Multisensory peripersonal space: Visual looming stimuli induce stronger response facilitation to tactile than auditory and visual stimulations.

Anticipating physical contact with objects in the environment is a key component of efficient motor performance. Peripersonal neurons are thought to play a determinant role in these predictions by enhancing responses to touch when combined with visual stimuli in peripersonal space (PPS). However, recent research challenges the idea that this visuo-tactile integration contributing to the prediction of tactile events occurs strictly in PPS. We hypothesised that enhanced sensory sensitivity in a multisensory context involves not only contact anticipation but also heightened attention towards near-body visual stimuli. To test this hypothesis, Experiment 1 required participants to respond promptly to tactile (probing contact anticipation) and auditory (probing enhanced attention) stimulations presented at different moments of the trajectory of a (social and non-social) looming visual stimulus. Reduction in reaction time as compared to a unisensory baseline was observed from an egocentric distance of around 2 m (inside and outside PPS) for all multisensory conditions and types of visual stimuli. Experiment 2 tested whether these facilitation effects still occur in the absence of a multisensory context, i.e., in a visuo-visual condition. Overall, facilitation effects induced by the looming visual stimulus were comparable in the three sensory modalities outside PPS but were more pronounced for the tactile modality inside PPS (84 cm from the body as estimated by a reachability judgement task). Considered together, the results suggest that facilitation effects induced by visual looming stimuli in multimodal sensory processing rely on the combination of attentional factors and contact anticipation depending on their distance from the body.

Comparison of peripersonal space in front and rear spaces.

The space immediately around the body, referred to as the peripersonal space (PPS), plays a crucial role in interactions with external objects and in avoiding unsafe situations. This study aimed to investigate whether the size of the PPS changes depending on direction, with a particular focus on the disparity between the front and rear spaces. A vibrotactile stimulus was presented to measure PPS while a task-irrelevant auditory stimulus (probe) approached the participant. In addition, to evaluate the effect of the probe, a baseline condition was used in which only tactile stimuli were presented. The results showed that the auditory facilitation effect of the tactile stimulus was greater in the rear condition than in the front condition. Conversely, the performance on tasks related to auditory distance perception and sound speed estimation did not differ between the two directions, indicating that the difference in the auditory facilitation effect between directions cannot be explained by these factors. These findings indicate that the strength of audio-tactile integration is greater in the rear space compared to the front space, suggesting that the representation of the PPS differed between the front and rear spaces.

Reshaping the peripersonal space in virtual reality.

Peripersonal space (PPS) is defined as the space that lies within reach. Previous research revealed that PPS can be dynamically reshaped with the use of tools extending the arm's reach. Here we investigated whether PPS reshaping depends on the kind of selected tool and/or the motor routine associated with its use. Participants carried out a visuo-tactile detection task in an immersive VR environment that allowed to measure the PPS size before and after a short period of tools use. In Experiment 1, participants had to pull or push objects towards or away from themselves using a shovel. In Experiment 2, they were required to either hammer or shoot an avatar placed in the Extrapersonal space. We found, for the first time in a VR environment, that a period of pull training was effective in enlarging the PPS, a result that replicates and expands previous findings carried out in real life conditions. However, no significant change in PPS size was achieved for training with other tools and motor routines. Our results suggest that the reshaping of PPS is a complex phenomenon in which the kind of interaction between the agent, the targets and the exploited motor routines all play a critical role.

Beyond peripersonal boundaries: insights from crossmodal interactions.

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

The left-right reversed visual feedback of the hand affects multisensory interaction within peripersonal space.

The interaction between vision and touch, known as the crossmodal congruency effect, has been extensively investigated in several research studies. Recent studies have revealed that the crossmodal congruency effect involves body representations. However, it is unclear how bodily information (e.g., location, posture, motion) is linked to visual and tactile inputs. Three experiments were conducted to investigate this issue. In Experiment 1, participants performed a crossmodal congruency task in which both their hand appearance and the motor trajectories were left-right reversed. The results showed that the crossmodal congruency effect was not observed in the reversal condition, whereas participants showed significant crossmodal congruency in the control condition, in which there was no visual manipulation of the hand. In Experiments 2 and 3, where either the hand appearance or motor trajectory was left-right reversed individually, a significant crossmodal congruency effect was observed. This study demonstrated that visual manipulation of hand appearance and motor trajectories both affected the crossmodal congruency effect, although neither showed a dominant effect that solely altered the crossmodal congruency effect. The present results provide insights into the relationship between visual-tactile interactions and bodily information.

Back and front peripersonal space: behavioural and EMG evidence of top-down and bottom-up mechanisms.

Previous studies have identified a 'defensive graded field' in the peripersonal front space where potential threatening stimuli induce stronger blink responses, mainly modulated by top-down mechanisms, which include various factors, such as proximity to the body, stimulus valence, and social cues. However, very little is known about the mechanisms responsible for representation of the back space and the possible role of bottom-up information. By means of acoustic stimuli, we evaluated individuals' representation for front and back space in an ambiguous environment that offered some degree of uncertainty in terms of both distance (close vs. far) and front-back egocentric location of sound sources. We aimed to consider verbal responses about localization of sound sources and EMG data on blink reflex. Results suggested that stimulus distance evaluations were better explained by subjective front-back discrimination, rather than real position. Moreover, blink response data were also better explained by subjective front-back discrimination. Taken together, these findings suggest that the mechanisms that dictate blink response magnitude might also affect sound localization (possible bottom-up mechanism), probably interacting with top-down mechanisms that modulate stimuli location and distance. These findings are interpreted within the defensive peripersonal framework, suggesting a close relationship between bottom-up and top-down mechanisms on spatial representation.

Space for power: feeling powerful over others' behavior affects peri-personal space representation.

We investigated whether and how social power affects the representation of peri-personal space (PPS). We applied a multisensory interaction task to assess PPS representation and the Personal Sense of Power Scale to assess participants' feelings of power over others' behaviors and over others' opinions. In Study 1, we probed PPS representation in a virtual social context. Participants with a higher sense of power showed a less defined differentiation between the close and far space as compared to participants with a lower sense of power. This effect was replicated in Study 2 when participants performed the task in a non-social context (with no person in the scene), but only after they were reminded of an episode of power. Thus, social power-the perception of power over others' behavior-affects the multisensory representation of the self in space by blurring the differentiation between one's own PPS and the space of others.

Paying attention to the outcome of others' actions has dissociated effects on observer's peripersonal space representation and exploitation.

The representation of peripersonal space (PPS representation) and the selection of motor actions within it (PPS exploitation) are influenced by action outcomes and reward prospects. The present study tested whether observing the outcome of others' actions altered the observer's PPS representation and exploitation. Participants (observers) performed a reachability-judgement task (assessing PPS representation) before and after having observed a confederate (actors) performing a stimuli-selection task on a touch-screen table. In the stimuli-selection task, the stimuli selected could either yield a reward or not, but the probability to select a reward-yielding stimulus was biased in space, being either 50%, 25% or 75% in the actor's proximal or distal space. After the observation phase, participants performed the stimuli-selection task (assessing PPS exploitation), but with no spatial bias in the distribution of reward-yielding stimuli. Results revealed an effect of actors' actions outcome on observers' PPS representation, which changed according to the distribution of reward-yielding stimuli in the actors' proximal and distal spaces. No significant effect of actors' actions outcome was found on observers' PPS exploitation. As a whole, the results suggest dissociated effects of observing the outcome of others' actions on PPS representation and exploitation.

Changes in perceived peripersonal space following the rubber hand illusion.

Peripersonal space (PPS), the region immediately surrounding the body is essential for bodily protection and goal directed action. Previous studies have suggested that the PPS is anchored to one's own body and in the current study we investigated whether the PPS could be modulated by changes in perceived body ownership. While theoretically important, this anchoring can also have implications for patients with altered body perception. The rubber hand illusion (RHI) is a way to manipulate body ownership. We hypothesized that after induction of a left hand RHI, the perceived space around the body shifts to the right. Sixty-five participants performed a landmark task before and after a left hand RHI. In the landmark task, participants had to determine whether a vertical landmark line was left or right from the center of a horizontal screen. One group of the participants was exposed to synchronous stroking, the other group experienced asynchronous stroking. Results showed a shift in space to the right (e.g. away from the own arm), but only for the 'synchronous stroking' group. These results suggest that the relevant action space becomes linked to the fake hand. Critically, subjective ownership experience did not correlate with this shift, but proprioceptive drift did. This suggests that multisensory integration of bodily information drives this shift in space around the body and not feelings of ownership.

Close to me but unreachable: spotting the link between peripersonal space and empathy.

The space surrounding the body [i.e. peripersonal space (PPS)] has a crucial impact on individuals' interactions with the environment. Research showed that the interaction within the PPS increases individuals' behavioral and neural responses. Furthermore, individuals' empathy is affected by the distance between them and the observed stimuli. This study investigated empathic responses to painfully stimulated or gently touched faces presented within the PPS depending on the presence vs absence of a transparent barrier erected to prevent the interaction. To this aim, participants had to determine whether faces were painfully stimulated or gently touched, while their electroencephalographic signals were recorded. Brain activity [i.e. event-related potentials (ERPs) and source activations] was separately compared for the two types of stimuli (i.e. gently touched vs painfully stimulated faces) across two barrier conditions: (i) no-barrier between participants and the screen (i.e. no-barrier) and (ii) a plexiglass barrier erected between participants and the screen (i.e. barrier). While the barrier did not affect performance behaviorally, it reduced cortical activation at both the ERP and source activation levels in brain areas that regulate the interpersonal interaction (i.e. primary, somatosensory, premotor cortices and inferior frontal gyrus). These findings suggest that the barrier, precluding the possibility of interacting, reduced the observer's empathy.

The role of early attachment experiences in modulating defensive peripersonal space.

Selecting appropriate defensive behaviours for threats approaching the space surrounding the body (peripersonal space, PPS) is crucial for survival. The extent of defensive PPS is measured by recording the hand-blink reflex (HBR), a subcortical defensive response. Higher-order cortical areas involved in PPS representation exert top-down modulation on brainstem circuits subserving HBR. However, it is not yet known whether pre-existing models of social relationships (internal working models, IWM) originating from early attachment experiences influence defensive responses. We hypothesized that organized IWM ensure adequate top-down regulation of brainstem activity mediating HBR, whereas disorganized IWM are associated with altered response patterns. To investigate attachment-dependent modulation on defensive responses, we used the Adult Attachment Interview to determine IWM and recorded HBR in two sessions (with or without the neurobehavioral attachment system activated). As expected, the HBR magnitude in individuals with organized IWM was modulated by the threat proximity to the face, regardless of the session. In contrast, for individuals with disorganized IWM, attachment system activation enhances HBR regardless of the threat position, suggesting that triggering emotional attachment experiences magnifies the threatening valence of external stimuli. Our results indicate that the attachment system exerts a strong modulation on defensive responses and the magnitude of PPS.

The relationship between action, social and multisensory spaces.

Several spaces around the body have been described, contributing to interactions with objects (peripersonal) or people (interpersonal and personal). The sensorimotor and multisensory properties of action peripersonal space are assumed to be involved in the regulation of social personal and interpersonal spaces, but experimental evidence is tenuous. Hence, the present study investigated the relationship between multisensory integration and action and social spaces. Participants indicated when an approaching social or non-social stimulus was reachable by hand (reachable space), at a comfortable distance to interact with (interpersonal space), or at a distance beginning to cause discomfort (personal space). They also responded to a tactile stimulation delivered on the trunk during the approach of the visual stimulus (multisensory integration space). Results showed that participants were most comfortable with stimuli outside reachable space, and felt uncomfortable with stimuli well inside it. Furthermore, reachable, personal and interpersonal spaces were all positively correlated. Multisensory integration space extended beyond all other spaces and correlated only with personal space when facing a social stimulus. Considered together, these data confirm that action peripersonal space contributes to the regulation of social spaces and that multisensory integration is not specifically constrained by the spaces underlying motor action and social interactions.

Object coding in peripersonal space depends on object ownership.

Previous studies have shown that objects located in the peripersonal space (PPS) receive enhanced attention, as compared with extrapersonal space (EPS), However, most objects in the environment belong to someone in particular and how object ownership influences object coding in relation to PPS representation is still unclear. In the present study, after having chosen their own mug, participants performed a reachability judgement task of self-owned and other-owned mugs presented at different distances while facing a virtual character. This task was followed, on each trial, by a localisation task in which participants had to indicate where the mug, removed from view, was previously located. The two tasks were separated by a 900-ms visual mask during which the virtual character was unnoticeably shifted by 3° to evaluate the spatial frame-of-reference used. The results showed that self-owned mugs were processed faster than other-owned mugs, but only when located in the PPS. Furthermore, reachability judgements were biased for self-owned mugs, leading to an extension of the PPS representation, especially for participants with a high score on the fantasy scale of Interpersonal Reactivity Index (IRI). Finally, the virtual character shift altered the localisation performance but only for the distant mugs, suggesting a progressive shift from egocentric to allocentric frame-of-reference when moving from the PPS to EPS, irrespective of object ownership. Overall, our data reveal that the representations of ownership and PPS interact to facilitate the processing of manipulable objects, to an extent that depends on individual sensitivity to the social presence of others.

Game theoretical trajectory planning enhances social acceptability of robots by humans.

Since humans and robots are increasingly sharing portions of their operational spaces, experimental evidence is needed to ascertain the safety and social acceptability of robots in human-populated environments. Although several studies have aimed at devising strategies for robot trajectory planning to perform safe motion in populated environments, a few efforts have measured to what extent a robot trajectory is accepted by humans. Here, we present a navigation system for autonomous robots that ensures safety and social acceptability of robotic trajectories. We overcome the typical reactive nature of state-of-the-art trajectory planners by leveraging non-cooperative game theory to design a planner that encapsulates human-like features of preservation of a personal space, recognition of groups, sequential and strategized decision making, and smooth obstacle avoidance. Social acceptability is measured through a variation of the Turing test administered in the form of a survey questionnaire to a pool of 691 participants. Comparison terms for our tests are a state-of-the-art navigation algorithm (Enhanced Vector Field Histogram, VFH) and purely human trajectories. While all participants easily recognized the non-human nature of VFH-generated trajectories, the distinction between game-theoretical trajectories and human ones were hardly revealed. Our results mark a strong milestone toward the full integration of robots in social environments.

Multisensory-driven facilitation within the peripersonal space is modulated by the expectations about stimulus location on the body.

Compelling evidence from human and non-human studies suggests that responses to multisensory events are fastened when stimuli occur within the space surrounding the bodily self (i.e., peripersonal space; PPS). However, some human studies did not find such effect. We propose that these dissonant voices might actually uncover a specific mechanism, modulating PPS boundaries according to sensory regularities. We exploited a visuo-tactile paradigm, wherein participants provided speeded responses to tactile stimuli and rated their perceived intensity while ignoring simultaneous visual stimuli, appearing near the stimulated hand (VTNear) or far from it (VTFar; near the non-stimulated hand). Tactile stimuli could be delivered only to one hand (unilateral task) or to both hands randomly (bilateral task). Results revealed that a space-dependent multisensory enhancement (i.e., faster responses and higher perceived intensity in VTNear than VTFar) was present when highly predictable tactile stimulation induced PPS to be circumscribed around the stimulated hand (unilateral task). Conversely, when stimulus location was unpredictable (bilateral task), participants showed a comparable multisensory enhancement in both bimodal conditions, suggesting a PPS widening to include both hands. We propose that the detection of environmental regularities actively shapes PPS boundaries, thus optimizing the detection and reaction to incoming sensory stimuli.

Interdigitated Columnar Representation of Personal Space and Visual Space in Human Parietal Cortex.

Personal space (PS) is the space around the body that people prefer to maintain between themselves and unfamiliar others. Intrusion into personal space evokes discomfort and an urge to move away. Physiologic studies in nonhuman primates suggest that defensive responses to intruding stimuli involve the parietal cortex. We hypothesized that the spatial encoding of interpersonal distance is initially transformed from purely sensory to more egocentric mapping within human parietal cortex. This hypothesis was tested using 7 Tesla (7T) fMRI at high spatial resolution (1.1 mm isotropic), in seven subjects (four females, three males). In response to visual stimuli presented at a range of virtual distances, we found two categories of distance encoding in two corresponding radially-extending columns of activity within parietal cortex. One set of columns (P columns) responded selectively to moving and stationary face images presented at virtual distances that were nearer (but not farther) than each subject's behaviorally-defined personal space boundary. In most P columns, BOLD response amplitudes increased monotonically and nonlinearly with increasing virtual face proximity. In the remaining P columns, BOLD responses decreased with increasing proximity. A second set of parietal columns (D columns) responded selectively to disparity-based distance cues (near or far) in random dot stimuli, similar to disparity-selective columns described previously in occipital cortex. Critically, in parietal cortex, P columns were topographically interdigitated (nonoverlapping) with D columns. These results suggest that visual spatial information is transformed from visual to body-centered (or person-centered) dimensions in multiple local sites within human parietal cortex.SIGNIFICANCE STATEMENT Recent COVID-related social distancing practices highlight the need to better understand brain mechanisms which regulate "personal space" (PS), which is defined by the closest interpersonal distance that is comfortable for an individual. Using high spatial resolution brain imaging, we tested whether a map of external space is transformed from purely visual (3D-based) information to a more egocentric map (related to personal space) in human parietal cortex. We confirmed this transformation and further showed that it was mediated by two mutually segregated sets of columns: one which encoded interpersonal distance and another that encoded visual distance. These results suggest that the cortical transformation of sensory-centered to person-centered encoding of space near the body involves short-range communication across interdigitated columns within parietal cortex.

A normative model of peripersonal space encoding as performing impact prediction.

Accurately predicting contact between our bodies and environmental objects is paramount to our evolutionary survival. It has been hypothesized that multisensory neurons responding both to touch on the body, and to auditory or visual stimuli occurring near them-thus delineating our peripersonal space (PPS)-may be a critical player in this computation. However, we lack a normative account (i.e., a model specifying how we ought to compute) linking impact prediction and PPS encoding. Here, we leverage Bayesian Decision Theory to develop such a model and show that it recapitulates many of the characteristics of PPS. Namely, a normative model of impact prediction (i) delineates a graded boundary between near and far space, (ii) demonstrates an enlargement of PPS as the speed of incoming stimuli increases, (iii) shows stronger contact prediction for looming than receding stimuli-but critically is still present for receding stimuli when observation uncertainty is non-zero-, (iv) scales with the value we attribute to environmental objects, and finally (v) can account for the differing sizes of PPS for different body parts. Together, these modeling results support the conjecture that PPS reflects the computation of impact prediction, and make a number of testable predictions for future empirical studies.

During the COVID-19 pandemic participants prefer settings with a face mask, no interaction and at a closer distance.

Peripersonal space is the space surrounding our body, where multisensory integration of stimuli and action execution take place. The size of peripersonal space is flexible and subject to change by various personal and situational factors. The dynamic representation of our peripersonal space modulates our spatial behaviors towards other individuals. During the COVID-19 pandemic, this spatial behavior was modified by two further factors: social distancing and wearing a face mask. Evidence from offline and online studies on the impact of a face mask on pro-social behavior is mixed. In an attempt to clarify the role of face masks as pro-social or anti-social signals, 235 observers participated in the present online study. They watched pictures of two models standing at three different distances from each other (50, 90 and 150 cm), who were either wearing a face mask or not and were either interacting by initiating a hand shake or just standing still. The observers' task was to classify the model by gender. Our results show that observers react fastest, and therefore show least avoidance, for the shortest distances (50 and 90 cm) but only when models wear a face mask and do not interact. Thus, our results document both pro- and anti-social consequences of face masks as a result of the complex interplay between social distancing and interactive behavior. Practical implications of these findings are discussed.

Tool-use Extends Peripersonal Space Boundaries in Schizophrenic Patients.

BACKGROUND AND HYPOTHESIS: A primary disruption of the bodily self is considered a core feature of schizophrenia (SCZ). The "disembodied" self might be underpinned by inefficient body-related multisensory integration processes, normally occurring in the peripersonal space (PPS), a plastic sector of space surrounding the body whose extent is altered in SCZ. Although PPS is a malleable interface marking the perceptual border between self and others, no study has addressed the potential alteration of its plasticity in SCZ. We investigated the plasticity of PPS in SCZ patients after a motor training with a tool in the far space. STUDY DESIGN: Twenty-seven SCZ patients and 32 healthy controls (HC) underwent an audio-tactile task to estimate PPS boundary before (Session 1) and after (Session 3) the tool-use. Parameters of PPS, including the size and the slope of the psychometric function describing audio-tactile RTs as a function of the audio-tactile distances, were estimated. STUDY RESULTS: Results confirm a narrow PPS extent in SCZ. Surprisingly, we found PPS expansion in both groups, thus showing for the first time a preserved PPS plasticity in SCZ. Patients experienced a weaker differentiation from others, as indicated by a shallower PPS slope at Session 1 that correlated positively with negative symptoms. However, at Session 3, patients marked their bodily boundary in a steeper way, suggesting a sharper demarcation of PPS boundaries after the action with the tool. CONCLUSIONS: These findings highlight the importance of investigating the multisensory and motor roots of self-disorders, paving the way for future body-centred rehabilitation interventions that could improve patients' altered body boundary.

Embodiment of supernumerary robotic limbs in virtual reality.

The supernumerary robotic limb system expands the motor function of human users by adding extra artificially designed limbs. It is important for us to embody the system as if it is a part of one's own body and to maintain cognitive transparency in which the cognitive load is suppressed. Embodiment studies have been conducted with an expansion of bodily functions through a "substitution" and "extension". However, there have been few studies on the "addition" of supernumerary body parts. In this study, we developed a supernumerary robotic limb system that operates in a virtual environment, and then evaluated whether the extra limb can be regarded as a part of one's own body using a questionnaire and whether the perception of peripersonal space changes with a visuotactile crossmodal congruency task. We found that the participants can embody the extra-limbs after using the supernumerary robotic limb system. We also found a positive correlation between the perceptual change in the crossmodal congruency task and the subjective feeling that the number of one's arms had increased (supernumerary limb sensation). These results suggest that the addition of an extra body part may cause the participants to feel that they had acquired a new body part that differs from their original body part through a functional expansion.