The prediction-confirmation account of the sense of body ownership: Evidence from a rubber hand illusion paradigm.

We investigated the contribution of multisensory predictions to body ownership, and beyond, to the integration of body-related signals. Contrary to the prevailing idea, according to which, to be integrated, cues necessarily have to be perceived simultaneously, we instead proposed the prediction-confirmation account. According to this account, a perceived cue can be integrated with a predicted cue as long as both signals are relatively simultaneous. To test this hypothesis, a standard rubber hand illusion (RHI) paradigm was used. In the first part of each trial, the illusion was induced while participants observed the rubber hand being touched with a paintbrush. In the subsequent part of the trial, (i) both rubber hand and the participant's real hand were stroked as before (i.e., visible/synchronous condition), (ii) the rubber hand was not stroke anymore (i.e., visible/tactile-only condition), or (iii) both rubber hand and the participant's real hand were synchronously stroked while the location where the rubber hand was touched was occulted (i.e., occulted/synchronous condition). However, in this latter condition, participants still perceived the approaching movement of the paintbrush. Thus, based on this visual cue, the participants can properly predict the timepoint at which the tactile cue should occur (i.e., visuotactile predictions). Our major finding was that compared with the visible/tactile-only condition, the occulted/synchronous condition did not exhibit a decrease of the RHI as in the visible/synchronous condition. This finding supports the prediction-confirmation account and suggests that this mechanism operates even in the standard version of the RHI.

Representations of the relative proportions of body part width.

Despite our wealth of experience with our bodies, our perceptions of our body size are far from veridical. For example, when estimating the relative proportions of their body part lengths, using the hand as a metric, individuals tend to exhibit systematic distortions which vary across body parts. Whilst extensive research with healthy populations has focused on perceptions of body part length, less is known about perceptions of the width of individual body parts and the various components comprising these representations. Across four experiments, representations of the relative proportions of body part width were investigated for both the self and other, and when using both the hand, or a hand-sized stick as the metric. Overall, we found distortions in the perceived width of body parts; however, different patterns of distortions were observed across all experiments. Moreover, the variability across experiments appears not to be moderated by the type of metric used or individuals' posture at the time of estimation. Consequently, findings suggest that, unlike perceptions of body part length, assessed using an identical methodology, our representations of the width of the body parts measured in this task are not fixed and vary across individuals and context. We propose that, as stored width representations of these parts are not necessarily required for navigating our environments, these may not be maintained by our perceptual systems, and thus variable task performance reflects the engagement of idiosyncratic guessing strategies.

Body image at the trunk: An investigation into externally referenced width perception and picture mapping.

Body image is a conscious representation of the body, encompassing how our body feels to us. Body image can be measured in a variety of ways, including metric and depictive measures. This study sought to assess body image at the trunk by investigating, and comparing, a metric and depictive measure. Sixty-nine healthy participants estimated their thorax, waist, and hip width by externally referencing mechanical calipers. Participants were also asked to select the true image of their trunk from a random display of nine images containing the true image and incrementally shrunken or enlarged images. Participants demonstrated evidence of thorax and waist width overestimation in the width perception task, with no evidence for hip misestimation. For the picture mapping task, the majority of participants were inaccurate. In participants who were inaccurate, approximately equal proportions underestimated and overestimated their trunk width. The two tasks were found to be independent of each other. Distortions, or inaccuracies, were apparent in a metric measure, and inaccuracies also present in a depictive measure, of body image at the trunk for healthy participants. An overestimation bias was apparent in the metric, but not depictive, task. No relationship was found between tasks..

Multisensory conflict affects body schema and reaching space.

Multisensory integration plays a crucial role in building the sense of body ownership, i.e., the perceptual status of one's body for which the body is perceived as belonging to oneself. Temporal and spatial mismatching of visual and tactile signals coming from one's body can reduce ownership feelings towards the body and its parts, i.e., produce disownership feelings. Here, we investigated whether visuo-tactile conflict also affects the sensorimotor representation of the body in space (i.e., body schema) and the perception of the space around the body in terms of action potentiality (i.e., reaching space). In two experiments, body schema (Experiment 1) and reaching space (Experiment 2) were assessed before and after either synchronous or asynchronous visuo-tactile stimulation. Results showed that the asynchronous condition, provoking multisensory conflict, caused disownership over one's hand and concurrently affected the body schema and the reaching space. These findings indicate that body schema and reaching space could be dynamically shaped by the multisensory regularities that build up the sense of body ownership.

What I think she thinks about my paralysed body: Social inferences about disability-related content in anosognosia for hemiplegia.

The neuropsychological disorder of anosognosia for hemiplegia (AHP) can offer unique insights into the neurocognitive processes of body consciousness and representation. Previous studies have found associations between selective social cognition deficits and anosognosia. In this study, we examined how such social cognition deficits may directly interact with representations of one's body as disabled in AHP. We used a modified set of previously validated Theory of Mind (ToM) stories to create disability-related content that was related to post-stroke paralysis and to investigate differences between right hemisphere damage patients with (n = 19) and without (n = 19) AHP. We expected AHP patients to perform worse than controls when trying to infer paralysis-related mental states in the paralysis-related ToM stories and explored whether such differences depended on the inference patients were asked to perform (e.g. self or other referent perspective-taking). Using an advanced structural neuroimaging technique, we expected selective social cognitive deficits to be associated with posterior parietal cortex lesions and deficits in self-referent perspective-taking in paralysis-related mentalising to be associated with frontoparietal disconnections. Group- and individual-level results revealed that AHP patients performed worse than HP controls when trying to infer paralysis-related mental states. Exploratory lesion analysis results revealed some of the hypothesised lesions, but also unexpected white matter disconnections in the posterior body and splenium of the corpus collosum associated with a self-referent perspective-taking in paralysis-related ToM stories. The study has implications for the multi-layered nature of body awareness, including abstract, social perspectives and beliefs about the body.

The neural representation of body part concepts.

Neuropsychological and neuroimaging studies provide evidence for a degree of category-related organization of conceptual knowledge in the brain. Some of this evidence indicates that body part concepts are distinctly represented from other categories; yet, the neural correlates and mechanisms underlying these dissociations are unclear. We expand on the limited prior data by measuring functional magnetic resonance imaging responses induced by body part words and performing a series of analyses investigating the cortical representation of this semantic category. Across voxel-level contrasts, pattern classification, representational similarity analysis, and vertex-wise encoding analyses, we find converging evidence that the posterior middle temporal gyrus, the supramarginal gyrus, and the ventral premotor cortex in the left hemisphere play important roles in the preferential representation of this category compared to other concrete objects.

Sense of ownership influence on tactile perception: Is the predictive coding account valid for the somatic rubber hand Illusion?

According to the predictive coding account, the attenuation of tactile perception on the hand exposed to the visuo-tactile Rubber Hand Illusion (vtRHI) relies on a weight increase of visual information deriving from the fake hand and a weight decrease of tactile information deriving from the individual's hand. To explore if this diametrical modulation persists in the absence of vision when adopting the somatic RHI (sRHI), we recorded tactile acuity measures before and after both RHI paradigms in 31 healthy individuals, hypothesizing a weight decrease for somatosensory information deriving from the hand undergoing the illusion and a weight increase for those deriving from the contralateral hand in the sRHI. Our results showed a significant overall decrease in tactile acuity on the hand undergoing the illusion whilst no changes emerged on the contralateral hand during sRHI. Since the sRHI was not accompanied by the hand spatial remapping, despite the generation of the feeling of ownership toward the fake hand, we hypothesized spatial remapping might play a pivotal role in determining sensory information weight attribution.

Alterations of tactile and anatomical spatial representations of the hand after stroke.

Stroke often causes long-term motor and somatosensory impairments. Motor planning and tactile perception rely on spatial body representations. However, the link between altered spatial body representations, motor deficit and tactile spatial coding remains unclear. This study investigates the relationship between motor deficits and alterations of anatomical (body) and tactile spatial representations of the hand in 20 post-stroke patients with upper limb hemiparesis. Anatomical and tactile spatial representations were assessed from 10 targets (nails and knuckles) respectively cued verbally by their anatomical name or using tactile stimulations. Two distance metrics (hand width and finger length) and two structural measures (relative organization of targets positions and angular deviation of fingers from their physical posture) were computed and compared to clinical assessments, normative data and lesions sites. Over half of the patients had altered anatomical and/or tactile spatial representations. Metrics of tactile and anatomical representations showed common variations, where a wider hand representation was linked to more severe motor deficits. In contrast, alterations in structural measures were not concomitantly observed in tactile and anatomical representations and did not correlate with clinical assessments. Finally, a preliminary analysis showed that specific alterations in tactile structural measures were associated with dorsolateral prefrontal stroke lesions. This study reveals shared and distinct characteristics of anatomical and tactile hand spatial representations, reflecting different mechanisms that can be affected differently after stroke: metrics and location of tactile and anatomical representations were partially shared while the structural measures of tactile and anatomical representations had distinct characteristics.

Visuotactile integration in individuals with fibromyalgia.

Our brain constantly integrates afferent information, such as visual and tactile information, to perceive the world around us. According to the maximum-likelihood estimation (MLE) model, imprecise information will be weighted less than precise, making the multisensory percept as precise as possible. Individuals with fibromyalgia (FM), a chronic pain syndrome, show alterations in the integration of tactile information. This could lead to a decrease in their weight in a multisensory percept or a general disruption of multisensory integration, making it less beneficial. To assess multisensory integration, 15 participants with FM and 18 pain-free controls performed a temporal-order judgment task in which they received pairs of sequential visual, tactile (unisensory conditions), or visuotactile (multisensory condition) stimulations on the index and the thumb of the non-dominant hand and had to determine which finger was stimulated first. The task enabled us to measure the precision and accuracy of the percept in each condition. Results indicate an increase in precision in the visuotactile condition compared to the unimodal conditions in controls only, although we found no intergroup differences. The observed visuotactile precision was correlated to the precision predicted by the MLE model in both groups, suggesting an optimal integration. Finally, the weights of the sensory information were not different between the groups; however, in the group with FM, higher pain intensity was associated with smaller tactile weight. This study shows no alterations of the visuotactile integration in individuals with FM, though pain may influence tactile weight in these participants.

Unlocking the potential of 'passive' modulation: How sensory stimulation shapes hand and face size.

Knowledge of the body size is intricately tied to multisensory integration processes that rely on the dynamic interplay of top-down and bottom-up mechanisms. Recent years have seen the development of passive sensory stimulation protocols aimed at investigating the modulation of various cognitive functions, primarily inducing perceptual learning and behaviour change without the need for extensive training. Given that reductions in sensory input have been associated with alterations in body size perception, it is reasonable to hypothesize that increasing sensory information through passive sensory stimulation could similarly influence the perception of the size of body parts. The primary aim of this study was to investigate the potential modulatory effects of passive sensory stimulation on the perception of hand and face size in a group of young adults. Passive sensory stimulation effectively modulated the size representation of the stimulated hand, supporting the notion that access to somatosensory and proprioceptive information is prioritised for the hands but may not extend to the face. Increased somatosensory input resulted in a reduction of distortion, providing evidence for bottom-up modulation of size representation. Passive sensory stimulation can induce subjective changes in body size perception without the need for extensive training. This paradigm holds promise as a potential alternative for modulating distorted size representation in individuals with body representational deficits.

Attachment and body representations in adolescents with personality disorder.

BACKGROUND: Attachment theory has served as an influential framework for understanding psychopathology, partly due to reliable assessment methodology. The influence of insecure attachment on attitudes toward the body and the impact this might have for the development of psychopathology is however less well elucidated. METHOD: A total of 123 adolescents (35 with borderline personality disorder or BPD, 25 with other personality disorders [OPD] and 63 comprising a normative control group) were interviewed with the Adult Attachment Interview (AAI) and the Mirror Interview (MI). The MI questions respondents about how they feel about their bodies, as they look in the mirror. RESULTS: The AAIs from the Borderline group were predominantly insecure-preoccupied and unresolved. These adolescents had significantly lower levels of a positive and integrated sense of self and body than the other groups. Regression results revealed a high loving relationship with fathers, low involving anger with father, high coherence of mind, slight derogation of mother & low levels of unresolved loss uniquely and additively predicted 55% of variance in the summary score assigned to MI responses, that is, the summary score for a Positive and Integrated Body Representation (PIBR). CONCLUSION: Unfavorable attachment experiences and current states of mind regarding attachment may give rise to problems with establishing PIBRs, and thus play a role in the development of psychopathology, especially BPD.

Experiencing an Elongated Limb in Virtual Reality Modifies the Tactile Distance Perception of the Corresponding Real Limb.

In measurement, a reference frame is needed to compare the measured object to something already known. This raises the neuroscientific question of which reference frame is used by humans when exploring the environment. Previous studies suggested that, in touch, the body employed as measuring tool also serves as reference frame. Indeed, an artificial modification of the perceived dimensions of the body changes the tactile perception of external object dimensions. However, it is unknown if such a change in tactile perception would occur when the body schema is modified through the illusion of owning a limb altered in size. Therefore, employing a virtual hand illusion paradigm with an elongated forearm of different lengths, we systematically tested the subjective perception of distance between two points [tactile distance perception (TDP) task] on the corresponding real forearm following the illusion. Thus, the TDP task is used as a proxy to gauge changes in the body schema. Embodiment of the virtual arm was found significantly greater after the synchronous visuotactile stimulation condition compared with the asynchronous one, and the forearm elongation significantly increased the TDP. However, we did not find any link between the visuotactile-induced ownership over the elongated arm and TDP variation, suggesting that vision plays the main role in the modification of the body schema. Additionally, significant effect of elongation found on TDP but not on proprioception suggests that these are affected differently by body schema modifications. These findings confirm the body schema malleability and its role as a reference frame in touch.

Biases in hand perception are driven by somatosensory computations, not a distorted hand model.

To sense and interact with objects in the environment, we effortlessly configure our fingertips at desired locations. It is therefore reasonable to assume that the underlying control mechanisms rely on accurate knowledge about the structure and spatial dimensions of our hand and fingers. This intuition, however, is challenged by years of research showing drastic biases in the perception of finger geometry.1,2,3,4,5 This perceptual bias has been taken as evidence that the brain's internal representation of the body's geometry is distorted,6 leading to an apparent paradox regarding the skillfulness of our actions.7 Here, we propose an alternative explanation of the biases in hand perception-they are the result of the Bayesian integration of noisy, but unbiased, somatosensory signals about finger geometry and posture. To address this hypothesis, we combined Bayesian reverse engineering with behavioral experimentation on joint and fingertip localization of the index finger. We modeled the Bayesian integration either in sensory or in space-based coordinates, showing that the latter model variant led to biases in finger perception despite accurate representation of finger length. Behavioral measures of joint and fingertip localization responses showed similar biases, which were well fitted by the space-based, but not the sensory-based, model variant. The space-based model variant also outperformed a distorted hand model with built-in geometric biases. In total, our results suggest that perceptual distortions of finger geometry do not reflect a distorted hand model but originate from near-optimal Bayesian inference on somatosensory signals.

The role of visuomotor synchrony on virtual full-body illusions in children and adults.

The present study explored the effects of visuomotor synchrony in virtual reality during the embodiment of a full human avatar in children (aged 5-6 years) and adults. Participants viewed their virtual bodies from a first-person perspective while they moved the body during self-generated and structured movement. Embodiment was measured via questions and psychophysiological responses (skin conductance) to a virtual body-threat and during both movement conditions. Both children and adults had increased feelings of ownership and agency over a virtual body during synchronous visuomotor feedback (compared to asynchronous visuomotor feedback). Children had greater ownership compared to adults during synchronous movement but did not differ from adults on agency. There were no differences in SCRs (frequency or magnitude) between children and adults, between conditions (i.e., baseline or movement conditions) or visuomotor feedback. Collectively, the study highlights the importance of visuomotor synchrony for children's ratings of embodiment for a virtual avatar from at least 5 years old, and suggests adults and children are comparable in terms of psychophysiological arousal when moving (or receiving a threat to) a virtual body. This has important implications for our understanding of the development of embodied cognition and highlights the considerable promise of exploring visuomotor VR experiences in children.

The multidimensional assessment of body representation and interoception in multiple sclerosis.

BACKGROUND: The mental representation of the body (or body representation, BR) derives from the processing of multiple sensory and motor inputs and plays a crucial role in guiding our actions and in how we perceive our body. Fundamental inputs for BR construction come also from the interoceptive systems which refer to the whole bidirectional processes between the brain and the body. People with Multiple sclerosis (MS) show an abnormal multisensory integration which may compromise BR and interoception integrity. However, no study has evaluated possible deficits on distinct and dissociable dimensions of body representation (i.e., action-oriented, aBR; and a nonaction-oriented body representation, NaBR) and interoception (i.e., interoceptive accuracy, interoceptive sensibility, and interoceptive awareness) in MS. OBJECTIVE: In the present study, we aimed to determine whether participants with MS present changes in BR and interoceptive dimensions. METHODS: We performed comparison analyses on tasks and questionnaires tapping all BR and interoceptive dimensions between 36 people with relapsing-remitting MS (RRMS) and 42 healthy controls, and between 23 people with progressive MS (PMS) and 33 healthy controls. RESULTS: Overall, patients with MS exhibited lower interoceptive accuracy than matched controls. The RRMS group also showed higher visceral interoceptive sensibility levels. No differences were found in BR accuracy measures, but the PMS reported longer response times when performing the aBR task. CONCLUSION: These findings open a new issue on the role of inner-signal monitoring in the body symptomatology of MS and highlight the need for an accurate BR and interoceptive assessment in a clinical setting.

The characteristics of the implicit body model of the trunk.

Knowing where the body is in space requires reference to a stored model of the size and shape of body parts, termed the body model. This study sought to investigate the characteristics of the implicit body model of the trunk by assessing the position sense of midline and lateral body landmarks. Sixty-nine healthy participants localised midline and lateral body landmarks on their thorax, waist and hips, with perceived positions of these landmarks compared to actual positions. This study demonstrates evidence of a significant distortion of the implicit body model of the trunk, presenting as a squatter trunk, wider at the waist and hips. A significant difference was found between perceived and actual location in the horizontal (x) and vertical (y) directions for the majority of trunk landmarks. Evidence of a rightward bias was noted in the perception of six of the nine body landmarks in the horizontal (x) direction, including all midline levels. In the vertical (y) direction, a substantial inferior bias was evident at the thorax and waist. The implicit body model of the trunk is shown to be distorted, with the lumbar spine (waist-to-hip region) held to be shorter and wider than reality.

Body representation after a stroke in the brainstem.

BACKGROUND: Lesion occurring in the brainstem may cause a postural tilt and balance disorders, which could be due to an inaccurate perception of the body orientation. The objective of this study was to determine the effects of a brainstem stroke on body representation in horizontal and frontal plane, and links with impaired posture and neuroanatomy. METHODS: Forty patients with stroke in left brainstem (L-BS) or right (R-BS) were compared with 15 matched control subjects (C). The subjective straight-ahead (SSA) was investigated using a method disentangling lateral deviation and tilt components of error. RESULTS: The L-BS patients had contralesional lateral deviation of SSA. In addition, they showed an ipsilesional tilt, more severe for the trunk than for the head. By contrast, in R-BS patients, the representation of the body midline was fairly accurate in both the horizontal and frontal planes and did not differ from that of control subjects. CONCLUSION: This work highlights an asymmetry of representation of body associated with left brainstem lesions extending to the right cerebral hemisphere. This deviation appears only after a left lesion, which may point to a vestibular dominance. These results open a new perspective of neuro-rehabilitation of postural disorders after a stroke, with the correction of the representation of body orientation.

Balancing the Senses: Electrophysiological Responses Reveal the Interplay between Somatosensory and Visual Processing During Body-Related Multisensory Conflict.

In the study of bodily awareness, the predictive coding theory has revealed that our brain continuously modulates sensory experiences to integrate them into a unitary body representation. Indeed, during multisensory illusions (e.g., the rubber hand illusion, RHI), the synchronous stroking of the participant's concealed hand and a fake visible one creates a visuotactile conflict, generating a prediction error. Within the predictive coding framework, through sensory processing modulation, prediction errors are solved, inducing participants to feel as if touches originated from the fake hand, thus ascribing the fake hand to their own body. Here, we aimed to address sensory processing modulation under multisensory conflict, by disentangling somatosensory and visual stimuli processing that are intrinsically associated during the illusion induction. To this aim, we designed two EEG experiments, in which somatosensory- (SEPs; Experiment 1; N = 18; F = 10) and visual-evoked potentials (VEPs; Experiment 2; N = 18; F = 9) were recorded in human males and females following the RHI. Our results show that, in both experiments, ERP amplitude is significantly modulated in the illusion as compared with both control and baseline conditions, with a modality-dependent diametrical pattern showing decreased SEP amplitude and increased VEP amplitude. Importantly, both somatosensory and visual modulations occur in long-latency time windows previously associated with tactile and visual awareness, thus explaining the illusion of perceiving touch at the sight location. In conclusion, we describe a diametrical modulation of somatosensory and visual processing as the neural mechanism that allows maintaining a stable body representation, by restoring visuotactile congruency under the occurrence of multisensory conflicts.

Tactile localization accuracy at the low back.

Localizing tactile stimulation is an important capability for everyday function and may be impaired in people with persistent pain. This study sought to provide a detailed description of lumbar spine tactile localization accuracy in healthy individuals. Sixty-nine healthy participants estimated where they were touched at nine different points, labelled in a 3 × 3 grid over the lumbar spine. Mislocalization between the perceived and actual stimulus was calculated in horizontal (x) and vertical (y) directions, and a derived hypotenuse (c) mislocalization was calculated to represent the direct distance between perceived and actual points. In the horizontal direction, midline sites had the smallest mislocalization. Participants exhibited greater mislocalization for left- and right-sided sites, perceiving sites more laterally than they actually were. For all vertical values, stimulated sites were perceived lower than reality. A greater inaccuracy was observed in the vertical direction. This study measured tactile localization for the low back utilizing a novel testing method. The large inaccuracies point to a possible distortion in the underlying perceptual maps informing the superficial schema; however, further testing comparing this novel method with an established tactile localization task, such as the point-to-point method, is suggested to confirm these findings.

The influence of hand posture on tactile processing: Evidence from a 7T functional magnetic resonance imaging study.

Although behavioral evidence has shown that postural changes influence the ability to localize or detect tactile stimuli, little is known regarding the brain areas that modulate these effects. This 7T functional magnetic resonance imaging (fMRI) study explores the effects of touch of the hand as a function of hand location (right or left side of the body) and hand configuration (open or closed). We predicted that changes in hand configuration would be represented in contralateral primary somatosensory cortex (S1) and the anterior intraparietal area (aIPS), whereas change in position of the hand would be associated with alterations in activation in the superior parietal lobule. Multivoxel pattern analysis and a region of interest approach partially supported our predictions. Decoding accuracy for hand location was above chance level in superior parietal lobule (SPL) and in the anterior intraparietal (aIPS) area; above chance classification of hand configuration was observed in SPL and S1. This evidence confirmed the role of the parietal cortex in postural effects on touch and the possible role of S1 in coding the body form representation of the hand.