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What is Peripersonal Space? An Examination of Unresolved Empirical Issues

and Emerging Findings
Article in Wiley interdisciplinary reviews. Cognitive science · July 2018

DOI: 10.1002/wcs.1472
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Received: 17 December 2017 Revised: 7 June 2018 Accepted: 8 June 2018
DOI: 10.1002/wcs.1472
ADVANCED REVIEW
What is peripersonal space? An examination of unresolved

empirical issues and emerging findings
Samuel B. Hunley | Stella F. Lourenco
Department of Psychology, Emory University,

Atlanta, Georgia Findings from diverse fields of study, including neuroscience, psychology, zoo-
Correspondence logy, and sociology, demonstrate that human and non-human primates maintain a
Stella F. Lourenco, Department of Psychology, representation of the space immediately surrounding the body, known as periperso-
Emory University, 36 Eagle Row, Atlanta,
nal space (PPS). However, progress in this field has been hampered by the lack of
GA 30322.
Email: stella.lourenco@emory.edu an agreed upon definition of PPS. Since the beginning of its formal study, scientists
Funding information have argued that PPS plays a crucial role in both defensive and non-defensive
Emory University actions. Yet consensus is lacking about the cognitive and neural instantiation of
these functions. In particular, researchers have begun to ask whether a single,
unified system of spatial-attentional resources supports both the defensive and non-
defensive functions of PPS or, rather, whether there are multiple, independent sys-
tems. Moreover, there are open questions about the specificity of PPS. For exam-
ple: Does PPS dissociate from other well-known phenomena such as personal
space and the body schema? Finally, emerging research has brought attention to
important questions about individual differences in the flexibility of PPS and the
distribution of PPS in front compared to behind the body. In this advanced review,
we shed light on questions about the nature of PPS, offering answers when the
research permits or providing recommendations for achieving answers in future
research. In so doing, we lay the groundwork for a comprehensive definition
of PPS.
This article is categorized under:
Cognitive Biology > Evolutionary Roots of Cognition
Psychology > Attention
Psychology > Perception and Psychophysics
Neuroscience > Plasticity
KEYWORDS
body schema, individual differences, peripersonal space, personal space, spatial

representation
1 | INTRODUCTION
Human and non-human primates maintain a representation of the space surrounding the body—a region of space known as
peripersonal space (hereafter, PPS) (Brain, 1941; Hall, 1966; Previc, 1998; Rizzolatti, Scandolara, Matelli, & Gentilucci,
1981). PPS, which is differentiated from the space farther away from the body (a separate region of space known as extraper-
sonal space), plays a crucial role in coordinating one's actions for both defensive and non-defensive purposes (Cléry, Guip-
poni, Wardak, & Hamed, 2015; de Vignemont & Iannetti, 2015; Graziano & Cooke, 2006; Graziano & Gross, 1998;
Lourenco, Longo, & Pathman, 2011). The successful coordination of these actions, whether they involve reaching for one's
WIREs Cogn Sci. 2018;e1472. wires.wiley.com/cogsci © 2018 Wiley Periodicals, Inc. 1 of 17

https://doi.org/10.1002/wcs.1472
2 of 17 HUNLEY AND LOURENCO
coffee mug or avoidance of an attacker, requires knowing the object's or other organism's location relative to one's self, specif-
ically one's body. Yet, despite decades of research from diverse fields such as neuroscience, psychology, zoology, and sociol-
ogy, there are open questions about the system (or systems) that undergird PPS with both defensive and non-defensive
functions; the relation between PPS and other representations of space and the body; and the influence of social-affiliative fac-
tors on PPS.
These questions arise in large part because of definitional issues that have plagued the PPS literature. Scientific progress
necessitates definitions that allow for accurate and consistent measurement across labs and studies (Tal, 2015). In the case of
PPS, achieving consensus on a definition has been difficult because of the complexity of the construct. Early research on PPS
tended to focus on isolated functions or individual characteristics of PPS. Although such an approach certainly has its advan-
tages, it can lead to confusion about what does, and does not, constitute PPS. More recent approaches have made greater
efforts to account for the multi-dimensional nature of PPS, and to this end, a comprehensive definition of PPS that specifies its
different functions and characteristics is warranted. Following a consideration of some of the unresolved issues in the PPS lit-
erature, and relevant emerging topics, we conclude the current advanced review with an updated definition of PPS that we
hope will allow for achieving consensus in this literature and that may shape the research agenda in future years.
The term “peripersonal space,” as first introduced by Rizzolatti and colleagues (Rizzolatti et al., 1981), described the
nearby representational space in terms of what was reachable—that is, within range of the arm's reach. From this perspective,
PPS emphasizes a map of objects surrounding the body as a guide to reaching behaviors. The emphasis on reaching was in
keeping with earlier conceptualizations of the nearby space surrounding the body; the earlier concepts of “grasping space”
(Brain, 1941) and “operational space” (Von Uexküll, 1992) were essentially the space in which objects could be acted upon
(because they were within reach). However, such a definition has clear limitations, given that PPS representations are not
strictly defined by the length and position of the arm. Although PPS in humans has been found to be related to arm length
(Longo & Lourenco, 2006; Lourenco et al., 2011), both behavioral and neural data suggest that the transition between PPS
and extrapersonal space is not an abrupt one that occurs at the end of arm's reach but, rather, occurs gradually across the length
of the arm and beyond (Longo & Lourenco, 2007; Longo, Trippier, Vagnoni, & Lourenco, 2015). In addition, body-part-
specific representations of PPS have also been described for non-reaching body parts, such as the face (Sambo & Iannetti,
2013). Such findings call for a broader definition of PPS.
Other early attempts to define the space around the body, such as “body buffer zone” (Horowitz, Duff, & Stratton, 1964)
and “flight zone” (Hediger, 1955), emphasized defensive, flight-or-fight behaviors. Cooke and Graziano (2003) (see also Gra-
ziano & Cooke, 2006) connected these concepts to PPS, arguing that PPS might serve as a defensive buffer around the body,
keeping the body far enough away from threats. Although initially described in monkeys, this definition has since been
extended to humans, where researchers have described PPS as providing a “safety margin” around the body (Sambo & Ian-
netti, 2013). However, this definition makes it difficult to disentangle PPS from other concepts such as “personal space” typi-
cally used to define the preferred interpersonal space between individuals. Thus, an accurate and comprehensive definition of
PPS may necessitate reference to personal space and the social-affiliative mechanisms that influence one's interactions with
conspecifics.
2 | T H E C UR REN T A DV AN CE D R E V IE W
The purpose of the current advanced review is to provide an updated and broad-based empirical conceptualization of PPS. To
this end, we revisit topics in the literature for which consensus has been lacking and we attempt to integrate emerging findings
in an effort to provide a comprehensive account of what PPS is, including its functional properties and characteristic features.
By investigating questions about how PPS is instantiated, the functions it performs, and its relation to other constructs, we lay
the groundwork for a definition of PPS that makes sense of unresolved issues and integrates emerging findings within the clas-
sic literature.
As is the case with most reviews on the topic of PPS, our own discussion is necessarily limited. Other extant reviews cover
important topics such as the role of tool use in the plasticity of PPS (Holmes, 2012), the multisensory nature of PPS (Van der
Stoep, Nijboer, Van der Stigchel, & Spence, 2015), and the neural correlates of PPS (Cléry et al., 2015; Pellegrino & Làdavas,
2015). We direct the reader to these reviews when appropriate throughout the current advanced review. Although we reference
different measures of PPS, we have covered less research with neurological patients and the measures that are more typical for
such patients (e.g., cross-modal extinction). A challenge with understanding PPS is that these different tasks may capture dis-
tinct functions or characteristics of PPS. It is, thus, crucial to consider a range of measures, which we have attempted to do in
this advanced review while also focusing on recent research with neurologically healthy individuals.
HUNLEY AND LOURENCO 3 of 17
3 | T W O FU NC T I ON S O F P P S
Recently, de Vignemont and Iannetti (2015) provided a formalization of the two primary functions of PPS: (1) the guidance of
non-defensive behaviors in relation to neutral, nonthreatening objects; and (2) the guidance of defensive behaviors in relation
to potentially noxious or harmful stimuli. Thus, in the following sections, we examine extant research on PPS in terms of these
two distinct functions, concluding with a discussion of the implications of these functions for the network underlying PPS.
3.1 | The non-defensive function

In seminal work on PPS, researchers examined spatial representations near the body of macaque monkeys for object-oriented
actions involving nonthreatening objects (Graziano & Gross, 1995; Iriki, Tanaka, & Iwamura, 1996; Rizzolatti et al., 1981;
Rizzolatti, Matelli, & Pavesi, 1983). Multisensory neurons sensitive to visual and tactile stimuli that responded to objects
placed at, or near, the skin were described in the monkey pariarcuate cortex, with maximal activation to objects placed a few
centimeters above the skin. Moreover, this mapping was anchored to particular body parts, such that specific groups of neu-
rons responded only to objects near the hand, whereas other neurons responded only to objects near the mouth (see Figure 1)
(Graziano et al., 1997; Graziano, Yap, & Gross, 1994). Based on these findings, Graziano and Gross (1998) suggested that
these neurons maintained a “body-part-centered” mapping of the space immediately surrounding the body, crucial for guiding
effector-specific movement in relation to nearby objects (for review, see Pellegrino & Làdavas, 2015).
Behavioral studies of neurological patients as well as neuroimaging studies employing functional magnetic resonance
imaging (fMRI) and transcranial magnetic stimulation (TMS) with healthy participants demonstrated that humans also main-
tain populations of neurons, located within the intraparietal sulcus (IPS), supramarginal gyrus (SMG), and regions of the pre-
motor cortex, that selectively represent the space near the body (Berti & Frassinetti, 2000; Brozzoli, Ehrsson, & Farnè, 2014;
Cowey, Small, & Ellis, 1994; Ferri et al., 2015; Serino, Canzoneri, & Avenanti, 2011; for reviews, see Cléry et al., 2015; Gri-
vaz, Blanke, & Serino, 2017). Much evidence also suggests that representations of PPS in humans, like those of the macaque
monkey, are multimodal in nature, responding to visual (Longo & Lourenco, 2006), visuo-tactile (Noel, Pfeiffer, Blanke, &
Serino, 2015; Serino et al., 2015), and audio-tactile (Canzoneri, Magosso, & Serino, 2012; Ricciardi et al., 2017) information
(for review, see Van der Stoep et al., 2015). Moreover, and crucially, as in the monkey, these representations in humans
appear to be body-part-centered, such that bodily dimensions influence the size of PPS. For example, individual differences in
arm length have been shown to predict differences in the extent (i.e., “size”) of PPS in humans, such that human participants
with longer arms have correspondingly larger PPSs as measured by a visual line bisection paradigm (see Box 1 and Figure 2;
Longo & Lourenco, 2007; Lourenco et al., 2011). There is also evidence that representations of PPS associated with the arm
are specifically affected by manipulations applied to the arm, not the torso (Lourenco & Longo, 2009), including hand-specific
FIGURE 1 Receptive fields for bimodal visual-tactile neurons associated with the macaque face and hand, depicting
the range for which these neurons were activated. Black wedge and dot indicate the region from which the researchers
conducted their recordings (Reprinted with permission from Graziano, Hu, and Gross (1997) Copyright 1997 by the
American Physiological Society)
BOX 1
VISUAL LINE BISECTION AS A MEASURE OF PPS
The line bisection paradigm takes advantage of lateral visual-attentional biases that differ as a function of distance from
the body. Specifically, neurologically healthy participants are biased to bisect lines slightly to the left of center when the
lines are presented close to the body (e.g., 30 cm), a phenomenon known as pseudoneglect (Jewell & McCourt, 2000).
Pseudoneglect has been characterized as increased right parietal activation when objects are presented in PPS, producing
a left-oriented attentional bias (Longo et al., 2015). Importantly, this bias shifts rightward as lines are presented at farther
distances from the body (i.e., extrapersonal space). Thus, the rate at which participants shift from left to right bias is
taken as an index of the size of PPS, when considered in terms of visual-attentional biases.
effects (for review, see Brozzoli et al., 2014), and evidence for a dissociation in cross-modal effects as applied to the lower
limbs (e.g., legs and feet) compared with the torso (Schicke, Bauer, & Röder, 2009; Stone, Kandula, Keizer, & Dijkerman,
2018), consistent with body-part-centered representations. Neural evidence corroborates the body-part specificity of PPS
(Cléry et al., 2015) and, thus, challenges the definition of PPS as exclusively the space that is reachable, since some of the
implicated body parts such as face and torso are not involved in reaching actions.
Evidence from both humans and monkeys also suggests that PPS flexibly adapts to changing action capabilities. For
example, research with paraplegic humans demonstrates that their representations of PPS expand after simply seeing their par-
alyzed legs moved by another person (Scandola, Aglioti, Bonente, Avesani, & Moro, 2016). Moreover, hand tools that expand
an animal's range of effective action have been shown to produce expansion of PPS associated with the arm. After manipulat-
ing a rake, monkeys demonstrated expansion of PPS, encompassing space previously outside of actionable range (Iriki et al.,
1996) (for reviews, see Holmes, 2012; Maravita & Iriki, 2004). Likewise, studies with humans found expansion following tool
use by neurological patients (Ackroyd, Riddoch, Humphreys, Nightingale, & Townsend, 2002; Berti & Frassinetti, 2000) and
healthy participants using sticks (Canzoneri et al., 2013; Canzoneri, Marzolla, Amoresano, Verni, & Serino, 2013; Farnè &
Làdavas, 2000; Longo & Lourenco, 2006), as well as by blind individuals using a cane (Serino, Bassolino, Farnè, & Làdavas,
2007). These effects have been taken as evidence that reachable space is enlarged by tool use. Importantly, though, the oppo-
site effect has also been found in humans, such that there is contraction of PPS when movement is impaired with arm weights
(Lourenco & Longo, 2009) or barriers (Morgado, Gentaz, Guinet, Osiurak, & Palluel-Germain, 2013), as well as following
the loss of a limb (Canzoneri, Marzolla, et al., 2013), providing support for symmetric plasticity of PPS. Of note, in the case
of arm weights, the effect was restricted to the PPS associated with the arm in that weighting the torso (vis-à-vis a backpack)
did not result in contraction of PPS (Lourenco & Longo, 2009). In other words, manipulations applied to specific effectors
(e.g., the arm) affected only representations associated with the specific effector, suggesting that plasticity is also body-part-
4 3
r(47) = –.32
3.5
2.5
Rightward shift (% line length / m)
Rightward shift (% line length / m)
3
2
2.5
1.5
2
1.5 1
1
0.5
0.5
0
0
–0.5 –0.5
–1 –1
55 65 75 85 30 60 90 120 150 180 210 240
Arm length (cm) Distance (cm)
FIGURE 2 Both figures include data from the studies of Longo and Lourenco (2007) and Lourenco et al. (2011). Left figure: Scatterplot of the relation
between arm length and the rate of participants' rightward shift on the visual line bisection task. Longer arms were associated with more gradual rightward
shifts in bias, indicative of larger peripersonal spaces. Right figure: Participants' mean estimates (squares) of center at each distance (error bars 1 SEM). The
black line is the mean slope across participants. The red line is the mean slope for participants with short arms (<72.5 cm), and the green line is the mean
slope for participants with long arms (≥72.5 cm)
centered. Taken together, these findings suggest representations of PPS that flexibly adapt to changing physical capabilities as
they relate to the arm or reaching behaviors, under non-defensive conditions. However, as described below, this plasticity is
not limited to the arm and it extends to defensive conditions. Thus, “reachable” space can be used to partially characterize
PPS, but PPS should not be defined exclusively in this manner.
3.2 | The defensive function

Perhaps the most important behavior for any animal is threat detection: although other behaviors, such as feeding and mating,
can typically be safely delayed, defending against or fleeing from an active, physical threat cannot be postponed without
deadly consequence (Hediger, 1955). The Swiss zoologist Hediger (1955) argued that a key goal in this endeavor was to keep
potential threats at a safe distance by maintaining a “flight distance” or “flight zone” around the animal's body that elicited
fight-or-flight behaviors when crossed by a threatening object (e.g., projectile) or other animal (e.g., predator). In his original
account, Hediger (1955) reported that this distance was determined by an animal's body size (i.e., larger animals had larger
flight zones) and defensive strategy (i.e., animals who relied on flight, rather than fight, had larger flight zones), suggesting
that animals' physical dimensions and defensive capabilities played central roles in defining the space close to the body—what
we now refer to as PPS.
Subsequent research with human and nonhuman primates has provided evidence for a “margin of safety” that is repre-
sented in the primate brain (Graziano & Cooke, 2006). Graziano and colleagues documented distinct startle responses in
macaque monkeys for sudden air puffs presented near, but not far from, the face, which suggested PPS that was sensitive to
defense-related information (Cooke & Graziano, 2003). Moreover, it was found that these startle responses were evoked when
stimulating the ventral intraparietal area (VIP) and polysensory zone (PZ), both of which are associated with the representation
of PPS in monkeys.
Behavioral work in humans dovetails with the findings from monkeys. More specifically, Sambo and Iannetti (2013) pro-
vided evidence for what the researchers refer to as a “safety margin” in humans, like that described in monkeys, by taking
advantage of a human defensive reflex known as the hand blink reflex (HBR; see also Bufacchi, Liang, Griffin, & Iannetti,
2016; Sambo, Forster, Williams, & Iannetti, 2012; Sambo, Liang, Cruccu, & Iannetti, 2012). In the case of the HBR, humans
produce defensive eye-blinks when the hand is stimulated, but this effect, as measured by muscular response using electromy-
ography (EMG), is strongest when the hand is in proximity to the face and weaker when farther away from the face (see
Figure 3). By noting the distance at which the HBR occurs in relation to the face, the researchers have argued that this
approach can be used as a measure of PPS. Similarly, Paepe, Crombez, Spence, and Legrain (2014) described hand-centered
representations of PPS related to pain detection that were spatially locked to the stimulated hand (see also Serino et al., 2015),
suggesting that defensive representations exist for multiple body parts.
There is evidence that individual differences in trait anxiety and fear are related to PPS. For example, it has been found
that higher trait claustrophobic fear is associated with larger PPSs, as measured by a visual line bisection task (see Box 1)
(Lourenco et al., 2011; Sambo, Liang, et al., 2012), suggesting individual differences in the size of PPS that reflect the defen-
sive mechanisms of claustrophobic fear. Similarly, individual differences in anxiety have been shown to predict PPS size as
measured by HBR, such that greater anxiety predicts larger PPSs (Sambo & Iannetti, 2013). Sambo and Iannetti (2013), how-
ever, failed to find a correlation between PPS and trait claustrophobic fear, which they suggested might have reflected task-
specific differences between their study and that of Lourenco et al. (2011). Whereas Lourenco et al. (2011) used a line bi-
section task to capture representations of PPS, which Sambo and Iannetti (2013) proposed may have primed non-defensive
mechanisms, Sambo and Iannetti (2013) used an HBR task, which may have strictly involved defensive mechanisms. Another
possibility worth considering is that these effects reflected body-part specificity. For instance, the line bisection task might
40
EMG (mV)
20
Ultra-far Far Near Ultra-Near
0
0 50 100 150 200
Time (ms)
FIGURE 3 Waveform depicting average muscular responses from participants at “ultra-far” (60 cm), “far” (40 cm), “near” (20 cm), and “ultra-near” (4 cm)
distances during the hand blink reflex (HBR) task. Greater waveform activity represents a stronger HBR. Source: Sambo and Iannetti (2013)
specifically tap PPS related to the arm (Lourenco & Longo, 2009), whereas the HBR task likely measures PPS associated with
the face (Sambo, Forster, et al., 2012). Given the accumulating evidence for body-part-centered representations of PPS, it
would follow that PPS associated with the arm, which may be more affected by restricted movement, should be more strongly
associated with claustrophobic fear. By contrast, PPS associated with the face, a particularly sensitive somatosensory region,
might be specifically associated with general anxiety. Yet another possibility is that the HBR may not be measuring PPS.
Given that the stimulus in HBR tasks is applied directly to the skin rather than near the skin, this task could instead involve
priming another spatial mechanism, such as the body schema. The extant data are consistent with PPS representations that are,
at least partially, defined by individual differences in sensitivity to threatening contexts (e.g., fear or anxiety); however, addi-
tional research may be needed to better understand the aforementioned effects related to the HBR task.
Like PPS representations that support non-defensive behaviors, there is evidence that representations of PPS are flexible in
defensive contexts. For example, Vagnoni, Andreanidou, Lourenco, and Longo (2017), Vagnoni, Lourenco, and Longo
(2012), Vagnoni, Lourenco, and Longo (2015) have reported that looming images of threatening stimuli (i.e., snakes and spi-
ders) were judged as arriving sooner than nonthreatening stimuli (i.e., rabbits and butterflies) on a time-to-contact (TTC) task.
One interpretation is that this effect could result from participants experiencing larger representations of PPS in the presence
of evolutionary threatening animals as compared to nonthreatening animals, leading them to judge the former as making con-
tact with themselves sooner. This interpretation is bolstered by findings indicating that participants demonstrate larger PPSs,
as measured by crossmodal tasks, in response to approaching threatening stimuli (e.g., dog growls and images of spiders) than
to nonthreatening stimuli (e.g., sheep bleats and images of butterflies [Taffou & Viaud-Delmon, 2014]; see also Haan, Smit,
Stigchel, & Dijkerman, 2016). Moreover, other research suggests that representations of PPS flexibly accommodate to tool
use, even in defensive contexts. For example, Rossetti, Romano, Bolognini, and Maravita (2015) reported that humans
showed earlier autonomic fear responses to an approaching dangerous stimulus (i.e., a needle) after using a tool that extended
their reach. These findings suggest that PPS, as related to defense of the body, responds to threat-related information, as well
as to the changing motor capabilities of the body.
3.3 | A network of PPS representation: integrating defensive and non-defensive functions

An unresolved issue in the literature on PPS is whether the representation of PPS, with its distinct defensive and non-defensive
functions, reflects a single, unified system, or rather, whether there are multiple, independent systems supporting these func-
tions. A single system of PPS would afford obvious advantages. A system that integrates information across defensive and
non-defensive functions would presumably enable an efficient coordination of behaviors related to both functions. Both types
of behaviors require the coordination of perception and action in relation to objects and other living entities surrounding the
body. And because there may be conditions in which this coordination includes both defensive and non-defensive behaviors
(e.g., grasping a non-threatening coffee cup to hurl at a threatening predator), there would be benefit to a system that maps
these behaviors to a shared representation of PPS (see Figure 4, de Vignemont & Iannetti, 2015).
However, it is also clear from existing research that there are dissociations between defensive and non-defensive functions
of PPS. Indeed, neural evidence from monkeys suggests that non-defensive behaviors are instantiated by a pathway that
includes the anterior parietal area AIP and 7b, as well as premotor area F5, whereas defensive behaviors are instantiated in a
parietal-premotor pathway that involves VIP and F4 (for review, see Cléry et al., 2015). There is also some evidence that dif-
ferentiation within these pathways may correspond to specific body parts. For example, areas AIP, 7b, and F5 are largely tied
to hand and arm representations, whereas areas VIP and F4 are linked to representations of the head and face (Cléry et al.,
2015). Taken together, the extant data suggest differentiation of PPS along defensive and non-defensive functions, both in
terms of the neural mechanisms and body-part specificity.
de Vignemont and Iannetti (2015) proposed that PPS is represented by two distinct, dissociable systems. One system of
PPS was specialized for the “goal-directed” (i.e., “working”) non-defensive behaviors, whereas the other system was special-
ized for the protective, defensive actions (see Figure 4). According to this proposal, these systems of PPS obey different prin-
ciples because non-defensive behaviors are organized quite distinctly from defensive behaviors. For example, non-defensive
behaviors are thought to be largely organized toward an object that is the intended recipient of an action, whereas defensive
actions are thought to be geared away from an object, such as ducking out of the way of a projectile or avoiding obstacles
(Bracha, 2004; Dosey & Meisels, 1969; Graziano & Cooke, 2006), though defensive actions would presumably involve
aggressive (i.e., fight) responses as well. Moreover, according to de Vignemont and Iannetti (2015), these behaviors necessi-
tate different levels of motor precision, hence the differentiation of systems. For example, grasping an object with the hand
requires slow, precise motor guidance. By contrast, protective actions, such as ducking one's head, must be rapid, not precise,
to keep the threat away from the body.
Consistent with a multiple systems perspective of PPS is research suggesting that particular traits (e.g., anxiety) and task
manipulations (e.g., restriction of movement) affect PPS differently under defensive and non-defensive contexts. For example,
Sensory stimulus Sensory stimulus
or
General PPS map Defensive PPS map Working PPS map
or
Defensive action Working action Defensive action Working action
FIGURE 4 According to de Vignemont and Iannetti (2015), a single system of peripersonal space (PPS) (left) would represent both threatening (e.g., a
spider) and nonthreatening (e.g., an apple) stimuli on a common map of PPS. By contrast, a multiple (two) systems perspective (right) would include separate
maps for defensive and non-defensive (i.e., “working”) functions (Reprinted with permission from de Vignemont and Iannetti (2015). Copyright 2014
Elsevier Ltd.)
research suggests that higher trait anxiety is associated with larger PPSs in defensive (HBR) contexts (Sambo & Iannetti,
2013). By contrast, other research in which participants are required to judge reachability to a neutral object has revealed that
when anxiety is induced experimentally, participants underestimate their perceived reach, suggesting a representation of PPS
that is smaller than would be appropriate for the required action (Graydon, Linkenauger, Teachman, & Proffitt, 2012). If PPS
were subsumed by a single system, it is unclear why anxiety would produce differential effects in the size of PPS. Yet, a chal-
lenge for this work will be to determine whether these effects reflect differences in the size of PPS per se or in more general
motivational factors. For example, in the reachability task, anxiety could decrease motivation to reach, independent of PPS
size. On a separate test of graspability, in which distance was held constant, participants also underestimated their ability to
grasp a block when anxiety was induced. One possibility is that grasping occurred in PPS and, thus, reflects the relation
between anxiety and PPS. Another possibility is that anxiety produces a general effect on task requirements, not specific to
reachability. More research is needed.
Recent work from our own lab, together with that of our colleagues, suggests that restriction of movement may affect PPS
differently depending on whether defensive or non-defensive functions are relevant to the task. As indicated above, we have
shown that when participants' wrists are weighted, there is contraction of PPS, as measured by a visual line bisection task
(Lourenco & Longo, 2009). More recently, a chin-rest manipulation, which also restricts movement, resulted in reduced TTC
judgments for looming objects, consistent with expansion of PPS (Vagnoni et al., 2017). By contrast, the chin-rest manipula-
tion in this study yielded no effect on participants' line bisection performance. A comparison across experiments with different
manipulations of immobilization suggests a dissociation between tasks that tap defensive versus non-defensive functions, con-
sistent with a multiple systems account of PPS. However, the observed dissociations in this research are far from straightfor-
ward given that performance in the line bisection task was not similarly affected by different restriction manipulations—wrist
weights were previously found to result in contraction (see Lourenco & Longo, 2009) whereas the chin-rest manipulation led
to no observable change (see Vagnoni et al., 2017).
The extant findings are suggestive of a dissociation between the defensive and non-defensive functions of PPS and, thus,
would seem consistent with an account that posits multiple systems of PPS. Importantly, however, there are also experimental
conditions that suggest interactions between these so-called systems. For example, following a tool use manipulation that
extended reach, human participants experienced earlier autonomic fear responses to a threatening object, namely a needle
(Rossetti et al., 2015). Moreover, under non-laboratory, real- world, conditions, one might argue that defensive and non-
defensive behaviors are often coordinated such as when tools (e.g., weapons) are used in precise actions to defend against a
predator. Are such behaviors best characterized by two separate systems of PPS? Although advocates of a multiple systems
account of PPS may not deny that interactions between defensive and non-defensive pathways are needed (indeed, de Vigne-
mont and Iannetti (2015) acknowledged such a possibility), it is unclear how such interactions would occur under such an
account. One possibility is that information between defensive and non-defensive pathways is readily translatable, but this
property would seem to differ from the perspective of de Vignemont and Iannetti (2015), in which there are different spatial
maps that feed to different actions. Thus, we would argue for an alternative model of PPS that includes two specialized
pathways for defensive and non-defensive functions, but that exists as a single, integrated network in which interactions read-
ily occur.
More specifically, we suggest that although defensive and non-defensive pathways are accompanied by distinct neural
regions, they likely represent the locations of objects using a common coordinate structure. A multiple systems account would
likely posit that other, relevant systems might be recruited to accomplish the coordination between defensive and non-
defensive behaviors. However, we propose that this coordination may be possible within the PPS network itself because the
structure of the different representations or “maps” (de Vignemont and Iannetti, 2015), which reference the body, likely
support similar computations. Evidence for such a possibility comes from much research showing that defensive and
non-defensive behaviors demonstrate common forms of plasticity (i.e., expansion and contraction) and that defensive and
non-defensive behaviors are impacted by common variables such as body size and trait anxiety. As in the multiple systems
account of de Vignemont and Iannetti (2015), we emphasize two distinct defensive and non-defensive pathways. However,
we also suggest that it may be more fruitful to consider how these pathways interact to produce coordinated actions. This per-
spective combines both the single and multiple systems perspectives by providing an account of how distinct pathways of
defensive and non-defensive functions may exist within a unified network.
4 | D I S T I N GU I S HI N G P P S FR O M O T H E R C O N S T R U C T S
In this next section, we ask about the relation between PPS and other well-known constructs in the literature—namely, body
schema and personal space. We take up body schema and personal space in particular because of the definitional overlap that
has existed between these constructs and PPS. We address differing perspectives on the nature of the relations between PPS
and these constructs, with the goal of resolving some of the ambiguities that exist in this literature about the relations between
PPS, the body schema, and personal space.
4.1 | PPS and body representations

Scientists have long reflected on the presence of an “organized model” of the body, referred to in the classic neurology litera-
ture as the body schema (Head & Holmes, 1911). More specifically, the body schema is considered a coherent representation
of the body's spatial properties, which includes the length of one's limbs and their hierarchical arrangement (Haggard & Wol-
pert, 2005). It is believed that the representation of the positions of one's body parts, which are updated during bodily move-
ment and are projected onto space, provide spatial organization for action (Haggard & Wolpert, 2005). Recent research from
neuroscience confirms the existence of a body representation relying on proprioceptive information from muscles, joints, and
skin relevant for controlling posture and action, as well as locating stimuli on the body (Maravita, Spence, & Driver, 2003;
Martel, Cardinali, Roy, & Farnè, 2016; Medina & Coslett, 2010).
An intriguing hypothesis is that PPS arises directly, in casual fashion, from the body schema (Martel et al., 2016). Such a
hypothesis follows straightforwardly from the fact that knowing one's position in space is necessary for coordinating reaching
and grasping behaviors to objects. Much of the work on PPS expansion after using a tool (Ackroyd et al., 2002; Canzoneri,
Ubaldi, et al., 2013; Longo & Lourenco, 2006; Rossetti et al., 2015) would seem to suggest that the body schema is integral to
PPS. Indeed, in this research, it is often suggested that expansion of PPS occurs because the tool is incorporated into the body
schema, thereby supporting expansion of PPS by way of the body schema (Maravita & Iriki, 2004). Effects of active tool use
on the body schema have been documented across a variety of studies. In these studies, tool use induces kinematic changes
(i.e., free-hand reaching kinematics), with participants adjusting the velocity of their reach as if they perceived their forearm
as longer (Cardinali, Brozzoli, & Farnè, 2009). Other research also shows that participants judge their forearms as longer (and
skinnier) after using a tool to complete a task, suggesting that the tool was incorporated into their body schema (Canzoneri,
Marzolla, et al., 2013; Cardinali et al., 2009; Sposito, Bolognini, Vallar, & Maravita, 2012). Moreover, the use of prosthetic
limbs by amputees has been shown to induce changes in both the body schema and PPS, such that wearing a prosthesis
increases the perceived length of the stump (Canzoneri, Ubaldi, et al., 2013; Mayer, Kudar, Bretz, & Tihanyi, 2008) and
appears to extend PPS beyond the stump (Canzoneri, Ubaldi, et al., 2013).
Classic work by Iriki et al. (1996) with monkeys found that, following training with a rake, bimodal neurons for the ani-
mal's arm and hand expanded to include the space accessible by the rake. However, this was not the case for neurons with
receptive fields on the animal's fingers. Although one could reasonably argue that expansion related to the fingers might be
more relevant for a precision grip, and, thus, would be more likely under conditions that require precise manipulations, it is
worth considering that the dissociation between arm/hand and fingers poses a challenge for PPS representations that build
directly on the body schema, since fingers would still be relevant for a coherent body representation that involves the hand
and that is implicated in grasping a tool.
Thus, an alternative possibility is that PPS is not causally related to the body schema. This alternative possibility is bol-
stered by findings using a cross-modal congruency task in which a visual stimulus appearing near the hand produces a speeded
response to tactile stimuli presented to the same hand (Spence, Pavani, & Driver, 1998; Spence, Pavani, & Driver, 2004;
Spence, Pavani, Maravita, & Holmes, 2004). The cross-modal congruency effect has been found in relation to the end of a tool
after the tool was used to complete a task, suggesting an expansion of PPS (Maravita et al., 2003). However, if PPS arises
from the body schema, then one would expect PPS to expand to include the entire length of the tool. That is, because tool use
results in a continuous extension of the arm (Canzoneri, Ubaldi, et al., 2013; Cardinali, Frassinetti, et al., 2009), PPS should
also incorporate the tool in a continuous manner. Yet Holmes, Calvert, and Spence (2007) found that this expansion included
only the tip of the tool that had been actively used, not the middle portion of the tool (see also Holmes, Sanabria, Calvert, &
Spence, 2007; Holmes, Spence, Hansen, Mackay, & Calvert, 2008). Moreover, although it has been found that PPS can be
made to expand to include the middle portions of a tool, this is only the case when the middle portions have a functional fea-
ture used to complete the task (Park, Strom, & Reed, 2013; but see Bonifazi, Farnè, Rinaldesi, & Làdavas, 2007). Thus,
whereas active use of a tool may lead to an expanded representation of the arm that mimics the full length of the tool
(Canzoneri, Ubaldi, et al., 2013; Cardinali, Brozzoli, & Farnè, 2009), there are conditions in which expansion of PPS appears
discontinuous, linked to the specific regions that support the relevant action in space (Holmes, 2012).
Further evidence in support of a dissociation between PPS and the body schema comes from studies using remote-
controlled devices such as a computer mouse. Using a cross-modal auditory-tactile task, Bassolino, Serino, Ubaldi, and Làda-
vas (2010) found that PPS expanded to include the space acted upon with a computer mouse. However, it is unclear how using
a computer mouse would alter the body schema in such a way that would produce the documented expansion of PPS. Thus,
an interpretation of PPS expansion in this case is that it resulted from some mechanism other than a change to the body
schema. Recently, evidence of wheelchair experience revealed that expansion of PPS was possible even under a passive move-
ment condition (i.e., the participant was moved around the environment by someone else), suggesting that visual experience,
such as optic flow, is sufficient to remap extrapersonal space as PPS (Galli, Noel, Canzoneri, Blanke, & Serino, 2015). More-
over, an active condition (in which the participant moved him- or herself around the environment in the wheelchair) did not
produce PPS expansion. The authors argued that “embodiment” of the tool did not lead to greater expansion, which would
seem inconsistent with expansion of PPS vis-à-vis the body schema.
Other research showing contraction of PPS during a visual line bisection task with weighted wrists (Lourenco & Longo,
2009) might be considered in light of the debate on the relation between PPS and the body schema. One possible account of
this result is that PPS contraction occurs vis-à-vis the body schema. For example, if arm weights caused participants to per-
ceive their arms as smaller in this condition, then PPS might shrink to accommodate this perceived change in the body
schema, similar to how individuals with shorter arms have been shown to have smaller PPSs than individuals with longer arms
(Longo & Lourenco, 2007). On this account, PPS and the corresponding contraction is causally related to the body schema.
However, another account, from a perspective in which PPS and the body schema are not causally related, is that weighting
the arm may affect PPS, not the body schema. Indeed, one way in which this could occur is that weighting the arm would alter
the anticipated action consequences and might shift attentional demands to the space closer to the body. The representation of
the arm instead might remain unaltered as a function of weighting. Such a possibility is especially important given the putative
anisotropy of the body schema. Although changes to the body schema that follow the direction of bodily growth are possible,
the evidence suggests resistance when the alterations are not biologically plausible for the body (De Vignemont, Ehrsson, &
Haggard, 2005).
Cardinali, Brozzoli, and Farnè (2009) proposed that the body schema provides necessary, but insufficient, support for the
representation of PPS. On this perspective, the body schema is the necessary “skeleton” for PPS. This argument is certainly
consistent with a definition of PPS in which the representation of nearby space is anchored to specific body parts
(e.g., Graziano & Gross, 1998). Indeed, as discussed earlier, there are numerous examples of body-part-centered effects of
PPS (see Section 3). Cardinali, Brozzoli, and Farnè (2009) noted that the body schema is insufficient for the representation of
PPS because additional inputs from modalities such as vision and audition must be incorporated within this representation.
We agree and would further argue that general attentional mechanisms may also be crucial in specifying the nearby space.
Importantly, however, we challenge the notion that the body schema per se is necessary for the representation of PPS. An
alternative possibility is that although the PPS representation includes reference to the body, including specific body parts, the
bodily reference point need not emerge from a fully organized model of the body, namely the body schema. Instead, the body-
centered quality of PPS could rely on somatosensory representations without the full, organized structure accompanying the
body schema. Such somatosensory representations would be necessary for anchoring relevant actions to the space surrounding
the body. Crucially, they might also be sufficient for mapping objects to the surrounding space that are acted upon in typical
non-threatening situations or avoided in the case of a threatening context.
4.2 | PPS versus personal space

Drawing from Hediger's concept of the “flight zone,” which Hediger used to explain an animal's defensive responses to
approaching predators (Hediger, 1955), proxemics defined personal space as the distance people maintain between themselves
and others (Hall, 1966; Sommer, 1959). Personal space has come to be conceived of as the zone of safety surrounding the
human body (Holmes & Spence, 2004), and it includes the management of interpersonal distances to others. Evidence for the
phenomenon of personal space begins with early research in social psychology and cultural anthropology, in which it was
shown that people maintained a consistent boundary between themselves and their social partners, referred to as their preferred
comfort distance (Bassolino et al., 2010) (for recent evidence, see Holt et al., 2014; Kennedy, Gläscher, Tyszka, & Adolphs,
2009). Incursions of this distance were found to produce feelings of discomfort and anxiety (Hayduk, 1981), with people cre-
ating larger distances to intruders in uncomfortable situations and smaller distances when in comfortable situations (Holt
et al., 2014; Kennedy et al., 2009). Recent evidence demonstrating that interpersonal distance expands after one listens to a
conversation with aggressive content compared to neutral content is consistent with personal space as crucial for providing a
protective “bubble” around the body (Vagnoni, Lewis, Tajadura-Jimenez, & Cardini, 2018).
These descriptions of personal space display more than a passing similarity to PPS as related to its defensive function.
Indeed, one may ask whether personal space is truly distinguishable from PPS or whether it is another way in which to
describe the defensive function of PPS. As indicated earlier (see Section 3), PPS is relevant to our interactions with neutral
objects as well as to our defensive behaviors toward potentially harmful stimuli, including predatory conspecifics. The fact
that definitions for both PPS and personal space have referenced a safety zone surrounding the body has perhaps, not surpris-
ingly, led to the hypothesis that these two constructs rely on shared mechanisms of representation. Indeed, parallels in perfor-
mance on tasks measuring PPS and personal space provide support for this possibility. For instance, higher trait anxiety and
fear are associated with a larger PPS (Lourenco et al., 2011; Sambo & Iannetti, 2013) and personal space (Bogovic, Mihano-
vic, Jokic-Begic, & Svagelj, 2013; Bracha, 2004), suggesting individual differences for both constructs that are predicted by
personality traits. Moreover, studies using fMRI have found that intrusions into personal space caused by social looming stim-
uli (i.e., faces) activate brain regions (i.e., dorsal IPS and ventral premotor cortex) that are similarly implicated in the represen-
tation of PPS (Holt et al., 2014; for review, see Pellegrino & Làdavas, 2015), suggestive of common neural mechanisms in the
representation of these spaces.
Recent research suggests that PPS and personal space recruit common motor mechanisms during processing. More specifi-
cally, Iachini, Coello, Frassinetti, and Ruggiero (2014) had participants complete two versions of a stop-approach task in
which the participants interacted with avatars in a virtual reality (VR) environment. In one condition, participants completed a
standard version of the stop-approach task as a measure of personal space. This task required that participants indicate the dis-
tance at which they first became uncomfortable with the avatar. In another condition, designed to measure PPS, participants
indicated at which distance they could physically reach the avatar with their hand. Both tasks involved a passive condition in
which participants waited while the avatar walked toward them and an active condition in which participants walked toward
the avatar. The authors found that, in the passive condition, participants' preferred comfort distance to the avatar was signifi-
cantly larger than the distance they estimated they could reach the avatar. In other words, in the passive condition, personal
space and PPS appeared to be significantly different. However, in the active condition, there was no difference between partic-
ipants' preferred comfort distance and the distance they estimated they could reach the avatar. This finding suggests that per-
sonal space and PPS may not differ under active conditions. According to Iachini et al., 2014, 2016, the size of one's personal
space is reduced when the individual perceives that he or she has control over the interaction. In other words, a sense of motor
control may induce a reduction in personal space, suggesting that personal space can be modulated by action-related mecha-
nisms, as has been found for PPS, and consistent with a network of PPS that interfaces with, or subsumes, interactions with
people (Canzoneri, Marzolla, et al., 2013; Farnè & Làdavas, 2000; Longo & Lourenco, 2006).
Consistent with the claim that mechanisms related to motor control are recruited in the representation of PPS and personal
spaces, Quesque et al. (2017) found that tool use produced expansion of personal space, as has been shown for PPS (Ackroyd
et al., 2002; Canzoneri, Ubaldi, et al., 2013; Longo & Lourenco, 2006; Rossetti et al., 2015). More specifically, participants in
this study reported larger preferred interpersonal distances between themselves and a virtual human-like walker (created by a
projected point-light display) when it crossed in front of them after they had experience manipulating a tool. This study pro-
vides further support for a common motor mechanism for establishing interpersonal distance with individuals as well as
perception-action guidance with objects.
Yet there is other research examining the effects of tool use on PPS and personal space that is suggestive of a dissociation
between these two constructs. For example, Patané, Iachini, Farnè, and Frassinetti (2016) found that tool use produced expan-
sion of PPS, as measured by a reachability task. By contrast, they found no effect on personal space using a task that measured
participants' preferred comfort distance to a social partner. In another study, Patané, Farnè, and Frassinetti (2017) found that
on a social tool-use task, in which participants engaged in cooperative tool use with a confederate, participants again reported
an increased range of reachability, suggesting expansion of PPS. In this task, though, participants reported smaller preferred
comfort distances between themselves and the confederate. In other words, in the context of a social tool-use task, there
appeared to be opposite effects on PPS and personal space, which could be interpreted as expansion in the case of PPS and
contraction in the case of personal space.
Important for interpreting the putative dissociation between PPS and personal space in the study of Patané et al. (2017) are
the different components of the task. Patané et al. (2017) argued that personal space was affected by social cooperation
between the participant and confederate, not the length of the tool used in the cooperative exchange, whereas PPS was affected
by tool use (specifically the length of the tool relevant for the task), not the nature of the exchange between the participant and
confederate. Thus, rather than a dissociation between representations of PPS and personal space, an alternative interpretation
of these findings is that the changes observed in the different conditions reflect differential relevance of social-affiliative fac-
tors (or tool use), which we suggest below may reflect the defensive and non-defensive functions of PPS.
Recent evidence suggests that PPS is sensitive to social-affiliative factors (Teneggi, Canzoneri, di Pellegrino, & Serino,
2013). For example, using a cross-modal visual-tactile task to assess the boundary of PPS, Teneggi et al. (2013) found that this
boundary expanded following a cooperative exchange with another person. On the one hand, this effect may seem surprising
because a cooperative other should be considered non-threatening and, thus, a defensive PPS ought to shrink, not expand. On
the other hand, if the overall context can be considered relatively non-threatening, then perhaps it is the non-defensive path-
way of PPS that is activated, so that PPS is likely to expand in the presence of a cooperative other because the primary goal is
coordination with this individual. Under such non-defensive conditions, the other individual may take on a functional role
(much like a tool) in assisting with actions. It has been suggested that such effects may be supported by neuronal populations,
referred to as “mirror neurons” that code for both one's own actions and that of another (Rizzolatti & Craighero, 2004). Such
neuronal populations, it is suggested, support the anticipation of others' behaviors and, thereby, aid in the coordination of joint
actions. It has also been suggested that social-affiliative mechanisms are relevant for joint actions and, thus, may impact the
PPS network (Brozzoli, Gentile, Bergouignan, & Ehrsson, 2013). These mechanisms may include brain regions implicated in
face processing such as the fusiform face area (FFA) (Holt et al., 2014) and fear such as the amygdala (Kennedy et al., 2009),
as well as processes related to theory of mind (i.e., “mentalizing”), joint attention (Brozzoli et al., 2013), and emotional
expression (Ruggiero et al., 2017).
Pellencin, Paladino, Herbelin, and Serino (2018) also found that the boundary of PPS, as assessed with a cross-modal audio-
tactile task, was expanded when in the presence of individuals who were rated as moral (vs. immoral). By contrast, the same partic-
ipants preferred interpersonal distances (i.e., personal spaces) that were closer to the moral individuals. These data would, again,
seem to suggest that PPS and personal space are distinct, dissociable constructs, because participants' judgments suggested larger
PPSs, but smaller personal spaces. However, the other possibility is that such effects reflect the defensive and non-defensive func-
tions of PPS, such that larger non-defensive, and smaller defensive, PPSs both facilitate interactions with a moral other.
Following from the extant research, we would argue that social-affiliative factors may be crucial in specifying the func-
tional (defensive vs. non-defensive) pathway of PPS necessary for interacting with people within one's PPS, as well as the
coordination across defensive and non-defensive pathways. For example, social-affiliative factors might be relevant for deter-
mining whether a conspecific is a friend or foe and, more generally, whether a scenario is threatening or non-threatening. For
example, in a cooperative scenario, non-defensive PPS would be relevant for facilitating actions such that non-defensive PPS
would increase in size because the primary goal is ongoing (or anticipated) interactions with another individual. PPS should
thus expand (rather than contract) in the presence of other individuals who are more cooperative (Teneggi et al., 2013), moral
(Pellencin et al., 2018), or perhaps even romantic. In correspondence, defensive PPS should decrease in size in such non-
threatening scenarios. Returning to the study of Patané et al. (2017), we would suggest that the cooperative tool use task pro-
vided a measure of non-defensive PPS and that PPS expanded following tool use because the goal of the task was coordinated
action in a relatively non-threatening context. We would further suggest that the coordination of action across a network of
PPS that includes both defensive and non-defensive pathways would involve decreasing the size of defensive PPS (i.e., the
interpersonal distance measure of personal space in the study by Patané et al., 2017) given the non-threatening context.
Following from the original definition of personal space, in which the defensive component is emphasized, we suggest that
the construct of personal space may be best subsumed under the defensive pathway of the PPS network, as the putative disso-
ciations between PPS and personal space are well accounted for under a network of PPS that includes both defensive and non-
defensive pathways for interacting with objects and other living entities. We emphasize that persons, like objects, are also rele-
vant for a variety of non-defensive interactions and, thus, may be mapped to space according to the non-defensive pathway of
PPS when relevant.
An interesting issue for future research is the extent to which we can compare research using VR, in which the other indi-
viduals are avatars, with real scenarios involving human confederates. Although recent research with VR confirms its appro-
priateness for assessing PPS (Serino et al., 2017), an important question for future research will be whether avatars may be
treated more like inanimate objects than real-life organisms, with implications for whether defensive or non-defensive path-
ways of PPS are implicated in the interactions.
5 | E M E RG I N G Q U E S T IO N S IN P P S R E S E A R C H
5.1 | Individual differences in the flexibility of PPS

As discussed earlier, the size of one's PPS adjusts flexibly to motor challenges (e.g., tool use and arm weights) (Ackroyd et al.,
2002; Lourenco & Longo, 2009) and the presence of threats (e.g., dog barks and images of spiders) (Haan et al., 2016; Taffou &
Viaud-Delmon, 2014). Flexible effects of expansion and contraction have been found to occur immediately in response to stim-
uli, which may be unsurprising given the often immediate need to coordinate approach and avoidance behaviors. There is also
evidence suggesting that such effects may be durable, as in the case of expert tool users (e.g., blind cane users; Serino et al.,
2007). However, given individual differences in the size of PPS, which have been found to relate to trait fear
(e.g., claustrophobic fear; Lourenco et al., 2011) and anxiety (e.g., general trait anxiety; Sambo & Iannetti, 2013), one may ask
whether there are meaningful individual differences in the flexibility of one's PPS as well. Moreover, there are open questions
about whether individual differences in flexibility affect the non-defensive and defensive pathways of PPS in comparable ways.
Individuals high in claustrophobic fear (Lourenco et al., 2011) and general anxiety (Sambo & Iannetti, 2013) have larger
PPSs, such that an intriguing possibility is that these individuals may also be hampered by limited flexibility (i.e., expansion
and/or contraction) in their PPSs. Research from our lab suggests that fear, specifically trait claustrophobic fear, is associated
with individual differences in the flexibility of PPS (Hunley, Marker, & Lourenco, 2017). In particular, Hunley et al. (2017)
found that individuals with higher trait claustrophobic fear experienced decreased expansion of PPS relative to individuals
lower in trait claustrophobic fear when the task involved bisecting lines with sticks (see Figure 5). Previous work had shown
that participants demonstrated expansion of PPS when using a stick to bisect lines compared to when using a laser pointer
(Longo & Lourenco, 2006). Researchers have argued that a stick, unlike a laser pointer, affords expansion because it has func-
tional consequences in the bisection task (but see, Holmes, 2012). However, Hunley et al. (2017) found that expansion of PPS
following tool use was not comparable across individuals. Participants higher in claustrophobic fear experienced less expan-
sion when using tools to bisect lines, suggesting relatively less flexible PPSs among these individuals.
Given these findings, one can begin to ask whether other cognitive (e.g., trait anxiety; Sambo & Iannetti, 2013) and physical
(e.g., arm length; Longo & Lourenco, 2007) factors that have been associated with the size of PPS also affect the flexibility of
these representations. Trait anxiety is associated with decreased attentional flexibility (Derryberry & Reed, 2002; Eysenck, Derak-
shan, Santos, & Calvo, 2007; Pacheco-Unguetti, Acosta, Callejas, & Lupiáñez, 2010), which, in turn, could lead to decreased
flexibility of PPS. Likewise, in the case of arm length, because individuals with longer arms can act on a broader range of space,
an intriguing possibility is that they might have more flexible representations of space. However, PPS is not limited to the arm,
4
3
Rate of rightward shift
2
(% line length/m)
–1
–2
0 20 40 60 80
CLQ (Total) score
FIGURE 5 Scatterplot of the relation between trait claustrophobic fear, as measured by the claustrophobia questionnaire (CLQ), and the rate of the rightward
shift of participants' estimates on the visual line bisection task when using sticks (Hunley et al., 2017). Larger rightward shifts are indicative of smaller
peripersonal space (PPS). When using a stick to bisect lines, individuals higher in claustrophobic fear had smaller PPS, suggesting decreased expansion of
PPS among these individuals and, thus, less flexibility in their PPS representations
and, thus, should not be considered exclusively in terms of the surrounding space that is reachable. Put differently, the extent of
PPS flexibility may depend on the body part (i.e., arm vs. head) as well as the specific function (i.e., defensive vs. non-defensive)
that is implicated in the task. For example, representations of PPS might exhibit greater flexibility under non-defensive conditions
when coordinating actions toward non-threatening objects and might exhibit less flexibility under defensive conditions when
attempting to maintain a safe distance to a threat. Likewise, PPS associated with the arm, which is implicated in reaching, may be
more flexible than that for the head, at least when the actions are non-defensive. Future studies will be crucial for testing the
extent to which such individual differences play a role in determining the flexibility of PPS and whether there are further differ-
ences depending on whether the defensive or non-defensive function is relevant to the task.
5.2 | How is PPS represented around the body?

In many studies of PPS, researchers examine only the space in front of the body. However, humans exist and act in a
three-dimensional world. Indeed, humans must interact with objects (and people) at all positions around the body. We sit
in chairs simply by using the feel of the chair against the back of our legs or the knowledge that the chair is behind
us. We navigate the physical environment while accommodating objects in front of the body (e.g., pushing a stroller) and
behind it (e.g., pulling wheeled luggage). We also anticipate the approach of others both directly in front of, as well as
behind, us. Such actions require a representation (or representations) of where these objects are in space relative to differ-
ent sectors surrounding the body in order to act. Likewise, threats are not limited to approaching from the front of the
body. As such, to defend against incoming attacks, humans must maintain vigilance, and organize defensive behaviors,
both in front of the body, which is mostly visible, as well as behind the body, which is not accessible to vision.
Empirical findings have shown that the PPS in human and nonhuman animals extends around the body (Farnè &
Làdavas, 2002; Graziano, Reiss, & Gross, 1999; Kitagawa, Zampini, & Spence, 2005; Van der Stoep et al., 2015). Yet
important questions about how PPS is represented around the body remain unanswered. Unlike the emphasis in the PPS
literature in characterizing PPS representations as related to specific body parts, less work has concerned comparisons of
the space as it surrounds the body. For example, we can ask whether PPS is symmetric, equally large in front of and
behind the body, or asymmetric, such that PPS could be larger in front compared to the rear or vice versa. Moreover, the
type of asymmetry (i.e., larger frontal PPS or larger rear PPS) could be function-dependent. For example, given that most
human perception-action capabilities are frontward-oriented, a larger frontward representation of PPS would be advanta-
geous under non-defensive conditions when guiding behaviors toward objects and other living entities that are non-threat-
ening. By contrast, under defensive conditions, it could be advantageous to be more vigilant toward the rear. Again,
because many human perception-action capabilities are frontward-oriented, any action in response to a threat approaching
from the rear would necessarily take more time, potentially making that threat more dangerous than if it had approached
from the front. Consequently, a larger representation of PPS behind the body could be adaptive in preparing for oncoming
threats.
As it stands, the extant evidence is mixed, with some evidence suggesting an asymmetrical representation that is larger in front
of the body than the rear, and other research suggesting a symmetrical representation around the body. For example, research
examining the processing of multisensory information, particularly audio-tactile stimuli, has found differences in how such infor-
mation is integrated in front of the body as compared to the rear under non-defensive circumstances (Gillmeister & Forster, 2012;
Kóbor, Füredi, Kovács, Spence, & Vidnyánszky, 2006; Occelli, Spence, & Zampini, 2011). More specifically, although interfering
information (e.g., crossing the arms) affects the processing of multisensory stimulation both in front, and behind, the body, this
effect is weaker behind the body (Kóbor et al., 2006). Moreover, attentional effects on performance are strongest behind the body
when the hands are held close to the body, whereas they are strongest in front of the body when the hands are held farther away
(Gillmeister & Forster, 2012), consistent with a larger front than rear PPS. These differences in the processing of multisensory
information for front and rear planes are suggestive of differences in PPS, such that greater attentional resources may be directed
toward the front of the body, with interfering attentional information having a stronger effect in the frontal plane.
The above conclusion is further supported by findings from research examining the defensive function of PPS. In particu-
lar, Bufacchi et al. (2016) examined PPS using an HBR task in which participants placed their hand at various locations rela-
tive to their face, including placing their hand behind their head. Again, participants demonstrated a larger PPS in front of,
compared to behind, the body. As noted earlier, however, it is possible that the HBR task may not be specific to PPS, with
effects instead related to the body schema for the stimulated hand. Nevertheless, together with research on multisensory inte-
gration, a preliminary interpretation is that PPS is represented asymmetrically under both defensive and non-defensive condi-
tions. Another explanation for these findings, though, is that these effects are dependent on the specific body part, such that
there may be specificity for face-centered PPS, which may be inherently frontward oriented given the anatomy of the head.
PPS representations may be more symmetrical for body parts (e.g., the arms or legs) that more readily interact with frontal and
rear planes relative to the head, as well as across both defensive and non-defensive contexts. Indeed, research by Galli
et al. (2015), which examined PPS in the front and rear of the body using an audio-tactile interaction task while participants
were seated in a wheelchair, found that PPS was represented symmetrically around the body. As such, more research is needed
to better understand this phenomenon.
Future research should also consider how individual differences in bodily dimensions (i.e., arm length) (Ferri et al., 2015),
as well as trait fear (Lourenco et al., 2011) and anxiety (Sambo & Iannetti, 2013) affect front and back sectors of PPS. The
current, albeit limited, research on individual differences has focused exclusively on PPS in the frontal plane. An open ques-
tion is whether individual differences in PPS exist behind the body as well. Work from Noel et al. (2015) demonstrating that
expansion of PPS in front of the body accompanied by contraction behind the body suggests that these representations may
rely on shared underlying resources. As such, individual differences in PPS may likewise exist behind the body. By examining
individual differences in how PPS is represented around the body, researchers can better understand the role they play in deter-
mining the layout of front and rear PPS, both of which are crucial for supporting our interactions with objects and other living
entities under defensive and non-defensive conditions.
6 | W H A T I S P P S ? A CO M P RE HE NS I V E D E F I N I T I O N
Having identified some of the limits of earlier definitions of PPS and having highlighted recent advances in the literature on PPS,
we are prepared to offer an updated, comprehensive definition of PPS. To answer the central question—What is PPS?—we draw on
classic and recent empirical findings from work with human and nonhuman animals and, importantly, we revisit questions about the
relations between PPS, body schema, and personal space. We also reference emerging issues such as individual differences and dif-
ferent sectors of PPS in an effort to provide a broad-based empirical definition of this multi-dimensional phenomenon.
We argue that PPS is a network of body-part-centered representations responsible for the coordination of actions toward, and
in avoidance of, objects and other living entities, including people. These actions include reaching and grasping objects and peo-
ple that surround the body, as well as maintaining vigilance for objects and people that may be approaching one's body.
Although the body-centered point of reference that characterizes PPS is often described as the coherent body schema, we have
suggested that the body reference frame within PPS may instead reflect more primitive somatosensory representations.
The PPS network is organized according to distinct defensive and non-defensive functions. Accumulating behavioral and
neural data provide support for dissociations between defensive and non-defensive functions, but there is shared organization
and common characteristics that may support an integrated network. The defensive and non-defensive functions of PPS have
been shown to dissociate according to the neural pathways and general action patterns, but importantly, both functions show
plasticity (i.e., expansion and contraction) and individual differences in relation to body size and anxiety. An important caveat
with respect to the “size” of PPS is that this size is not defined a priori. The extent of PPS depends on bodily dimensions, var-
ies according to personality traits, and both expands and contracts across context, though the degree of flexibility may vary
depending on function, body part, and personality traits.
The PPS network is multimodal, in that information from different sensory modalities, including vision and audition, is
relevant when mapping objects and people onto one's surrounding space. We have argued that the well-known construct of
personal space, which is often considered one's safety zone in relation to other people, may be best subsumed under the defen-
sive pathway of PPS, and systems relevant for processing social-affiliative mechanisms may interface with the PPS network,
playing a central role in recruiting defensive and non-defensive pathways when interacting with people.
And yet, there is certainly more work to be done. One challenge for this research, as mentioned at the start of the current
review, is that numerous tasks have been used to assess PPS, and these tasks may be tied to different functions or features of
PPS. Given the complexity of PPS, it will be important to consider how measures of reachability versus cross-modal sensitiv-
ity versus attentional bias, affect our current conceptualization of PPS, including how it relates to constructs such a body
schema and personal space, and how best to consider the influences of social-affiliative mechanisms.
ACKNOWLEDGMENTS
This work was supported by generous funding from Emory University (Laney Graduate School). The authors are grateful to
the reviewers for their helpful feedback.
CONFLICT OF INTEREST
The authors have declared no conflicts of interest for this article.
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Perception and action
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How to cite this article: Hunley SB, Lourenco SF. What is peripersonal space? An examination of unresolved empiri-
cal issues and emerging findings. WIREs Cogn Sci. 2018;e1472. https://doi.org/10.1002/wcs.1472
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What is Peripersonal Space? An Examination of Unresolved Empirical Issues

Article in Wiley interdisciplinary reviews. Cognitive science · July 2018

Sam Hunley Stella F Lourenco

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What is peripersonal space? An examination of unresolved

Department of Psychology, Emory University,

body schema, individual differences, peripersonal space, personal space, spatial

WIREs Cogn Sci. 2018;e1472. wires.wiley.com/cogsci © 2018 Wiley Periodicals, Inc. 1 of 17

3.1 | The non-defensive function

VISUAL LINE BISECTION AS A MEASURE OF PPS

Rightward shift (% line length / m)

3.2 | The defensive function

3.3 | A network of PPS representation: integrating defensive and non-defensive functions

Sensory stimulus Sensory stimulus

General PPS map Defensive PPS map Working PPS map

Defensive action Working action Defensive action Working action

4.1 | PPS and body representations

4.2 | PPS versus personal space

5.1 | Individual differences in the flexibility of PPS

5.2 | How is PPS represented around the body?

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