Peripersonal Space Representation Research Articles

Vestibular modulation of peripersonal space boundaries.

Human-environment interactions are mediated through the body and occur within the peripersonal space (PPS), the space immediately adjacent to and surrounding the body. The PPS is taken to be a critical interface between the body and the environment, and indeed, body-part specific PPS remapping has been shown to depend on body-part utilization, such as upper limb movements in otherwise static observers. How vestibular signals induced by whole-body movement contribute to PPS representation is less well understood. In a series of experiments, we mapped the spatial extension of the PPS around the head while participants were submitted to passive whole-body rotations inducing vestibular stimulation. Forty-six participants, in three experiments, executed a tactile detection reaction time task while task-irrelevant auditory stimuli approached them. The maximal distance at which the auditory stimulus facilitated tactile reaction time was taken as a proxy for the boundary of peri-head space. The present results indicate two distinct vestibular effects. First, vestibular stimulation speeded tactile detection indicating a vestibular facilitation of somatosensory processing. Second, vestibular stimulation modulated audio-tactile interaction of peri-head space in a rotation direction-specific manner. Congruent but not incongruent audio-vestibular motion stimuli expanded the PPS boundary further away from the body as compared to no rotation. These results show that vestibular inputs dynamically update the multisensory delineation of PPS and far space, which may serve to maintain accurate tracking of objects close to the body and to update spatial self-representations.

Tool use changes the spatial extension of the magnetic touch illusion.

Characterizing the brain mechanisms that allow humans to use tools to interact with the environment is a major goal in neuroscience. It has been proposed that handheld tools are incorporated into the multisensory representation of the body and its surrounding (peripersonal) space, underlying our remarkable tool use ability. One single-cell recording study in tool-using monkeys provided qualitative support for this hypothesis, and the results from a vast number of human studies employing different experimental paradigms have been ambiguous. Here, we made use of the recently reported magnetic touch illusion-a perceptual correlate of peripersonal space-to examine the effect of tool use on the representation of visuotactile peripersonal space. The results showed that active tool use leads to an extension of the "illusion volume" around the entire length of a tool, which was significantly greater compared with a manual control task. These findings support the notion that the multisensory representation of peripersonal space is extended to incorporate handheld tools and provide a three-dimensional estimation of this remapping process. (PsycINFO Database Record

Anisotropy of lateral peripersonal space is linked to handedness.

The space immediately surrounding our bodies, i.e., peripersonal space (PPS), is a critical area for the interaction with the external world, be it to deal with imminent threat or to attain objects of interest. In the brain, a dedicated system codes PPS in motor terms for the purpose of action. Yet, humans have asymmetric motor abilities: the dominant hand has an advantage in term of movements' precision and reaction time. Furthermore, spatial attention is asymmetric and seems to be linked to a right hemispheric dominance for spatial processing. Here, we tested whether handedness and attentional asymmetries impact the detection of a tactile stimulus when an irrelevant auditory stimulus is looming towards the individual from the right or left hemispace. We examined the distance at which sound started speeding up tactile detection to estimate the morphometry of peri-trunk PPS. Our results show that right-handers' PPS is larger in the left than in the right hemispace, whereas left-handers' PPS is symmetric. The expansion of right-handers' PPS on the side of the non-dominant hand is coherent with a protective function of PPS. Left-handers' symmetric PPS can be related to the symmetric request of their motor abilities induced by living in a right-handers' world. These findings reveal that PPS is not uniform and suggest that general mechanisms of spatial processing as well as motor skills could play a role in the representation of peri-trunk PPS.

The power of the imagination to affect peripersonal space representations

ABSTRACTIt is known that visual processing is altered for objects near the hands as well as for objects near the imagined position of the hands, indicating that the imagination can be used to remap peri-hand space. Little is known, however, about the physical conditions that allow for this remapping. In the present study, participants in one experiment performed visual searches through displays that were beyond reach while imagining that their hands were near the display. This imagined impossible posture slowed visual search rates, showing that imagination effectively remapped peri-hand space to be near the display. In another experiment, participants searched displays they were holding, but sometimes imagined their hands to be far away. This produced faster search rates, revealing a remapping of peri-hand space away from the monitor. The results provide new insights into the mechanisms involved in the representation of peri-hand space and about the power of imagined actions to influence body representations.

Peripersonal space representation develops independently from visual experience

Our daily-life actions are typically driven by vision. When acting upon an object, we need to represent its visual features (e.g. shape, orientation, etc.) and to map them into our own peripersonal space. But what happens with people who have never had any visual experience? How can they map object features into their own peripersonal space? Do they do it differently from sighted agents? To tackle these questions, we carried out a series of behavioral experiments in sighted and congenitally blind subjects. We took advantage of a spatial alignment effect paradigm, which typically refers to a decrease of reaction times when subjects perform an action (e.g., a reach-to-grasp pantomime) congruent with that afforded by a presented object. To systematically examine peripersonal space mapping, we presented visual or auditory affording objects both within and outside subjects’ reach. The results showed that sighted and congenitally blind subjects did not differ in mapping objects into their own peripersonal space. Strikingly, this mapping occurred also when objects were presented outside subjects’ reach, but within the peripersonal space of another agent. This suggests that (the lack of) visual experience does not significantly affect the development of both one’s own and others’ peripersonal space representation.

Peripersonal space boundaries around the lower limbs

Neurophysiological investigations in non-human primates have shown that bi- and tri-modal fronto-parietal neurons exist that respond to touch on the body and visual (and/or auditory) stimuli near the body. The receptive fields of these neurons extend into space around the body, producing a zone wherein multisensory information is readily integrated. This space around the body, known as peripersonal space (PPS), has also been investigated behaviourally in humans. Some studies have focused on how far into depth the spatial boundaries of PPS extend. Most of these investigations have focused on the upper body (e.g., hands, face, trunk), while little is known about the size of PPS for the lower body (i.e. legs and feet). Thus, the aim of the current study was to delineate a PPS boundary around the lower limbs in healthy participants using a multisensory interaction task. Participants made speeded responses to the presence of vibrations applied to the toes while a task-irrelevant visual stimulus approached towards (Experiment 1) or receded from (Experiment 2) the feet. Participants responded significantly faster to tactile stimuli when the visual stimulus was within approximately 73 cm from the feet, but only when it approached (and not receded from) the legs. This is the first study, to our knowledge, to outline the size of PPS for the lower limbs. These findings could provide insight into the mechanisms underlying multisensory integration in the lower limbs, and add to the current body of knowledge on PPS representations.

Social perception of others shapes one's own multisensory peripersonal space

The perception of our self is not restricted to our physical boundaries, but it extends beyond the body to incorporate the space where individual–environment interactions occur, i.e., the peripersonal space (PPS). PPS is generally conceived as a low-level multisensory-motor interface mediating hand–object interactions. Recent studies, however, showed that PPS representation is affected by higher-level cognitive factors. Here we asked whether the multisensory representation of PPS is influenced by high-level mechanisms implied in social interactions, such as the social perception of others. To this aim, in Experiment 1, we developed and validated a new multisensory interaction task in mixed reality (i.e., the Social PPS task). This task allows measuring the boundaries of PPS between one self and another person in a fully controlled, yet highly ecological, set-up. In the Experiment 2, we used this task to measure how participants' PPS varied when facing another person. The social perception of this person was manipulated via a classic social psychology procedure, so that, in two conditions, she was perceived either as a moral or an immoral character. We found that PPS representation is sensitive to the social perception of the other, being more extended when participants were facing a moral than when facing an immoral person. This effect was specific for social context, as no change in PPS was found if participants were facing an object, instead of the person. Interestingly, the social manipulation affected also attitude, identification, willingness to interact with the other, so as interpersonal distance. Together these findings show that social perception of others affects both the psychological representation of the others in relation to oneself and the multisensory representations of the space between oneself and the other, offering new insights about the role of social cognition in body representation.

Reachability judgement in optic ataxia: Effect of peripheral vision on hand and target perception in depth

The concept of peripersonal space was first proposed by Rizzolatti, Scandolara, Matelli, and Gentilucci (1981), who introduced the term to highlight the close links between somatosensory and visual processing for stimuli close to the body and suggested that this near-body space could in fact be characterized as an action space (Rizzolatti, Fadiga, Fogassi, & Gallese, 1997). Supporting this idea, patients with right hemisphere lesions have been described as impaired in performing actions towards objects and in perceiving their location − but only when the objects were presented within arm's reach (Bartolo, Carlier, Hassaini, Martin, & Coello, 2014; Brain, 1941). Whether the deficit of optic ataxia patients in processing target locations for action has an effect on the representation of peripersonal space has never been explored. The present study highlights optic ataxia patients' specific difficulties in processing hand-to-target distances in a motor task and in a perceptual task requiring identification of what is reachable in the visual environment. The difficulties are especially evident when both the target and the hand are perceived in the visual periphery. Indeed, when patient I.G. was able to fixate the target, her reaching accuracy and her perception of reachable space both largely improved. Furthermore, the difficulties were enhanced when the hand and the target were both in the lower visual field (in a fixed-far condition vs a fixed-near condition). This novel up-down dimension of optic ataxia fits with the larger representation of the lower visual field in the posterior parietal cortex (Pitzalis et al., 2013; Previc, 1990).

Learning peripersonal space representation in a humanoid robot with artificial skin

The peripersonal space (PPS) is a model of the nearby space that is of special relevance for the life of any complex animal. It is defined as the space surrounding our bodies, and is best described as a multisensory and sensorimotor interface with a fundamental role in the sensory guidance of actions. Owing to it is the ability to perform timely and appropriate actions toward objects located in the nearby space, which is critical for the survival of every animal. Depending on context, these actions may constitute an approaching or an avoidance behavior. In the case of defensive behavior, this creates a "margin of safety" around the body, such as the flight zone of grazing animals or the multimodal attentional space that surrounds the skin in humans (Graziano & Cooke, 2006). Peripersonal space thus deserves special attention and probably justifies the specific neural circuitry devoted to its representation. The brain has to dynamically integrate information coming from several modalities being these motoric, visual, auditory or somatosensory (Graziano & Cooke, 2006; Holmes & Spence, 2004). In animals and humans, PPS representations are gradually formed through physical interaction with the environment and in a complex interplay of body growth and neural maturation processes. Self-touch (also called double-touch) is presu mably one of the behaviors that impact the formation of multimodal body representations (Rochat, 1998). They occur across different motor and sensory modalities, with the motor/proprioceptive and tactile starting already in prenatal stage. Vision is presumably incorporated later, during the first months after birth hand in hand with the maturation of the visual system. Perhaps even later, contingencies will encompass external objects.

Learning peripersonal space representation in a humanoid robot with artificial skin

Learning peripersonal space representation in a humanoid robot with artificial skin

Mechanical pinprick pain in patients with unilateral spatial neglect: The influence of space representation on the perception of nociceptive stimuli.

Crossing the hands over the midline can reduce the perceived intensity of nociceptive stimuli applied onto the hands. It remains unclear to what extent intact representation of peripersonal space influences this effect. Here we used the crossed-hands paradigm in patients with unilateral spatial neglect, a neuropsychological condition characterized by the inability to detect, attend and respond to contralesional (most often left) stimuli, and spared ability to process stimuli in the non-affected space. Sixteen post-stroke patients without unilateral neglect and 11 patients with unilateral spatial neglect received punctate mechanical pinprick stimuli onto their crossed or uncrossed hands. We tested: (i) whether deficits in space representation reduce the possibility of observing 'crossed-hands analgesia', and; (ii) whether placing the contralesional hand, normally lying in the affected space in the healthy space would increase the number of detected stimuli. Our results showed that neglect patients did not exhibit 'crossed-hands' analgesia, but did not provide strong evidence for an improvement in the number of detected stimuli when the contralesional hand was in the healthy space. These findings uphold the notion that the perception of nociceptive stimuli is modulated by the relative position of the hands in space, but raise questions about the conditions under which these effects may arise. We show that deficits in space representation can influence the processing of mechanical pinprick stimuli. Our results raise several questions on the mechanisms underlying these effects, which are relevant for the clinical practice.

Peripersonal Space and Margin of Safety around the Body: Learning Visuo-Tactile Associations in a Humanoid Robot with Artificial Skin.

This paper investigates a biologically motivated model of peripersonal space through its implementation on a humanoid robot. Guided by the present understanding of the neurophysiology of the fronto-parietal system, we developed a computational model inspired by the receptive fields of polymodal neurons identified, for example, in brain areas F4 and VIP. The experiments on the iCub humanoid robot show that the peripersonal space representation i) can be learned efficiently and in real-time via a simple interaction with the robot, ii) can lead to the generation of behaviors like avoidance and reaching, and iii) can contribute to the understanding the biological principle of motor equivalence. More specifically, with respect to i) the present model contributes to hypothesizing a learning mechanisms for peripersonal space. In relation to point ii) we show how a relatively simple controller can exploit the learned receptive fields to generate either avoidance or reaching of an incoming stimulus and for iii) we show how the robot can select arbitrary body parts as the controlled end-point of an avoidance or reaching movement.

Keeping you at arm's length: modifying peripersonal space influences interpersonal distance.

Peripersonal space represents the area around the body where objects are coded in motor terms for the purpose of voluntary goal-directed actions. Previous studies have suggested that peripersonal space is also a safe space linked with our private area, influencing interpersonal space in social contexts. However, whether these two spaces rely on similar embodied processes remains an open issue. In the present study, participants observed a point-light walker (PLW) approaching them from different directions and passing near them at different distances from their right or left shoulder. While approaching, the PLW disappeared at a distance of 2m and the task for the participants was to estimate if the interpersonal distance, at the time the PLW would have reached their level, was comfortable or not. Between two sessions of comfort judgments, the participants manipulated a 70cm tool entailing an extension of peripersonal space, or a 10cm tool entailing no extension of peripersonal space. The results revealed that the comfortable interpersonal distance was larger when the PLW crossed the mid-sagittal plane of the participants than when it approached them laterally, with a concomitant increase of response time. After participants manipulated the long tool, comfortable interpersonal distance increased, but predominantly when the PLW trajectory implied crossing the participants' mid-sagittal plane. This effect was not observed when participants manipulated the short tool. Two control tasks showed that using the long tool modified the reachability (control 1), but not the time to passage (control 2) estimates of PLW stimuli, suggesting that tool use extended peripersonal space without changing perceived visual distances. Overall, the data show that comfortable interpersonal distance is linked to the representation of peripersonal space. As a consequence, increasing peripersonal space through tool use has the immediate consequence that comfortable interpersonal distance from another person also increases, suggesting that interpersonal-comfort space and peripersonal-reaching space share a common motor nature.

Multisensory simultaneity judgment and proximity to the body

The integration of information across different sensory modalities is known to be dependent upon the statistical characteristics of the stimuli to be combined. For example, the spatial and temporal proximity of stimuli are important determinants with stimuli that are close in space and time being more likely to be bound. These multisensory interactions occur not only for singular points in space/time, but over “windows” of space and time that likely relate to the ecological statistics of real-world stimuli. Relatedly, human psychophysical work has demonstrated that individuals are highly prone to judge multisensory stimuli as co-occurring over a wide range of time—a so-called simultaneity window (SW). Similarly, there exists a spatial representation of peripersonal space (PPS) surrounding the body in which stimuli related to the body and to external events occurring near the body are highly likely to be jointly processed. In the current study, we sought to examine the interaction between these temporal and spatial dimensions of multisensory representation by measuring the SW for audiovisual stimuli through proximal–distal space (i.e., PPS and extrapersonal space). Results demonstrate that the audiovisual SWs within PPS are larger than outside PPS. In addition, we suggest that this effect is likely due to an automatic and additional computation of these multisensory events in a body-centered reference frame. We discuss the current findings in terms of the spatiotemporal constraints of multisensory interactions and the implication of distinct reference frames on this process.

Body part-centered and full body-centered peripersonal space representations.

Dedicated neural systems represent the space surrounding the body, termed Peripersonal space (PPS), by integrating visual or auditory stimuli occurring near the body with somatosensory information. As a behavioral proxy to PPS, we measured participants’ reaction time to tactile stimulation while task-irrelevant auditory or visual stimuli were presented at different distances from their body. In 7 experiments we delineated the critical distance at which auditory or visual stimuli boosted tactile processing on the hand, face, and trunk as a proxy of the PPS extension. Three main findings were obtained. First, the size of PPS varied according to the stimulated body part, being progressively bigger for the hand, then face, and largest for the trunk. Second, while approaching stimuli always modulated tactile processing in a space-dependent manner, receding stimuli did so only for the hand. Finally, the extension of PPS around the hand and the face varied according to their relative positioning and stimuli congruency, whereas the trunk PPS was constant. These results suggest that at least three body-part specific PPS representations exist, differing in extension and directional tuning. These distinct PPS representations, however, are not fully independent from each other, but referenced to the common reference frame of the trunk.

Tactile representation of the head and shoulders assessed by fMRI in the nonhuman primate.

In nonhuman primates, tactile representation at the cortical level has mostly been studied using single-cell recordings targeted to specific cortical areas. In this study, we explored the representation of tactile information delivered to the face or the shoulders at the whole brain level, using functional magnetic resonance imaging (fMRI) in the nonhuman primate. We used air puffs delivered to the center of the face, the periphery of the face, or the shoulders. These stimulations elicited activations in numerous cortical areas, encompassing the primary and secondary somatosensory areas, prefrontal and premotor areas, and parietal, temporal, and cingulate areas as well as low-level visual cortex. Importantly, a specific parieto-temporo-prefrontal network responded to the three stimulations but presented a marked preference for air puffs directed to the center of the face. This network corresponds to areas that are also involved in near-space representation, as well as in the multisensory integration of information at the interface between this near space and the skin of the face, and is probably involved in the construction of a peripersonal space representation around the head.

Investigating the spatial and temporal modulation of visuotactile interactions in older adults.

Previous research has shown that spatially and temporally disparate multisensory events are more likely to interact for older adults. For visuotactile interactions, this suggests that the representation of peripersonal space is expanded and temporal perception within this space is less precise. Previously, visuotactile space has been found to expand horizontally into the opposite hemispace, and here we sought to replicate and extend this by exploring both horizontal and vertical space from the hand. Moreover, we investigated whether both spatial and temporal domains are affected for an individual, which have previously been measured using distinct tasks and different participants. We presented a modified cross-modal congruency task (Poole et al. in Multisens Res. doi: 10.1163/22134808-00002475 , 2015a) to thirty older participants (age range 65-85 years), with unisensory tactile performance equated for each individual. For the temporal manipulation, the timings of visual distractors and tactile targets were offset. For the spatial manipulation, visual distractors were presented from multiple positions in ipsilateral and contralateral hemispaces. Whilst the temporal modulation of visuotactile interactions for older adults was equivalent to that observed in young adults, spatial modulation was reduced; significant visuotactile interactions were observed for visual distractors presented in the same and opposite hemispace to the stimulated hand, in the lower visual field. This suggests an expanded representation of visuotactile space surrounding the hand in older adults, which occurs horizontally into the contralateral hemispace only, rather than expanding both vertically and horizontally. This is likely to have consequences for perception of space and goal-directed action in ageing.

Editorial: Taking a hands-on approach: current perspectives on the effect of hand position on vision.

Over the past 10 years, perception scientists have uncovered a surprising connection between people's vision and their hands. There is now compelling evidence that how people perceive, attend to, think about, and remember visual information depends on how close they have their hands to that information. With their hands near, people perform figure-ground assignment more efficiently, parse temporally adjacent events more precisely, and hold more information in visual working memory. Near their hands, people also detect sudden visual onsets more quickly, but search through arrays of items more slowly, and take longer to switch between different ways of interpreting the same perceptual content (e.g., “seeing the forest” vs. “seeing the trees”). These are but some of the ways in which visual processing changes when people's hands are in proximity of viewed information—a host of effects that we refer to here, collectively, as hand-altered vision (HAV). The first decade of research into HAV has generated a substantial amount of new knowledge, which we recently reviewed in contemporaneous papers (Tseng et al., 2012; Brockmole et al., 2013). We subsequently established this Research Topic as a bridge to the next era of HAV research, through which we aimed to gather perspectives from across the research literatures on human action and peripersonal space representation. All told, the work here consists of 12 articles from 34 researchers who represent 23 institutions worldwide. Thanks to the efforts of our contributors, our scientific understanding of HAV has progressed along several major channels.

The wheelchair as a full-body tool extending the peripersonal space

Dedicated multisensory mechanisms in the brain represent peripersonal space (PPS), a limited portion of space immediately surrounding the body. Previous studies have illustrated the malleability of PPS representation through hand-object interaction, showing that tool use extends the limits of the hand-centered PPS. In the present study we investigated the effects of a special tool, the wheelchair, in extending the action possibilities of the whole body. We used a behavioral measure to quantify the extension of the PPS around the body before and after Active (Experiment 1) and Passive (Experiment 2) training with a wheelchair and when participants were blindfolded (Experiment 3). Results suggest that a wheelchair-mediated passive exploration of far space extended PPS representation. This effect was specifically related to the possibility of receiving information from the environment through vision, since no extension effect was found when participants were blindfolded. Surprisingly, the active motor training did not induce any modification in PPS representation, probably because the wheelchair maneuver was demanding for non-expert users and thus they may have prioritized processing of information from close to the wheelchair rather than at far spatial locations. Our results suggest that plasticity in PPS representation after tool use seems not to strictly depend on active use of the tool itself, but is triggered by simultaneous processing of information from the body and the space where the body acts in the environment, which is more extended in the case of wheelchair use. These results contribute to our understanding of the mechanisms underlying body–environment interaction for developing and improving applications of assistive technological devices in different clinical populations.

Extending peripersonal space representation without tool-use: evidence from a combined behavioral-computational approach.

Stimuli from different sensory modalities occurring on or close to the body are integrated in a multisensory representation of the space surrounding the body, i.e., peripersonal space (PPS). PPS dynamically modifies depending on experience, e.g., it extends after using a tool to reach far objects. However, the neural mechanism underlying PPS plasticity after tool use is largely unknown. Here we use a combined computational-behavioral approach to propose and test a possible mechanism accounting for PPS extension. We first present a neural network model simulating audio-tactile representation in the PPS around one hand. Simulation experiments showed that our model reproduced the main property of PPS neurons, i.e., selective multisensory response for stimuli occurring close to the hand. We used the neural network model to simulate the effects of a tool-use training. In terms of sensory inputs, tool use was conceptualized as a concurrent tactile stimulation from the hand, due to holding the tool, and an auditory stimulation from the far space, due to tool-mediated action. Results showed that after exposure to those inputs, PPS neurons responded also to multisensory stimuli far from the hand. The model thus suggests that synchronous pairing of tactile hand stimulation and auditory stimulation from the far space is sufficient to extend PPS, such as after tool-use. Such prediction was confirmed by a behavioral experiment, where we used an audio-tactile interaction paradigm to measure the boundaries of PPS representation. We found that PPS extended after synchronous tactile-hand stimulation and auditory-far stimulation in a group of healthy volunteers. Control experiments both in simulation and behavioral settings showed that the same amount of tactile and auditory inputs administered out of synchrony did not change PPS representation. We conclude by proposing a simple, biological-plausible model to explain plasticity in PPS representation after tool-use, which is supported by computational and behavioral data.