This study aimed to identify neuronal markers of bodily self-consciousness using the rubber hand illusion and EEG recordings. In the first part, participants underwent the rubber hand illusion while their brain activity was recorded using EEG during rest with eyes open and closed. Correlational analyses found that greater alpha power in central electrodes correlated with stronger feelings of embodiment and loss of control over one's real hand as reported on questionnaires following the illusion, but only in the eyes open condition. The findings suggest that ongoing brain activity, particularly alpha oscillations, may be a neuronal marker related to experiences of bodily self-consciousness induced by the rubber hand illusion.

1. 1
Identifying Neuronal Markers to Determine Bodily Self-
Consciousness using the Rubber Hand Illusion and EEG
By: Jasrina Kaushal
Supervised off-campus by: Dr. Georg Northoff
Supervised on-campus by: Dr. Iain McKinnel
Thesis submitted in partial fulfillment of the requirements for the degree of Bachelor of
Science with Honours in Integrated Science
Carleton University
Ottawa, Ontario, Canada
April 2015
© 2015 Jasrina Kaushal

2. 2
IDENTIFYING NEURONAL MARKERS TO DETERMINE BODILY SELF-
CONSCIOUSNESS USING THE RUBBER HAND ILLUSION AND EEG
Jasrina Kaushal*1
, Marcello Costantini 2
, and Georg Northoff 2
1
Department of Integrated Science, Carleton University, Ottawa, Ontario Canada K1S 5B6
2
Mind, Brain Imaging, and Neuroethics Unit, Royal Ottawa Mental Health Centre, Ottawa,
Canada K1Z 7K4
Abstract
The ability to experience one's own body is referred to as bodily self-consciousness. The rubber
hand illusion (RHI), whereby a test subject develops a feeling of ownership of a fake hand, can
be used to study embodiment, one component of bodily self-consciousness. The purpose of this
study is to identify neuronal markers that determine bodily self-consciousness in healthy
individuals. In the first session, participants watched a rubber hand being stroked while their
hidden hand was stroked simultaneously. An introspective self-report was completed by
participants after the RHI to rate their agreement or disagreement with 27 statements
corresponding to their subjective experience of the RHI. The second session consisted of
recording participants’ brain activity during rest using an EEG. Correlational analyses revealed a
linear relation between alpha power, recorded from central electrodes, and embodiment of the
rubber hand. These findings suggest that bodily self-consciousness can ultimately be traced to
the intrinsic activity of the brain and that the Rubber Hand Illusion provides a strong foundation
for studying bodily self-consciousness.

3. 3
Table of Contents
Introduction......................................................................................................................................3
Methods............................................................................................................................................9
Results............................................................................................................................................14
Discussion......................................................................................................................................18
References......................................................................................................................................23

4. 4
Introduction
In everyday life we do not doubt that our body belongs to us. When we shake hands with
someone, we know which hand belongs to whom. However, this is not always the case for
certain individuals. There are indeed, clinical conditions such as schizophrenia, and eating
disorders in which bodily self-consciousness is altered. For instance, an individual suffering from
anorexia may perceive that their body is slightly larger than it actually is, displaying an altered
sense of bodily self-consciousness. Thus, finding neural markers for bodily self-consciousness
may be clinically relevant.
Consciousness involves the ability to experience or to feel, subjectivity, sentience,
wakefulness, having a sense of the self, and the executive control system of the brain (Farthing,
1992). Simply put, we are conscious when we are awake, aware of the self and the world around
us (Laureys & Tononi, 2009). Consciousness involves complex interconnected and organized
neural mechanisms diffused in several regions of the brain. Although recent neuroscientific and
psychological discoveries have led to the possible basis of consciousness, research remains
unfinished.
The “I” of conscious experience can be explained by self-consciousness. Being aware of
oneself, or of one’s own thoughts and actions are the defining characteristics for self-
consciousness (Costanini, 2014). For one to have an understanding of who they are as an
individual is crucial in determining one's identity. Recent approaches to study self-consciousness
have been to target mechanisms in the brain that are responsible for processing bodily signals
(such as, bodily self-consciousness). Bodily self-consciousness is the representation and
processing of bodily information, and the overall experience of owning a body (Costantini, 2014

5. 5
and Blanke, 2012). However, it has been difficult to test for the processes underlying bodily self-
consciousness because the body is always present and we are unable to dissociate the body from
the mind.
Through interoception and exteroception, the brain constantly receives information from
the body (Costantini, 2014). The interoceptive system is associated with autonomic motor
control and receives signals from bodily organs such as the heart, stomach, lungs, etc.
Interoceptive signals represent the perception of the body from the inside and include regulation
of homeostasis as well as feelings such as emotions, drives etc. (Suzuki et al, 2013). Whereas the
exteroceptive system represents perception of the body from the outside, guides somatic motor
activity and receives information from the external environment through the five senses: vision,
olfaction, touch, gustatory, and auditory (Costantini, 2014). Through elaboration of the
interoceptive system representations and their incorporation with the exteroceptive system
signals, a sense of self emerges (Suzuki et al, 2013). However, this interaction of exteroceptive
and interoceptive systems in determining the experience of bodily self-consciousness remains
poorly understood today (Suzuki et al, 2013). Bodily self-consciousness is especially difficult to
study because of the constant information flow between the two systems, the distinction between
the physical body, and the experienced subjective body becomes nearly impossible (Costantini,
2014). Thus, neuroscientists have recently began to employ the use of bodily illusions in order to
examine the neural mechanisms underlying bodily self-consciousness. The objective of these
illusions are for participant's to feel a sense of ownership over a synthetic body-part. They have
been used to explore the complex relationships that exist between the brain's representation of
the body and the physical body itself (Moseley, Gallace & Spence, 2012).

6. 6
An illusion is described as the brain's failure to produce a subjective experience which
corresponds to reality. Thus, by understanding how the brain fails in this function, gives us a
better understanding on how the basic function is performed as well as the perceptual processes
responsible for the function. Through the use of bodily illusions, it also becomes possible to
influence perceived integration of an external object (such as a synthetic body part) into the
representation of the physical body itself (Longo, 2008). The Rubber Hand Illusion (RHI) is an
example of a bodily illusion that provides a powerful experimental tool to illustrate an
individual's bodily self (Costantini, 2012). In the RHI, participants watch a rubber hand being
stroked while their hidden hand is stroked simultaneously (known as the induction period). This
sensation creates a sense of ownership over the rubber hand, and participants perceive the rubber
hand as actually being part of their own body (Longo, 2008). Although the RHI can be robust,
approximately 30% of participant's do not experience it (Zhou et al, 2014). After the induction
period, most participants also perceive the location of their hand to be closer to the rubber hand
than it actually is. If the rubber hand is stroked asynchronously with regards to the participant's
own hand, then this illusion does not occur (Costantini, 2012). The RHI provides an excellent
approach for studying and manipulating embodiment, and has been so used in several recent
studies (Longo, 2008).
Information from different sensory modalities such as sound, smell, touch, sight and taste
are incorporated together in a phenomenon known as multisensory integration (Stein, Stanford &
Rowland, 2009). This representation of combined modalities allows for meaningful perceptual
experiences and is critical for adaptive behaviours, ultimately increasing the chances of survival
(Costantini, 2008). It has been suggested that bodily self-consciousness is mediated by
multisensory integration (Ehrsson, Holmes & Passingham, 2005). Under normal circumstances,

7. 7
bodily self-consciousness appears to be stable, however there is growing data indicating that this
feature is dependent on dynamic multisensory integration of self-related signals and is capable of
change (Suzuki et al, 2013). This is demonstrated in the RHI, as the attribution of the seen rubber
hand to the self is dependent on the integration between the somatic and visual signals from the
hand (Ehrsson, Holmes & Passingham, 2005). In a study conducted by Tsakiris & Haggard
(2005), after synchronous visuotactile stimulation, subjects perceived their hand to be located
significantly closer to the rubber hand compared to asynchronous visuotactile stimulation
(Tsakiris et al, 2007). This also suggests multisensory integration between the tactile experience
of the paintbrush on the participant’s own hand and visually perceived rubber hand occurs in
order for the RHI to elude bodily self-consciousness (Tsakiris et al, 2007).
Other studies demonstrating multisensory integration in humans are the McGurk Effect
(McGurk & MacDonald, 1976) and the cross-modal congruency effect (Spence, Pavani, &
Driver, 1998). The common characteristics within these studies are that they follow the spatial
rule and the temporal rule to exhibit multisensory integration. In the spatial rule, the constituent
unisensory stimuli arise from approximately the same location for a stronger effect of
multisensory integration (Costantini, 2014). While in the temporal rule, the two stimuli must be
exhibited at approximately the same time in order for multisensory integration to be enabled
(Costantini, 2014). This spatiotemporal limitation holds true in the RHI, as the paintbrushes must
be stroked at the same time and the distance between the real hand and rubber hand cannot be too
large in order for the illusion to occur. Because the RHI follows the same perceptual rules
necessary for multisensory integration, it is proposed that this mechanism is essential for bodily
self-consciousness to occur (Costantini, 2014).

8. 8
Another important aspect to consider is that oscillary activity is necessary for
multisensory integration and perceptual processing to occur. In the body, functioning cells
create chemical, mechanical, thermal, and faint electrical energy (Hulbert, 1947). Cortical cells
in the brain act in clusters, and their actions such as: perceiving, thinking, initiating voluntary
movements, etc. create faint electrical energy which can be measured using an
electroencephalogram (EEG) (Hulbert, 1947). In EEG experiments, "awake resting state" is one
of the most frequently used experimental condition because it defines a 'baseline' of brain
activity (Laufs et al, 2003). Thus, we can use awake rest-state to measure deviations from
baseline brain activity. In this study, we measured ongoing activity using an EEG during awake
rest state. Neural oscillations, specifically in the alpha wavelength, during baseline represent a
tool that can be used in multisensory integration and perceptual processing. During rest, EEG
oscillary activity has significant effects on perceptual processing, and in representing a measure
to study bodily self-consciousness.
In this study, we investigate neuronal markers that determine bodily self-consciousness
using the RHI, introspective self-reports given after induction period, and recording resting-state
EEG activity in two conditions: eyes closed (EO) and eyes open (EO). Drawing from previous
evidence we predicted that there will be a positive correlation between alpha power, recorded
from central electrodes in both eyes open (EO) and eyes closed (EC) and ownership of the rubber
hand. The EEG component is associated to neuronal markers that can be found when the brain is
at rest. There is variability in the overall sense and feeling of owning a body, and this variability
in bodily self-consciousness can be traced at rest.

9. 9
Methods
Participants
Thirty-three healthy undergraduate students were included in the present study.
Participants were recruited through online advertisement and were paid to partake in the
experiment. Three participants reported taking anti-depressant drugs during the time of study and
their results were excluded. All students were right handed as reported by the Edinborough
Handiness Questionnaire, 1975 (EHQ).
Materials
Participants were seated across from the experimenter, facing a table containing an open
ended box with two compartments (Figure 1). One compartment had a transparent cover, and the
other was enclosed with an opaque black cover. Subjects placed one hand in the transparent
compartment with a mirror, and their other hand in the opaque compartment. The life-sized
rubber hand was also visible in the transparent compartment. The rubber hand was aligned with
the participant's hand, approximately shoulder width apart. The participants wore a cape, which
was attached to the front of the box in order to keep their arms out of sight throughout the
experiment.

10. 10
Figure 1. Rubber Hand Illusion experimental set-up.
Procedure
The experiment consisted of two sessions. Prior to the experimental trial, one
measurement of proprioceptive drift was taken, in which a ruler was placed above the box and
participants were asked to verbally report the position of their perceived index finger in the
opaque compartment. The purpose of the proprioceptive drift measurement is to identify if a
change of perceived finger location towards the rubber hand correlates with the illusion. To
prevent participants from indicating the same number for each measurement, the ruler was offset
at a different length each trial.
Following the proprioceptive drift, the first session began. The experimenter stroked the
index finger of the rubber hand and the subject's unseen hand simultaneously with two identical

11. 11
paintbrushes. The participant was instructed to watch the rubber hand being stroked and
verbalize any sensations they might be feeling. The fingers were stroked for 3 minutes, at a rate
of approximately 1 stroke per second. Following this induction period, a second measure of
proprioceptive drift was recorded.
Subjects were instructed to remove their hand from the box and complete a standard
questionnaire. The subjects were instructed to rate their disagreement or agreement to 27
statements, using a 7-item Likert Scale corresponding to their subjective experience of the RHI.
Responses varied from -3 to +3 in which +3 indicated that the participant "strongly agreed", - 3
that they "strongly disagree" and 0 that they "neither agree nor disagree" with the statements.
The second part of the experiment consisted of two sessions of rest conditions: eyes open
(EO) and eyes closed (EC) measured using an EEG. An EEG cap with 32 channels was used to
study this resting state. Figure 2 illustrates ongoing activity located in the central electrodes (CZ,
CPZ, and PZ), which were used in the correlational analyses with the self-report questionnaires
given after the RHI. The participants were told to rest for approximately 5 minutes while an EEG
cap was placed on their scalp, connected to Neuroscan with 32 channels. In the EO condition,
participants were instructed to resist the urge to blink to the best of their abilities.

12. 12
Figure 2. An EEG cap with 32 Channels, the red region (CZ, CPZ, and PZ or channels 15, 20,
and 25) illustrates on-going activity.
Data Analysis
Pre-processing of information involved importing EEG data from Neuroscan to Matlab.
The files were condensed by down sampling to 500 kHz, removing EKG components
(electrocardiogram; electrical activity of the heart), low-pass 0.5 and high pass 30 filtering, and
individually removing artifacts by eye (this includes eye blinks, horizontal and vertical eye
movements, etc.). In the EO condition an extra step included rejecting components using ICA
(independent component analysis). The ICA function aids in separating additional artifacts
embedded in the data.

13. 13
Alpha power was computed using the Fast Fuorier Trasform (FFT). The FFT was used to
decompose complex signals from the EEG into frequencies of interest, namely 5 Hz, 10 Hz, and
15 Hz (theta, alpha, beta). Then, the power in each frequency was correlated with the answers
from the questionnaire. Correlational analysis between scores obtained from the questionnaire
with alpha power were calculated using SPSS Statistics.

14. 14
Results
Self-report RHI Questionnaire
Results are displayed in Figure 3 and 4. Questions used in the self-report RHI
questionnaire, with significant questions highlighted, are displayed in Table 1. Results in the
eyes open (EO) condition indicate that alpha power (10 Hz) in channel PZ correlate with
question 2 in the self-report measure, which states: 'it seemed like the rubber hand began to
resemble my real hand' (p= 0.047). Also in the EO condition, alpha power in channel CZ
correlates with question 16: 'it seemed like my hand was out of my control' (p= 0.024). In the EO
condition, alpha power in channel PZ correlates with question 7, 'it seemed like the rubber hand
was in the location where my hand was' (p= 0.039). However, this result was no longer
significant once data from the participant's who recorded taking anti-depressant drugs had been
removed from the study.
There were no significant results found in the eyes closed (EC) condition of the
experiment. There were also no significant results found in the theta (5 Hz) or beta (15 Hz)
powers in both EO and EC conditions. Furthermore, there was no significance found in the
remaining questions from the self-report. Demonstrating that ongoing activity in alpha
wavelength in CZ and PZ regions (Figure 1) during rest in EO condition predicts the strength of
the RHI.

15. 15
Figure 3. Channel CZ- Alpha Power Correlation with Question 16.
Figure 2. Channel 15 - Alpha Power Correlation with
Question 16
Alpha Power (Hz)
Responseto#16

16. 16
Figure 4. Channel PZ- Alpha Power Correlation with Question 2
Proprioceptive Drift
Measurements of proprioceptive drift were quantified as the difference between the
perceived index finger location before and after the RHI. Drift towards the rubber hand are
indicated by positive numbers. Results indicate no correlation between proprioceptive drift and
neuronal measures. Thus, proprioceptive drift is insignificant in regards to identifying neuronal
markers to determine bodily self-consciousness.
Figure 4. Channel 25 - Alpha Power Correlation with
Question 2
ResponsetoQuestion2
Alpha Power (Hz)

17. 17
Table 1. Statements and corresponding item number used in Self-Report RHI Questionnaire,
adopted from Longo, et al
Item Corresponding Statement
1 It seemed like I was looking directly at my own hand, rather than at the rubber hand
2 It seemed like the rubber hand began to resemble my real hand
3 It seemed like the rubber hand belonged to me
4 It seemed like the rubber hand was my hand
5 It seemed like the rubber hand was part of my body
6 It seemed like my hand was in the location where the rubber hand was
7 It seemed like the rubber hand was in the location where my hand was
8 It seemed like the touch I felt was caused by the paintbrush touching the rubber hand
9 It seemed like I could have moved the rubber hand if I had wanted
10 It seemed like I was in control of the rubber hand
11 It seemed like my own hand became rubbery
12 It seemed like I was unable to move my hand
13 It seemed like I could have moved my hand if I had wanted to
14 It seemed like I couldn't really tell where my hand was
15 It seemed like my hand had disappeared
16 It seemed like my hand was out of my control
17 It seemed like my hand was moving towards the rubber hand
18 It seemed like the rubber hand was moving towards my hand
19 It seemed like I had three hands
20 I found that experience enjoyable
21 I found that experience interesting
22 The touch of the paintbrush on my finger was pleasant
23 I had the sensation of pins and needles in my hand
24 I had the sensation that my hand was numb
25 It seemed like the experience of my hands was less vivid than normal
26 I found myself liking the rubber hand
27 It seemed like I was feeling the touch of paintbrush in the location where I saw the rubber
hand being touched

18. 18
Discussion
The present findings provide a systematic attempt to recognize neuronal markers that
determine bodily self-consciousness in healthy individuals. By combining an experimental
manipulation of the experience of one's own body through the use of a bodily illusion, and a
structured neurological approach to measure overall experience through recordings from a 32-
channel EEG cap, we were able to examine the experience of bodily self-consciousness. Our
findings suggest that psychometric methods can be a useful tool in determining the underlying
neuronal mechanisms responsible for the experience of bodily self-consciousness. In particular,
we demonstrated that ongoing activity in alpha wavelength in CZ and PZ regions during rest in
the EO condition predicts the strength of the RHI. The CZ region is located in the central zone,
or midpoint of the brain, and the PZ region is located in the upper tempoparitetal area of the
brain. If an individual displays increased alpha activity in those cortical brain regions during rest
with eyes open, they are more likely to experience the RHI, and subsequently have heightened
awareness of bodily self-consciousness.
Healthy participants viewed the experimenter stoke the rubber hand and their unseen
hand simultaneously. Phenomenology of the RHI was measured by self-report questionnaires
adapted from Longo et al. (Longo et al, 2008). Results indicated that participants experienced the
illusion and gained ownership of the rubber hand. These findings are consistent with previous
studies that suggest alpha power is associated with self-relatedness and inhibition of attention to
the external world, leading to increased limb ownership (Yeh et al, 2014). Increased alpha
powers are correlated to stronger self focus processing. Alpha power is increased when an
individual does not pay attention to their environment but pay attention to themselves.
Furthermore, alpha activity over central areas of the brain have been correlated to sensorimotor

19. 19
processing such as motion perception, overall movements, and motor imagery. This also
demonstrates why there were only significant results in the alpha power as opposed to the theta
(5 Hz) or beta (15 Hz) powers. Thus, alpha power frequency (which can be recorded from an
EEG) is a key feature in the process underlying bodily self-consciousness.
There were no significant results found in the EC condition, which is rather surprising as
most literature exhibits an association between greater alpha activity during awake-rest in EC.
The 'Berger Effect' states that there is either an increase or disappearance of alpha band
oscillations during eyes open (Yet et al, 2014). However, in the current study we found the exact
opposite, with an increase of alpha during EO. There are a few plausible reasons behind the
increased alpha activity during EO rather than EC condition. Alpha activity is blocked when an
individual is attentively processing mental operations, processing external information, or is
entering deep sleep (Laufs et al, 2013). Perhaps during the EC phase, the participants were in
deep thought or were in deep relaxation, entering the first stage of sleep. Because the participants
are undergraduate students, it is an assumption that they are busy with school work, possibly
stressed and/or tired. Simply put, it is possible that some of these students were ruminating,
ultimately producing a decrease in alpha activity during EC. Whereas in EO, they are more
awake and inhibiting the processing of external stimuli. Undergraduate students do not
accurately depict an entire population, and results may have been varied had a different sub-set
population were used this study.
Our data has broad clinical implications, as results from this study can be used to
compare to alpha activity among psychiatric disorders in which individuals have distorted
perceptions of their own body.

20. 20
There has been an association between disruptions in bodily self consciousness and
individuals suffering from disorders of body representation and illusory body perceptions
(Heydrich et al, 2010). Recent approaches for the clinical implications of bodily self-
consciousness can be found in reports of patients which had brain damage resulting in deficits in
processing bodily signals (Blanke, 2012). A well-known example was conducted by neurologist
Josef Gerstmann whom described two patients with damage to their right temporoparietal cortex,
which ultimately lead to a loss of ownership for their left arm and hand (Blanke, 2012). They
were diagnosed as somatoparaphrenia; a disorder in which patients have somatosensory and/or
motor deficits and deny ownership of the entire side of their body, or a limb (Valler & Ronchi,
2008). Other patients with somatoparaphrenia may display the opposite pattern and mis-attribute
their limb as belonging to someone else, or self-attribute someone else’s limb as their own
(Blanke, 2012). This suggests that the brain regions of the right temporoparietal cortex plays a
key role in the processing of bodily self-consciousness. It is also a region in which on-going
alpha activity is prominent, as displayed in our data.
Other research highlights the importance of multisensory bodily processing in patients
suffering from various forms of conditions such as strokes, tumors, migraines, and psychiatric
disorders such as anorexia, body dysmorphic disorder, and schizophrenia (Heydrich et al, 2010).
Symptoms associated with these disorders in which patients display alterations in perceptual
bodily self-consciousness include: body part displacement, experience of absence of body part,
misidentification of one’s own body part, disconnection of a body part from the body, and
phantom limbs (Heydrich et al, 2010). In regards to schizophrenia, research suggests that
patients display a weaker sense of body ownership over a rubber hand compared to healthy
individuals. More specifically, it has been found that ownership of the rubber hand correlated

21. 21
with the negative symptoms associated with schizophrenia (Ferri et al, 2013). The underlying
mechanisms responsible for the negative symptoms may have common features with
mechanisms responsible for processing the RHI (Ferri et al, 2013). As the RHI provides a strong
basis in determining what processes may be involved in these symptoms, much research must be
done until an exact basis can be found. However, it is apparent that disturbed body ownership
displayed in individuals suffering from schizophrenia may contribute to some of the psychotic
symptoms exhibited (Thakkar et al, 2011). Thus, it is vital that an individual is able to recognize
themselves, and process bodily self-consciousness otherwise they may suffer from the symptoms
indicated.
Transcranial Magnetic Stimulation (TMS) is a non-invasive tool used to stimulate nerve
cells through magnetic fields from a coil placed adjacent to the scalp, which carry short lasting
electrical currents to the brain (Siebner, 2008). TMS is most commonly used to treat individuals
suffering from major depressive disorder; in which electrical currents stimulate nerve cells
primarily in the frontal region which is associated with mood regulation. Particular clinical
applications to the current study are the possibility to investigate the therapeutic potential of
TMS as a unique treatment modality for individuals suffering from bodily perception disorders.
More specifically, through the use of Repetitive TMS (rTMS), which utilizes group pulses of
stimulation at specific frequencies to attain a constant train of activation power over a short
period of time (Jin et al, 2005). The current study examined alpha power in relation to exhibiting
higher processing of bodily self-consciousness. Patients suffering from various bodily perception
disorders such as schizophrenia, have been documented to have reduced alpha activity,
contributing to their symptoms and overall weakened sense of bodily self-consciousness (Jin et
al, 2005). RTMS can be used to modulate alpha powers, and it is plausible that as alpha power is

22. 22
altered, this can ultimately manipulate and create a heightened sense of bodily self-
consciousness. RTMS has been intensively studied and employed for the treatment of depressive
disorders, and unfortunately there is only a small number of reports for its clinical implications
on bodily perception disorders. For a future study it would be fascinating to examine if rTMS
could be used to modify alpha activity in patients with schizophrenia, and if this ultimately
increased their bodily self-consciousness by administrating the RHI before and after rTMS. It is
important for an individual to have a sense of bodily self-consciousness in order for them to
determine their own unique identity. If methods such as rTMS could increase alpha powers, and
ultimately enhance an individuals' bodily self-consciousness this could help improve the quality
of the life for that individual by alleviating their symptoms and aiding in establishing a sole
identity. While various parameters for the efficacy of this form of therapy for bodily perception
disorders most be fully investigated, hopefully the current study provides a step in the right
direction for further research examining bodily self-consciousness.

23. 23
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