Vogeley,

 K.  and  Gallagher,  S.  2011.  The  self  in  the  brain.  In  S.  Gallagher  (ed.),  The  Oxford  Handbook  of  the  
Self  (111-­‐36).  Oxford:  Oxford  University  Press.    
 

Self in the brain1

Kai Vogeley and Shaun Gallagher

1. Introduction
In 1977 the philosopher Karl Popper and the Nobel Prize-winning neuroscientist John
Eccles published The Self and its Brain, one of the first contributions in modern
neurophilosophy. In that book they defended a dualism that viewed the self as an
autonomous entity that interacted with, and in fact controlled brain processes. This
view, which Eccles (1989, 1994) further defended and developed, was not at all
representative of either the philosophical or neuroscientific communities of the time,
and it was, in terms of its general, non-reductionist philosophical position, comparable
to Descartes’s famous doctrine of the pineal gland as the site of interaction between
mind and brain. According to Popper, “the action of the mind on the brain may consist
in allowing certain fluctuations to lead to the firing of neurones” (p. 541). Eccles,
famous for his work on synaptic mechanisms, proposed that the probabilistic operations
of synaptic connections could be the place of interaction. “The self-conscious mind acts
upon … neural centres, modifying the dynamic spatio-temporal patterns of the neural
events” (Popper & Eccles, 1977, p. 495).
The work by Popper and Eccles motivated a debate in the journal Neuroscience
between the philosopher Mario Bunge, who defended an emergentist view, and the
neuroscientist Donald McKay, who staked out a position between materialism and
dualism (Bunge 1977, 1979; McKay 1978, 1979, 1980). McKay argued that we should
“start from our immediate experience of what it is like to be a person” (MacKay, 1978,
p. 601). He proposed that since each change in our experience corresponds to a change
in brain activity, we have both first-person data and third-person data about the same
entity, the conscious self, but that one set of data was not reducible to the other set,
despite the high, possibly even perfect, degree of correlation. It is, he suggested, like

                                                                                                               
1   Shaun Gallagher’s work on this chapter was, in part, supported by the National Science
Foundation under Grant No. 0639037, and the European Science Foundation’s Eurocore
program: Consciousness in a Natural and Cultural Context, BASIC project. Any opinions,
findings, and conclusions or recommendations expressed in this chapter are those of the
authors and do not necessarily reflect the views of NSF or ESF.
   112  

the relation between a computer and a mathematical equation being solved by the
operations of the computer. Without being able to identify the precise part of the
computer in which an equation interacts with the mechanism, we can nonetheless say
that the nature of the equation determines what the computer will do, while the laws of
physics that govern the processes of the computer chip will determine the solution to
the equation.
It is important to notice that in these discussions the concept of self was not clearly
defined, and this is one thing that leads these theorists into the murky metaphysical
corners of dualism and materialism. If, since 1977 cognitive neuroscience has
significantly advanced our understanding of certain brain functions, during the same
time philosophy has further clarified and complicated the notion of the self, as one can
see from the chapters in this present volume. If one were to oversimplify things and to
define self as having three clearly distinct aspects, for instance A, B, and C, or to define
n clearly distinct conceptions of self (1, 2, … n), then it is an important principle for
work in the neuroscience of self to say precisely which aspect or conception of self is at
stake when we go looking in the brain for “the self.” Our intention in this chapter is not
to provide an exhaustive review of recent work on the neuroscience of self (given the
amount of recent research in this area, this is likely an impossible task), but in part to
show why this principle is important and that it potentially allows the operationalisation
of key features of the self (Vogeley et al. 1999; Gallagher 2000).

2. Conceptual issues: The self in the brain: everywhere

and nowhere
Joseph LeDoux (2002, 31) points out that theories of the self ‘are not usually framed in
ways that are compatible with our understanding of brain function’.2 Indeed, it is
frequently the case that the neuroscience of self stays on a very general level by
including in the concept of self a variety of different self-related aspects. For example,
autobiographical knowledge, personal beliefs, self-conceptions, and the recognition of
our own face have been grouped together as related to left hemisphere activity (Turk et
al., 2003; see also Kircher et al., 2000). Alternatively, Platek et al. (2003, p. 147) claim
that there “is growing evidence that processing of self-related information (e.g.,
autobiographical memory, self face identification, theory of mind) is related to activity
in the right frontal cortex” (see also Devinksy, 2000; Miller et al., 2001).
Representations of both the physical and phenomenological aspects of self, and “self-
representation in general” apparently involves the right lateral parietal cortex (Lou et
al., 2004, p. 6831), while the “self model . . . a theoretical construct comprising
essential features such as feelings of continuity and unity, experience of agency, and
body-centered perspective” (Fossati et al., 2003, p. 1943) involves activation of the
                                                                                                               
2
Northoff et al. (2006, 453) put it this way: “Neither the historical nor modern psychological
approaches are obviously isomorphic with any known brain analysis.”  
   113  

medial prefrontal cortex in both hemispheres.

Functional neuroimaging and neuropsychological studies have recently revealed the
importance of cortical midline structures (CMS) for processing information related to
the self. In particular, activation of the ventromedial prefrontal cortex (VMPFC) has
been repeatedly observed when subjects think about themselves (Northoff and
Bermpohl, 2004; Northoff et al., 2006) and specifically when they make judgements
regarding their own personality traits (D’Argembeau et al., 2007; Gutchess et al., 2007;
Heatherton et al., 2006; Johnson et al., 2002; Kelley et al., 2002; Ruby et al., 2009). In
addition, lesion studies suggest that damage to the VMPFC leads to deficits in self-
awareness (Stuss et al., 2001a,b).
Other studies that explore the cerebral areas associated with self-referential processing
in normal aging observe that younger and older adults engaged the VMPFC to a similar
extent when judging personality traits in reference to the self versus another person
(Gutchess et al. 2007). The only cerebral differences between the groups in self versus
other personality assessment were found in somatosensory and motor-related areas.3
On the one hand, on the basis of such studies, some theorists posit a unitary system
responsible for all self-related phenomena. For example Northoff has argued for a self-
system in the CMS (Northoff et al. 2006; Northoff & Bermpohl 2004). We note,
however, that this putative self-system corresponding to CMS covers a quite large set
of brain regions (including ventro- and dorsomedial prefrontal, anterior and posterior
cingulate, retrosplenial and medial parietal cortices) and involves, in addition, the
multitude of connections between CMS and subcortical areas, and the possible role of
subcortical areas in a sense of embodied self (see Northoff and Panksepp 2008). On the
other hand, the widespread nature of CMS seems to suggest that there is no specialized
brain area responsible for generating the self, just as there is no candidate region or
system at hand that could be made fully responsible for the constitution of
consciousness. In a number of recent reviews, a variety of conceptual and
methodological issues have been highlighted that qualify any claims for the notion of
an overall, consistent self-system in the brain.4

                                                                                                               
3  Rubyet al. (2009) explored the neural substrate of self-personality assessment from different
perspectives. Young and older adults evaluated personality traits in reference to the self
versus a close friend or relative. Moreover, they were asked to make those judgements from
either their own (first-person) perspective or the perspective of their friend or relative (third-
person perspective).  
4
Complications in the interpretation of the data are quite common in this literature. For
example, Northoff et al. (2006) cite the study by Kelley et al. (2002) as showing that the
medial prefrontal cortex is involved in processing self-related stimuli. Gillihan and Farah
(2005), in contrast, citing that study along with others, claim that processing of the traits of
others also activated this area and that may signify that it is involved in “person processing”
rather than specifically self-processing. The question is further complicated, as noted by
Kelley et al. and Northoff et al., by the fact that the MPFC normally deactivates (relative to
   114  

Gillihan and Farah (2005) demonstrate that even when studies focus on a specific self-
related representation, such as self-face recognition (judged in contrast with another
person’s face, or morphed faces), different methodologies and different subject groups
will identify different areas of the brain for this function:
• right hemisphere (Platek & Gallup, 2002, in neurologically intact subjects;
Preilowski 1979; Keenan et al. 2003, in studies of split-brain patients; Keenan et
al. 1999; 2000; Keenan et al. 2001, using TMS in normal subjects and the “Wada
test” in patients undergoing epilepsy surgery)
• left hemisphere (Turk et al. 2003, in studies of split-brain patients)
• left anterior insula, putamen, and pulvinar, the right anterior cingulate cortex, and
globus pallidus, in subjects asked to identify a picture of their own face from a
series of photos (Sugiura et al. 2000, using PET);
• left fusiform gyrus, anterior cingulate cortex, right supramarginal gyrus, superior
parietal lobule, and precuneus (Sugiura et al. 2000, using PET); right middle,
superior, and inferior frontal gyri (Platek et al. 2004, using fMRI); and right
insula, hippocampal formation, and lenticular and subthalamic nuclei, the left
prefrontal cortex (inferior and middle frontal gyri), right middle temporal gyrus,
left cerebellum, as well as parietal lobe and lingual gyrus (Kircher et al. 2000,
using fMRI), in subjects asked to judge orientation in a set of photos that include
their own.
Using a similar diversity of methods, studies of self-agency show the involvement of a
further variety of brain areas. For example, when the sense of self-agency fails, as in
cases of schizophrenic delusions of control, studies show hyperactivation of the right
parietal cortex (Spence et al. 1997). If non-pathological subjects are asked to imagine
themselves acting, however, we find activation in left inferior parietal, posterior insula,
post-central gyrus, and inferior occipital gyrus, bilaterally (Ruby and Decety 2001).
When subjects are asked to distinguish actions that are self-generated (accompanied by
a sense of self-agency) from actions generated by others, activation is seen in the right
inferior parietal area for our actions (Chaminade and Decety 2002) as well as in the
anterior insula bilaterally (Farrer and Frith 2002).
In their review Gillihan and Farah (2005) show that studies of self-trait descriptions
(part of what they refer to as the psychological component of self) yielded no clear
results for specialized brain areas because of various confounds. For example, although
in a number of studies the medial prefrontal cortex was activated for self-judgments,
the same area was also activated for other-judgments. “It is therefore possible that
                                                                                                                                                                                                                                                                                                                                                         
baseline) in contexts of task-dependent activities and external stimulation. The MPFC is part
of what Raichle et al. (2001) call the “default mode” of brain function. Kelley et al. (2002)
suggest that the normal baseline activation of the MPFC might be interpreted as a region
responsible for a default self-referentiality when the person is not engaged in some task or
attending to some stimulus. See below for further discussion.  
   115  

medial PFC plays a role in person processing in general…. [or that] it is possible that
the activation of these areas is a function of amount and type of knowledge rather than
self versus other knowledge per se” (p. 89). Other confounds prevent any clear results
from studies of autobiographical memory.
On various definitions of first-person perspective, experiments show activation of
different areas of the brain. (1) Often in psychology and cognitive neuroscience first-
person perspective simply means that the object of my attention is myself rather than
someone else (e.g., Ruby and Decety, 2001). For example, first-person perspective may
be operationalized in terms of a narrative perspective as what I can say when I occupy
the role of self-narrator. While narrating about self or another person activates the right
prefrontal cortex, narrating about self but not the other activates the right
temporoparietal junction and bilateral anterior cingulate cortex (Vogeley et al. 2001).
(2) In philosophy, first-person perspective is frequently thought of as the view from the
place of the observer (whether in introspection or in perception). Accordingly, if I
describe the world from the perspective of an observer in the egocentric spatial
framework, then I am taking a first-person perspective; likewise if I give a
phenomenological report on my own experience. In this sense, processing in early
sensory areas may be considered correlated to first-person experience (Gillihan and
Farah 2005, p. 92; Vogeley and Fink 2003). When subjects are asked to report on their
own emotional reactions to stimuli, in contrast to objective spatial properties of the
stimuli, significantly more activation can be found in the anterior cingulate cortex
(Lane et al. 1997) or medial prefrontal areas (Gusnard et al. 2001).5
It thus looks like the entire cortex is specialized for self-referential processing, or, in
other words, as the review by Gillihan and Farah (2005) demonstrates, there is no
specialized or common area responsible for self-related representations.6 Even theorists
like Northoff who set out to identify a circumscribed self-system end up pointing to an
extensive set of brain areas that come close to the diversity summarized by Gillihan and
Farah (see Figure 1). Thus, LeDoux seems correct to maintain that “different
components of the self reflect the operation of different brain systems, which can be but
are not always in sync” (2002, 31). Perhaps a more critically pessimistic view on the
general inconsistency of results is summarized by Craik et al. (1999, p. 30): “every
significant activation in the [self condition] was also found in either the [other person
condition] or the [general semantic] condition, or both” (cited by Gillihan and Farah
2005, p. 94). One issue that needs to be resolved in this regard is whether we should
                                                                                                               
5
As Gillihan and Farah (2005) point out, although both the Lane and Gusnard studies involve
first-person reports on affective experience, the third-person conditions involve no affective
component, so the contrast results of brain activation may be emotion-related rather than
precisely for first-person perspective.  
6   “The different ways in which the self–nonself distinction is confounded with other

distinctions across studies are likely to account for the different patterns of activation in
different studies … even when the same aspect of the self is under study” (Gillihan and
Farah 2005, 94).    
   116  

think of the self exclusively in terms of self-referential processing. It could be argued

that the self is not exhausted in just such processing, since even as I respond to stimuli
that are not self-referential, my response is itself always self-specific. For example, my
response is made on the basis of information that is already framed from the
perspective in which I perceive the stimulus – a first-person (egocentric) perspective.
In this case, it is not the self as an object of reference (e.g., I report on recognizing my
own photo, or judging whether I think or feel thus and thus) but the self-as-subject (as
the one doing the reporting in ways that may not be self-referential) that is at stake.
Legrand and Ruby (2009) have followed up on just this idea in a clarifying way.

A
B

u = face
X = agency
O = traits
p = memory
+ = first-person perspective

Figure 1. (A) Self-related activation in a “glass brain” view (all areas of activation are
visible regardless of whether they fall on the midline). From Gillihan and Farah (2005).
(B) Activation in CMS observed in imaging studies during self-related tasks in
different domains (from Northoff et al. 2006)
   117  

In a meta-analysis of neuroimaging studies, Legrand and Ruby (2009) detected a

certain pattern, the so-called “E-network” (“evaluation network”), within the variety of
differential self-related activations. They point out that four brain areas are repeatedly
activated in self versus non-self contrasts: medial prefrontal cortex, precuneus/posterior
cingulate gyrus, temporoparietal junction, and temporal pole. Rather than conclude that
this network is a self-network, however, they raise another question. They note that
since the areas of this network are activated in tasks that involve a contrast between self
and non-self, they are not really self-specific, i.e., specialized only for self. Numerous
studies in fact show that the areas of the E-network are also the areas activated in
“theory of mind” (ToM) tasks which require the modeling of the internal experiences or
mental states of others (see below). Thus, “the main brain regions recruited for others’
mind representation are also and precisely the main brain regions reported in self
studies and … this overlap extends beyond the brain areas usually pointed out, that is, it
comprises the medial prefrontal cortex, the precuneus/posterior cingulate, the
temporoparietal junction, and the temporal pole” (254-58). Legrand and Ruby go on to
postulate and to cite multiple studies that show that the lowest common denominator
involved in all of the tasks designed for self-referential or other-referential activation is
inferential processing using memory recall, which they term ‘evaluation’. Thus, the
brain areas in the E-network are activated for certain cognitive processes that apply not
just to self, but to other persons, and even to objects. They are not specialized for self-
referential processing. For example, fMRI studies by Fangmeier et al.(2006) and
Fonlupt (2003) show very clearly that the medial prefrontal cortex is activated in
logical (deductive reasoning) processes that involve premise integration where the
premises are not about self or other, but about an array of single letters, billiard-ball
causality, or color.7
It might seem at this point that we are making only little progress in identifying any
consistent brain-related activity responsible for self. First, rather than finding a
common, well-circumscribed brain area of self-referential activation, we found a large
number of diverse areas activated for a variety of self-related experience. Now,
however, on the Legrand-Ruby interpretation, the diverse areas implicated in self-
referential experience are in fact not areas of activation exclusively for self. At first it
seemed the self was almost everywhere in the brain; now it seems to be nowhere.

                                                                                                               
7
Activation of the medial prefrontal cortex (MPFC) for evaluative inference would be
consistent with data cited by both Northoff et al. (2006) and Gillihan and Farah (2005)
without deciding the issue about self-reference. Legrand and Ruby point to a different
interpretation of the default mode activation of the MPFC. Rather than self-specific activity,
the MPFC may be busy inferring or predicting future events. Bar (2007) makes this proposal,
which “posits that rudimentary information is extracted rapidly from the input to derive
analogies linking that input with representation in memory. The linked stored representations
then activate the associations that are relevant in the specific context, which provides focused
predictions” (p. 280).
 
   118  

3. Key aspects of self

3.1 Self-agency and self-ownership

This perplexity about the brain’s role in self-specific processes may stem from the lack
of any clear correspondence between theoretical conceptions of self and how cognitive
neuroscience operationalizes such conceptions. To get a grasp on the difficulties
involved in this, let’s take a look at one area in which a seemingly clear distinction with
regard to self-experience helps to structure experimental design. Gillihan and Farah
(2005) suggest that one of the most clear and specialized instances of self-specified
brain functions have to do with body ownership and agency for action. They suggest
that asomatognosia (the lack of recognition of one’s body, e.g., following stroke)
“demonstrates both the functional independence and the neural localization of limb
ownership; our sense of a limb as being our own is distinct from our sense that other
people’s limbs belong to them. [In addition] studies of the experience of agency in limb
movements indicate a similar functional independence and neural localization” (93).
One important and widely-cited experimental brain-imaging study cited by Gillihan and
Farah, as well as by Norhoff, and Legrand and Ruby, is Farrer and Frith (2002). The
study by Farrer and Frith, like a number of other studies (e.g., Chaminade and Decety
2002; Farrer et al. 2003; Tsakiris and Haggard 2005), refers to a phenomenological
distinction between the sense of ownership and the sense of agency (Gallagher 2000).
The distinction is made as part of a delineation of what a number of theorists call the
minimal or core self. The minimal self is constituted by a pre-reflective consciousness
of oneself as an immediate, embodied subject of experience, and is distinguished from
the narrative (extended) self, the “more or less coherent self (or self-image) that is
constituted with a past and a future in the various stories that we and others tell about
ourselves” (Gallagher 2000, 16). The minimal self involves a number of aspects,
including the sense of ownership (SoO) -- the sense that I am the one who is undergoing
an experience, for example, the sense that my body is moving, regardless of whether
the movement is voluntary or involuntary -- and the sense of agency (SoA) -- the sense
that I am the one who is causing or generating an action. These are pre-reflective,
implicit, or tacit aspects of our experience. That is, one’s SoA, for example, is not
something generated in a reflective or conceptual thinking about what I am doing.
Rather, it is built into the doing, the action, itself (Marcel 2003; Gallagher and Marcel
1999).
Recent conceptual developments distinguish between different levels in the
phenomenology of agency comprising an implicit or first-order level of an experiential
feeling of agency as opposed to an explicit higher-level of judgment or attribution of
agency (Gallagher 2007; Synofzik et al., 2008). As such, SoO and SoA are
distinguished from a reflective level of intention formation or attribution that may be
involved in reporting about or evaluating our actions. Thus, Stephens and Graham
   119  

(2000) make a corresponding distinction on the reflective level between attributions of

subjectivity (ownership) and attributions of agency. Whereas SoO and SoA are first-
order (phenomenal) experiences related to my bodily movement and action, attributions
are reflective judgments that I make about my actions. If someone asks me whether I
bought a car yesterday, my affirmative response is an attribution of agency to myself.
As such, attributions of ownership (“this happened to me”) or agency (“I did it”)
involve the narrative self, although they may normally be reports on, or based on, my
actual pre-reflective experience of agency or ownership (Gallagher 2010).
It may be a helpful oversimplification to think of the SoA as generated in efferent
signals or motor control processes that occur just prior to or in the initial stages of the
action. The SoO, may be tied to sensory feedback, its attenuation or cross-modal
integration (see e.g., Tsakiris and Haggard 2005; Tsakiris, this volume). SoA is
missing in the case of involuntary movement where there is no efferent motor
command, but the SoO is still present (“My body moved”). One can also think of
pathologies that may involve a loss of the SoA (as in schizophrenic delusions of control
or anarchic hand) or SoO (as in asomatognosia following stroke, or alien hand
syndrome).
The fMRI study by Farrer and Frith (2002) sets out to identify the neural correlates of
the SoA, which they distinguish from the SoO.
Subjects manipulated a joystick [to drive a colored circle moving on a screen to specific
locations on the screen]. Sometimes the subject caused this movement and sometimes
the experimenter. This paradigm allowed us to study the sense of agency without any
confounding from the sense of ownership. To achieve this, subjects were requested to
execute an action during all the different experimental conditions. By doing so the
effect related to the sense of ownership (I am performing an action) would be present in
all conditions and would be canceled in the various contrasts (p. 597).
One might wonder why the SoA varies across tasks in this experiment, since in each
case the subject is required to move the joystick. If one conceives of the SoA as a pre-
reflective experience that is built into action, perhaps tied to efferent motor commands,
then one would expect that the subject’s action of moving the joystick would be
accompanied by SoA for that action. Farrer and Frith, however, although citing the
distinction between SoA and SoO as defined above, operationalize the SoA according
to a different definition. In the experiment the SoA is not tied to motor control
processes, but to what is accomplished by the action. Does my action have an effect in
the world, or, in this case, on the computer screen? If so, then I have a sense of agency
for that action. We will refer to this as a SoA tied to action consequence.
One should, in fact, welcome this as a useful clarification of the SoA, although it is not
presented as such in the study. That is, the suggestion that the SoA may involve more
than just motor control processes, and that it may include aspects tied to intentional
fulfillment of the action (action consequence) is important, especially for fully
embodied and embedded versions of cognitive experience (see Gallagher 2008; 2010).
   120  

Farrer and Frith, however, fail to note their shift of definition or the complexity
involved in the SoA.
But things are more complicated than this. In some trials the subject is informed ahead
of each task whether the movement will be his or not his, and he is asked to perform the
task regardless of this. In the cases when the action represented on the computer screen
is not the subject’s action, and because the subject knows it, one might suspect that his
SoA short-circuits in the control of the joystick. That is, if he does not have SoA tied to
the intentional aspect of the task, his SoA may be redirected specifically to his bodily
movement in a way that would be consistent with the original definition of SoA. In the
case where the subject is not the agent for the task represented on the computer screen,
the right inferior parietal cortex (IPC) is activated. Consistent with other studies
(Spence et al., 1997; Ruby and Decety, 2001), Farrer and Frith interpret this to mean
that the IPC activation correlates to a sense of other-agency, rather than to the subject’s
own SoA for control of the joystick. When the subject does know that he is causing the
action on the screen, his anterior insula is activated bilaterally. Thus, the experimenters
identify activation in the anterior insula as the correlate of SoA.
If we set aside some of these complexities and accept the operational definition of SoA
in terms of the experiment, another problem appears when Farrer and Frith turn to
theoretical considerations about why the anterior insula should be involved in
generating SoA. In effect, they revert from the definition of SoA in terms of action
consequences to the original definition in terms of motor control.
Why should the parietal lobe have a special role in attributing actions to others while
the anterior insula is concerned with attributing actions to the self? The sense of agency
(i.e., being aware of causing an action) occurs in the context of a body moving in time
and space. Damasio (1999) has suggested that the sense of agency critically depends
upon the experience of such a body. There is evidence that both the inferior parietal
lobe and the anterior insula are representations of the body.... the anterior insula, in
interaction with limbic structures, is also involved in the representation of body
schema.... One aspect of the experience of agency that we feel when we move our
bodies through space is the close correspondence between many different sensory
signals. In particular there will be a correspondence between three kinds of signal:
somatosensory signals directly consequent upon our movements, visual and auditory
signals that may result indirectly from our movements, and last, the corollary discharge
associated with motor commands that generated the movements. A close
correspondence between all these signals helps to give us a sense of agency. (601-602)
A number of things can be said about this explanation. (1) Note the introduction of the
issue of attribution, i.e., judging rather than experiencing agency. Indeed, although
starting with the distinction between the pre-reflective, experiential SoO and SoA, the
authors state: “The main purpose of this experiment was to compare brain activity
associated with attributing an action to oneself and attributing an action to another
person” (597). Indeed, as part of the protocol, the subjects, who were informed prior to
   121  

the task that they would be controlling the icon or someone else would be, were asked,
in the former case, “to drive the circle, to be aware that they drove the circle, and thus
to mentally attribute the visualized action to themselves” (597). As a result we could
rightly ask whether the results reflected a pre-reflective SoA tied to motor control, a
pre-reflective SoA tied to action consequence, or a reflective attribution of agency. (2)
Farrer and Frith, while asking why the insula might be responsible for attribution of
self-agency go on to decide this issue on theoretical grounds that are more related to
SoA understood in terms of motor control – that is, it seems tied to a body-schematic,
pre-reflective experience of the body and sensory feedback integration with “corollary
discharge associated with motor commands”. One interpretation that might make sense
out of this is to say that attribution of agency is based on (is a report on) the pre-
reflective SoA tied to motor control, which is associated with increased neural
activation of the insula. Yet the experiment was set up to ignore the SoA as tied to
motor control, and to focus on the SoA as tied to action consequence. (3) The
mention of the integration of “somatosensory signals directly consequent upon our
movements, [and] visual and auditory signals that may result indirectly from our
movements” may suggest some involvement of the SoO if the latter is based on such
integration. This raises a further question about how Farrer and Frith explain the
attribution of agency to the other person and activation of the IPC. They cite well-
known evidence that suggests the IPC may be responsible for SoO --"Lesions in the
right parietal cortex have been associated with disturbances in the feeling of belonging
of the patient’s limbs…. patients with right parietal lesion do not recognize their limbs
as their own and perceive them as belonging to others" (p. 601). So one question that
remains unanswered is whether IPC can be involved in both the SoO for one’s body,
and the sense of other-agency.
We can learn some valuable lessons about research on the self from this kind of study.
First, experimental brain-imaging can help to clarify the meaning of concepts like the
sense of agency. Theoretical models may suggest that SoA is closely tied to motor
control; but when operationalizing SoA in the context of an experiment, we can start to
see that it may be a much more complex concept. Second, if we are to obtain any clear
results from such experiments, we definitely have to make careful distinctions between
reflective and pre-reflective aspects of the self.

3.2 Self in action

The predominant account on explaining the SoA of our own actions has been the
“central monitoring theory” or “comparator model” as a theory of motor learning and
motor control (Frith, 1992; von Holst & Mittelstaedt, 1950; Wolpert et al., 1995). Two
types of internal models are implemented in the central motor system: so-called inverse
and forward models. The sense of agency particularly hinges on the forward model
which uses an efference copy, that is, a copy of a motor command, to predict its sensory
consequences. We are usually unaware of the results of this comparison as long as the
   122  

desired state is successfully achieved (Blakemore & Frith, 2003). As outlined above a
good opportunity to study this phenomenon is the capacity to monitor one’s own
movement to false visual feedback without being aware of the adjustment (David et al.,
2007; 2008; Farrer & Frith, 2002; Fourneret & Jeannerod, 1998). In pathological states
in which patients experience so-called delusions of control by attributing their own
actions to external sources (Daprati et al., 1997; Franck et al., 2001; Haggard et al.,
2003) the central monitoring mechanisms might be disturbed (Blakemore et al. 2002) or
errors might arise from a disturbed judgment or attribution of agency (e.g., Fourneret et
al., 2001). The posterior parietal cortex (PPC) represents a very likely candidate for
providing reference to the agent of an action as this region seems to monitor the
concordance between self-produced actions and their visual consequences, being
especially involved in the detection of visual-motor incongruence (Chaminade &
Decety, 2002; Farrer et al., 2003; Farrer & Frith, 2002; Fink et al., 1999). Similarly, the
cerebellum has been implicated in signaling discrepancies between predicted and actual
sensory consequences of movements (e.g., an associated tone; Blakemore et al., 2001).
Recently, a region in visual association cortex called the extrastriate body area (EBA)
has been reported to show greater activity during self-generated movements (Peelen  
and   Downing   2005). This finding opened the possibility that the EBA may also be
involved in the SoA, as suggested by Jeannerod (2004). Indeed, activity in the EBA
could be shown to be differentially modulated by a manipulation of agency responding
to the perception of visuo-motor incongruence. Furthermore, the EBA showed a similar
response pattern as the PPC as well as an increased functional connectivity to the PPC
underlining a close functional relationship between them (David et al., 2007).
Another account was presented by Haggard and colleagues with the phenomenon of so-
called “intentional binding” showing that voluntary—but not involuntary or passive—
movements and movement consequences are temporally bound together in conscious
awareness. Subjects judged the perceived time of voluntary movements as occurring
later and the sensory consequences as earlier than it was actually the case (Haggard et
al., 2002; Haggard & Clark, 2003; Tsakiris & Haggard, 2003). The phenomenon of
intentional binding appears to be related to increased activation of the SMA or pre-
SMA and insula. Recruitment of these regions has specifically been associated with
awareness and execution of self-generated actions, action preparation and the subject’s
own intention-to-act (Cunnington et al., 2006; Farrer et al., 2003a; Farrer & Frith, 2002;
Haggard & Clark, 2003; Haggard & Whitford, 2004; Lau, Rogers & Passingham,
2006).
In summary, findings from cognitive neuroscience suggest that essentially the brain
regions of ventral premotor cortex (vPMC), the supplementary motor area (SMA and
pre-SMA), the cerebellum and the dorsolateral prefrontal cortex (DLPFC), the posterior
parietal cortex (PPC), the posterior segment of the superior temporal sulcus (pSTS),
and the insula are associated with experiences of agency (Blakemore et al., 2001; Farrer
et al., 2003; Farrer & Frith, 2002; Fink et al., 1999; Jeannerod, 2004). This list of brain
regions appears to subserve a network of sensorimotor transformations and motor
   123  

control, whereas a second component represents heteromodal association cortices

associated with a variety of cognitive functions (e.g. prefrontal cortex [Fuster 1997;
2001]). One could speculate that motor system-related regions may subserve
“executive” functions whereas heteromodal associative regions subserve “supervisory”
functions (David et al. 2008). At the current stage, a multifactorial and multilevel
model appears to provide the most helpful and comprehensible framework for
integrating divergent theories and findings (Synofzik et al., 2008; Wegner & Sparrow,
2004).

3.3 Self in space

In the context of spatial cognition the self refers to the experience of the centeredness of
the subjective multidimensional and multimodal experiential space upon one´s own
body. During spatial cognition, as “a means of representing the locations of entities in
space” (Klatzky 1998), we operate in an egocentric reference frame, constituted by
subject-to-object relations. Egocentric reference frames can be differentiated with
respect to the midline of the visual field, the head, the trunk, or the longitudinal axis of
the limb involved in the execution of a certain action (Behrmann 1999). In contrast, an
allocentric reference frame, sometimes also referred to as “exocentric” or “geocentric”,
is constituted by object-to-object relations (best described in a Cartesian coordinate
system). It refers to a framework that is independent of the agent´s position (Klatzky
1998; Aguirre & D´Esposito 1999; Vogeley & Fink 2003).
Cognitive operations when perceiving a visual scene from one´s own perspective (1PP)
differ from taking a view of the same scene from another person´s viewpoint (3PP),
although both tasks are centered on the body of the respective agent, the self or the
other. To clearly separate these two levels of descriptions, the perspective-related terms
1PP and 3PP indicate the phenomenal level, whereas the terms egocentric and
allocentric reference frames refer to the cognitive or neural level as conceptualized by
the onlooking (scientific) observer. The crucial difference between 1PP and 3PP is that
3PP necessitates a translocation of the egocentric viewpoint.
A number of studies have focused on the issue of perspective taking in space. Taking
1PP appears to rely at least in part on temporo-parietal processing as assessed in
navigational tasks (Maguire et al. 1998, 1999; Zacks 2008). For example, Maguire et al.
(1998) demonstrated that a right inferior parietal region was activated whenever
egocentric calculations (i.e. computing body turns toward the target) were necessary in
addition to the processing of allocentric spatial information (mediated via the
hippocampus). Subtracting a static condition from “ego-movement” conditions
including trail-following or way-finding also involved bilateral medial parietal cortex
(Maguire et al. 1998). These findings have been corroborated by other studies (Maguire
et al. 1999).
   124  

A simple spatial cognitive task to be solved from 1PP and 3PP was employed in a 3D-
visuospatial task in which a virtual scene consisting of a virtual character surrounded by
red objects was presented. Subjects were asked to assess the number of red balls as seen
from either their own (1PP) or the virtual character´s perspective (3PP). Both
conditions were based on egocentric operations, as the objects have to be located in
relation to an agent in both conditions, either the test person or the virtual character. In
case of 3PP, however, additional use of allocentric operations is necessary to generate
egocentric coordinates for the agent. Neural correlates revealed differentially increased
neural activity during 1PP (as opposed to 3PP) in the left medial prefrontal cortex and
the posterior cingulate cortex bilaterally. In contrast, 3PP was associated with
differentially increased neural activity in the region of the superior parietal lobe
bilaterally, predominantly on the right side, and right premotor cortex (Vogeley et al.,
2004). The relevance of the right parietal cortex for spatial cognition can also be
inferred from lesion studies exploring patients with right parietal lesions leading to
extinction or spatial neglect (Behrmann 1999; Marshall & Fink 2001) and from mental
rotation tasks (Vogeley & Fink 2003; Zacks 2008, Wolbers & Hegarty 2010).
It has been hypothesized that 1PP generates a spatial model of one’s own body in the
brain, upon which the experiential space is centered (Berlucchi & Aglioti 1997). This
conjecture is in good accordance with reports on increased neural activity of right
inferior parietal cortex involving visuo-spatial attention not only in navigation tasks
(Maguire et al., 1998) but also the assessment of the subjective mid-sagittal plane
(Vallar et al., 1999, Galati et al. 2001). Another important source of information of
bodily states is obviously the reference to a gravitational vertical as upright orientation
as primary reference. Andersen et al. (1999) reviewed evidence for the fact that
vestibular information is used by the posterior parietal cortex for the perception of self-
motion. A highly relevant study in this respect was performed by Ruby and Decety
(2001) who studied perspective taking in a motor imagery task. Subjects were asked to
imagine that either themselves (1PP) or the experimenter (3PP) manipulates an object.
During 1PP simulation of action, only regions in the left hemisphere were activated,
including the inferior parietal lobe, precentral gyrus, superior frontal gyrus, occipito-
temporal junction and anterior insula. During 3PP simulation of action, the right
hemisphere was activated, namely the inferior parietal cortex, precuneus, posterior
cingulate and frontopolar cortex.

3.4 Social self

Studies on the neural mechanisms of self-related processes and social cognitive
processes have become a key topic in cognitive neuroscience over the past decade.
More recently, social (cognitive) neuroscience has emerged as a new subdiscipline in
neuroscience, and has recently developed into an autonomous scientific discipline
(Cacioppo et al. 2004). Social neuroscience focuses on processes that are related to the
adequate ascription of mental states to others for the purpose of successful
   125  

communication or interaction between personal agents. A key constituent is the

capacity to distinguish between one’s own and others’ mental states, and this seems to
involve some form of self-consciousness (Newen & Vogeley 2000, 2009).
Closely related to the ability to attribute and maintain a self-perspective is the capacity
to discriminate between self and other in intersubjective interaction. The ability to
attribute mental states (beliefs, desires, intentions) to others, is often referred to as
"mindreading" (e.g., Baron-Cohen, 1995). This ability has been studied extensively in
so-called "theory of mind" (ToM) paradigms (e.g. Fletcher et al., 1995; Premack and
Woodruff 1978), in which mental states or propositional attitudes of an agent are
inferred or simulated. The concept of ToM is used to refer to both the attribution of
mental states to oneself and to others.
In a typical ToM paradigm, a subject has to model the knowledge, attitudes or beliefs of
another person. ToM capacities may be related to the ability to assign and maintain a
self-perspective. In classical ToM paradigms (e.g. Fletcher et al. 1995), in which mental
states or propositional attitudes of an agent need to be modeled (e.g. "Person A knows,
believes, etc., that p" with p being a physical event), self-perspective refers to the
special situation, in which I am the agent myself (e.g. "I know, believe, etc., that p"),
and to the subjective experiential multi-dimensional space centered around one’s own
person. These studies have repeatedly demonstrated increased neural activity associated
with ToM conditions in the anterior cingulate cortex. Vogeley et al. (2001) were able to
replicate these findings and to demonstrate additional differential brain activation when
the test persons themselves were involved as an agent in a particular story. The capacity
for taking 1PP in such ToM contexts showed differential activation in the right
temporo-parietal junction and the medial aspects of the superior parietal lobe, i.e. the
precuneus (Vogeley et al. 2001). Neural activations common to 1PP and 3PP in ToM
contexts were observed, however, in the anterior cingulate cortex. Similar activations
patterns were also found in two other studies involving self-referential processing.
Anterior cingulate activations were found during judgments about trait adjectives that
were related to oneself as opposed to others (Kelley et al. 2002) and during a study in
which volunteers were asked to think intensely on how they would describe the
personality traits and physical appearance of themselves as opposed to others (Kjaer et
al. 2002).
Ruby & Decety (2003) studied perspective taking by asking subjects to respond to a list
of health-related questions, either from one´s own or someone else´s perspective.
During 1PP the postcentral gyrus was activated, whereas 3PP relevant activations
comprised the anterior medial prefrontal cortex, the left superior temporal sulcus and
temporal pole and the right inferior parietal lobe. The results of this particular study are
somewhat different from the studies reported above as the right inferior parietal lobule
is activated during 3PP but not during 1PP. This could be integrated in the hypothesis
that the right temporo-parietal region is crucial for the successful differentiation
between 1PP and 3PP.
   126  

However, the fact, that differential brain loci in different brain lobes are activated in
association with the attribution of 1PP relative to "mind-reading" (reviewed in
Gallagher & Frith 2003), of or ascription of trait adjectives to others, suggests that these
components constitute distinct psychological processes.

3.5. Note on the default mode of brain function

Empirical evidence for the recruitment of medial cortical activation sites during
experiences of self-reference is provided by the concept of a so-called “default mode of
the brain” put forward recently (Gusnard et al. 2001; Raichle et al. 2001). According to
this hypothesis, resting states, stimulus-independent thoughts and the like, which are
experienced as a “state of self” correlate with “the default mode of the brain”
characterized by certain cortical activation patterns, predominantly in the anterior and
posterior cingulate and medial parietal cortices. If a cognitive activity requires a higher
demand, neural activation is “shifted” towards the recruited neuronal network; medial
frontal and parietal regions in turn then tend to decrease their activity (Raichle et al.
2001). According to the speculative interpretation of Gusnard et al. (2001), this is not
merely a noisy signal, but might reflect a “continuous simulation of behavior” or “an
inner rehearsal as well as an optimization of cognitive and behavioral serial programs
for the individual´s future”, in short: a state of the “multifaceted self” (p. 4263). What
appears as “state of self” on the phenomenal level, appears as “default brain state” on
the neuronal level. Similarly, Andreasen et al. (1995) described a posterior cingulate
deactivation during situations in which subjects were not engaged in a focused
cognitive task, attributing this deactivation to spontaneous, probably purely associative
mentation processes. This ongoing purely associative mentation would then be
suspended when the subject becomes engaged in an experimental task requiring
specific cognitive activities. In the same sense, Burgess et al. (2001) argue that the
precuneus supports the inspection of internal images.
The association of the default mode of the brain with self-related mentation processes,
however, is controversial. The resting state is not well described in cognitive terms.
Furthermore, the areas involved in the default network overlap with the E-network
(Legrand and Ruby 2009), and the absence of external stimulation tells us nothing
about what kind of cognitive operations may be taking place. They may be self-related
or non-self-related, ruminating or forward looking, reflective or day-dreamy.

4. Levels of processing: Reflective and prereflective

aspects of self
To understand individual behavior requires a conceptualization of the self in the context
of perceiving and acting in its environment. This relation of the subject itself with
surrounding objects and pragmatic contexts, was conceptualized as the “core self” by
   127  

Damasio (1999), and postulated to be a transient relation, in need of re-instantiation

from moment to moment, a process which in turn constantly refers to the so-called
“proto self”, which remains unconscious and represents bodily states. Medial cortical
regions are hypothetically recruited if such a state of “core self” is instantiated, a
prediction in accord with medial cortical activation sites, which comprise anterior
medial prefrontal, medial parietal and posterior cingulate cortex (Damasio 1999).
Gillihan and Farah 2005, reflecting broad trends in the literature, take a
“commonsensical approach to the meaning of self” and include three broad categories
of self-related information under this heading. (1) Physical self, which focuses on
either specific body parts, including one’s own face, or the body as a whole; (2)
psychological self, which includes conceptual knowledge about oneself,
autobiographical memory, semantic memory (facts about oneself), first-person
perspective, and self-other differentiation; (3) agency, which combines “elements of the
physical and the psychological, in that agency concerns the role of the psychological
self in causing the actions of the physical self” (p. 77). We can see that these
distinctions cut across others made between minimal and narrative aspects of self, as
well as pre-reflective and reflective. Getting clear on this latter distinction, however,
suggests a different way of thinking about the relation between brain and self.
It is possible to challenge what seems to be the primacy of explicit self-referential
processing in the neuroscientific experimental studies of self. Self-referential
processing involves the kind of evaluation that is involved in most experimental tasks
in studies of the self – “e.g., one refers to ‘this face’ as one’s own by evaluating the
resemblance of this face to one’s representation of one’s face” (Legrand and Ruby
2009, p. 271). Such tasks, however, are clearly reflective tasks that require subjects to
reflectively evaluate information about themselves. In such tasks, the self operates as
an intentional object that comes under attentive consideration. My concern, or my task,
is about, or is directed to my self. In many, if not most of one’s everyday experiences,
however, this is not the case. My attention is most often directed at the environment, at
others, at accomplishing some task that does not call for any explicit self-consciousness
or attention to the self. If we think of it in terms of reflective regard, then we can say
that the connection between the self-as-subject, i.e., the one who is doing the reflecting
(or who is engaged in the task of reflecting), and the self-as-object, i.e., the self that is
reflected on or referred to (the self as intentional object), or, more generally, whatever I
learn or can report about myself, is neither exclusive nor non-contingent, and is
therefore not self-specific, in the sense of the term defined by Legrand and Ruby.8 The
one aspect that they define as self-specific is the embodied first-person perspective,
which constitutes being the self-as-subject. That is, a first-person perspective is
                                                                                                               
8
“We define the notion of self-specificity according to the two criteria of exclusivity and
noncontingency. … a given self S is constituted by a self-specific component C only if C
characterizes S exclusively (i.e., C does not characterize non-S) and noncontingently (i.e.,
changing or losing C would amount to changing or losing the distinction between S and non-
S)” (Legrand and Ruby 2009, 272).
   128  

something only selves have; and absent a first-person perspective there is no self. First-
person perspective is basic for the self/non-self distinction, is the point of orientation
for every perception and action, and is to be taken in a constitutive sense.9
The first-person perspective is relational or intentional insofar as having a first-person
perspective on anything relates that thing (some object in the world, another person,
even the self-as-object) to the experiencing subject. Legrand and Ruby (2009) thus
suggest that this concept can be cashed out at a basic neurophysiological level, namely,
the level of sensorimotor integrative processes involving efference and reafference.
The fact that one can find activation in sensorimotor areas not only when one is
perceiving and acting, but also in tasks related to language, emotion, and
intersubjectivity strengthens their suggestion. One challenge is to discover any activity
that involves first-person perspective that does not activate such areas.
What, however, does the first-person perspective amount to? One possibility is that it
is an ecological phenomenon, in the Gibsonian sense. Neisser’s (1988) conception of
the ecological aspect of self suggests that any perception or action engaged in by the
subject always provides some information about the subject, as well as about the
perceived or acted upon object or environment. In terms of perception and action, at
least, this embodied, ecological, pre-reflective self-awareness is more basic (and
developmentally more primary) than any form of self-recognition (see Rochat, this
volume, for the developmental evidence; and Gallup, Anderson,   and   Platek, this
volume, for the neural correlates of self-recognition).

5. Conclusion
To achieve clarity in neuroscientific studies of self it is incumbent on researchers to
define the precise aspect of self under study. Selves are experiential, ecological, and
agentive; they are often engaged in reflective evaluations and judgments; they are
capable of various forms of self-recognition, self-related cognition, self-narrative, and
self-specific perception and movement. In many of these activities, selves are more
“in-the-world” than “in-the-brain,” and they are in-the-world as-subject more so than
as-object. In this regard we endorse Dennett’s warning: “it  is  a  category  mistake  to  
start   looking   around   for   [selves]   in   the   brain”(1992,   p.109).     The project rather
should be the study of what happens in the brain when the self-as-subject is engaged in
                                                                                                               
9
We offer the phrase ‘in a constitutive sense’ to avoid a certain ambiguity in Legrand and
Ruby’s account. One may have the impression that when they discuss the self/non-self
distinction they may mean the usual relative distinction between one’s self and everything
else. In that respect, another person would count as part of the non-self. But in a constitutive
sense the distinction between self/non-self means that there are certain aspects that define
what would count as a self – e.g., first-person perspective – and fail to define everything that
is not a self. In that case another person counts as a self (rather than non-self) because they
also have a first-person perspective.
   129  

the world, in specific actions and in specific social contexts. This requires not only the
sophisticated tools of the neuroimaging lab, and the brilliant experiments of
neuropsychology, but also the subtle conceptual tools of philosophy.

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