Neuropsychologia ] (]]]]) ]]]–]]]

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Neuropsychologia
journal homepage: www.elsevier.com/locate/neuropsychologia

Vestibular stimulation modifies the body schema

Christophe Lopez a,b,n,1, Helene-Marianne Schreyer a,2, Nora Preuss a,2, Fred W. Mast a
a
Department of Psychology, University of Bern, Bern, Switzerland
b
Laboratoire de Neurosciences Intégratives et Adaptatives, UMR 7260, Centre National de la Recherche Scientifique (CNRS), Aix-Marseille Université, Marseille, France

a r t i c l e i n f o abstract

Article history: Mental body representations are flexible and depend on sensory signals from the body and its surrounding.
Received 1 December 2011 Clinical observations in amputees, paraplegics and brain-damaged patients suggest a vestibular contribu-
Received in revised form tion to the body schema, but studies using well-controlled psychophysical procedures are still lacking. In
21 March 2012
Experiment 1, we used a tactile distance comparison task between two body segments (hand and
Accepted 10 April 2012
forehead). The results showed that objects contacting the hand were judged longer during caloric vestibular
stimulation when compared to control thermal stimulation. In Experiment 2, participants located four
Keywords: anatomical landmarks on their left hand by pointing with their right hand. The perceived length and width
Vestibular system of the left hand increased during caloric vestibular stimulation with respect to a control stimulation. The
Body schema
results show that the body schema temporarily adjusts as a function of vestibular signals, modifying the
Body image
internal representation of the hand size. The data provide evidence that vestibular functions are not limited
Touch
Caloric vestibular stimulation to postural and oculomotor control, and extend the contribution of the vestibular system to bodily
Bodily consciousness cognition. The findings from this study suggest the inclusion of vestibular signals into current models of
Multisensory integration body representations and bodily self-consciousness.
& 2012 Elsevier Ltd. All rights reserved.

1. Introduction visually enlarged arm; Taylor-Clarke, Jacobsen, & Haggard, 2004)

Mental body representations have long been considered as
rather rigid maps, but recent studies provide firm evidence about
their malleability (Berlucchi & Aglioti, 2009; Serino & Haggard,
2010). In neurological patients, lesions and seizures in brain regions
involved in body representations can temporarily or permanently
modify the perceived size and shape of the entire body and
individual body segments (Dieguez, Staub, & Bogousslavsky,
2007). There is also a large amount of data indicating that in
neurologically normal population, mental body representations
continuously adjust as a function of exteroceptive and propriocep-
tive sensory signals (reviews in Serino & Haggard, 2010; de
Vignemont, 2010). Multisensory conflicts have been used as a
particularly interesting means to demonstrate the malleability of
mental body representations. For example, the perceived size of
body parts, measurable by the comparison of tactile distances
applied over various body segments, can be manipulated by view-
ing distorted visual feedback from the body (e.g. when looking at a
n
Corresponding author at: Universität Bern, Institut für Psychologie, Abteilung
für Kognitive Psychologie, Wahrnehmung und Methodenlehre, Muesmattstrasse,
45, 3012 Bern, Switzerland.
E-mail address: christophe.lopez@univ-amu.fr (C. Lopez).
1
Current address: Laboratoire de Neurosciences Intégratives et Adaptatives,
UMR 7260 CNRS – Aix-Marseille Université, Center St Charles, Pôle 3C – Case B 3,
Place Victor Hugo 13331 Marseille Cedex 03, France. Tel.: þ 33 4 13 55 08 41;
fax: þ 33 4 13 55 08 44.
2
Helene-Mariane Schreyer and Nora Preuss contributed equally to the
present study.
and proprioceptive-tactile conflicts (e.g. during the Pinocchio illu-
sion; de Vignemont, Ehrsson, & Haggard, 2005).
One fundamental aspect of mental body representations con-
cerns the so-called body schema (e.g. Gallagher, 2005; de Vignemont,
2010). According to Serino and Haggard (2010) the body schema
‘‘represents the positions of body parts in space, relative to each
other. It is of primarily proprioceptive origin, short-lived, and
updated as our bodies move. It serves to guide our actions and
our interactions with the external world’’ (p. 229). This type of
representation of the body configuration includes the size and shape
of body segments, i.e. body’s metric properties that need to be taken
into account during grasping and reaching objects. The multisensory
foundations of the body schema have been investigated a long time
ago, but a strong emphasis has been put on the proprioceptive
system. Holmes and Spence (2004), for example, wrote ‘‘that the
body schema should not necessarily be restricted to proprioceptive
and somatosensory modalities alone, but should also incorporate
visual and perhaps auditory information’’. Recently, cognitive neu-
roscience has made substantial advances in understanding the
multisensory mechanisms that underlie the body schema by manip-
ulating signals from the visual, tactile and proprioceptive systems
and, more recently, from the auditory system (Botvinick & Cohen,
1998; de Vignemont et al., 2005; Dieguez, Mercier, Newby, & Blanke,
2009; Tajadura-Jimenez et al., 2009; Taylor-Clarke et al., 2004;
Tsakiris & Haggard, 2005). However, the majority of studies on the
multisensory foundations of the body schema did not take into
account a contribution of vestibular signals. This is indeed surprising
given the fact that the body schema is continuously updated when

0028-3932/$ - see front matter & 2012 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.neuropsychologia.2012.04.008

Please cite this article as: Lopez, C., et al. Vestibular stimulation modifies the body schema. Neuropsychologia (2012), http://dx.doi.org/
10.1016/j.neuropsychologia.2012.04.008
2 C. Lopez et al. / Neuropsychologia ] (]]]]) ]]]–]]]
the body moves. The vestibular system is specialized in coding
three-dimensional body rotations and translations and body orien-
tations with respect to gravity (Angelaki & Cullen, 2008). Conse-
quently, relatively little is known about the influence of vestibular
signals on mental body representations. Exclusively focusing on
visual and tactile perception may hamper the elaboration of more
comprehensive multisensory models of the body schema. Although
some recent work in philosophy of mind has noted this shortcoming
(Macpherson, 2011), and despite recent attempts to include vestib-
ular signals in neuroscientific models of the bodily self (Blanke &
Metzinger, 2009), the interactions between the vestibular system
and other sensory systems remain largely underappreciated.
The contribution of the vestibular signals to the body schema
has nevertheless been recognized by some neurophysiologists like
Paillard (1991), for whom ‘‘the ubiquitous geotropic constraint
[i.e. gravitational acceleration, which is detected and coded by
vestibular receptors] dominates the [body-, world-, object- and
retina-centered] reference frames that are used in the visuomotor
control of actions and perceptions, and thereby becomes a crucial
factor in linking them together’’ (p. 472, our italics). According to
Paillard, gravity signals would therefore help merging and giving
coherence to the various references frames underpinning the body
schema. Human beings have evolved under a constant gravita-
tional field on Earth, and this physical constraint has shaped our
body representations. Therefore, grasping and reaching actions
performed under gravity are constrained by gravito-inertial forces
and preconscious internal models of gravity (McIntyre, Zago,
Berthoz, & Lacquaniti, 2001). Reaching a target during body
rotations generates Coriolis and centrifugal torques that tend to
deviate the hand from its optimal trajectory. It has been suggested
that vestibular signals generated during body movements are used
to correct the hand location and trajectory (Guillaud, Simoneau, &
Blouin, 2011). In conclusion, there is evidence that vestibular
signals update the body schema during hand actions (Bresciani
et al., 2002) and that they shape the way we perform actions and
interact with objects in the environment.
While the influence of vestibular signals on hand location and
motion is well-documented, the role of vestibular signals on the
representation of the body’s metric properties, i.e. the perceived
shape and size of the body, is less clear. Some reports from
parabolic flight missions (creating temporary weightlessness) point
out that the lack of gravitational input disorganizes the body
schema to the point that some participants experience a ‘‘tele-
scoping motion of the feet down and the head up internally
through the body’’, reversing the participant’s body orientation
(Lackner, 1992). Animal data confirm an influence of vestibular
signals on body representations since microgravity can perma-
nently disorganize the somatosensory maps in the primary soma-
tosensory cortex (Zennou-Azogui, Bourgeon, & Xerri, 2011). Clinical
observations provide compelling evidence that vestibular informa-
tion contributes to the body schema. Acute vestibular vertigo and
total vestibular deafferentation induce a drastic mismatch between
vestibular, visual and somatosensory signals, leading to distortions
of the body schema (Lopez & Blanke, 2007; Lopez, Halje, & Blanke,
2008; Sang, Jauregui-Renaud, Green, Bronstein, & Gresty, 2006;
Schilder, 1935). About 100 years ago, Bonnier (1905, 1907)
described distortions of the perceived shape and size of the body
in patients with vestibular disorders. One of his patients ‘‘felt that
he was divided into two persons’’, while another patient ‘‘felt his
head became enormous, immense, losing itself in the air; his body
disappeared and his whole being was reduced to only his face’’.
Schilder (1935) described in his monograph The image and appear-
ance of the human body some forms of alteration of the body
schema in vestibular patients who experienced that the ‘‘neck
swell during dizziness’’, ‘‘extremities had become larger’’, ‘‘hands
became larger and moved in different directions’’ and ‘‘feet seem to
Please cite this article as: Lopez, C., et al. Vestibular stimulation modifies the body schema. Neuropsychologia (2012), http://dx.doi.org/
10.1016/j.neuropsychologia.2012.04.008
elongate’’ (p. 117). Interestingly, Bonnier introduced the term
‘‘aschématie’’ (meaning ‘‘loss’’ of the schema) to account for the
various disorders of topographic body representations, including
the volume, shape and position of the body and body segments,
and he suggested that vestibular disorders evoke distortions of this
(body) ‘‘schema’’ (see Vallar & Rode, 2009). Recently, Rode et al.
(2012) described a patient suffering from vestibular disorders due
to a brainstem lesion with the concomitant sensation of macro-
somatognosia restricted to the face. The authors showed that
vestibular stimulation temporarily alleviated the distorted percep-
tion of the face, suggesting that vestibular signals can act on the
neural basis of the body schema. Moreover, it was found that
caloric vestibular stimulation (CVS, the irrigation of the auditory
canal with cold or warm water or air) can change the morpholo-
gical, postural and kinetic characteristics of phantom limbs in
paraplegics, and thus their body schema (Le Chapelain, Beis,
Paysant, & Andre, 2001). Similarly, in amputees, CVS evoked the
perception of a phantom limb in patients who did not experience
phantoms before, or altered the phantom perception in those who
experienced phantoms already, indicating that CVS can influence
mental representations of a non-existing body segment (André,
Martinet, Paysant, Beis, & Le Chapelain, 2001). In addition to this,
there is evidence for a beneficial influence of CVS on various bodily
disorders of neurological origin (e.g. disownership for body parts,
anosognosia and hemisanesthesia; see Bottini et al., 1995; Rode
et al., 1992). These findings have in common that they all suggest a
vestibular influence on the brain regions representing the body
structure. Whereas previous studies relied on self-reports there
still is a paucity of well-controlled psychophysical experimental
procedures. To our knowledge, tactile detection and illusory own-
ership for body parts were measured in healthy participants during
vestibular stimulation (Ferre, Sedda, Gandola, & Bottini, 2011;
Lopez, Lenggenhager, & Blanke, 2010), but these studies were not
designed to assess changes of the perceived size and shape of body
segments. Since previous reports did not investigate large popula-
tions of patients or did not use well-controlled paradigms, the
present study aims at testing vestibular-induced modifications of
the perceived size of body parts in a more controlled manner when
compared to previous studies. The hypothesis that vestibular
signals contribute to the body schema (more specifically the body’s
metric properties) is based on electrophysiological recordings in
animals showing that vestibular signals project to somatosensory
areas such as the hand and neck representations of the primary
somatosensory cortex (Schwarz, Deecke, & Fredrickson, 1973;
Schwarz & Fredrickson, 1971, review in Lopez & Blanke, 2011).
The secondary somatosensory cortex, insula and retroinsular
cortex also receive vestibular afferents (Bottini et al., 2001, 1995;
Lopez, Blanke, & Mast, in press). The anatomical overlap of tactile
and proprioceptive maps with vestibular maps provides the neural
basis for vestibular influence on body representations (Lopez et al.,
2008). In a first experiment, we investigated the body schema by
means of tactile signals in order to infer the relative metric
properties of two body parts. Importantly, it has been proposed
that ‘‘tactile input must be scaled by a pre-existing body model in
order to compute the size of the objects touching the skin’’ (Longo
et al., 2010; p. 659). Therefore, the potential influence of vestibular
stimulation on the body schema can be measured by a change in
the perceived size of objects touching the body surface. We used a
well-controlled paradigm based on the comparison of tactile
distances applied to separate body parts (de Vignemont et al.,
2005; de Vignemont, Majid, Jola, & Haggard, 2008; Taylor-Clarke
et al., 2004). This procedure has been shown to provide a sensitive
measure to assess modifications of the body schema due to visual
and proprioceptive stimulation. We manipulated tactile signals
from the palm (left hand) because its neurophysiological properties
have been well-described (Knibestöl & Vallbo, 1980), and we
 C. Lopez et al. / Neuropsychologia ] (]]]]) ]]]–]]] 3

compared this representation with that of a non-lateralized body
part (the sagittal line of the forehead). In a second experiment, we
assessed the body schema without involving tactile signals (Longo
& Haggard, 2010). This task allowed to measure directly the
representations of the body’s metric properties and did not involve
comparisons between two body parts. Participants localized four
anatomical landmarks on their left hand by pointing in darkness on
a digitizing tablet in order to obtain measurements of the
perceived length and width of the participant’s hand. During both
experiments, we provided exactly the same binaural CVS known to
activate the right cerebral hemisphere, in which the left hand is
represented. Based on overlapping neural networks between
tactile, proprioceptive and vestibular signals (Bottini et al., 2005,
1995; Lopez & Blanke, 2011), and in combination with clinical
findings showing the effects of CVS on the body schema (André
et al., 2001; Le Chapelain et al., 2001), we hypothesized that CVS
would modify the hand representation when compared to a
control thermal stimulation at body temperature.
2. Methods and results
2.1. Experiment 1: tactile distance judgments
2.1.1. Participants
The data were obtained from 18 students (age: 21  35 years; mean age7 SD:
25 74 years) who received credits for their participation. All of them were right-
handed (mean laterality quotient, Edinburgh Handedness inventory (Oldfield,
1971): þ 777 21%). Experimental procedures were approved by the Ethics
Committee of the Faculty of Human Sciences, University of Bern, and participants
gave written informed consent.
2.1.2. Caloric vestibular stimulation
We used caloric vestibular stimulation (CVS), which consists of injecting a
constant air flow into the auditory canals through a short plastic tube (Airmatic II,
GN Otometrics, Taastrup, Denmark). During CVS, warm air (47 1C) was injected in
the right ear and at the same time cold air (20 1C) was injected in the left ear. This
stimulation induces a perceived body motion to the right, and activates mostly the
right cerebral hemisphere, in which the left hand is predominantly represented
(Lopez & Blanke, 2011). In order to control for unspecific effects caused by CVS
(tactile and auditory stimulation), we applied a sham stimulation that consists of
injecting a constant air flow at body temperature (37 1C) simultaneously in both
ears. This stimulation produces exactly the same test conditions, with the only
difference that there is no vestibular stimulation.
2.1.3. Materials for the tactile distance judgment
The participants were blindfolded and they positioned their hand, palms up,
on padded armchairs. Experimental procedures were adapted from the studies by
Taylor-Clarke et al. (2004) and de Vignemont et al. (2005), in which tactile stimuli
were applied manually to the skin using solenoids or spheres, a condition in which
the pressure applied to the skin can vary across trials. We improved the procedure
by constructing a device that allows for controlling the pressure acting on the skin
surface (for similar procedures, see Ferre et al., 2011). Tactile stimuli were delivered
using a pair of plastic rods separated by distances of 10 mm, 20 mm, 30 mm and
40 mm (Fig. 1A). The plastic rods were screwed onto a rectangular piece of wood,
allowing for the precise adjustment of the distance between the participant’s hand
and this support by varying the length of each plastic rod under the support
(distance h1). A rectangular box was attached to the participant’s left hand and
another rectangular box was attached to their forehead. The shape of the boxes was
designed to fit the shape of the hand and the curvature of the forehead, and they
were attached to the body using Velcros strips (Fig. 1B). Two screws were mounted
vertically on the boxes and were designed to receive the support for the tactile
stimuli (see Fig. 1C for an illustration of the hand box; similar procedures were used
for the forehead box). On each of these screws, a screw nut served to adapt for each
participant the distance (distance h2) between the support for the tactile stimuli
and their left hand or the midline of their forehead. Before the experiment proper,
each tactile stimulus was tested and the distances h1 and h2 were adapted to the
morphology of the participant’s left hand and forehead. Importantly, due to the

adjustable
height (h2)
8c
m

12 cm
4.5 cm

adjustable
height (h1)

4 tactile distances

10 mm

20 mm

30 mm

40 mm

Fig. 1. Experimental set-up for the tactile distance comparisons task. (A) Wooden support for the tactile stimuli. Two plastic rods separated by distances ranging from 10
to 40 mm are screwed on a rectangular piece of wood. (B) Schematic representation of the hand box designed to receive the tactile stimuli. (C) The support for the tactile
stimuli can be inserted in the hand box and is held in place by two screws. The adjustment of distances h1 and h2 for each participant allows for applying constant tactile
stimuli on the palmar surface of the left hand.

Please cite this article as: Lopez, C., et al. Vestibular stimulation modifies the body schema. Neuropsychologia (2012), http://dx.doi.org/
10.1016/j.neuropsychologia.2012.04.008
4 C. Lopez et al. / Neuropsychologia ] (]]]]) ]]]–]]]

presence of the screw nuts on the boxes, the pressure applied on the hand was
maintained constant across trials.
2.1.4. Experimental procedures
The tactile distance comparison task was conducted during two conditions:
CVS and sham stimulation. There were two blocks of CVS and two blocks of sham
stimulation, presented in a counterbalanced order across participants. Each block
consisted of the following procedures: Participants were initially seated on a
tiltable chair with their head pitched forward by 301 and they received CVS or
sham for 2 min. Participants were subsequently pitched backward by 901 until
their head was tilted 601 with respect to the vertical. In this position, the
horizontal semicircular canals were oriented vertically, providing a maximal effect
of CVS (for a detailed description see Mast, Merfeld, and Kosslyn (2006)). During
CVS, participants performed the tactile distance comparison task. The experi-
menter applied one tactile stimulus on the left hand immediately followed by one
tactile stimulus on the forehead (this procedure was used in half of the
participants, while in the other half the tactile stimulus on the left hand followed
the stimulus applied to the forehead). The participants were required to answer as
quickly and spontaneously as possible whether the longer distance was felt on
their hand or forehead. For each condition (CVS and sham) 28 tactile stimuli were
applied in a randomized order: Similar distances between the hand and the
forehead were applied 16 times in total (the pairs 10–10 mm, 20–20 mm, 30–
30 mm, 40–40 mm were each applied 4 times) and trials with the hand stimulus
longer than the forehead stimulus (20–10 mm, 30  20 mm, 40–30 mm), or the
forehead stimulus longer than the hand stimulus (10–20 mm, 0–30 mm, 30–
40 mm) were applied 12 times in total. The 28 tactile stimuli were presented in
two separate blocks lasting for about 3 min each. After the completion of each
block, participants were turned back to the upright position for 5 min of rest and
they filled out questionnaires.
2.1.6. Eye movements recording
After the experiment proper, eye movements were recorded by videonystag-
mography (eVNG, BioMed, Jena, Germany) during 45s of CVS and sham stimula-
tion. We computed the mean slow phase velocity of the nystagmus in degrees
per second as in previous reports (Lopez, Borel, Magnan, & Lacour, 2005) and
included in the final analysis only those participants who exhibited a clear caloric
nystagmus (n¼15; 3 participants were excluded due to the absence of a clear
caloric nystagmus). The nystagmus served as a control for effective vestibular
stimulation by means of CVS.
2.1.7. Results
We quantified the percentage of times participants experienced the tactile
stimulus applied on the hand as longer than the same tactile distance applied on the
forehead. This analysis included only trials presenting same tactile distances on the
hand and forehead. A repeated-measures ANOVA revealed a higher occurrence of
the perception of the hand being longer than the forehead during CVS (mean7
SEM: 58.373.1%) when compared to sham stimulation (48.872.9%; F1,14 ¼ 8.9;
Po0.05) (Fig. 2B). A distance applied on the hand was perceived as longer than the
same distance applied on the forehead during CVS in 12 out of 15 participants
(Fig. 2A). Two participants had comparable results during CVS and sham and only
one participant showed the opposite pattern. This result indicates that CVS can
induce a change in the perceived length of external objects touching the left hand.
Additional recordings indicate that binaural CVS evoked a nystagmus with a
mean slow phase eye velocity of 4.1 7 3.41/s. No nystagmus was evoked during
sham stimulation. The analysis of the questionnaire data revealed a stronger
illusory self-motion during CVS (mean strength of illusory motion: 3.17 0.5) when
compared to sham stimulation (0.77 0.2; F1,14 ¼ 28; P o 0.001). The analysis of the
Cox and Swinson questionnaire (two-sided paired t-tests) revealed stronger
ratings after CVS than sham stimulation for several items related to bodily
sensations (Fig. 2C). Particularly, during CVS, participants reported feeling ‘spacy’,
being detached from their own body and the surroundings, and not being in
control of their self.

2.1.5. Subjective reports

Participants filled-out a questionnaire designed to evaluate self-motion during
CVS and sham stimulation (adapted from (Lenggenhager, Lopez, & Blanke, 2008)
and (Lopez et al., 2010)). They reported the magnitude of the sensation of body
motion using a 7-point scale ranging from 0 (‘‘no motion’’) to 6 (‘‘strong motion’’).
Participants also filled out the Cox & Swinson (2002) questionnaire to evaluate
bodily sensations resembling possible depersonalization symptoms immediately
after CVS or sham stimulation (Sang et al., 2006). This questionnaire contains 28
items and we extracted those items related to bodily sensations such as ‘‘Body
feels strange/different in some way’’, ‘‘Body feels numb’’ and ‘‘Feeling detached or
separated from body’’. Participants rated the intensity of each item using a 5-point
scale ranging from 0 (‘‘does not occur’’) to 4 (‘‘very severe’’).
2.2. Experiment 2. localization of anatomical landmarks
Results from Experiment 1 indicate a change in tactile size judgment on the
left hand relative to the forehead. The representation of the left hand is lateralized
in the right cerebral hemisphere, whereas the representation of the forehead’s
medial skin is poorly lateralized (Bittar, Nandi, Carter & Aziz, 2005; Iannetti et al.,
2003). Given the fact that CVS activates the right hemisphere, we hypothesized
that it has modified the representation of the left hand. Experiment 2 was
designed to directly test the hypothesis according to which CVS has led to an
increase in the perceived hand size. We used a paradigm to obtain an objective
measure of the representation of the body’s metric properties without involving

CVS CVS%
80 CVS 80 CVS 1.8 sham sham% 80
sham sham
1.6

Frequency of reported sensation (%)

70
70 70
mean % hand longer than forehead

1.4
60
% hand longer than forehead

Intensity of subjective report

60 60 1.2
50
1.0
50 50 40
0.8
30
40 40 0.6
20
0.4
30 30
0.2 10

20 20 0.0 0
Q1 Q2 Q3 Q4 Q5

Fig. 2. Influence of vestibular stimulation on the tactile distance comparison task and bodily sensations. (A) The percentage of trials for which the tactile stimulus applied
on the left hand is reported as longer than the same stimulus applied on the forehead is illustrated for each participant (P1  P15). (B) Histograms depict the average
data7 SEM (Po 0.05). (C) Rating of 5 item questionnaires from the depersonalization-derealization questionnaire after CVS and sham stimulation (Cox & Swinson, 2002).
Leftward y-axis and histograms depict the mean intensity ( 7 SEM) of each sensation. Rightward y-axis and lines depict the occurrence (in %) of each bodily sensation. Q1:
Feel ‘‘spacy’’ or ‘‘spaced out’’; Q2: Feeling of not being in control of self; Q3: Surroundings seem strange or unreal; Q4: Feeling of detachment or separation from
surroundings; Q5: Feeling detached or separated from your body (P o0.05, t-tests on the intensity of the rating).

Please cite this article as: Lopez, C., et al. Vestibular stimulation modifies the body schema. Neuropsychologia (2012), http://dx.doi.org/
10.1016/j.neuropsychologia.2012.04.008
 C. Lopez et al. / Neuropsychologia ] (]]]]) ]]]–]]] 5

tactile stimulation. In addition, this task did not involve comparisons between two
body parts.
2.2.1. Participants
The data were obtained from 17 participants (age: 22  29 years; mean
age7 SD: 25 7 2 years). All of them were right-handed (mean laterality quotient,
Edinburgh Handedness inventory (Oldfield, 1971): þ79 7 20%).
increase in the perceived length of the left hand (P o0.05, one-sided paired t-test),
as hypothesized from Experiment 1. The data also indicate a significant increase in
the perceived width of the left hand (P o0.05, one-sided paired t-test). As for
Experiment 1, questionnaire analyses showed a stronger illusory self-motion
during CVS when compared to sham stimulation (F1,16 ¼ 7.54; Po 0.05). The Cox
and Swinson’s questionnaire revealed that participants reported stronger changes
in bodily perceptions (e.g. ‘‘body feels strange or different in some way’’) during
CVS than sham stimulation (Po 0.05, two-sided paired t-test).

2.2.2. Materials for the localization task

Experimental procedures were adapted from Longo and Haggard (2010).
Participants were blindfolded and they positioned their left hand, palm down,
on a table in front of them so that their hand was aligned with their body mid-
sagittal plane. The hand was positioned under a digitizing tablet. The participants
were asked to locate four anatomical landmarks on the dorsal surface of their left
hand: the knuckle of the little finger, the knuckle of the index finger, the tip of
the middle finger and the wrist. Each anatomical landmark was initially shown to
the participant and stickers were applied to the skin on the corresponding location
(see Fig. 3B and C). Participants used a stylus to indicate with their right hand the
location of the landmarks. The digitizing tablet automatically recorded the x and y
coordinates of the location indicated by the participant.
2.2.3. Experimental procedures
The localization of anatomical landmarks was conducted during one block of
CVS and one block of sham stimulation, presented in a counterbalanced order
across participants. CVS and sham stimulation were applied in the same way as for
Experiment 1 (see Mast et al., 2006). Participants performed the localization task
when lying backwards and while receiving CVS in their outer ear canals. For each
condition (CVS and sham), participants judged the location of the four anatomical
landmarks in four blocks. Each block consisted of 10 consecutive pointings to the
same anatomical landmark. The order in which they had to locate the landmarks
was randomized across participants. For each participant, we calculated the mean
x and y coordinates of the perceived position of each anatomical landmark. Hence,
it was possible to obtain an objective measure of the perceived length of the hand
(Euclidean distance between the tip of the middle finger and the wrist) and of the
width of the hand (Euclidean distance between the knuckle of the little finger and
the knuckle of the index finger). After completion of each block, participants were
turned back to the upright position for 5 min of rest and they filled out the same
questionnaires as for Experiment 1.
2.2.4. Results
Fig. 3A illustrates the perceived length and width of the left hand during CVS
and sham stimulation. The analysis of the pointing task revealed a significant
3. Discussion
The results indicate that tactile stimuli applied on the left hand
were perceived as longer during CVS than during a control
stimulation not activating vestibular receptors (Experiment 1) and
that CVS increased the perceived length and width of the left hand
during a task requiring to locate anatomical landmarks on the hand
(Experiment 2). Altogether, the data indicate that vestibular stimu-
lation was able to modify the instantaneous representation of body
segments, suggesting an influence of vestibular signals on the
neural mechanisms underlying the body schema. The present
study provides further evidence that body representations are
not rigid but rather adapt to sensory signals originating from the
body and its environment (review in Serino & Haggard, 2010). The
data from this study extend previous findings showing changes in
perceived size and posture of body segments during experimental
manipulation of bodily signals. This is the case, for example, during
muscular vibrations and activation of proprioceptive receptors (de
Vignemont et al., 2005; Ehrsson, Kito, Sadato, Passingham, & Naito,
2005; Lackner, 1988), during presentation of distorted visual
feedback from the body (Taylor-Clarke et al., 2004), and during
tactile illusions (Bruno & Bertamini, 2010; Dieguez et al., 2009).
Here, we show distortions of the body schema during vestibular
stimulation. Although the contribution of vestibular signals to the
coding of whole-body movements and orientation is obvious, its
contribution to body segments perception may appear less evident.
However, it has been suggested that vestibular signals are

Fig. 3. Influence of vestibular stimulation on the localization of anatomical landmarks. (A) The mean perceived width (distance between the knuckle of the little finger and
the knuckle of the index finger) and length (distance between the tip of the middle finger and the wrist) of the left hand is represented for CVS and sham stimulation. Error
bars represent SEM. (B, C) The perceived location of the four anatomical landmarks is illustrated for each participant during CVS (part B) and sham stimulation (part C). The
photograph of the hand represents the hand of one participant with the four anatomical landmarks and does not reflect the average hand size or hand position in the
population.

Please cite this article as: Lopez, C., et al. Vestibular stimulation modifies the body schema. Neuropsychologia (2012), http://dx.doi.org/
10.1016/j.neuropsychologia.2012.04.008
6 C. Lopez et al. / Neuropsychologia ] (]]]]) ]]]–]]]
important to code for fine arm reaching movements (Bresciani
et al., 2002). Importantly, the central nervous system uses vestib-
ular signals to elaborate an internal model of gravity, referencing
arm movements and position (McIntyre et al., 2001).
In addition to this, vestibular signals control the tonus and
contractions of limb muscles through vestibulospinal reflexes
(Britton et al., 1993) and this information is fed back to the brain
to update the current posture of the body. Therefore, vestibular
signals are involved in sensorimotor circuits between the brain and
body parts.
The data demonstrate that a tactile distance applied on the
skin is perceived as longer during CVS than during sham stimula-
tion. The question is whether this change in the perception of
external objects reflects a perceived enlargement or reduction of
the body part that is in contact with these objects. A previous
study by de Vignemont et al. (2005) used similar experimental
procedures and provides a clear conclusion. They evoked the
experience of an elongation of the index finger by vibrating biceps
muscles and asked participants to perform a tactile distance
comparison task (between the finger and the forehead). They
showed that ‘‘a tactile distance feels bigger when the stimulated
body part feels temporarily elongated’’ (de Vignemont et al.,
2005; p. 1288). In another study by Bruno and Bertamini
(2010), the participant’s hand was stroked synchronously with a
rubber hand, the size of which was enlarged or reduced. When
their hand was synchronously stroked with an enlarged rubber
hand, participants judged an external object touching the stroked
hand to be larger than an identical object touching the contral-
ateral (unstimulated) hand. Conversely, they judged an external
object to be smaller when they were exposed to a reduced rubber
hand. The authors concluded that the altered mental representa-
tion of the hand modified the interpretation of active touch
(Bruno & Bertamini, 2010). Data from Experiment 1 showed that
CVS modifies a tactile size judgment on the left hand relative to
the forehead. However, the results do not permit to draw firm
conclusions about whether CVS has led to an increase in the
perceived hand size or to a decrease in the perceived forehead
size. The latter is unlikely because the representation of the
medial forehead on which the tactile stimuli were applied is
poorly lateralized. Neuroimaging studies and deep brain stimula-
tion showed no lateralization of the forehead neither in the
primary nor in the secondary somatosensory cortex (Iannetti
et al., 2003; Bittar et al., 2005). The paradigm we used activates
the right cerebral hemisphere (see Lopez et al., in press) in which
the left hand is more dominantly represented (Ruben et al., 2001).
In Experiment 2, we demonstrated that CVS led to an increase in
the perceived hand size with respect to a control stimulation.
Previous psychophysical experiments demonstrated that the
body’s metric properties (such as the distance between the
knuckles of the little and index fingers) are based on a body
model (Longo & Haggard, 2010). Therefore, the fact that the
perceived length and width of the left hand were increased
during CVS demonstrates the influence of vestibular signals on
the body schema.
To date, the influence of vestibular signals upon body schema
has been suggested by subjective reports in various clinical
populations. For example, in lower limb amputees, CVS evoked
a change in the shape or position of the phantom limb (André
et al., 2001). Disturbance in the coding of vestibular signals can
also lead to abnormal bodily perceptions (fore reviews, see Lopez
& Blanke, 2007; Lopez et al., 2008). Oto-neurological patients
suffering from vestibular dysfunctions (such as Menie re’s disease)
may report changes in the perceived shape and size of the body
(Bonnier, 1905, 1907; Schilder, 1935; Rode et al., 2012). These
authors have described enlargement, elongation, or increase in
the volume at the level of the hands, neck, feet and face,
Please cite this article as: Lopez, C., et al. Vestibular stimulation modifies the body schema. Neuropsychologia (2012), http://dx.doi.org/
10.1016/j.neuropsychologia.2012.04.008
indicating that vestibular signals interact with the body model
of various segments. In the same vein, several authors reported
the co-occurrence of vestibular illusions and distortions of the
body schema in brain-damaged and epileptic patients (Blanke,
Landis, Spinelli, & Seeck, 2004; Hécaen & de Ajuriaguerra, 1952;
Heydrich, Lopez, Seeck, & Blanke, 2011). Thus, distortions in
mental body representations can either be caused by pathological
conditions affecting the peripheral vestibular sensors or at cor-
tical levels processing vestibular signals.
Vestibular signals project to multiple multisensory brain regions
(review in Lopez & Blanke, 2011), and the cortical processing of
vestibular information can possibly account for the behavioral results
we found in this study. Several brain areas are associated with
vestibular–somatosensory interactions. First, electrophysiological
recordings in monkeys showed convergence of tactile, proprioceptive
and vestibular signals in the primary somatosensory cortex. For
example, areas 3aHv and 3aNv are two vestibular regions located in
the hand and neck representations of the primary somatosensory
cortex (Schwarz et al., 1973; Schwarz & Fredrickson, 1971). Second,
interactions of tactile, proprioceptive (from the legs and neck) and
vestibular signals were reported in the parieto-insular vestibular
cortex (Grüsser, Pause, & Schreiter, 1990) and in the intraparietal
cortex (Bremmer, Klam, Duhamel, Ben Hamed, & Graf, 2002).
Neuroimaging studies in humans also revealed that vestibular regions
overlap with somatosensory regions in the insular, retroinsular and
secondary somatosensory cortex, as well as in the premotor cortex
and supramarginal gyrus (Bottini et al., 2001, 1995). The temporo-
parietal junction is yet another candidate because it is the site of
convergence of various sensory signals from the body and its
environment (Lopez et al., 2008). Indeed, electrical stimulations of
the temporo-parietal junction in a conscious epileptic patient mod-
ified body representations (e.g., the patient’s arm appeared shorter)
and evoked vestibular sensations (Blanke, Ortigue, Landis, & Seeck,
2002). This study provided a direct demonstration of a brain region
commonly involved in the conscious experience of the shape and size
of body segments and in vestibular processing. However, more work
is needed to precisely determine vestibular-somatosensory interac-
tions, but the wide range of vestibular projections at the cortical level
suggests that such interactions can occur in several brain regions
simultaneously.
We believe that the present research will have theoretical and
conceptual impacts by pointing out the importance to include
vestibular signals and vestibular representations in current multi-
sensory models of the body schema. However, not much is known
about the exact mechanisms by which vestibular signals con-
struct the body schema. On the basis of the effects of CVS on the
shape and size of phantom limbs, André et al. (2001) proposed
that the vestibular system ‘‘triggers the procedure of reconstruc-
tion of the global body schema’’ (p. 190). Others have proposed
that vestibular signals link the different spatial references frames
on which bodily processing is based on (Lopez & Blanke, 2007).
We have proposed that the multisensory vestibular cortex (that
overlaps with the somatosensory cortex) should be important for
the body schema, because it integrates signals from the personal
(tactile, proprioceptive) space, the allocentric (visual) space, and
the gravitational, or geocentric, space (vestibular otolithic sig-
nals). Therefore, this cortex has the capacity to combine several
reference frames, to maintain the unity of the spatial experience,
and therefore the potential to contribute to the neural represen-
tation of the body schema and the body’s metric properties
(the shape, size and weight of the different body segments).
In conclusion, our data indicate that vestibular signals influence
the body schema in healthy participants. Importantly, the data
provide further evidence that vestibular functions are not limited
to postural and oculomotor reflexes, and extend the contribution
of the vestibular system to bodily cognition. We hope that our
 C. Lopez et al. / Neuropsychologia ] (]]]]) ]]]–]]]
findings will inspire other researchers to include vestibular
information into current models of body representations and
bodily self-consciousness.
Acknowledgments
We thank M. Hunziker, H. Hutmacher, F. Meyer, B. Schreyer,
K. Wegmüller, V. Gashaj and C. Mozzini Vellen for technical
support. This study was supported by the Swiss National Science
Foundation (SINERGIA project ‘Balancing Self and Body’: CRSII1-
125135). C. Lopez is supported by the Volkswagenstiftung’s
European Platform for Life Sciences, Mind Sciences, and the
Humanities (‘The (Un)bound Body Project. Exploring the constraints
of embodiment & the limits of body representation’).
References
André, J. M., Martinet, N., Paysant, J., Beis, J. M., & Le Chapelain, L. (2001).
Temporary phantom limbs evoked by vestibular caloric stimulation in ampu-
tees. Neuropsychiatry, Neuropsychology, and Behavioral Neurology, 14, 190–196.
Angelaki, D. E., & Cullen, K. E. (2008). Vestibular system: The many facets of a
multimodal sense. Annual Review of Neuroscience, 31, 125–150.
Berlucchi, G., & Aglioti, S. M. (2009). The body in the brain revisited. Experimental
Brain Research, 200, 25–35.
Bittar, R. G., Nandi, D., Carter, H., & Aziz, T. Z. (2005). Somatotopic organization of
the human periventricular gray matter. Journal of Clinical Neuroscience, 12,
240–241.
Blanke, O., & Metzinger, T. (2009). Full-body illusions and minimal phenomenal
selfhood. Trends in Cognitive Science, 13, 7–13.
Blanke, O., Landis, T., Spinelli, L., & Seeck, M. (2004). Out-of-body experience and
autoscopy of neurological origin. Brain, 127, 243–258.
Blanke, O., Ortigue, S., Landis, T., & Seeck, M. (2002). Stimulating illusory own-
body perceptions. Nature, 419, 269–270.
Bonnier, P (1905). L’Aschématie. Revue de Neurologie (Paris), 12, 605–609.
Bonnier, P (1907). Le vertige. Paris: Masson.
Bottini, G., Karnath, H. O., Vallar, G., Sterzi, R., Frith, C. D., Frackowiak, R. S., et al.
(2001). Cerebral representations for egocentric space: Functional–anatomical
evidence from caloric vestibular stimulation and neck vibration. Brain, 124,
1182–1196.
Bottini, G., Paulesu, E., Gandola, M., Loffredo, S., Scarpa, P., Sterzi, R., et al. (2005).
Left caloric vestibular stimulation ameliorates right hemianesthesia. Neurol-
ogy, 65, 1278–1283.
Bottini, G., Paulesu, E., Sterzi, R., Warburton, E., Wise, R. J., Vallar, G., et al. (1995).
Modulation of conscious experience by peripheral sensory stimuli. Nature, 376,
778–781.
Botvinick, M., & Cohen, J. (1998). Rubber hands ‘feel’ touch that eyes see. Nature,
391, 756.
Bremmer, F., Klam, F., Duhamel, J. R., Ben Hamed, S., & Graf, W. (2002). Visual-
vestibular interactive responses in the macaque ventral intraparietal area
(VIP). European Journal of Neuroscience, 16, 1569–1586.
Bresciani, J. P., Blouin, J., Popov, K., Bourdin, C., Sarlegna, F., Vercher, J. L., et al.
(2002). Galvanic vestibular stimulation in humans produces online arm
movement deviations when reaching towards memorized visual targets.
Neuroscience Letters, 318, 34–38.
Britton, T. C., Day, B. L., Brown, P., Rothwell, J. C., Thompson, P. D., & Marsden, C. D.
(1993). Postural electromyographic responses in the arm and leg following
galvanic vestibular stimulation in man. Experimental Brain Research, 94,
143–151.
Bruno, N., & Bertamini, M. (2010). Haptic perception after a change in hand size.
Neuropsychologia, 48, 1853–1856.
Cox, B. J., & Swinson, R. P. (2002). Instrument to assess depersonalization–
derealization in panic disorder. Depression and Anxiety, 15, 172–175.
de Vignemont, F. (2010). Body schema and body image—Pros and cons. Neurop-
sychologia, 48(3), 669–680.
de Vignemont, F., Ehrsson, H. H., & Haggard, P. (2005). Bodily illusions modulate
tactile perception. Current Biology, 15, 1286–1290.
de Vignemont, F., Majid, A., Jola, C., & Haggard, P. (2008). Segmenting the body into
parts: Evidence from biases in tactile perception. The Quaterly Journal of
Experimental Psychology, 62, 500–512.
Dieguez, S., Mercier, M. R., Newby, N., & Blanke, O. (2009). Feeling numbness for
someone else’s finger. Current Biology, 19, R1108–R1109.
Dieguez, S., Staub, F., & Bogousslavsky, J. (2007). Asomatognosia. In:
J. Bogousslavsky, & O. Godefroy (Eds.), The behavioral and cognitive neurology
of stroke (pp. 215–253). Cambridge University Press.
Ehrsson, H. H., Kito, T., Sadato, N., Passingham, R. E., & Naito, E. (2005). Neural
substrate of body size: Illusory feeling of shrinking of the waist. PLoS Biology, 3,
e412.
Please cite this article as: Lopez, C., et al. Vestibular stimulation modifies the body schema. Neuropsychologia (2012), http://dx.doi.org/
10.1016/j.neuropsychologia.2012.04.008
7
Ferre, E. R., Sedda, A., Gandola, M., & Bottini, G. (2011). How the vestibular system
modulates tactile perception in normal subjects: A behavioural and physiolo-
gical study. Experimental Brain Research, 208, 29–38.
Gallagher, S. (2005). How the body shapes the mind. New York: Oxford University
Press.
Grüsser, O. J., Pause, M., & Schreiter, U. (1990). Vestibular neurones in the parieto-
insular cortex of monkeys (Macaca fascicularis): Visual and neck receptor
responses. Journal of Physiology, 430, 559–583.
Guillaud, E., Simoneau, M., & Blouin, J. (2011). Prediction of the body rotation
induced torques on the arm during reaching movements: Evidence from a
proprioceptively deafferented subject. Neuropsychologia, 49, 2055–2059.
Hécaen, H., & de Ajuriaguerra, J. (1952). Méconnaissances et hallucinations corpor-
elles. Intégration et désintégration de la somatognosie. Paris: Masson.
Heydrich, L., Lopez, C., Seeck, M., & Blanke, O. (2011). Partial and full own-body
illusions of epileptic origin in a child with right temporoparietal epilepsy.
Epilepsy & Behavior, 20, 583–586.
Holmes, N. P., & Spence, C. (2004). The body schema and the multisensory
representation(s) of peripersonal space. Cognitive Processing, 5(2), 94–105.
Iannetti, G. D., Porro, C. A., Pantano, P., Romanelli, P. L., Galeotto, F., & Cruccu, G.
(2003). Representation of different trigeminal divisions within the primary
and secondary human somatosensory cortex. Neuroimage, 19, 906–912.
Knibestöl, M., & Vallbo, A. B. (1980). Intensity of sensation related to activity of
slowly adapting mechanoreceptive units in the human hand. Journal of
Physiology, 300, 251–267.
Lackner, J. R. (1988). Some proprioceptive influences on the perceptual represen-
tation of body shape and orientation. Brain, 111, 281–297.
Lackner, J. R. (1992). Spatial orientation in weightless environments. Perception,
21, 803–812.
Le Chapelain, L., Beis, J. M., Paysant, J., & Andre, J. M. (2001). Vestibular caloric
stimulation evokes phantom limb illusions in patients with paraplegia. Spinal
Cord, 39, 85–87.
Lenggenhager, B., Lopez, C., & Blanke, O. (2008). Influence of galvanic vestibular
stimulation on egocentric and object-based mental transformations. Experi-
mental Brain Research, 184, 211–221.
Longo, M. R., Azañón, E., & Haggard, P. (2010). More than skin deep: body
representation beyond primary somatosensory cortex. Neuropsychologia,
48(3), 655–668.
Longo, M. R., & Haggard, P. (2010). An implicit body representation underlying
human position sense. Proceedings of the National Academy of Sciences USA,
107(26), 11727–11732.
Lopez, C., & Blanke, O. (2007). Neuropsychology and neurophysiology of self-
consciousness. Multisensory and vestibular mechanisms. In: A. Holderegger,
B. Sitter-Liver, C. W. Hess, & G. Rager (Eds.), Hirnforschung und Menschenbild.
Beiträge zur interdisziplinären Verständigung (pp. 183–206). Fribourg, Basel:
Academic Press, Schwabe.
Lopez, C., & Blanke, O. (2011). The thalamocortical vestibular system in animals
and humans. Brain Research Reviews, 67, 119–146.
Lopez, C., Borel, L., Magnan, J., & Lacour, M. (2005). Torsional optokinetic
nystagmus after unilateral vestibular loss: Asymmetry and compensation.
Brain, 128, 1511–1524.
Lopez, C., Halje, P., & Blanke, O. (2008). Body ownership and embodiment:
Vestibular and multisensory mechanisms. Neurophysiologie Clinique/Clinical
Neurophysiology, 38, 149–161.
Lopez, C., Lenggenhager, B., & Blanke, O. (2010). How vestibular stimulation
interacts with illusory hand ownership. Consciousness & Cognition, 19,
33–47.
Lopez, C., Blanke, O., & Mast F. W.. The vestibular cortex in the human brain
revealed by coordinate-based activation likelihood estimation meta-analysis.
Neuroscience, http://dx.doi.org/10.1016/j.neuroscience.2012.03.028, in press.
Macpherson, F. (2011). The senses: Classical and contemporary philosophical per-
spectives. Oxford University Press.
Mast, F. W., Merfeld, D. M., & Kosslyn, S. M. (2006). Visual mental imagery during
caloric vestibular stimulation. Neuropsychologia, 44, 101–109.
McIntyre, J., Zago, M., Berthoz, A., & Lacquaniti, F. (2001). Does the brain model
Newton’s laws?. Nature Neuroscience, 4, 693–694.
Oldfield, R. C. (1971). The assessment and analysis of handedness: The Edinburgh
inventory. Neuropsychologia, 9, 97–113.
Paillard, J. (1991). Knowing where and knowing how to get there. In: J. Paillard
(Ed.), brain and space (pp. 461–481). Oxford University Press.
Rode, G., Charles, N., Perenin, M. T., Vighetto, A., Trillet, M., & Aimard, G. (1992).
Partial remission of hemiplegia and somatoparaphrenia through vestibular
stimulation in a case of unilateral neglect. Cortex, 28, 203–208.
Rode, G., Vallar, G., Revol, P., Tilikete, C., Jacquin-Courtois, S., Rossetti, Y., et al.
(2012). Facial macrosomatognosia and pain in a case of Wallenberg’s syn-
drome: Selective effects of vestibular and transcutaneous stimulations. Neu-
ropsychologia, 50(2), 245–253.
Ruben, J., Schwiemann, J., Deuchert, M., Meyer, R., Krause, T., Curio, G., et al.
(2001). Somatotopic organization of human secondary somatosensory cortex.
Cerebral Cortex, 11(5), 463–473.
Sang, F. Y., Jauregui-Renaud, K., Green, D. A., Bronstein, A. M., & Gresty, M. A.
(2006). Depersonalisation/derealisation symptoms in vestibular disease. Jour-
nal of Neurolology, Neurosurgery & Psychiatry, 77, 760–766.
Schilder, P. (1935). The image and appearance of the human body. New York:
International Universities Press.
Schwarz, D. W. F., & Fredrickson, J. M. (1971). Rhesus monkey vestibular cortex:
A bimodal primary projection field. Science, 172, 280–281.
8 C. Lopez et al. / Neuropsychologia ] (]]]]) ]]]–]]]

Schwarz, D. W. F., Deecke, L., & Fredrickson, J. M. (1973). Cortical projection of
group I muscle afferents to areas 2, 3a, and the vestibular field in the rhesus
monkey. Experimental Brain Research, 17, 516–526.
Serino, A., & Haggard, P. (2010). Touch and the body. Neuroscience and Biobeha-
vioral Reviews, 34, 224–236.
Tajadura-Jimenez, A., Kitagawa, N., Valjamae, A., Zampini, M., Murray, M. M., &
Spence, C. (2009). Auditory–somatosensory multisensory interactions are
spatially modulated by stimulated body surface and acoustic spectra. Neu-
ropsychologia, 47, 195–203.
Taylor-Clarke, M., Jacobsen, P., & Haggard, P. (2004). Keeping the world a constant
size: Object constancy in human touch. Nature Neuroscience, 7, 219–220.
Tsakiris, M., & Haggard, P. (2005). The rubber hand illusion revisited: Visuotactile
integration and self-attribution. Journal of Experimental Psychology: Human
Perception and Performance, 31, 80–91.
Vallar, G., & Rode, G. (2009). Commentary on Bonnier P. L’aschématie. Rev Neurol
(Paris) 1905;13:605-9. Epilepsy & Behavior, 16(3), 397–400.
Zennou-Azogui, Y., Bourgeon, S., & Xerri, C. (2011). Hypergravity experience during
development alters forepaw somatosensory maps and influences cortical experi-
ence-dependent plasticity in adult rat. In Proceedings of the conference on
development and plasticity of thalamocortical systems. Arolla, Switzerland.

Please cite this article as: Lopez, C., et al. Vestibular stimulation modifies the body schema. Neuropsychologia (2012), http://dx.doi.org/
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