prediction error') were related to heightened anxiety, in line with previous studies. No significant correlations were found between DASS-21 scores and heartbeat counting accuracy. In conclusion, this study suggests that individual differences in thermosensory perception in different areas of the body are associated with self-reported anxiety and depression.

Updating Prospective Self-Efficacy Beliefs About Cardiac Interoception in Anorexia Nervosa: An Experimental and Computational Study.

Patients with anorexia nervosa (AN) typically hold altered beliefs about their body that they struggle to update, including global, prospective beliefs about their ability to know and regulate their body and particularly their interoceptive states. While clinical questionnaire studies have provided ample evidence on the role of such beliefs in the onset, maintenance, and treatment of AN, psychophysical studies have typically focused on perceptual and 'local' beliefs. Across two experiments, we examined how women at the acute AN (N = 86) and post-acute AN state (N = 87), compared to matched healthy controls (N = 180) formed and updated their self-efficacy beliefs retrospectively (Experiment 1) and prospectively (Experiment 2) about their heartbeat counting abilities in an adapted heartbeat counting task. As preregistered, while AN patients did not differ from controls in interoceptive accuracy per se, they hold and maintain 'pessimistic' interoceptive, metacognitive self-efficacy beliefs after performance. Modelling using a simplified computational Bayesian learning framework showed that neither local evidence from performance, nor retrospective beliefs following that performance (that themselves were suboptimally updated) seem to be sufficient to counter and update pessimistic, self-efficacy beliefs in AN. AN patients showed lower learning rates than controls, revealing a tendency to base their posterior beliefs more on prior beliefs rather than prediction errors in both retrospective and prospective belief updating. Further explorations showed that while these differences in both explicit beliefs, and the latent mechanisms of belief updating, were not explained by general cognitive flexibility differences, they were explained by negative mood comorbidity, even after the acute stage of illness.

The Role of the Skin in Interoception: A Neglected Organ?

In the past 2 decades, interoception has received increasing attention in the fields of psychology and cognitive science, as well as neuroscience and physiology. A plethora of studies adopted the perception of cardiac signals as a proxy for interoception. However, recent findings have cast doubt on the methodological and intrinsic validity of the tasks used thus far. Therefore, there is an ongoing effort to improve the existing cardiac interoceptive tasks and to identify novel channels to target the perception of the physiological state of the body. Amid such scientific abundancy, one could question whether the field has been partially neglecting one of our widest organs in terms of dimensions and functions: the skin. According to some views grounded on anatomical and physiological evidence, skin-mediated signals such as affective touch, pain, and temperature have been redefined as interoceptive. However, there is no agreement in this regard. Here, we discuss some of the anatomical, physiological, and experimental arguments supporting the scientific study of interoception by means of skin-mediated signals. We argue that more attention should be paid to the skin as a sensory organ that monitors the bodily physiological state and further propose thermosensation as a particularly attractive model of skin-mediated interoception.

Changes in self-other boundaries modulate children's body image attitudes.

One's own face is a key distinctive feature of our physical appearance, yet multisensory visuo-tactile stimulation can alter self-other boundaries, eliciting changes in adult's self-face representation and social cognition processes. This study tested whether changing self-face representation by altering self-other boundaries with the enfacement illusion modulates body image attitudes toward others in 6-11-year-old children (N = 51; 31 girls; predominantly White). Across all ages, congruent multisensory information led to stronger enfacement (η2p = 0.06). Participants who experienced a stronger enfacement illusion showed preference for larger body size, suggesting increased positive body size attitudes. This effect was stronger in 6-7-year-olds compared to 8-9-year-olds. Thus, blurring self-other boundaries successfully modulates self-face representation and body image attitudes toward others in children. Our results suggest that increased self-resemblance through self-other blurring resulting from the enfacement illusion may reduce social comparisons between self and other and result in positive body size attitudes.

The perception of affective and discriminative touch in blind individuals.

Enhanced tactile acuity in blindness is among the most widely reported results of neuroplasticity following prolonged visual deprivation. However, tactile submodalities other than discriminative touch are profoundly understudied in blind individuals. Here, we examined the influence of blindness on two tactile submodalities, affective and discriminative touch, the former being vital for social functioning and emotional processing. We tested 36 blind individuals and 36 age- and sex-matched sighted volunteers. In Experiment 1, we measured the perception of affective tactile signals by asking participants to rate the pleasantness of touch delivered on the palm (nonhairy skin, sparsely innervated with C tactile [CT] fibers) or the forearm (hairy skin, densely innervated with CT fibers) in a CT-optimal versus a CT-nonoptimal manner using a paradigm grounded in studies on tactile sensory neurophysiology. In Experiment 2, we implemented a classic task assessing discriminative touch abilities, the grating orientation task. We found that blind individuals rated the touch as more pleasant when delivered on the palm than on the forearm, while the opposite pattern was observed for sighted participants, who rated stimulation on the forearm as more pleasant than stimulation on the palm. We also replicated the previous findings showing enhanced discriminative tactile acuity in blind individuals. Altogether, our results suggest that blind individuals might experience affective touch differently than sighted individuals, with relatively greater pleasantness perceived on the palm. These results provide a broader insight into somatosensory perception in blind individuals, for the first time taking into consideration the socioemotional aspect of touch.

Heartbeat counting accuracy is enhanced in blind individuals.

Blind individuals have superior abilities to perform perceptual tasks that rely on exteroceptive information, since visual deprivation is associated with heightened cross-modal plasticity. However, it is unknown whether neuroplasticity after visual loss also affects interoception, that is, the sensations arising from one's inner organs that convey information about the physiological state of the body. Herein, we examine the influence of blindness on cardiac interoception, which is an interoceptive submodality that has important links to emotional processing and bodily self-awareness. We tested 36 blind and 36 age- and sex-matched sighted volunteers and examined their cardiac interoceptive ability using the heartbeat counting task. The results showed that blind individuals had significantly higher accuracy in perceiving their heartbeat than did individuals in a matched sighted control group. In contrast, there were no significant differences between the groups in the metacognitive dimensions of cardiac interoception or the purely physiological measurement of heart rate, thereby underscoring that the improved accuracy likely reflects a superior perceptual sensitivity to cardiac interoceptive signals in blind individuals. We conclude that visual deprivation leads to an enhanced ability to count one's own heartbeats, which has important implications for the study of the extent of cross-modal plasticity after visual loss, understanding emotional processing in blind individuals, and learning how bodily self-awareness can develop and be sustained in the absence of visual experience. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Exploring multisensory integration of non-naturalistic sounds on body perception in young females with eating disorders symptomatology: a study protocol.

BACKGROUND: Bodily illusions can be used to investigate the experience of being in a body by manipulating the underlying processes of multisensory integration. Research suggests that people with eating disorders (EDs) may have impairments in visual, interoceptive, proprioceptive, and tactile bodily perception. Furthermore, people with EDs also show abnormalities in integrating multisensory visuo-tactile and visual-auditory signals related to the body, which may contribute to the development of body image disturbances. Visuo-auditory integration abnormalities have been observed also in people with subthreshold ED symptomatology. However, it remains unclear whether these impairments are specific to bodily signals or if they extend to any auditory signals. METHODS: We will recruit 50 participants (aged 18-24; females assigned at birth) with ED symptomatology (subthreshold group) and 50 control participants. The Eating Disorder Examination Questionnaire will be administered to screen for ED symptomatology and divide the sample into two groups accordingly (control and subthreshold group using a clinical cut-off score of 2.8). The strength of both illusions will be measured implicitly with estimations of body part position and size, and explicitly with self-report questionnaires. As a secondary aim, regression analysis will be run to test the predictive role of susceptibility for both illusions on interoceptive body awareness (measured by the Multidimensional Assessment of Interoceptive Awareness Scale) and sensory-processing sensitivity (measured by the Highly Sensitive Person Scale). DISCUSSION: Our study may contribute to our understanding of the mechanisms underlying body image disturbances. The results may pave the way for novel clinical interventions targeting early symptoms prior to the development of the disorder in young females.

This study aims to investigate whether individuals with subthreshold eating disorder (ED) symptoms show differences in the integration of proprioceptive and auditory cues in relation to body representation, as compared to healthy controls. This will be examined by focusing on two different body parts (finger and waist) with different levels of emotional significance for individuals with EDs. Participants will be screened for ED symptoms and divided into groups accordingly. The strength of the illusions will be measured using estimations of body part position and size, as well as self-report questionnaires. Regression analysis will be used to assess the predictive role of susceptibility to the body illusions on interoceptive body awareness and sensory-processing sensitivity. The results of this study may have implications for understanding the underlying mechanisms of body image disturbances in EDs, and inform the development of clinical interventions for individuals with subthreshold EDs.

Visuo-thermal congruency modulates the sense of body ownership.

Thermosensation has been redefined as an interoceptive modality that provides information about the homeostatic state of the body. However, the contribution of thermosensory signals to the sense of body ownership remains unclear. Across two rubber hand illusion (RHI) experiments (N = 73), we manipulated the visuo-thermal congruency between the felt and seen temperature, on the real and rubber hand respectively. We measured the subjectively experienced RHI, the perceived hand location and temperature of touch, and monitored skin temperature. We found that visuo-thermal incongruencies between the seen and felt touch reduced the subjective and behavioural RHI experience (Experiment 1). Visuo-thermal incongruencies also gave rise to a visuo-thermal illusion effect, but only when the rubber hand was placed in a plausible position (Experiment 2) and when considering individual differences in interoceptive sensibility. Thus, thermosensation contributes to the sense of body ownership by a mechanism of dynamic integration of visual and thermosensory signals.

Interoception as independent cardiac, thermosensory, nociceptive, and affective touch perceptual submodalities.

Interoception includes signals from inner organs and thin afferents in the skin, providing information about the body's physiological state. However, the functional relationships between interoceptive submodalities are unclear, and thermosensation as skin-based interoception has rarely been considered. We used five tasks to examine the relationships among cardiac awareness, thermosensation, affective touch, and nociception. Thermosensation was probed with a classic temperature detection task and the new dynamic thermal matching task, where participants matched perceived moving thermal stimuli in a range of colder/warmer stimuli around thermoneutrality. We also examined differences between hairy and non-hairy skin and found superior perception of dynamic temperature and static cooling on hairy skin. Notably, no significant correlations were observed across interoceptive submodality accuracies (except for cold and pain perception in the palm), which indicates that interoception at perceptual levels should be conceptualised as a set of relatively independent processes and abilities rather than a single construct.

Limits of cross-modal plasticity? Short-term visual deprivation does not enhance cardiac interoception, thermosensation, or tactile spatial acuity.

In the present study, we investigated the effect of short-term visual deprivation on discriminative touch, cardiac interoception, and thermosensation by asking 64 healthy volunteers to perform four behavioral tasks. The experimental group contained 32 subjects who were blindfolded and kept in complete darkness for 110 min, while the control group consisted of 32 volunteers who were not blindfolded but were otherwise kept under identical experimental conditions. Both groups performed the required tasks three times: before and directly after deprivation (or control) and after an additional washout period of 40 min, in which all participants were exposed to normal light conditions. Our results showed that short-term visual deprivation had no effect on any of the senses tested. This finding suggests that short-term visual deprivation does not modulate basic bodily senses and extends this principle beyond tactile processing to the interoceptive modalities of cardiac and thermal sensations.

Identifying with the beautiful: Facial attractiveness effects on unisensory and multisensory self-other distinction.

People tend to evaluate their own traits and abilities favourably and such favourable self-perceptions extend to attractiveness. However, the exact mechanism underlying this self-enhancement bias remains unclear. One possibility could be the identification with attractive others through blurring of self-other boundaries. Across two experiments, we used the enfacement illusion to investigate the effect of others' attractiveness in the multisensory perception of the self. In Experiment 1 (N = 35), participants received synchronous or asynchronous interpersonal visuo-tactile stimulation with an attractive and non-attractive face. In Experiment 2 (N = 35), two new faces were used and spatial incongruency was introduced as a control condition. The results showed that increased ratings of attractiveness of an unfamiliar face lead to blurring of self-other boundaries, allowing the identification of our psychological self with another's physical self and specifically their face, and this seems to be unrelated to perceived own attractiveness. The effect of facial attractiveness on face ownership showed dissociable mechanisms, with multisensory integration modulating the effect on similarity but not identification, an effect that may be purely based on vision. Overall, our findings suggest that others' attractiveness may lead to positive distortions of the self. This research provides a psychophysical starting point for studying the impact of others' attractiveness on self-face recognition, which can be particularly important for individuals with malleable, embodied self-other boundaries and body image disturbances.

Body weight distortions in an auditory-driven body illusion in subclinical and clinical eating disorders.

Previous studies suggest a stronger influence of visual signals on body image in individuals with eating disorders (EDs) than healthy controls; however, the influence of other exteroceptive sensory signals remains unclear. Here we used an illusion relying on auditory (exteroceptive) signals to manipulate body size/weight perceptions and investigated whether the mechanisms integrating sensory signals into body image are altered in subclinical and clinical EDs. Participants' footstep sounds were altered to seem produced by lighter or heavier bodies. Across two experiments, we tested healthy women assigned to three groups based on self-reported Symptomatology of EDs (SED), and women with Anorexia Nervosa (AN), and used self-report, body-visualization, and behavioural (gait) measures. As with visual bodily illusions, we predicted stronger influence of auditory signals, leading to an enhanced body-weight illusion, in people with High-SED and AN. Unexpectedly, High-SED and AN participants displayed a gait typical of heavier bodies and a widest/heaviest visualized body in the 'light' footsteps condition. In contrast, Low-SED participants showed these patterns in the 'heavy' footsteps condition. Self-reports did not show group differences. The results of this pilot study suggest disturbances in the sensory integration mechanisms, rather than purely visually-driven body distortions, in subclinical/clinical EDs, opening opportunities for the development of novel diagnostic/therapeutic tools.

The Anticipation and Perception of Affective Touch in Women with and Recovered from Anorexia Nervosa.

Disruptions in reward processing and anhedonia have long been observed in Anorexia Nervosa (AN). Interoceptive deficits have also been observed in AN, including reduced tactile pleasure. However, the extent to which this tactile anhedonia is specifically liked to an impairment in a specialised, interoceptive C-tactile system originating at the periphery, or a more top-down mechanism in the processing of tactile pleasantness remains debated. Here, we investigated differences between patients with and recovered from AN (RAN) and healthy controls (HC) in the perception of pleasantness of touch delivered in a CT-optimal versus a CT-non-optimal manner, and in their top-down, anticipatory beliefs about the perceived pleasantness of touch. To this end, we measured the anticipated pleasantness of various materials touching the skin and the perceived pleasantness of light, dynamic touch applied to the forearm of 27 women with AN, 24 women who have recovered and 30 HCs using C Tactile (CT) afferents-optimal (slow) and non-optimal (fast) velocities. Our results showed that both clinical groups anticipated tactile experiences and rated delivered tactile stimuli as less pleasant than HCs, but the latter difference was not related to the CT optimality of the stimulation. Instead, differences in the perception of CT-optimal touch were predicted by differences in top-down beliefs, alexithymia and interoceptive sensibility. Thus, tactile anhedonia in AN might persist as a trait even after otherwise successful recovery of AN and it is not linked to a bottom-up interoceptive deficit in the CT system, but rather to a learned, defective top-down anticipation of tactile pleasantness.