The role of object ownership on online inhibition in peripersonal space.

Peripersonal space (PPS), as opposed to extrapersonal space (EPS), refers to the area surrounding the body within which individuals interact with objects or conspecifics. However, objects in PPS can belong to oneself or to others, which was found to influence how these objects are encoded. We analyzed the performances of motor responses in a reachability judgment task concerning self-owned and other-owned objects (cups) presented in PPS or EPS. EMG activities were recorded on the thumbs (flexor pollicis brevis) to detect correct and erroneous motor activations. Behavioral data showed that motor responses were shorter and longer for self-owned cups compared to other-owned cups in PPS and EPS, respectively. Ten percent of trials showed initial response errors, which were higher in the EPS for self-owned cups and in the PPS for other-owned cups. Eighty-two percent of these errors were corrected online, with corrections being more efficient for self-owned cups in the PPS. Overall, the data revealed that reachability judgments were faster and more accurate in the PPS, with more efficient inhibition processes in the presence of motor errors. Motor selection and correction are thus modulated by the social context of object ownership, highlighting the specific role of the PPS in encoding self-relevant objects for action.

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.

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.

Perception of visual and audiovisual trajectories toward and away from the body in the first postnatal year.

Perceiving motion in depth is important in everyday life, especially motion in relation to the body. Visual and auditory cues inform us about motion in space when presented in isolation from each other, but the most comprehensive information is obtained through the combination of both of these cues. We traced the development of infants' ability to discriminate between visual motion trajectories across peripersonal space and to match these with auditory cues specifying the same peripersonal motion. We measured 5-month-old (n = 20) and 9-month-old (n = 20) infants' visual preferences for visual motion toward or away from their body (presented simultaneously and side by side) across three conditions: (a) visual displays presented alone, (b) paired with a sound increasing in intensity, and (c) paired with a sound decreasing in intensity. Both groups preferred approaching motion in the visual-only condition. When the visual displays were paired with a sound increasing in intensity, neither group showed a visual preference. When a sound decreasing in intensity was played instead, the 5-month-olds preferred the receding (spatiotemporally congruent) visual stimulus, whereas the 9-month-olds preferred the approaching (spatiotemporally incongruent) visual stimulus. We speculate that in the approaching sound condition, the behavioral salience of the sound could have led infants to focus on the auditory information alone, in order to prepare a motor response, and to neglect the visual stimuli. In the receding sound condition, instead, the difference in response patterns in the two groups may have been driven by infants' emerging motor abilities and their developing predictive processing mechanisms supporting and influencing each other.

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.

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.

Demonstrative systems: From linguistic typology to social cognition.

This study explores the connection between language and social cognition by empirically testing different typological analyses of various demonstrative systems. Linguistic typology classifies demonstrative systems as distance-oriented or person-oriented, depending on whether they indicate the location of a referent relative only to the speaker, or to both the speaker and the listener. From the perspective of social cognition, speakers of languages with person-oriented systems must monitor their listener's spatial location in order to accurately use their demonstratives, while speakers of languages with distance-oriented systems can use demonstratives from their own, egocentric perspective. Resolving an ongoing controversy around the nature of the Spanish demonstrative system, the results of Experiment 1 confirmed that this demonstrative system is person oriented, while the English system is distance oriented. Experiment 2 revealed that not all three-way demonstrative systems are person oriented, with Japanese speakers showing sensitivity to the listener's spatial location, while Turkish speakers did not show such an effect in their demonstrative choice. In Experiment 3, Catalan-Spanish bilinguals showed sensitivity to listener position in their choice of the Spanish distal form, but not in their choice of the medial form. These results were interpreted as a transfer effect from Catalan, which revealed analogous results to English. Experiment 4 investigated the use of demonstratives to redirect a listener's attention to the intended referent, which is a universal function of demonstratives that also hinges on social cognition. Japanese and Spanish speakers chose between their proximal and distal demonstratives flexibly, depending on whether the listener was looking closer or further from the referent, whereas Turkish speakers chose their medial form for attention correction. In conclusion, the results of this study support the view that investigating how speakers of different languages jointly use language and social cognition in communication has the potential to unravel the deep connection between these two fundamentally human capacities.

Peripersonal Space from a multisensory perspective: the distinct effect of the visual and tactile components of Visuo-Tactile stimuli.

Peripersonal Space (PPS) is defined as the space close to the body where all interactions between the individual and the environment take place. Behavioural experiments on PPS exploit multisensory integration, using Multisensory Visuo-Tactile stimuli (MVT), whose visual and tactile components target the same body part (i.e. the face, the hand, the foot). However, the effects of visual and tactile stimuli targeting different body parts on PPS representation are unknown, and the relationship with the RTs for Tactile-Only stimuli is unclear. In this study, we addressed two research questions: (1) if the MVT-RTs are independent of Tactile-Only-RTs and if the latter is influenced by time-dependency effects, and (2) if PPS estimations derived from MVT-RTs depend on the location of the Visual or Tactile component of MVTs. We studied 40 right-handed participants, manipulating the body location (right hand, cheek or foot) and the distance of administration. Visual and Tactile components targeted different or the same body parts and were delivered respectively at five distances. RTs to Tactile-Only trials showed a non-monotonic trend, depending on the delay of stimulus administration. Moreover, RTs to Multisensory Visuo-Tactile trials were found to be dependent on the Distance and location of the Visual component of the stimulus. In conclusion, our results show that Tactile-Only RTs should be removed from Visuo-Tactile RTs and that the Visual and Tactile components of Visuo-Tactile stimuli do not necessarily have to target the same body part. These results have a relevant impact on the study of PPS representations, providing new important methodological information.

Peripersonal space around the upper and the lower limbs.

Peripersonal space (PPS), the space closely surrounding the body, is typically characterised by enhanced multisensory integration. Neurophysiological and behavioural studies have consistently shown stronger visuo-tactile integration when a visual stimulus is presented close to the tactually stimulate body part in near space (within PPS) than in far space. However, in the majority of these studies, tactile stimuli were delivered to the upper limbs, torso and face. Therefore, it is not known whether the space surrounding the lower limbs is characterised by similar multisensory properties. To address this question, we asked participants to complete two versions of the classic visuo-tactile crossmodal congruency task in which they had to perform speeded elevation judgements of tactile stimuli presented to the dorsum of the hand and foot while a simultaneous visual distractor was presented at spatially congruent or incongruent locations either in near or far space. In line with existing evidence, when the tactile target was presented to the hand, the size of the crossmodal congruency effect (CCE) decreased in far as compared to near space, suggesting stronger visuo-tactile multisensory integration within PPS. In contrast, when the tactile target was presented to the foot, the CCE decreased for visual distractors in near than far space. These findings show systematic differences between the representation of PPS around upper and lower limbs, suggesting that the multisensory properties of the different body part-centred representations of PPS are likely to depend on the potential actions performed by the different body parts.

Contribution of motor and proprioceptive information to visuotactile interaction in peripersonal space during bike riding.

The space immediately around the body, known as the peripersonal space (PPS), plays an important role in interactions with the environment. Specific representations are reported to be constructed in the brain. PPS expansion reportedly occurs during whole-body self-motions, such as walking; however, little is known regarding how dynamic cues in proprioceptive/motor information contribute to such phenomena. Thus, we investigated this issue using a pedaling bike situation. We defined PPS as the maximum distance at which a visual probe facilitated tactile detection at the chest. Experiment 1 compared two conditions where participants did or did not pedal the bike at a constant speed while observing an optic flow that simulated forward self-motion (pedaling and no pedaling). Experiment 2 investigated the effect of pedal resistances (high and low) while presenting the same optic flow as in Experiment 1. The results revealed that the reaction time (RT) difference (probe RT - baseline RT) was larger for the pedaling than for the no-pedaling condition. However, pedal resistance differences hardly affected the visuotactile interaction, although the participants clearly experienced differences in force. These results suggest that proprioceptive/motor cues can contribute to the modulation of PPS representation, but dynamic information included in these cues may have little influence.

Empathy as a predictor of peripersonal space: Evidence from the crossmodal congruency task.

To investigate whether individual differences in Empathy predict the characteristics of Peripersonal Space (PPS) representations, we asked participants to complete the IRI questionnaire and a visuo-tactile crossmodal congruency task (CCT) as an index of PPS. In the CCT, they responded to the elevation of a tactile target while ignoring a visual distractor presented at the same (i.e. congruent) or different (i.e. incongruent) elevation. The target-distractor distance was also manipulated in depth, with visual distractors randomly presented at near, middle or far locations (0 cm, 25 cm or 50 cm). The near and middle crossmodal congruency effects (CCE) were inversely related to participants' scores on the Empathic Concern sub-scale (EC). Furthermore, the slope of participants' CCE across locations was related to EC scores, with flatter slopes for higher EC individuals. Thus, higher EC individuals showed reduced visuo-tactile integration responses within PPS and a reduced differentiation between PPS and extra-personal space (EPS).

Associative learning in peripersonal space: fear responses are acquired in hand-centered coordinates.

Space coding affects perception of stimuli associated to negative valence: threatening stimuli presented within the peripersonal space (PPS) speed up behavioral responses compared with nonthreatening events. However, it remains unclear whether the association between stimuli and their negative valence is acquired in a body part-centered reference system, a main feature of the PPS coding. Here we test the hypothesis that associative learning takes place in hand-centered coordinates and can therefore remap according to hand displacement. In two experiments, we used a Pavlovian fear-learning paradigm to associate a visual stimulus [light circle, the conditioned stimulus (CS)] with an aversive stimulus (electrocutaneous shock) applied on the right hand only when the CS was displayed close (CS+) but when not far from it (CS-). Measuring the skin conductance response (SCR), we observed successful fear conditioning, with increased anticipatory fear responses associated with CS+. Crucially, experiment I showed a remapping of these responses following hand displacement, with a generalization to both types of CS. Experiment II corroborated and further extended our findings by ruling out the novelty of the experimental context as a driving factor of such modulations. Indeed, fear responses were present only for stimuli within the PPS but not for new stimuli displayed outside the PPS. By revealing a hand-centered (re)mapping of the conditioning effect, these findings indicate that associative learning can arise in hand-centered coordinates. They further suggest that the threatening valence of an object also depends on its basic spatial relationship with our body.NEW & NOTEWORTHY Associative fear learning takes place in hand-centered coordinates. Using a Pavlovian fear-learning paradigm, we show that the anticipatory skin conductance response indicating the association between the negative value and an initially neutral stimulus is acquired and then remapped in space when the stimulated body part moves to a different position. These results demonstrate the relationship between the representation of peripersonal space and the encoding of threatening stimuli. Hypotheses concerning the underlying neural network are discussed.

Time-to-contact perception in the brain.

Time-to-contact (TTC) perception refers to the ability of an observer to estimate the remaining time before an object reaches a point in the environment, and is of crucial importance in daily life. Noninvasive correlational approaches have identified several brain areas sensitive to TTC information. Here we report the results of two studies, including one during an awake brain surgery, that aimed to identify the specific areas causally engaged in the TTC estimation process. In Study 1, we tested 40 patients with brain tumor in a TTC estimation task. The results showed that four of the six patients with impaired performance had tumors in right upper parietal cortex, although this tumoral location represented only six over 40 patients. In Study 2, 15 patients underwent awake brain surgery electrostimulation mapping to examine the implication of various brain areas in the TTC estimation process. We acquired and normalized to MNI space the coordinates of the functional areas that influenced task performance. Our results seem to demonstrate that the early stage of the TTC estimation process involved specific cortical territories in the ventral region of the right intraparietal sulcus. Downstream processing of TTC could also involve the frontal eye field (middle frontal gyrus) related to ocular search. We also found that deactivating language areas in the left hemisphere interfered with the TTC estimation process. These findings demonstrate a fine grained, cortical representation of TTC processing close to the ventral right intraparietal sulcus and complement those described in other human studies.

Disconnected hand avatar can be integrated into the peripersonal space.

Several studies have shown that space immediately surrounding the body, or the peripersonal space is represented differently in the brain from the more distant extra-personal space. Moreover, the boundary of peripersonal space can be extended to space surrounding the tip of a tool held by the hand. However, it is not known if tools need to be connected to the body to modulate the peripersonal space. We used a line bisection task to investigate whether peripersonal space representation surrounds a virtual hand avatar that is disconnected from the body. Healthy participants conducted a line bisection task by responding with either a virtual hand avatar or a laser pointer. The to-be-bisected lines were presented either in peripersonal or extra-personal space. When the lines were placed in extra-personal space, the virtual hand avatar was presented near the line such that the hand avatar was far from participants and disconnected from their bodies. Results indicated a shift in the line bisection bias from the left to the right as the line presentation distance increased when using the laser pointer, whereas no shift in bias was observed when using the virtual hand avatar. This result indicates that objects resembling human hands presented even at a distance and disconnected from the body can be integrated into the peripersonal space, which suggests that peripersonal space representation is more flexible than previously reported.

Does response facilitation to visuo-tactile stimuli around a remote-controlled hand avatar reflect peripersonal space or attentional bias?

People react faster to visuo-tactile stimuli presented near the body (i.e., in peripersonal space) than to tactile stimuli presented alone. This multi-sensory facilitation effect has been used as a measurement of peripersonal space. Previous research has reported that peripersonal space representations can be modulated by actively using hand-held tools or disconnected hand avatars. However, previous research has ignored the possibility that the attentional effect of active tool use could affect multi-sensory facilitation. In the present study, we delivered tactile stimuli to participants' left or right hand concurrently with visual stimuli presented near a virtual hand avatar operated by the movements of participants' left or right hand, which was shown far in a virtual environment and disconnected from the body. Participants reacted to tactile stimuli while ignoring the visual stimuli. The results indicated a multi-sensory facilitation effect when tactile stimuli were delivered to the hand used to operate the hand avatar. In contrast, the facilitation was not observed when the tactile stimuli were delivered to the hand that is not operating the hand avatar. These results suggest that the strength of the multi-sensory facilitation effect differed across conditions, even though the visual attention captured around the hand avatar was controlled across conditions. We concluded that the modulation of peripersonal space resulting from using tools or avatars is nearly independent of visual attention captured around tools or avatars.

Opposing force fields induce direction-specific sensorimotor adaptation but a non-specific perceptual shift consistent with a contraction of peripersonal space representation.

Most of our daily interactions with objects occur in the space immediately surrounding the body, i.e. the peripersonal space. The peripersonal space is characterized by multisensory processing of objects which are coded in terms of potential actions, specifying for instance whether objects are within reach or not. Our recent work suggested a link between exposure to a new force field, which changed the effector dynamics, and the representation of peripersonal space. To better understand the interplay between the plasticity of the motor system and peripersonal space representation, the present study examined whether changing the direction of the force field specifically modified the perception of action boundaries. Participants seated at the centre of an experimental platform estimated visual targets' reachability before and after adapting upper-limb reaching movements to the Coriolis force generated by either clockwise or counter clockwise rotation of the platform (120°/s). Opposite spatial after-effects were observed, showing that force-field adaptation depends on the direction of the rotation. In contrast, perceived action boundaries shifted leftward following exposure to the new force field, regardless of the direction of the rotation. Overall, these findings support the idea that abrupt exposure to a new force field results in a direction-specific updating of the central sensorimotor representations underlying the control of arm movements. Abrupt exposure to a new force field also results in a nonspecific shift in the perception of action boundaries, which is consistent with a contraction of the peripersonal space. Such effect, which does not appear to be related to state anxiety, could be related to the protective role of the peripersonal space in response to the uncertainty of the sensorimotor system induced by the abrupt modification of the environment.

Expansion of space for visuotactile interaction during visually induced self-motion.

Peripersonal space (PPS), which refers to space immediately around an individual's body, plays an important role in interacting with external objects and avoiding unsafe situations. Studies suggest that, during self-motion perception, PPS expands in the direction in which a person perceives himself/herself to be traveling. In the present study, we built on this by investigating, using visually induced self-motion (vection), how visual self-motion information modulates PPS representation. In our experiment, large-field visual motion was presented through a head-mounted display that caused observers to perceive themselves as moving forward in a tunnel (LF condition). To clarify the effects of self-motion information, we compared the findings for this condition with those of another condition, in which small-field visual motion was presented; here, only the central visual field represented motion, which caused the observers to perceive relatively little self-motion (SF condition). Two speeds were tested for both conditions: 1.5 m/s and 6.0 m/s. For measurement, we used a visuotactile-interaction task in which participants, while observing a visual probe object approaching from various distances, were instructed to press a response key as soon as they detected tactile stimuli delivered to their chest. We measured the distance at which the visual approaching probe object facilitated tactile detection (visual-facilitation effect); this was determined through comparisons with trials when no probe was presented. The results showed that the visual facilitation effects were observed for larger distance in the LF than SF conditions, irrespective of tested speeds. These results suggest that visual self-motion information can modulate PPS representation. This finding fits well with the view that PPS representation contributes to protecting the body from potential threats in the environment.

Can I put myself in your shoes? Sharing peripersonal space reveals the simulation of the action possibilities of others.

The representation of the reachable space near the body or peripersonal space (PPS) plays an important role in action with objects and interaction with other people. Here we explored whether the representation of other people's PPS involves a simulation of their action possibility depending on the distance between the observer's body and the other person. We hypothesized that a simulation of action possibilities of others should be activated when they share the PPS of observers. In two experiments using Immersive Virtual Reality, the avatars were shown with their arms free or blocked (motor interference) at a reachable (i.e., Near/shared PPS) or not-reachable (i.e., Far/not-shared PPS) distance by the participants. Participants had to locate a glass in relation to the midline of the avatar's body, i.e., assuming the perspective of the avatar. To directly compare the representation of one's own and others' PPS, in Experiment 1 participants had to locate the glass in relation to themselves with their arms free or blocked. The glass was always within the PPS of either participants or avatars. The results showed that the localization of the glass from the avatar's perspective was slower with the blocked than free arm only when the avatar was in the near shared PPS, similar to the localization from one's own first-person perspective. Instead, there was no effect of motor interference in the far, not-shared PPS. This suggests that the representation of other people's PPS can involve motor simulation mechanisms when we share a common peripersonal area.

Hand blink reflex in virtual reality: The role of vision and proprioception in modulating defensive responses.

Our research focused on the role of vision and proprioception in modulating a defensive reflex (hand blink reflex, HBR) whose magnitude is enhanced when the threatened hand is inside the peripersonal space of the face. We capitalized on virtual reality, which allows dissociating vision and proprioception by presenting a virtual limb in congruent/incongruent positions with respect to the participants' limb. In experiment 1, participants placed their own stimulated hand in far/near positions with respect to their face (postural manipulation task), while observing a virtual empty scenario. Vision was not informative, but the HBR was significantly enhanced in near compared with far position, suggesting that proprioception is sufficient for the HBR modulation to occur. In experiment 2, participants did not perform the postural manipulation but they (passively) observed the avatar's virtual limb performing it. Proprioceptive signals were not informative, but the HBR was significantly enhanced when the observed virtual limb was near to the face, suggesting that visual information plays a role in modulating the HBR. In experiment 3, both participants and avatar performed the postural manipulation, either congruently (both of them far/near) or incongruently (one of them far, the other near). The HBR modulation was present only in congruent conditions. In incongruent conditions, the conflict between vision and proprioception confounded the system, abolishing the difference between far and near positions. Taken together, these findings promote the view that observing a virtual limb modulates the HBR, providing also new evidence on the role of vision and proprioception in modulating this subcortical reflex.

The spatial effect of fearful faces in the autonomic response.

Peripersonal space (PPS) corresponds to the space around the body and it is defined by the location in space where multimodal inputs from bodily and external stimuli are integrated. Its extent varies according to the characteristics of external stimuli, e.g., the salience of an emotional facial expression. In the present study, we investigated the psycho-physiological correlates of the extension phenomenon. Specifically, we investigated whether an approaching human face showing either an emotionally negative (fearful) or positive (joyful) facial expression would differentially modulate PPS representation, compared to the same face with a neutral expression. To this aim, we continuously recorded the skin conductance response (SCR) of 27 healthy participants while they watched approaching 3D avatar faces showing fearful, joyful or neutral expressions, and then pressed a button to respond to tactile stimuli delivered on their cheeks at three possible delays (visuo-tactile trials). The results revealed that the SCR to fearful faces, but not joyful or neutral faces, was modulated by the apparent distance from the participant's body. SCR increased from very far space to far and then to near space. We propose that the proximity of the fearful face provided a cue to the presence of a threat in the environment and elicited a robust and urgent organization of defensive responses. In contrast, there would be no need to organize defensive responses to joyful or neutral faces and, as a consequence, no SCR differences were found across spatial positions. These results confirm the defensive function of PPS.