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.

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.

Multiple representations of the body schema for the same body part.

Purposeful motor actions depend on the brain's representation of the body, called the body schema, and disorders of the body schema have been reported to show motor deficits. The body schema has been assumed for almost a century to be a common body representation supporting all types of motor actions, and previous studies have considered only a single motor action. Although we often execute multiple motor actions, how the body schema operates during such actions is unknown. To address this issue, I developed a technique to measure the body schema during multiple motor actions. Participants made simultaneous eye and reach movements to the same location of 10 landmarks on their hand. By analyzing the internal configuration of the locations of these points for each of the eye and reach movements, I produced maps of the mental representation of hand shape. Despite these two movements being simultaneously directed to the same bodily location, the resulting hand map (i.e., a part of the body schema) was much more distorted for reach movements than for eye movements. Furthermore, the weighting of visual and proprioceptive bodily cues to build up this part of the body schema differed for each effector. These results demonstrate that the body schema is organized as multiple effector-specific body representations. I propose that the choice of effector toward one's body can determine which body representation in the brain is observed and that this visualization approach may offer a new way to understand patients' body schema.

Morphology, Connectivity, and Encoding Features of Tactile and Motor Representations of the Fingers in the Human Precentral and Postcentral Gyrus.

Despite the tight coupling between sensory and motor processing for fine manipulation in humans, it is not yet totally clear which specific properties of the fingers are mapped in the precentral and postcentral gyrus. We used fMRI to compare the morphology, connectivity, and encoding of the motor and tactile finger representations (FRs) in the precentral and postcentral gyrus of 25 5-fingered participants (8 females). Multivoxel pattern and structural and functional connectivity analyses demonstrated the existence of distinct motor and tactile FRs within both the precentral and postcentral gyrus, integrating finger-specific motor and tactile information. Using representational similarity analysis, we found that the motor and tactile FRs in the sensorimotor cortex were described by the perceived structure of the hand better than by the actual hand anatomy or other functional models (finger kinematics, muscles synergies). We then studied a polydactyly individual (i.e., with a congenital 6-fingered hand) showing superior manipulation abilities and divergent anatomic-functional hand properties. The perceived hand model was still the best model for tactile representations in the precentral and postcentral gyrus, while finger kinematics better described motor representations in the precentral gyrus. We suggest that, under normal conditions (i.e., in subjects with a standard hand anatomy), the sensorimotor representations of the 5 fingers in humans converge toward a model of perceived hand anatomy, deviating from the real hand structure, as the best synthesis between functional and structural features of the hand.SIGNIFICANCE STATEMENT Distinct motor and tactile finger representations exist in both the precentral and postcentral gyrus, supported by a finger-specific pattern of anatomic and functional connectivity across modalities. At the representational level, finger representations reflect the perceived structure of the hand, which might result from an adapting process harmonizing (i.e., uniformizing) the encoding of hand function and structure in the precentral and postcentral gyrus. The same analyses performed in an extremely rare polydactyly subject showed that the emergence of such representational geometry is also found in neuromechanical variants with different hand anatomy and function. However, the harmonization process across the precentral and postcentral gyrus might not be possible because of divergent functional-structural properties of the hand and associated superior manipulation abilities.

Alteration of body representation in typical and atypical motor development.

Developmental coordination disorder (DCD) impacts the quality of life and ability to perform coordinated actions in 5% of school-aged children. The quality of body representations of individuals with DCD has been questioned, but never assessed. We hypothesize that children with DCD have imprecise body representations in the sensory and motor domains. Twenty neurotypical children, seventeen children with DCD (8-12 years old) and twenty neurotypical adults (25-45 years old) performed both sensory and motor body representation tasks: a limb identification and a limb movement task. We observed lower accuracy in the sensory task but not in the motor task. In both tasks, we observe a larger amplitude of errors, or synkinesis, in children with DCD than in neurotypical children. In neurotypical children, accuracy was lower than in neurotypical adults in the motor and sensory task, and the amplitude of sensory errors and synkinesis was higher than in neurotypical adults. Using a linear regression model, we showed that sensory accuracy is a good predictor of synkinesis production, and that synkinesis production is a good predictor of sensory accuracy, as can be expected by the perception-action loop. Results support the hypothesis of an imprecision of body representation in DCD. We suggest that this imprecision arises from noise in the body representation used at the level of internal models of action. Future studies may assess whether slower plasticity of body representations, initial imprecision, or both may account for this observation. At the clinical level, prevention strategies targeting body representation in early childhood are strategically important to limit such impairments. RESEARCH HIGHLIGHTS: Body representation is impaired in children with DCD and has a significant cost in terms of the accuracy of sensory identification of body parts and associated movements. Inaccuracies in the body representation measured in perception and in action (error amplitude and synkinesis) are related in both NT children and adults. In typical development, we provide evidence of a strong link between body schema and body image.

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.

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..

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.

Multisensory Integration in Body Representation.

To be aware of and to move one's body, the brain must maintain a coherent representation of the body. While the body and the brain are connected by dense ascending and descending sensory and motor pathways, representation of the body is not hardwired. This is demonstrated by the well-known rubber hand illusion in which a visible fake hand is erroneously felt as one's own hand when it is stroked in synchrony with the viewer's unseen actual hand. Thus, body representation in the brain is not mere maps of tactile and proprioceptive inputs, but a construct resulting from the interpretation and integration of inputs across sensory modalities.

Spatial tuning of electrophysiological responses to multisensory stimuli reveals a primitive coding of the body boundaries in newborns.

The ability to identify our own body and its boundaries is crucial for survival. Ideally, the sooner we learn to discriminate external stimuli occurring close to our body from those occurring far from it, the better (and safer) we may interact with the sensory environment. However, when this mechanism emerges within ontogeny is unknown. Is it something acquired throughout infancy, or is it already present soon after birth? The presence of a spatial modulation of multisensory integration (MSI) is considered a hallmark of a functioning representation of the body position in space. Here, we investigated whether MSI is present and spatially organized in 18- to 92-h-old newborns. We compared electrophysiological responses to tactile stimulation when concurrent auditory events were delivered close to, as opposed to far from, the body in healthy newborns and in a control group of adult participants. In accordance with previous studies, adult controls showed a clear spatial modulation of MSI, with greater superadditive responses for multisensory stimuli close to the body. In newborns, we demonstrated the presence of a genuine electrophysiological pattern of MSI, with older newborns showing a larger MSI effect. Importantly, as for adults, multisensory superadditive responses were modulated by the proximity to the body. This finding may represent the electrophysiological mechanism responsible for a primitive coding of bodily self boundaries, thus suggesting that even just a few hours after birth, human newborns identify their own body as a distinct entity from the environment.

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.

Voxelwise encoding models of body stimuli reveal a representational gradient from low-level visual features to postural features in occipitotemporal cortex.

Previous research on body representation in the brain has focused on category-specific representation, using fMRI to investigate the response pattern to body stimuli in occipitotemporal cortex. But the central question of the specific computations involved in body selective regions has not been addressed so far. This study used ultra-high field fMRI and banded ridge regression to investigate the computational mechanisms of coding body images, by comparing the performance of three encoding models in predicting brain activity in occipitotemporal cortex and specifically in the extrastriate body area (EBA). Our results indicate that bodies are encoded in occipitotemporal cortex and in the EBA according to a combination of low-level visual features and postural features.

The distorted hand: systematic but 'independent' distortions in both explicit and implicit hand representations in young female adults.

It has long been assumed that an accurate representation of the size and shape of one's body is necessary to successfully interact with the environment. Previous research has shown accurate representations when healthy participants make overt judgments (i.e. explicit) about the size of their bodies. However, when body size is judged implicitly, studies have shown systematic distortions. One suggestion for these differences, is that explicit and implicit representations are informed by different sensory modalities. Explicit representations rely on vision whereas implicit representations are informed by haptics. We designed an experiment to investigate if explicit representations that are informed by haptics are more like implicit representation featuring systematic distortions. We asked female participants to estimate the size of their fingers and hands in three different tasks: an explicit-haptic, an implicit, and an explicit-vision task. The results showed that all three representations were distorted and furthermore, the distortions for each representation were different from one another. These results suggest that inaccurate finger and hand length are a stereotypical feature of body representation that is present in both visual and haptic domains. We discuss the results in relation to theories of body representation.

More implicit and more explicit motor imagery tasks for exploring the mental representation of hands and feet in action.

The mental representation of the body in action can be explored using motor imagery (MI) tasks. MI tasks can be allocated along a continuum going from more implicit to more explicit tasks, where the discriminant is the degree of action monitoring required to solve the tasks (which is the awareness of using the mental representation of our own body to monitor our motor imagery). Tasks based on laterality judgments, such as the Hand Laterality Task (HLT) and the Foot Laterality Task (FLT), provide an example of more implicit tasks (i.e., less action monitoring is required). While, an example of a more explicit task is the Mental Motor Chronometry task (MMC) for hands and feet, where individuals are asked to perform or imagine performing movements with their limbs (i.e., more action monitoring is required). In our study, we directly compared hands and feet at all these tasks for the first time, as these body districts have different physical features as well as functions. Fifty-five participants were asked to complete an online version of the HLT and FLT (more implicit measure), and an online version of the MMC task for hands and feet (more explicit measure). The mental representation of hands and feet in action differed only when the degree of action monitoring decreased (HLT ≠ FLT); we observed the presence of biomechanical constraints only for hands. Differently, when the degree of action monitoring increased hands and feet did not show any difference (MMC hands = MMC feet). Our results show the presence of a difference in the mental representation of hands and feet in action that specifically depends on the degree of action monitoring.

Self-touch facilitates the recognition of the dis-owned left hand in somatoparaphrenia: a single case study.

We investigated whether self-administered tactile stimulation could act as a temporary restorative mechanism for body ownership disorders, both implicitly and explicitly. We tested this hypothesis in a patient with somatoparaphrenia, who displayed increased accuracy in explicitly recognizing their left hand during self-touch. Furthermore, the patient implicitly perceived their hand and the experimenter's hand as more belonging to their own body compared to conditions where vision was the sole sensory input. These findings highlight the importance of self-touch in maintaining a coherent body representation, while also demonstrating the potential dissociation between the recovery of explicit and implicit perceptions of body ownership.

A gait-based paradigm to investigate central body representation in cervical dystonia patients.

BACKGROUND: Cervical dystonia (CD) is a common adult-onset idiopathic form of dystonia characterized by an abnormal head posture caused by an excessive activity of the neck muscles. The position of the head is important to direct viewpoint in the rounding environment, and the body orientation, during gait, must be coherent with the subjective straight ahead (SSA). An alteration of the SSA, as in the case of CD patients, could affect gait when visual input is not available. The aim of this study was to probe the behavior of patients with CD during blindfolded walking, investigating the ability to walk straight ahead based only on somatosensory and vestibular information. METHODS: In this observational cross-sectional study, patients with CD and healthy control subjects (HC) were compared. All participants were evaluated through a gait analysis during blindfolded walking on a GAITRite carpet, relying on their own sense of straightness. RESULTS: Patients with CD showed lower values of path length (p < 0.001), a lower number of steps on the carpet (p < 0.001). A higher number of CD patients deviated during the task, walking out of the carpet, (p < 0.005) compared to HS. No relation was found between the dystonic side and the gait trajectory deviation. A significant correlation was found between pain symptom and gait performance. CONCLUSIONS: CD patients showed dysfunctions in controlling dynamic body location during walking without visual afferences, while the dystonic side does not seem to be related to the lateral deviation of the trajectory. Our results would assume that a general proprioceptive impairment could lead to an improper body position awareness in patients with CD.

Altered processing of conflicting body representations in women with restrictive anorexia nervosa.

Cognitive and affective impairments in processing body image have been observed in patients with Anorexia Nervosa (AN) and may induce the hypercontrolled and regulative behaviors observed in this disorder. Here, we aimed to probe the link between activation of body representations and cognitive control by investigating the ability to resolve body-related representational conflicts in women with restrictive AN and matched healthy controls (HC). Participants performed a modified version of the Flanker task in which underweight and overweight body images were presented as targets and distractors; a classic version of the task, with letters, was also administered as a control. The findings indicated that performance was better among the HC group in the task with bodies compared to the task with letters; however, no such facilitation was observed in AN patients, whose overall performance was poorer than that of the HC group in both tasks. In the task with body stimuli, performance among patients with AN was the worst on trials presenting underweight targets with overweight bodies as flankers. These results may reflect a dysfunctional association between the processing of body-related representations and cognitive control mechanisms that may aid clinicians in the development of optimal individualized treatments.

Hand-Selective Visual Regions Represent How to Grasp 3D Tools: Brain Decoding during Real Actions.

Most neuroimaging experiments that investigate how tools and their actions are represented in the brain use visual paradigms where tools or hands are displayed as 2D images and no real movements are performed. These studies discovered selective visual responses in occipitotemporal and parietal cortices for viewing pictures of hands or tools, which are assumed to reflect action processing, but this has rarely been directly investigated. Here, we examined the responses of independently visually defined category-selective brain areas when participants grasped 3D tools (N = 20; 9 females). Using real-action fMRI and multivoxel pattern analysis, we found that grasp typicality representations (i.e., whether a tool is grasped appropriately for use) were decodable from hand-selective areas in occipitotemporal and parietal cortices, but not from tool-, object-, or body-selective areas, even if partially overlapping. Importantly, these effects were exclusive for actions with tools, but not for biomechanically matched actions with control nontools. In addition, grasp typicality decoding was significantly higher in hand than tool-selective parietal regions. Notably, grasp typicality representations were automatically evoked even when there was no requirement for tool use and participants were naive to object category (tool vs nontools). Finding a specificity for typical tool grasping in hand-selective, rather than tool-selective, regions challenges the long-standing assumption that activation for viewing tool images reflects sensorimotor processing linked to tool manipulation. Instead, our results show that typicality representations for tool grasping are automatically evoked in visual regions specialized for representing the human hand, the primary tool of the brain for interacting with the world.

Differences in working memory coding of biological motion attributed to oneself and others.

The question how the brain distinguishes between information about self and others is of fundamental interest to both philosophy and neuroscience. In this functional magnetic resonance imaging (fMRI) study, we sought to distinguish the neural substrates of representing a full-body movement as one's movement and as someone else's movement. Participants performed a delayed match-to-sample working memory task where a retained full-body movement (displayed using point-light walkers) was arbitrarily labeled as one's own movement or as performed by someone else. By using arbitrary associations we aimed to address a limitation of previous studies, namely that our own movements are more familiar to us than movements of other people. A searchlight multivariate decoding analysis was used to test where information about types of movement and about self-association was coded. Movement specific activation patterns were found in a network of regions also involved in perceptual processing of movement stimuli, however not in early sensory regions. Information about whether a memorized movement was associated with the self or with another person was found to be coded by activity in the left middle frontal gyrus (MFG), left inferior frontal gyrus (IFG), bilateral supplementary motor area, and (at reduced threshold) in the left temporoparietal junction (TPJ). These areas are frequently reported as involved in action understanding (IFG, MFG) and domain-general self/other distinction (TPJ). Finally, in univariate analysis we found that selecting a self-associated movement for retention was related to increased activity in the ventral medial prefrontal cortex.

Chimpanzees (Pan troglodytes) detect strange body parts: an eye-tracking study.

This study investigated chimpanzee body representation by testing whether chimpanzees detect strangeness in body parts. We tested six chimpanzees with edited chimpanzee body pictures in eye-tracking tasks. The target body parts were arms or legs. For either target, there were four conditions: "normal" condition as control, where all bodies were normal; "misplaced" condition, where one arm or one leg was misplaced to an incorrect body location in each picture; "replaced by a chimpanzee part" condition, where one arm or one leg was replaced by a chimpanzee leg or arm, respectively, in its original place in each picture; and "replaced by a human part" condition, where one arm or one leg was replaced by a human arm or leg in each picture. Compared to the looking times toward the normal parts, chimpanzees had significantly longer looking times toward the human arms or legs. The looking times toward the misplaced parts were also longer than the normal parts, but the difference just failed to meet significance. These results indicate more interests toward strange body parts, compared to typical parts, suggesting that chimpanzees might have a body representation that is sufficiently sensitive to detect these aspects of strangeness.