Peripersonal Space Research Articles

Neural correlates of disturbance in the sense of agency in schizophrenia: An fMRI study using the ‘enfacement’ paradigm

An altered sense of self-awareness and agency has been proposed to underlie symptoms of schizophrenia. In this study, we used the enfacement illusion paradigm – in which perception of another person's face leads to changes in perception of one's own peri-personal space – to examine the brain correlates of the sense of agency and its potential disruption in schizophrenia. Thirty-three schizophrenic patients and 27 healthy controls underwent fMRI scanning during performance of a task designed to elicit the enfacement illusion. Activations were examined using whole-brain analysis and also in an a priori identified region of interest (ROI) in the temporoparietal junction (TPJ), a region that has been described as involved in self/other differentiation and sense of agency. Both groups showed a pattern of cortical activation involving the pre and postcentral cortex, Rolandic operculum, insula, parietal, temporal and occipital cortex bilaterally as well as TPJ (but only right-side in patients). Examination of the TPJ ROI revealed significantly reduced activation on the left in the patients that was associated with poorer insight. The findings suggest brain functional abnormality in schizophrenia related to the formation or maintenance of processes related to self and/or agency. Decreased function in the TPJ may have a role in the impaired insight seen in patients with the disorder.

Peripersonal and reaching space differ: Evidence from their spatial extent and multisensory facilitation pattern

Peripersonal space (PPS) is a multisensory representation of the space near body parts facilitating interactions with the close environment. Studies on non-human and human primates agree in showing that PPS is a body part-centered representation that guides actions. Because of these characteristics, growing confusion surrounds peripersonal and arm-reaching space (ARS), that is the space one’s arm can reach. Despite neuroanatomical evidence favoring their distinction, no study has contrasted directly their respective extent and behavioral features. Here, in five experiments (N = 140) we found that PPS differs from ARS, as evidenced both by participants’ spatial and temporal performance and by its modeling. We mapped PPS and ARS using both their respective gold standard tasks and a novel multisensory facilitation paradigm. Results show that: (1) PPS is smaller than ARS; (2) multivariate analyses of spatial patterns of multisensory facilitation predict participants’ hand locations within ARS; and (3) the multisensory facilitation map shifts isomorphically following hand positions, revealing hand-centered coding of PPS, therefore pointing to a functional similarity to the receptive fields of monkeys’ multisensory neurons. A control experiment further corroborated these results and additionally ruled out the orienting of attention as the driving mechanism for the increased multisensory facilitation near the hand. In sharp contrast, ARS mapping results in a larger spatial extent, with undistinguishable patterns across hand positions, cross-validating the conclusion that PPS and ARS are distinct spatial representations. These findings show a need for refinement of theoretical models of PPS, which is relevant to constructs as diverse as self-representation, social interpersonal distance, and motor control.

Affective sounds entering the peripersonal space influence the whole-body action preparation

The peripersonal space (PPS), the space surrounding us, is found to have enhanced multisensory-motor representation in the brain. In this study, we investigate how approaching sounds stopping at different distances within the peripersonal space, and carrying emotional content (positive, negative, and neutral), modulate the preparation of action as performing a Step. Premotor reaction times were measured by means of anticipatory forces and muscular activations to capture action preparation, the kinematics of stepping was considered for defining action performance, and for each stimulus, the individual perceived level of arousal and valence was evaluated. In general, we found a prompter premotor reaction for closer sounds compared to the farther ones and the fastest reactions detected for the neutral sound at each distance. We interpreted this time facilitation for neutral sound due to the large frequency spectrum of the stimuli and the absence of affective component and semantical content to decode. Interestingly, while at the close distance, none difference was found between positive and negative emotional stimuli, at the far distance faster reactions were present for negative compared to the positive sounds indicating that when arousal is less enhanced individuals are able to differentiate the emotional content of a sound. The kinematics observed after action initiation sustained the anticipatory results by showing that larger steps were performed when reacting to close compared to far sounds, being perceived as more arousing, and this happened particularly for neutral and negative sounds. Altogether, the results showed that action preparation is influenced by the vicinity and by the valence carried by looming auditory stimuli. For discriminating the stimuli valence, a certain distance, still within the PPS, is necessary; when instead stimuli are too close to the body valence discrimination is not performed.

Sharpening of peripersonal space during the COVID-19 pandemic

Our social world has been transformed by the COVID-19 pandemic. Beyond the direct impact of the pandemic on physical health, the social distancing measures implemented worldwide to slow down disease transmission have dramatically impacted social interactions1,2. These measures, including orders to stay at home and to maintain a social distance of at least 2 meters, have been essential to limit the spread of the disease, but they have had severe costs for humans as social animals2. Right before and right after the adoption of the most stringent measures in Switzerland in Spring 2020, we were conducting a series of experiments to measure the representation of the so-called peripersonal space — the space immediately surrounding our body, where we normally interact with objects and other individuals3. We found that the introduction of social distancing measures led to a reduction in the extent of the peripersonal space and enhanced its segregation between individuals, as if the presence of others in close space would activate an implicit form of freezing response.

Body size perception in stroke patients with paresis.

Recent studies have suggested that people’s intent and ability to act also can influence their perception of their bodies’ peripersonal space. Vice versa one could assume that the inability to reach toward and grasp an object might have an impact on the subject’s perception of reaching distance. Here we tested this prediction by investigating body size and action capability perception of neurological patients suffering from arm paresis after stroke, comparing 32 right-brain-damaged patients (13 with left-sided arm paresis without additional spatial neglect, 10 with left-sided arm paresis and additional spatial neglect, 9 patients had neither arm paresis nor neglect) and 27 healthy controls. Nineteen of the group of right hemisphere stroke patients could be re-examined about five months after initial injury. Arm length was estimated in three different methodological approaches: explicit visual, explicit tactile/proprioceptive, and implicit reaching. Results fulfilled the working hypothesis. Patients with an arm paresis indeed perceived their bodies differently. We found a transient overestimation of the length of the contralesional, paretic arm after stroke. Body size and action capability perception for the extremities thus indeed seem to be tightly linked in humans.

Peripersonal space in the front, rear, left and right directions for audio-tactile multisensory integration

Peripersonal space (PPS) is important for humans to perform body–environment interactions. However, many previous studies only focused on the specific direction of the PPS, such as the front space, despite suggesting that there were PPSs in all directions. We aimed to measure and compare the peri-trunk PPS in four directions (front, rear, left, and right). To measure the PPS, we used a tactile and an audio stimulus because auditory information is available at any time in all directions. We used the approaching and receding task-irrelevant sounds in the experiment. Observers were asked to respond as quickly as possible when a tactile stimulus was applied to a vibrator on their chest. We found that peri-trunk PPS representations exist with an approaching sound, irrespective of the direction.

VR Setup to Assess Peripersonal Space Audio-Tactile 3D Boundaries

Many distinct spaces surround our bodies. Most schematically, the key division is between peripersonal space (PPS), the close space surrounding our body, and an extrapersonal space, which is the space out of one’s reach. The PPS is considered as an action space, which allows us to interact with our environment by touching and grasping. In the current scientific literature, PPS’ visual representations are appearing as mere bubbles of even dimensions wrapped around the body. Although more recent investigations of PPS’ upper body (trunk, head, and hands) and lower body (legs and foot) have provided new representations, no investigation has been made yet concerning the estimation of PPS’s overall representation in 3D. Previous findings have demonstrated how the relationship between tactile processing and the location of sound sources in space is modified along a spatial continuum. These findings suggest that similar methods can be used to localize the boundaries of the subjective individual representation of PPS. Hence, we designed a behavioral paradigm in virtual reality based on audio-tactile interactions, which has enabled us to infer a detailed individual 3D audio-tactile representation of PPS. Considering that inadequate body-related multisensory integration processes can produce incoherent spatio–temporal perception, the development of a virtual reality setup and a method to estimate the representation of the subjective PPS volumetric boundaries will be a valuable addition for the comprehension of the mismatches occurring between body physical boundaries and body schema representations in 3D.

Remote hand: Hand-centered peripersonal space transfers to a disconnected hand avatar

The space surrounding our body is called peripersonal space (PPS). It has been reported that visuo-tactile facilitation occurs more strongly within PPS than outside PPS. Furthermore, previous research has revealed several methods by which PPS can be extended. The present study provides the first behavioral evidence of the transfer of PPS in a virtual environment by a novel technique. PPS representation was investigated using a remote-controlled hand avatar presented far from the body in a virtual environment. Participants showed strongest visuo-tactile facilitation at the far space around the remote hand and no facilitation at the near space around the real hand, suggesting that PPS transfers from near the body to the space around the hand avatar. The present results extend previous findings of the plasticity of PPS and demonstrate flexibility of PPS representation beyond the physical and anatomical limits of body representation.

On the content of Peripersonal visual experience

In a recent paper, ‘Peripersonal perception in action’ (Synthese, 2018), Frederique de Vignemont tackles the problem of defining what is peculiar to the visual perception of objects falling within the peripersonal space of the observer, i.e. the space immediately surrounding the body, and which is commonly described as the space in which action takes place. In this paper, I first discuss the proposal offered by de Vignemont about what characterizes peripersonal perception. Then, I suggest an extension of this account that offers a meticulous description of the nature of the Content of Peripersonal Visual Experience - a topic never explicitly considered in the philosophical literature on vision - by discussing some peculiar features of it that, as recognized also by de Vignemont’s account, still need to be explained. In particular, I offer a philosophical examination of the specificity of peripersonal visual experience, in relation to its phenomenological dimension, its optical mechanisms and its neurophysiological underpinnings, in the light of our best theories from vision science, and in comparison to the visual experience of other visual spaces.

Characterising sensorimotor adaptation in Complex Regional Pain Syndrome

It has been suggested that sensorimotor conflict contributes to the maintenance of some pathological pain conditions, implying that there are problems with the adaptation processes that normally resolve such conflict. We tested whether sensorimotor adaptation is impaired in people with Complex Regional Pain Syndrome (CRPS) by characterising their adaption to lateral prismatic shifts in vision. People with unilateral upper-limb CRPS Type I (n = 17), and pain-free individuals (n = 18; matched for age, sex, and handedness) completed prism adaptation with their affected/non-dominant and non-affected/dominant arms. We examined 1) the rate at which participants compensated for the optical shift during prism exposure (i.e., strategic recalibration), 2) endpoint errors made directly after prism adaptation (sensorimotor realignment) and the retention of these errors, and 3) kinematic markers associated with strategic control. Direct comparisons between people with CRPS and controls revealed no evidence of any differences in strategic recalibration, including no evidence for differences in a kinematic marker associated with trial-by-trial changes in movement plans during prism exposure. All participants made significant endpoint errors after prism adaptation exposure, indicative of sensorimotor realignment. Overall, the magnitude of this realignment did not differ between people with CRPS and pain-free controls. However, when endpoint errors were considered separately for each hand, people with CRPS made greater errors (indicating more rather than less realignment) when using their affected hand than their non-affected hand. No such difference was seen in controls. Taken together, these findings provide no evidence of impaired strategic control or sensorimotor realignment in people with CRPS. In contrast, they provide some indication that there could be a greater propensity for sensorimotor realignment in the CRPS-affected arm, consistent with more flexible representations of the body and peripersonal space. Our study challenges an implicit assumption of the theory that sensorimotor conflict might underlie some pathological pain conditions.

Inflexible Updating of the Self-Other Divide During a Social Context in Autism: Psychophysical, Electrophysiological, and Neural Network Modeling Evidence

BackgroundAutism spectrum disorder (ASD) affects many aspects of life, from social interactions to (multi)sensory processing. Similarly, the condition expresses at a variety of levels of description, from genetics to neural circuits and interpersonal behavior. We attempt to bridge between domains and levels of description by detailing the behavioral, electrophysiological, and putative neural network basis of peripersonal space (PPS) updating in ASD during a social context, given that the encoding of this space relies on appropriate multisensory integration, is malleable by social context, and is thought to delineate the boundary between the self and others. MethodsFifty (20 male/30 female) young adults, either diagnosed with ASD or age- and sex-matched individuals, took part in a visuotactile reaction time task indexing PPS, while high-density electroencephalography was continuously recorded. Neural network modeling was performed in silico. ResultsMultisensory psychophysics demonstrates that while PPS in neurotypical individuals shrinks in the presence of others—as to “give space”—this does not occur in ASD. Likewise, electroencephalography recordings suggest that multisensory integration is altered by social context in neurotypical individuals but not in individuals with ASD. Finally, a biologically plausible neural network model shows, as a proof of principle, that PPS updating may be inflexible in ASD owing to the altered excitatory/inhibitory balance that characterizes neural circuits in animal models of ASD. ConclusionsFindings are conceptually in line with recent statistical inference accounts, suggesting diminished flexibility in ASD, and further these observations by suggesting within an example relevant for social cognition that such inflexibility may be due to excitatory/inhibitory imbalances.

Integration of predictions and afferent signals in body ownership

We aimed at investigating the contribution of sensory predictions triggered by the sight of an object moving towards the body for the sense of body ownership. We used a recently developed psychophysical discrimination task to assess body ownership in the rubber hand illusion. In this task, the participants had to choose which of the two right rubber hands in view felt most like their own, and the ownership discriminations were fitted to psychometric curves. In the current study, we occluded the visual impressions of the object moving towards one of the rubber hands (during the first two-thirds of the path) and only revealed the final third of the object's movement trajectory when it touched the rubber hand (approach-occluded condition). Alternatively, we occluded only the final part so that the main part of the movement towards the model hand was visible (touch-occluded). We compared these two conditions to an illusion baseline condition where the object was visible during the entire trajectory and contact (no-occlusion). The touch-occluded condition produced equally strong hand ownership as the baseline condition with no occlusion, while ownership perception was significantly reduced when vision of the object approaching the rubber hand was occluded (approach-occluded). Our results show that tactile predictions generated from seeing an object moving towards the body are temporally exact, and they contribute to the rubber hand illusion by integrating with temporally congruent afferent sensory signals. This finding highlights the importance of multisensory predictions in peripersonal space, object permanence, and the interplay between bottom-up sensory signals and top-down predictions in body ownership.

Altered Peripersonal Space and the Bodily Self in Schizophrenia: A Virtual Reality Study.

Self-disturbances such as an anomalous perception of one's own body boundary are central to the phenomenology of schizophrenia (SZ), but measuring the spatial parameters of the hypothesized self-other boundary has proved to be challenging. Peripersonal space (PPS) refers to the immediate zone surrounding the body where the self interacts physically with the environment; the space that corresponds to hypothesized self-other boundary. PPS is represented by enhanced multisensory integration and faster reaction time (RT) for objects near the body. Thus, multisensory RT tasks can be used to estimate self-other boundary. We aimed to quantify PPS in SZ using an immersive virtual reality visuotactile RT paradigm. Twenty-four participants with SZ and 24 demographically matched controls (CO) were asked to detect tactile vibration while watching a ball approaching them, thrown by either a machine (nonsocial condition) or an avatar (social condition). Parameters of PPS were estimated from the midpoint of the spatial range where the tactile RT decreased most rapidly (size) and the gradient of the RT change at this midpoint (slope). Overall, PPS was smaller in participants with SZ compared with CO. PPS slope for participants with SZ was shallower than CO in the social but not in nonsocial condition, indicating an increased uncertainty of self-other boundary across an extended zone in SZ. Social condition also increased false alarms for tactile detection in SZ. Clinical symptoms were not clearly associated with PPS parameters. These findings suggest the context-dependent nature of weakened body boundary in SZ and underscore the importance of reconciliating objective and subjective aspects of self-disturbances.

Motor Inhibition to Dangerous Objects: Electrophysiological Evidence for Task-dependent Aversive Affordances.

Previous work suggests that perception of an object automatically facilitates actions related to object grasping and manipulation. Recently, the notion of automaticity has been challenged by behavioral studies suggesting that dangerous objects elicit aversive affordances that interfere with encoding of an object's motor properties; however, related EEG studies have provided little support for these claims. We sought EEG evidence that would support the operation of an inhibitory mechanism that interferes with the motor encoding of dangerous objects, and we investigated whether such mechanism would be modulated by the perceived distance of an object and the goal of a given task. EEGs were recorded by 24 participants who passively perceived dangerous and neutral objects in their peripersonal, boundary, or extrapersonal space and performed either a reachability judgment task or a categorization task. Our results showed that greater attention, reflected in the visual P1 potential, was drawn by dangerous and reachable objects. Crucially, a frontal N2 potential, associated with motor inhibition, was larger for dangerous objects only when participants performed a reachability judgment task. Furthermore, a larger parietal P3b potential for dangerous objects indicated the greater difficulty in linking a dangerous object to the appropriate response, especially when it was located in the participants' extrapersonal space. Taken together, our results show that perception of dangerous objects elicits aversive affordances in a task-dependent way and provides evidence for the operation of a neural mechanism that does not code affordances of dangerous objects automatically, but rather on the basis of contextual information.

When two worlds collide: the influence of an obstacle in peripersonal space on multisensory encoding

Multisensory coding of the space surrounding our body, the peripersonal space, is crucial for motor control. Recently, it has been proposed that an important function of multisensory coding is that it allows anticipation of the tactile consequences of contact with a nearby object. Indeed, performing goal-directed actions (i.e. pointing and grasping) induces a continuous visuotactile remapping as a function of on-line sensorimotor requirements. Here, we investigated whether visuotactile remapping can be induced by obstacles, e.g. objects that are not the target of the grasping movement. In the current experiment, we used a cross-modal obstacle avoidance paradigm, in which participants reached past an obstacle to grasp a second object. Participants indicated the location of tactile targets delivered to the hand during the grasping movement, while a visual cue was sometimes presented simultaneously on the to-be-avoided object. The tactile and visual stimulation was triggered when the reaching hand passed a position that was drawn randomly from a continuous set of predetermined locations (between 0 and 200 mm depth at 5 mm intervals). We observed differences in visuotactile interaction during obstacle avoidance dependent on the location of the stimulation trigger: visual interference was enhanced for tactile stimulation that occurred when the hand was near the to-be-avoided object. We show that to-be-avoided obstacles, which are relevant for action but are not to-be-interacted with (as the terminus of an action), automatically evoke the tactile consequences of interaction. This shows that visuotactile remapping extends to obstacle avoidance and that this process is flexible.

Testosterone administration in women increases the size of their peripersonal space.

Peripersonal space (PPS) is the space immediately surrounding the body, conceptualised as a sensory-motor interface between body and environment. PPS size differs between individuals and contexts, with intrapersonal traits and states, as well as social factors having a determining role on the size of PPS. Testosterone plays an important role in regulating social-motivational behaviour and is known to enhance dominance motivation in an implicit and unconscious manner. We investigated whether the dominance-enhancing effects of testosterone reflect as changes in the representation of PPS in a within-subjects testosterone administration study in women (N = 19). Participants performed a visuo-tactile integration task in a mixed-reality setup. Results indicated that the administration of testosterone caused a significant enlargement of participants' PPS, suggesting that testosterone caused participants to implicitly appropriate a larger space as their own. These findings suggest that the dominance-enhancing effects of testosterone reflect at the level of sensory-motor processing in PPS.

Fearful faces modulate spatial processing in peripersonal space: An ERP study

Peripersonal space (PPS) represents the region of space surrounding the body. A pivotal function of PPS is to coordinate defensive responses to threat. We have previously shown that a centrally-presented, looming fearful face, signalling a potential threat in one's surroundings, modulates spatial processing by promoting a redirection of sensory resources away from the face towards the periphery, where the threat may be expected – but only when the face is presented in near, rather than far space. Here, we use electrophysiological measures to investigate the neural mechanism underlying this effect. Participants made simple responses to tactile stimuli delivered on the cheeks, while watching task-irrelevant neutral or fearful avatar faces, looming towards them either in near or far space. Simultaneously with the tactile stimulation, a ball with a checkerboard pattern (probe) appeared to the left or right of the avatar face. Crucially, this probe could either be close to the avatar face, and thus more central in the participant's vision, or further away from the avatar face, and thus more peripheral in the participant's vision. Electroencephalography was continuously recorded. Behavioural results confirmed that in near space only, and for fearful relative to neutral faces, tactile processing was facilitated by the peripheral compared to the central probe. This behavioural effect was accompanied by a reduction of the N1 mean amplitude elicited by the peripheral probe for fearful relative to neutral faces. Moreover, the faster the participants responded to tactile stimuli with the peripheral probe, relative to the central, the smaller was their N1. Together these results, suggest that fearful faces intruding into PPS may increase expectation of a visual event occurring in the periphery. This fear-induced effect would enhance the defensive function of PPS when it is most needed, i.e., when the source of threat is nearby, but its location remains unknown.

Reaching Out for Food: How Food Incentives Modulate Peripersonal Space Perception.

Two experiments were conducted to determine, first, whether food items influence participants’ estimations of the size of their subjective peripersonal space. It was of particular interest whether this representation is influenced by satiated/hungry states and is differentially affected by valence and calorie content of depicted stimuli. Second, event-related brain potentials (ERPs) were used, in order to obtain information about the time course of the observed effects and how they depend on the spatial location of the food pictures. For that purpose, participants had to decide whether food items shown at various distances along a horizontal plane in front of them, were reachable or not. In Experiment 1, when participants were hungry, they perceived an increase of their peripersonal space modulated by high-calorie items which were experienced as being more reachable than low-calorie items. In Experiment 2, the reachability findings were replicated and early and late components of ERPs showed an attentional enhancement in far space for food items when participants were hungry. These findings suggest that participants’ subjective peripersonal space increased while being hungry, especially for high-calorie contents. Attention also seems to be oriented more strongly to far space items due to their expected incentive-related salience, expanding the subjective representation of peripersonal space.

Phase-coupling of neural oscillations contributes to individual differences in peripersonal space

The peripersonal space (PPS) is a multisensory and sensorimotor interface between our body and the environment. The location of PPS boundary is not fixed. Rather, it adapts to the environmental context and differs greatly across individuals.Recent studies have started to unveil the neural correlates of individual differences in PPS extension; however, this picture is not clear yet. Here, we used approaching auditory stimuli and magnetoencephalography to capture the individual boundary of PPS and examine its neural underpinnings. In particular, building upon previous studies from our own group, we investigated the possible contribution of an intrinsic feature of the brain, that is the “resting state” functional connectivity, to the individual differences in PPS extension and the frequency specificity of this contribution. Specifically, we focused on the activity synchronized to the premotor cortex, where multisensory neurons encoding PPS have been described. Results showed that the stronger the connectivity between left premotor cortex (lPM) and a set of fronto-parietal, sensorimotor regions in the right and left hemisphere, the wider the extension of the PPS. Strikingly, such a correlation was observed only in the beta-frequency band. Overall, our results suggest that the individual extension of the PPS is coded in spatially- and spectrally-specific resting state functional links.

The Interrelation Between Peripersonal Action Space and Interpersonal Social Space: Psychophysiological Evidence and Clinical Implications.

The peripersonal space is an adaptive and flexible interface between the body and the environment that fulfills a dual-motor function: preparing the body for voluntary object-oriented actions to interact with incentive stimuli and preparing the body for defensive responses when facing potentially harmful stimuli. In this position article, we provide arguments for the sensorimotor rooting of the peripersonal space representation and highlight the variables that contribute to its flexible and adaptive characteristics. We also demonstrate that peripersonal space represents a mediation zone between the body and the environment contributing to not only the control of goal-directed actions but also the organization of social life. The whole of the data presented and discussed led us to the proposal of a new theoretical framework linking the peripersonal action space and the interpersonal social space and we highlight how this theoretical framework can account for social behaviors in populations with socio-emotional deficits.