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Professionnel Documents
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DOI 10.3758/s13414-014-0748-6
K. T. Graham : M. T. Martin-Iverson
Pharmacology, Pharmacy & Anaesthesiology Unit, School of
Medicine and Pharmacology, University of Western Australia,
Crawley, WA, Australia
K. T. Graham : M. T. Martin-Iverson
Statewide Department of Neurophysiology, Clinical Research Unit,
North Metro Area Mental Health, Graylands Hospital, Perth,
Australia
N. P. Holmes
Centre for Integrative Neuroscience and Neurodynamics & School of
Psychology & Clinical Language Sciences, University of Reading,
Reading, Berkshire, UK
F. A. Waters (*)
Clinical Research Centre, North Metro Health Service Mental
Health, Graylands Hospital, Claremont, WA, Australia
e-mail: flavie.waters@health.wa.gov.au
F. A. Waters
School of Psychiatry and Clinical Neurosciences, The University of
Western Australia, 35 Stirling Highway, Crawley, WA 6009,
Australia
Introduction
In the past 16 years, the Rubber Hand Illusion (RHI,
Botvinick & Cohen, 1998) has received much interest as an
experimental paradigm for investigating mechanisms of selfperception, which may be defined as the perception of ones
self, including thoughts and actions, as residing within the
physical body (Arzy et al., 2006a, b). In the illusion, a fake
hand is placed in front of a participant and stroked with a
paintbrush while the participants real hand, out of their vision,
is stroked simultaneously (synchronous condition). In many
healthy individuals, this produces a striking illusory sensation
of displacement of the felt location of ones own hand, sensation of touch through the rubber hand, and embodiment (or
ownership) of the rubber hand which refers to the sensation
208
that the rubber hand belongs to ones body (that the rubber
hand is ones own hand; e.g., Heed et al., 2011; Longo et al.,
2008; Newport, Pearce, & Preston, 2010). Embodiment of the
rubber hand arises from a combination of processes, including
the correlation and reconciliation of conflicting temporal and
spatial information from the visual and tactile systems, and
from visual capture, or increased weighting of visual over
tactile and proprioceptive stimuli (Holmes, Snijders, &
Spence, 2006; Pavani, Spence, & Driver, 2000; Rossetti,
Desmurget, & Prablanc, 1995; Spence, Pavani, & Driver,
2000; Tipper et al., 1998). The RHI thus provides valuable
insight into how multiple sensory modalities are integrated to
contribute to self-perception.
The illusion can be abolished or diminished by introducing
a temporal delay between brush strokes on each hand (asynchronous condition; Botvinick & Cohen, 1998). Given that
embodiment of a body part requires synchronous input between sensory modalities, such as touch and vision, introduction of such a delay between brush strokes often is sufficient to
prevent the occurrence, or reduce the strength, of the illusion.
This asynchronous condition often is used as a comparison
condition to the synchronous condition (Botvinick & Cohen,
1998; Shimada, Fukuda, & Hiraki, 2009).
Qualitative changes in perception on the RHI are typically
rated using a questionnaire administered at the end of each
condition. In addition, quantitative shifts in perceived hand
location can be measured with an intermanual pointing task,
or by a verbal response. This measure, termed proprioceptive
drift, is the change in the perceived location of the participants hand after stroking. It provides insight into internal
adjustments to body perception that occur as a result of the
integration of visual and proprioceptive information (Tsakiris
& Haggard, 2005; Tsakiris, Prabhu, & Haggard, 2006).
In view of the recently emerging evidence that selfperception (as assessed using the RHI and other methodological tools) is a multicomponent process that comprises several
semi-dissociable elements (Coslett, Saffran, & Schwoebel,
2002; de Vignemont, 2010; Kammers et al., 2010), Longo
and colleagues in 2008 devised a 27-item questionnaire to
assess qualitative changes in the RHI to overcome the limited
range of domains assessed by the original 9-item questionnaire (Botvinick & Cohen, 1998). A principal components
analysis (PCA) was conducted to describe the underlying
structure of the questionnaire as completed by 131 university
students (75 females). Four major components were identified
in each of the synchronous and asynchronous conditions:
Embodiment of the Rubber Hand, Loss of Own Hand
(describing a reduction in perception of and in the participants
actual hand), Movement (sensations of the rubber hand and
the participants hand moving closer together), and Affect
(sensations of pleasantness and enjoyment of the illusion). A
fifth component, Deafference (pins and needles or numbness in the participants hand), was identified in the
asynchronous condition only. It was found that the first component, Embodiment, was further divisible into three subcomponents: Ownership (the rubber hand belonging to the participant), Location (feelings of the participants hand being in the
location of the rubber hand), and Agency (a sense of control
over the rubber hand). Further work has demonstrated that
these components are dissociable and may be seen as partially
independent (Kalckert & Ehrsson, 2012).
A common contemporary variant of the RHI includes a
projection of the participants own hand onto a videoscreen
embedded into a table, instead of the use of a rubber hand.
This digital version also allows the introduction of a delay in
image feedback. Similar to the RHI, the Projected Hand
Illusion (PHI) creates the illusory sensation of dissociation
of ones hand from ones body (Ijsselsteijn, de Kort, & Haans,
2006; Kammers et al., 2009a, b; Longo & Haggard, 2009;
Shimada et al., 2009; Shimada, Qi, & Hiraki, 2010; Tsakiris,
Longo, & Haggard, 2010; Tsakiris et al., 2006). The PHI has
the advantages of allowing precise timing of synchronous and
asynchronous stroking, allowing a more convincing illusion.
Despite a number of studies using the PHI (Ijsselsteijn, de
Kort, & Haans, 2006; Kammers et al., 2009a, b; Longo &
Haggard, 2009; Shimada et al., 2009; Shimada et al., 2010;
Tsakiris et al., 2006, 2010), the component structure of this
digitised version has not been investigated. Therefore, the first
objective of our study was to establish the component structure of the PHI in a community sample using Longo et al.s
(2008) questionnaire, subject to minor modifications in the
wording (such that rubber hand was replaced by image on
the screen).
Given the lack of research on the association between
individual characteristics of a general community sample
and proneness to the illusion (RHI or PHI), the secondary
goal of the study was to identify sources of variation that
contribute to individual differences in the experience of the
illusion. This is important for establishing the properties of
control samples to be used when examining the illusion in
clinical populations.
The first demographic variable of interest was age. One
study failed to find significant effect of age in young adults
(age range 1724; Dummer et al., 2009). Other studies using
the RHI found that children have more proprioceptive drift
towards the rubber hand than do young adults but show little
difference in ownership (Bremner et al., 2013; Cowie, Makin,
& Bremner, 2013). Similarly, Tajadura-Jimnez et al. (2012),
using a face-illusion where participants viewed an image of
another face being stroked simultaneously as their own, found
that age (range 1738 years) was associated with a decrease in
endorsement of the other face (analogous to embodiment
ratings). In other words, increasing age was linked to a decrease in proneness to the face-illusion. This finding was
explained as normal developmental changes in selfperception in younger adults to accommodate changes in
209
Materials
Demographics
A questionnaire assessing age, sex, handedness, years of
education, and self-reported medical problems was administered to all participants.
Projected hand illusion
Participants sat in front of a custom-made apparatus with their
hands resting on top of a table. Their right hand was hidden
from view behind a removable curtain. In front of them was a
Fujistu 17 colour monitor embedded horizontally in the table,
which displayed a live image of their right hand. The image
was captured by an analogue 1/3 Colour Charged-coupled
Device Camera (AVC-561, AVTECH, Taiwan) and transmitted to the monitor via an analogue DelayLine (DL1B-5379,
Ovation Systems Ltd., UK). The total delay in image transmission was <10 ms, much less than the 150 ms that is
required to detect visuomotor delays (Blakemore, Wolpert,
& Frith, 1999) and less than the 85-100 ms of video delay
reported by other studies (Ijsselsteijn et al., 2006; Longo &
Haggard, 2009; Shimada et al., 2009, 2010; Tsakiris et al.,
2006). The image of the hand was displaced 15 cm to the left
of the participants right hand. A tactile marker indicated
where the participants right middle finger should remain for
the duration of the experiment. A cloth was placed over the
shoulder and upper arm of the participant such that their arm
was obscured during each block.
Methods
Participants
210
Procedure
Demographic information, forward and backward digit spans,
and the PLE questionnaire ratings were all collected before the
illusion procedure. For the illusion, a paintbrush was used to
stroke the participants right finger and hand at a rate of 12
brushstrokes/second, out of view, for 3 minutes. Participants
were instructed to focus on the image of the hand on the
monitor and on the sensation of the paintbrush on their hand.
There were two conditions; in the synchronous condition, the
video input to the monitor was instantaneous (apart from the
<10 ms intrinsic delay of the system), and in the asynchronous
condition, a delay of 500 ms was imposed on the video input.
The order of presentation was counter-balanced across participants; participants were assigned a presentation order in a
pseudorandom manner.
At the end of each condition, a questionnaire assessing the
subjective experience of the illusion was administered. The
items were rated on a 7-point Likert scale ranging from 3
(strongly disagree), 0 (neutral), to +3 (strongly agree). The 20items were obtained from Longo et al. (2008), including items
relating to the components termed Embodiment of the Rubber
Hand, Loss of Own Hand, Movement, and Deafference. The
items in the Affect component were not included, because
these questions do not pertain to experiences elicited by the
illusion. In the questionnaire, references to the rubber hand
were replaced with the image of my hand. Item 12 was
changed such that it read I couldnt have moved my hand if I
had wanted. Proprioceptive drift also was obtained by asking
participants to indicate the felt location of their middle
finger. This was achieved by the participant moving a ruler
with their left hand, with eyes closed, to where they felt their
right middle finger to be. The distance from the middle finger
was recorded (positive values indicated a location closer to the
image of the hand). Two proprioceptive judgements were
made; one before and one after the stroking period. Proprioceptive drift was calculated in cm by subtracting the
prestroking judgement from the poststroking judgement.
Results
The items means were very similar to those reported by Longo
et al. (2008). The only major differences were that items 18,
19, and 20 were rated slightly lower in the current study.
211
Table 1 Component loadings, eigenvalues, and variance from PCA performed on questionnaire data. Questions with no component loadings had values
<0.32 (<10 % overlap in variance) and were not interpreted as part of the component
It seemed like
1
2
3
4
0.40
0.35
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
0.40
0.34
0.18
0.13
0.11
0.15
0.38
0.48
0.51
- 0.49
- 0.42
0.39
0.47
0.35
0.50
0.48
6.65
33.3
3.82
19.1
- 0.46
- 0.58
- 0.50
1.72
8.60
1.21
6.07
0.16
0.11
0.09
0.12
0.29
0.29
0.29
0.23
0.16
0.24
0.14
0.26
0.23
0.22
0.34
0.25
Total: 67.1 %
212
0.003) where the component scores for females were consistently higher than for males (Fig. 2). To investigate this
further, sex and component were included in a further
ANOVA as independent variables. This ANOVA again found
a main effect of sex (F(1, 46) = 7.63, p = 0.008, 2G = 0.04) as
well as an interaction between sex and component (F(3, 330)
= 3.35, p = 0.02, 2G = 0.03). Post-hoc pairwise t-tests
indicated higher component scores on Embodiment in females
compared with males (p = 0.01), but these were not significant
for Disembodiment (p = 0.51), Deafference (p = 0.80), or
Agency (p = 0.51). From these results, the component scores
were not increased uniformly across all components between
males and females.
There was a significant main effect of age (F(1, 44) = 6.40,
p = 0.04, slope = 0.02), whereby increasing age was associated with a decrease in component scores of all components.
Similarly, there was a main effect of PLE score (F(1, 44) =
9.66, p = 0.003, slope = 0.13), showing that an increase in
PLE score was associated with an increase in component
scores of all components (Fig. 3). There was no significant
association of years of education or DS in this ANCOVA
(both p > 0.05).
To investigate what was driving the interaction in the
previous ANCOVA, separate ANCOVAs were conducted
for each component.
Embodiment: there was a significant main effect of delay
condition on embodiment scores (F (1, 47) = 6.94, p = 0.01,
2G = 0.03) with embodiment scores higher in the synchronous than the asynchronous condition. Furthermore, there
was a significant effect of sex (F (1, 45) = 6.16, p = 0.02)
with the embodiment scores higher for females than for
males (Fig. 2). There was a nonsignificant effect of PLE
score (F (1, 45) = 3.54, p = 0.07, slope = 0.25) and
variables, such as age, forwards or backwards digit span,
and years of education, all failed to reach significance (all p
> 0.1). Our results showed that females rated Embodiment
Discussion
The first objective of our study was to describe the component
structure of the PHI in a community sample, as assessed using
213
214
Deafference
The third component extracted from our questionnaire was
Deafference; it consisted of three questions relating to a reduction in sensation from the participants own hand (items
215
expectation that the video image finger will move when one
moves ones own finger, which does not exist initially with an
external object such as a rubber hand. These expectations
likely lead to more initial agency with an image of ones
own hand than with a rubber hand or an image of anothers
hand.
Together, these findings demonstrate that in situations
where there is no explicit movement, and hence little opportunity for agency over actions to occur, the expectation of
control is highly correlated with embodiment. However, as
our current findings demonstrate, this relationship does not
necessarily hold true when actions are possible and have been
observed and/or when the object is an image of ones own
hand.
Interestingly, the experience of agency over the image
loaded onto the same component as agency over the participants hand. This is consistent with participants having an
initial expectation that a real-time video image of their own
hand will move as they move; there is initial recognition that
image is associated with their own hand. This differs from the
RHI, where one might expect that feelings of agency over the
other hand may be linked to a decrease in sense of agency
over the real hand, similar to how embodiment of the other
hand is accompanied by disembodiment of the own hand
(Albrecht et al., 2011; Ehrsson, 2009; Longo et al., 2008).
This separation of Agency and Embodiment is consistent
with the findings of Kalckert & Ehrsson (2012), who found a
double-dissociation of agency and embodiment in a version of
the hand illusion that allowed for willed movements of a
finger of the rubber hand; incongruent postures of the rubber
hand disrupted embodiment but not agency, and passive (as
oppose to active) movements of the rubber hand disrupted
sensations of agency but not embodiment. Functional imaging
evidence also supports a distinction in the neural pathways
responsible for embodiment (Gentile, Petkova, & Ehrsson,
2011) and agency (Leube et al., 2003; Spengler, von Cramon,
& Brass, 2009). Our findings therefore provide further psychometric evidence for this distinction.
Age and the illusion
Our study found evidence that the perception of the hand
illusion decreases with age. Specifically, increasing age was
associated with a decrease in ratings of the illusion on the four
component scores. One possibility is that increasing age may
be linked to a bias towards rating items lower. Evidence
against this suggest is provided by Tajadura-Jimnez et al.
(2012), who found effects of age only on embodiment ratings,
and not across all items, as would be expected if a bias was at
play. These authors suggested instead that people in late
adolescence and early adulthood are more likely to display
flexible self-perception, compared with later age-periods, because there is greater change in appearance and so a greater
216
Cognitive functioning
The current study found no association of attention or working
memory, as assessed with the Digit Span forward and backward respectively, with the four components (Embodiment,
Disembodiment, Deafference, Agency, or proprioceptive
drift), extending previous work (Holmes et al., 2006). It is
possible however that lower level processes required for sensation and perception, such as visual selective attention may
be involved.
Psychotic-like experiences
The current study showed that higher PLE scores were linked
to an increase in questionnaire ratings. These results extend
previous findings on measures of schizotypy (Asai et al.,
2011; Germine et al., 2013; Thakkar et al., 2011). Germine
et al. (2013) reported that positive, but not negative,
schizotypy was associated with increased ratings of embodiment, but only in the synchronous condition. They concluded
that this indicated an actual increase in the experience of the
illusion in those with greater levels of schizotypy. By contrast,
both the current study and Thakkar et al. (2011), found an
association of PLE with higher ratings across both synchronous and asynchronous conditions.
Some evidence suggests that increased PLE confers an
increased susceptibility to hand illusions. First, Germine
et al. (2013) found that increased PLE scores were associated
with embodiment of the other hand but not with other
measures of the illusion, such as proprioceptive drift or reduced afference. Second, Albrecht et al. (2011) found that
healthy volunteers administered the indirect dopamine agonist
dexamphetamine reported stronger illusions (particularly, on
Embodiment and Loss of Own Hand) but did not rate other
measures, such as deafference, higher. There is a wellestablished association of increased dopamine transmission
with schizophrenia-spectrum disorders, including increased
dopamine binding in neural pathways associated with the
development of psychosis in people with PLE (Gray, Pickering, & Gray, 1994; Soliman et al., 2007). The manipulation of
a neurotransmitter pathway strongly implicated in
schizophrenia-spectrum disorders resulting in changes in
self-perception consistent with those seen in people with
PLE is supportive evidence that PLE are associated with a
stronger illusion.
People with PLE are thought by some (Claridge, 1997;
Claridge & Beech, 1995) to lie on a continuum of psychosis
from the absence of psychotic-like experiences, to subclinical
manifestations of PLE, to clinically relevant psychosis. It is
already known that people with schizophrenia are more susceptible to the illusion (Peled et al., 2000, 2003); the current
results demonstrate that people with subclinical PLEs also
report a stronger illusion. Typically, symptoms of
217
Conclusions
Psychometric analysis of a hand illusion in a community
sample demonstrates that self-perception during the illusion
is divisible into multiple components that are common to both
synchronous and asynchronous conditions. These components are Embodiment of the Other Hand, Disembodiment
of Own Hand, Deafference, and Agency. Of particular
218
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