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Keywords: self-reference; attention; meditation; fMRI; insula; prefrontal cortex; somatosensory; plasticity
Since William James’ early conceptualization, the ‘self ’ has cortical midline processes may be characterised as support-
been characterised as a source of permanence beneath the ing narrative self-reference that maintains continuity of
constantly shifting set of experiences that constitute identity across time (Gallagher, 2004).
conscious life. This permanence is often related to the Narrative self-reference stands in stark contrast to the
construction of narratives that weave together the threads immediate, agentic ‘I’ supporting the notion of momentary
of temporally disparate experiences into a cohesive fabric. experience as an expression of selfhood. Most examinations
To account for this continuity, William James posited an of self-reference ignore mechanisms of momentary con-
explanatory ‘me’ to make sense of the ‘I’ acting in the present sciousness, which may represent core aspects of self-
moment (James, 1890). Recently, progress has been made in experience achieved earlier in development (Damasio,
characterizing the neural bases of the processes supporting 1999; Zelazo and Frye, 1998; Gallagher, 2004) and may
William James’ ‘me’ in the form of ‘narrative’ self-reference have evolved in earlier animal species (Panksepp, 2005).
(Gallagher, 2004), highlighting the role of the medial Indeed, little is known about whether the neural substrates
prefrontal cortices (mPFC) in supporting self awareness by underlying momentary self-reference are one and the same,
linking subjective experiences across time (Neisser, 1997; or distinct from, cortical midline structures supporting
Northoff and Bermpohl, 2004). The mPFC has been shown narrative experience. One hypothesis suggests that awareness
to support an array of self-related capacities, including of momentary self-reference is neurally distinct from
memory for self-traits (Craik et al., 1999; Kelley et al., 2002; narrative self-reference and is derived from neural markers
Fossati et al., 2003; Macrae et al., 2004), traits of similar of transient body states, in particular, right lateralised
others (Mitchell et al., 2006), reflected self-knowledge exteroceptive somatic and interoceptive insular cortices
(Lieberman et al., 2004; Ochsner et al., 2005), and (Damasio, 1999; Craig, 2004; Critchley et al., 2004). In the
aspirations for the future (Johnson et al., 2006). As such, present study, we examined this thesis.
Received 26 April 2007; Accepted 23 June 2007
We investigated these hypothesised dual modes of self-
Advance Access publication 13 August 2007 reference by employing functional magnetic resonance
This research was funded by grants from the National Science and Engineering Research Council and the imaging (fMRI) during attention to two temporally distinct
Canadian Institute of Health Research.
Correspondence should be addressed to Adam K. Anderson, Department of Psychology, University of foci of attention: the self as experienced across time and in
Toronto, 100 St George Street, Toronto, ON M5S 3G3, Canada. E-mail: anderson@psych.utoronto.ca. the immediate moment. One obstacle to investigating
ß The Author (2007). Published by Oxford University Press. For Permissions, please email: journals.permissions@oxfordjournals.org
314 SCAN (2007) N. A. S. Farb et al.
present-centred awareness is the well-established tendency person or (ii) an EF mode, monitoring their moment-to-
for the mind to wander and become distracted from moment experience in response to the adjectives. Trait-
the present moment in favour of temporally distant, related adjectives were chosen for their ability to promote
stimulus-independent thought (SIT) (Smallwood and self-reflection as evidenced by previous research (e.g. Fossati
Schooler, 2006). SIT consists of automatically generated et al., 2003).
verbal or visual experiences (McGuire et al., 1996; Teasdale
et al., 1995) akin to William James’ notion of a running MATERIALS AND METHODS
‘stream of consciousness’ (James, 1890), the basis of the Participants
narrative form of self-reference described above. SIT-related Participants were recruited upon enrolment in the mind-
neural activation has been shown to reflect an automatic fulness-based stress reduction (MBSR) program at
tendency to engage in narrative processes in the absence of a St Joseph’s Hospital in Toronto, an 8 week intensive
body state, without purpose or goal, other than noticing how standard quadrate birdcage head coil. Functional
things are from one moment to the next; if a participant MRI (fMRI) was conducted using T2-weighted single-shot
noticed themselves getting distracted by a particular thought spiral in–out k-space trajectories optimised for sensitivity
or memory, they were asked to calmly return their atten- to the blood-oxygenation-level-dependent (BOLD)effect
tion to their current experiences whenever they noticed (TE/TR/flip angle ¼ 30 ms/2000 ms/708, 20 cm field-of-view
such distraction. NF was characterised as judging what is (FOV), 5 mm slice thickness, 64 64 matrix, 26 slices in
occurring, trying to figure out what that trait word means to axial oblique orientation. Spiral in–out technique provided
the participant, whether it describes the participant, and improved capability to acquire fMRI signals in regions of
allowing oneself to become caught up in a given train of high magnetic susceptibility (Glover and Law, 2001).
thought. Participants were given multiple examples of each The first 15 TRs of each run were discarded due to scanner
mental state and were invited to ask questions and generate equilibration effects. Structural imaging was conducted with
Fig. 1 Cortical midline areas associated with the narrative self focus condition
(Narrative > time-series baseline) collapsed across MT (following 8 weeks of MT) and
novice (pre MT) groups. VMPFC, ventromedial prefrontal cortex; DMPFC, dorsomedial
present-centred EF. As such, the above results suggest that relative to NF, including the rostral subregions of the dorsal
moment-by-moment self-experience may rely simply on mPFC (BA 9/10/32; 87 voxels, 12 56 40; Z ¼ 3.77) and
task-related suppression of midline cortical representations ventral mPFC (BA 10; 86 voxels, 8 68 8; Z ¼ 3.98)
very similar to those supporting narrative self-focus. Another (Figure 3A), as well as reduced activity in the left dorsal
possibility is that dissociable neural markers of self-reference amygdala (BA 34; 9 voxels, 24 8 8; Z ¼ 3.20), suggest-
may be more evident following extensive training in present- ing that moment-by-moment self-experience may rely on
centred self-focus in the MT group, where engaging distinct suppression of mPFC cortical representations supporting
modes of self reference may be more effortless. narrative self-focus. In addition to the mPFC reductions, EF
In the MT group, experiential self-focus did result in resulted in increased recruitment of a right lateralised
pervasive deactivations along the anterior cortical midline cortical network, including the dorsal and inferolateral
318 SCAN (2007) N. A. S. Farb et al.
PFC (BA 46/45; 77 voxels, 48 56 8; Z ¼ 3.50), insular novices demonstrated strong coupling with midline cortical
cortex (6 voxels, 40 8 16; Z ¼ 3.24), a region along the regions, including the orbital sectors of the ventromedial
supramarginal gyrus consistent with the secondary somato- prefrontal cortices (vmPFC) (BA 11; 6 voxels, 4 56 24;
sensory cortex (SII, BA 40; 24 voxels, 40 40 20; Z ¼ 3.61) Z ¼ 3.68, P < 0.001) (R ¼ 0.609, P < 0.001), suggesting viscer-
extending posteriorly and dorsally into the angular gyrus of osomatic signals are by default associated with activation in
the inferior parietal lobule (BA 39; 7 voxels, 36 76 48; the vmPFC. Such a strong correlation may preclude
Z ¼ 3.11) (Figure 3B–D). This pattern is consistent with the dissociation of these modes of self-focus in novice
MT group demonstrating a marked shift from midline participants. Successfully maintaining present-moment
mPFC-centred recruitment towards right lateralised pre- awareness may require MT individuals to decouple the
frontal cortical and paralimbic structures, in particular automatic responsiveness of the vmPFC to insular activation.
demonstrating recruitment of hypothesised posterior viscer- Supporting this hypothesis, the right insular and vmPFC
osomatic representations. cortices were rendered uncorrelated in the MT group
To examine whether the distinct EF regions revealed in the (R ¼ 0.056, reflecting a significant decrease relative to
MT group were novel areas of recruitment compared to novices, Fisher’s r to Z ¼ 13.36, P < 0.001). This decoupling
novices rather than the product of subthreshold but similar was replaced by an increased coupling of the right insula
patterns of activation, we further performed ROI analyses with the dorsolateral prefrontal cortices (BA 9/44; 17 voxels,
of activation patterns from each participant in both the 56 20 32; Z ¼ 3.18, P < 0.001), as seen in the change from a
MT and novice groups (Figure 3, right panel). Experiential modest correlation in the novice group (R ¼ 0.283) to a very
relative to narrative self-focus was associated with significant strong correlation (R ¼ 0.783, Fisher’s r to Z ¼ 12.09,
reductions in the ventral (F ¼ 11.88, P < 0.003) and dorsal P < 0.001) in the MT group. This pattern of results suggests
mPFC (F ¼ 19.09, P < 0.001), in the MT but not the novice MT may afford greater access to distinct modes of self-focus
group (P’s > 0.3). In contrast, experiential self-focus resulted by promoting a shift away from viewing viscerosomatic
in significantly greater recruitment of the right lateral activity through the lens of the mPFC towards a distinct
PFC (F ¼ 14.75, P < 0.001), right insula (F ¼ 14.41, mode of sensory awareness supported by the lateral PFC.
P < 0.001), and SII/inferior parietal lobule (F > 10.28,
P < 0.003) in the MT but not the novice group (P’s > 0.3). DISCUSSION
Functional connectivity analyses. One reason that novices Consistent with a theory of self-reference as mentalising
may have difficulty in separately attending to distinct modes (Zelazo and Frye, 1998), linguistically mediated (Rochat,
of self-awareness is the habitual coupling of viscero-somatic 1995) and of higher order executive origin (Craik et al.,
and mPFC representations. To examine this more directly, 1999; Johnson et al., 2002; Kelley et al., 2002; Fossati et al.,
we assessed functional connectivity between the visceroso- 2003; Amodio and Frith, 2004; Macrae et al., 2004; Ochsner
matic cortices and the mPFC across the entire time series et al., 2005; Northoff and Heinzel, 2006), participants
(Figure 4). Out of the potential posterolateral seed regions, engaged midline prefrontal cortices (ventral and dorsal
the right insula was chosen as being the best documented in mPFC) and a left lateralised linguistic-semantic network
supporting viscerosomatic awareness (Damasio, 1999; (inferior lateral PFC, middle temporal and angular gyri)
Adolphs, 2002; Craig, 2004; Critchley et al., 2004). An during NF. Demonstrating a default bias towards NF as
analysis of right insular functional connectivity revealed that previously revealed in ‘resting’ mind wandering states
Dissociable neural modes of self-reference SCAN (2007) 319
(Mason et al., 2007), relatively restricted reductions in the cortex) sensory events, rather than their affective or
cortical midline network were found when attention was subjective self-referential value. This pattern mirrors neural
explicitly directed towards a moment-to-moment EF in dissociations between the affective and sensory components
novice participants with little training in this form of of pain sensation, with the former supported by anterior
self-reflection. These individuals revealed increased left midline structures, and latter supported by posterior lateral
lateralised prefrontal-parietal activations during EF likely sensory cortices (Rainville et al., 1997). This detached or
reflecting greater task-related linguistic processing that has objective mode of self-focus may be aided by the recruitment
been shown to be associated with decreased medial of the right angular gyrus of the inferior parietal lobule.
prefrontal recruitment (Gusnard et al., 2001). In addition to studies showing that this region is involved
Participant inexperience with different forms of self-focus in switching between first and third person perspectives
might limit the ability to accurately reveal functionally and (Ruby and Decety, 2004), stimulation of this region has been
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