Reflecting upon Feelings: An fMRI Study of Neural Systems Supporting the Attribution of Emotion to Self and Other

Article (PDF Available)inJournal of Cognitive Neuroscience 16(10):1746-72 · January 2005with471 Reads
DOI: 10.1162/0898929042947829 · Source: PubMed
Abstract
Understanding one's own and other individual's emotional states is essential for maintaining emotional equilibrium and strong social bonds. Although the neural substrates supporting ref lection upon one's own feelings have been investigated, no studies have directly examined attributions about the internal emotional states of others to determine whether common or distinct neural systems support these abilities. The present study sought to directly compare brain regions involved in judging one's own, as compared to another individual's, emotional state. Thirteen participants viewed mixed valence blocks of photos drawn from the International Affective Picture System while whole-brain fMRI data were collected. Preblock cues instructed participants to evaluate either their emotional response to each photo, the emotional state of the central figure in each photo, or (in a baseline condition) whether the photo was taken indoors or outdoors. Contrasts indicated (1) that both self and other judgments activated the medial prefrontal cortex (MPFC), the superior temporal gyrus, and the posterior cingulate/precuneus, (2) that self judgments selectively activated subregions of the MPFC and the left temporal cortex, whereas (3) other judgments selectively activated the left lateral prefrontal cortex (including Broca's area) and the medial occipital cortex. These results suggest (1) that self and other evaluation of emotion rely on a network of common mechanisms centered on the MPFC, which has been hypothesized to support mental state attributions in general, and (2) that medial and lateral PFC regions selectively recruited by self or other judgments may be involved in attention to, and elaboration of, internally as opposed to externally generated information.
16.10
Reflecting upon Feelings: An fMRI Study of Neural
Systems Supporting the Attribution of Emotion
to Self and Other
Kevin N. Ochsner
1
, Kyle Knierim
2
, David H. Ludlow
2
, Josh Hanelin
1
,
Tara Ramachandran
2
, Gary Glover
2
, and Sean C. Mackey
2
Abstract
& Understanding one’s own and other individual’s emotional
states is essential for maintaining emotional equilibrium and
strong social bonds. Although the neural substrates supporting
reflection upon one’s own feelings have been investigated, no
studies have directly examined attributions about the internal
emotional states of others to determine whether common or
distinct neural systems support these abilities. The present
study sought to directly compare brain regions involved in
judging one’s own, as compared to another individual’s, emo-
tional state. Thirteen participants viewed mixed valence blocks
of photos drawn from the International Affective Picture
System while whole-brain fMRI data were collected. Preblock
cues instructed participants to evaluate either their emotional
response to each photo, the emotional state of the central
figure in each photo, or (in a baseline condition) whether the
photo was taken indoors or outdoors. Contrasts indicated (1)
that both self and other judgments activated the medial
prefrontal cortex (MPFC), the superior temporal gyrus, and
the posterior cingulate/precuneus, (2) that self judgments
selectively activated subregions of the MPFC and the left tem-
poral cortex, whereas (3) other judgments selectively activated
the left lateral prefrontal cortex (including Broca’s area) and
the medial occipital cortex. These results suggest (1) that self
and other evaluation of emotion rely on a network of common
mechanisms centered on the MPFC, which has been hypothe-
sized to support mental state attributions in general, and (2)
that medial and lateral PFC regions selectively recruited by
self or other judgments may be involved in attention to and
elaboration of internally as opposed to externally generated
information. &
INTRODUCTION
At various moments in our daily lives we migh t have
cause t o r eflect on how we feel. We mig ht reflect
because someone asks ‘‘How are you?’’, or we might
take stock when we’ve accomplished a goal (I got the
job and I feel great!), experienced a significant life event
(My aunt died and I feel awful), or are anticipating one
(We’re expecting a child and are excited but nervous!).
Whatever it is that we reflect, research suggests that the
ability to readily and specifically answer the question,
‘‘How do I feel?’’ helps us to identify situations worth
seeking or avoiding, engage in behaviors that promote
desired affective states, and effectively regulate our
emotions (Barrett, Gross, Christensen, & Benvenuto,
2001; Lane & Schwartz, 1987). But our social world
prompts us to reflect not just upon our own feelings,
but the feelings of those around us as well. As we
interact with colleagues, compete with opponents, or
watch our friends and family experience their own ups
and downs, having insight into the feelings of others
enables us to understand what they value, how they feel
about us, to offer appropriate support or gain compet-
itive advantage, and to predict their future behavior
(Blair, 2003; Baron-Cohen, 1995).
Despite the importance of these abilities for our social
and emotional well-being, our understanding of the
underlying neurocognitive mechanisms has only just
begun to take shape. The goal of the present study was
to use func tional magnetic resonance imaging (fMRI) to
directly compare the neural processes supporting infer-
ences about one’s own and other individuals’ emotional
states. By determining whether and how these pro-
cesses are similar and different, we might gain insight
into the question of how we reflect upon feelings,
knowing that we feel good or bad, and that others feel
good or bad as well.
On one hand, there are reasons to believe that
common psychological and neural processes mediate
understanding of one’s own and other people’s emo-
tions. Proponents of this view might argue that the
perception of emotion in self and other involves drawing
inferences and making attributions about the nature of
internal mental states, a capacity referred to as theory of
mind (TOM) (Lane & McRae, in press; Gallagher & Frith,
1
Columbia University,
2
Stanford University
D 2004 Massachusetts Institute of Technology Journal of Cognitive Neuroscience 16:10, pp. 1–27
2003; Frith & Frith, 1999; Premack & Woodruff, 1978).
TOM is used to infer the intentions and beliefs that
motivate and guide goal-directed behavior, and has been
conceived as a centra l ‘‘mentalizing’’ ability that can be
broadly applied to understanding many kinds of mental
states, whether those states are one’s own or those of
another person (Gallagher & Frith, 2003; Frith & Frith,
1999; Premack & Woodruff, 1978).
Empirical support for a central mentalizing mecha-
nism comes from functional imaging studies observing
activation of a particular brain region, the medial pre-
frontal cortex (MPFC), both when individuals judge
some aspect of their own or someone else’s mental
states. When perceiving one’s self, MPFC activation has
been observed in a variety of conditions listed in Table 5
whose accompanying activation foci are visualized
in Figure 4A. Self-referential judgments activating the
MPFC include evaluating one’s internal state of arousal
or pleasantness (a–f in Table 5), judging likes and dis-
likes for externally presented stimuli (g–k), recognition
of one’s own face or voice (l–o), perspective taking (p–r),
judging one’s personality traits and attributes (s–y), or
when self-generating thoughts/associations i n a goal-
directed (z–ee) of spontaneous fashion (ff; cf. Gusnard
& Raichle, 2001). When perceiving others, MPFC activa-
tion has been observed in a variety of tasks and con-
ditions, also listed in Table 5 with accompanying
activation foci visualized in Figure 4B. Conditions pro-
ducing social-cognition-related activations of MPFC in-
clude judging the goodness/badness of actions or
images (1–6 in Table 5), perceiving eye gaze (7–10),
judgments of social t argets (represented by photos,
words, or cartoons of moving shapes) that may require
mental state inferences and/or recruitment of social
knowledge (11–19), taking a third-person perspective
(20–21), tasks requiring explicit TOM judgments of
intention (22–27), and games that participants believe
are being played interactively in real time with another
participant (28–29). Taken together, data from studies of
self-reflection and social-cognition-related judgments
implicate MPFC in the general process of ‘‘mentalizing’’
about internal states (Gallagher & Frith, 2003).
On the other hand, the cross-study comparisons cited
above provide only indirect support for the hypothesis
that the same MPFC-based system is recruited by self- or
other-focused mental state attributions. As vividly illus-
trated i n Figure 5, the variety of self-referential and
social-cognitive judgments employed thus far have acti-
vated virtually the entire extent of the medial frontal
cortex, spanning the dorsal (including Brodmann’s areas
[BAs] 8–10 and dorsal portions of BA 32) and ventral
(BAs 10, 11, 14, 25, and the ventral anterior cingulate)
MPFC as well as the anterior cingula te cortex (BAs 24–32).
The variability in activation could reflect overlap of self-
reflective and social-cognitive processing, but it also
reflects recruitment of a number of distinct processes
in distinct medial frontal subregions not yet well differ-
entiated across tasks, cross-study variability in functional
localization across participants, and/or variations intro-
duced by cross-experimenter differences in spatial or
statistical processing of data. In the absence of within-
study comparisons, it is quite difficult to know whether
apparent overlap in fact reflects common recruitment of
underlying processes. To date, only one study has ad-
dressed this issue in the context of TOM. Using story
vignettes, Vogeley et al. (2001) found MPFC involvement
both when making TOM attributions about descriptions
of another individuals’ behavior and when making in-
tentional judgments about descriptions of one’s own
behavior, but did not find differential MPFC activation
between the two conditions. No studies have examined
this issue in the context of emotion.
More generally, to date, no studies have attempted to
isolate processes supporting judgments about the ‘‘inter-
nal emotional’’ states of others. Although TOM-related
studies may sometimes use cues with affective connota-
tions (e.g., Wicker, Perrett, Baron-Cohen, & Decety, 2003),
they may more commonly require judgments about non-
affective, cognitive states, and in any event, the extent to
which emotional inferences are required in such tasks
has not been systematically manipulated or measured
distinct from the need for drawing inferences about
cognitive states. Studies showing MPFC activation in
response to em otional films (e.g., Lane, Fink, Chau,
& Dolan, 1997), when recal ling emotional memories
(Reiman et al., 1997), during visual imagery of traumatic
events (Shin et al., 1997), or when perceiving happy,
angry, or sad emotional facial expressions (Kesler-West
et al., 2001; Blair, Morris, Frith, Perrett, & Dolan, 1999;
Phillips et al., 1998), also are ambiguous because they do
not control the extent to which participants reflect upon
their own emotional state as compared to that experi-
enced, perceived, or imagined by others. Two studies
have found greater MPFC activat ion when judging how
well positive or negative trait words described them-
selves as compared to another famous individual (Kelley
et al., 2002; Craik et al., 1999), but it is not clear how
much dispositional trait judgments are related to judg-
ments of one’s current emotional state.
Beyond the possible recruitment of the MPFC, a
number of other processing systems might be similarly
or differentially recruited during self- and/or other-
focused emotion perception. Indeed, the mechanisms
supporting ‘‘mentalizing’’ are complex, and when fully
unpacked may include a larger network of neural sys-
tems that are thought to play a role in TOM and/or
self-reflective judgments, including: frontal regions im-
portant for language and working memory; superior
temporal regions implicated in processing nonverbal
cues with social significance; the posterior cing ulate
cortex, which has been associated with affective evalua-
tion; and the parietal cortex, which may be involved in the
representation of spatial perspectives that help distin -
guish self and other (Frith & Frith, 1999, 2003; Gallagher
2 Journal of Cognitive Neuroscience Volume 16, Number 10
& Frith, 2003; Maddock, Garrett, & Buonocore, 2003;
Meltzoff & Decety, 2003; Saxe & Kanwisher, 2003; Vogeley
& Fink, 2003; Johnson et al., 2002; Kelley et al., 2002;
Brunet, Sarfati, Hardy-Bayle, & Decety, 2000; Castelli,
Happe, Frith, & Frith, 2000; Gal lagher, Happe, et al.,
2000; Kircher, Senior, Phillips, Benson, et al., 2000; Mad-
dock & Buonoco re, 1997). Attributions of emotion to
self and other could commonly or differentially recruit
such systems independently of, or in concert with,
MPFC. This possibility is supported by the study of
Vogeley et al. (2001), who found greater temporal–
parietal activation when participants made intentional
TOM attributions about their own as compared to
another person’s behavior.
To identify the common and distinct neural systems
supporting the evaluation of emotion in self and other,
the present study employed a variation of a paradigm
developed by Lane et al. (1997; cf. Gusnard et al., 2001).
In this task, participants were presented with a series of
blocks of photographic images and for each block were
asked to judge either their own emotional response to
each photo (pleasant, unpleasant or neutral), or to
judge where the image had been taken (indoors, out-
doors, or not sure). The present study modified this
paradigm through the inclusion of a third condition,
which asked participants to judge the emotional re-
sponse of the central character in each image (pleasant,
unpleasant, or neutral). The inclusion of this condition
allows (1) identification of regions commonly activated
when judging one’s own (self blocks) or another per-
son’s (other blocks) feelings as compared to a percep-
tual judgment controlling for spatial attention to images
(in–out blocks), and (2) direct comparison of regions
implicated in evaluating one’s own (self blocks) or
another person’s emotional experience (other blocks)
to identify regions uniquely activated by each type
of judgment.
RESULTS
Behavioral Results
An ANOVA on proportion of affect judgments with type
of judgment (self vs. other) and valence of judgment
(pleasant, unpleasant, or neutral) as within-subject fac-
tors revealed a main effect of a valence [F(2,12) = 27,48,
p < .0001], and no significant effects involving type of
judgment (Figure 1A). Planned contrasts demonstrated
that the greatest proportion of affect judgments were
positive [F(1,12) > 9.0, p < .006 for comparisons to
neutral and negative] and the smallest proportion were
neutral [F(1,12) > 53.94, p < .001 for comparisons to
negative and positive]. An ANOVA on response times
using the same factors again revealed a main effect of a
valence [F(2,12) = 36.92, p < .0001], and no significant
effects involving type of judgme nt (Figure 1B). Planned
contrasts demonstrated that response times were lon-
gest when participants judged an emotional response to
be neutral [F(1,12) > 63.80, p < .001 for comparisons to
negative and positive]. Positive and negative judgments
were made with equal speed [F(1,12) < 1]. Analyses of
judgments made on in–out trials were conducted sepa-
rately and revealed a nonsignificant tendency for par-
ticipants to judge more photos as having been taken
outside rather than inside [t(12) = 2.08, p < .06], whereas
unsure judgments were made least often [t(12) > 5.16,
p < .001 for comparisons with inside and outside].
Similar trends were found for response times, with
outside judgments made most rapidly [t(12) > 3.31,
p < .008 for comparisons with inside and unsure],
whereas unsure judgments tended to be made most
slowly [t(12) = 1.60, p < .14 vs. inside]. Overall, response
times for emotional self and other as compared to
spatial in–out judgments were made with equal speed
( p > .47).
Imaging Results
Regions associated with the evaluation of one’s own
emotional experience were identified in the contrast of
self and in–out blocks, whereas regions associated with
the evaluation of another person’s emotional experience
were identified in the cont rast of other and in–out
blocks.
1
These two contrasts revealed similar patterns
of activation, as shown in Tables 1 and 2, and the blue
circles in Figure 4A and B. Activation was found in
overlapping regions of the MPFC, including a peak
activated voxel (6, 52, 32) that was found for clusters
in both contrasts, and in similar and/or overlapping
regions of the posterior cingulate cortex/precuneus,
and the superior temporal gyrus (STG). Dissimilar acti-
vations included medial prefrontal activations that ex-
tended dorsally into BA 8, the left superior lateral
prefrontal cortex, the anterior cingulate cortex, and
the left inferior parietal lobe for self-focused emotion
attributions, and the left ventral lateral prefrontal (over-
lapping Broca’s area) and parahippocampal cortices for
other-focused emotion attributions.
Regions commonly activated by both self and other
attributions were identified formally by masking th e
contrast of other and in–out blocks with the contrast
of self and in–out blocks, and were confirmed by per-
form ing planned comparison t tests on measures of
percent signal change to verify that self and other blocks
in showed similar patterns of activation that were signif-
icantly different from activation shown in–out blocks. As
shown in Table 3 and Figure 2, common activation was
identified in regions of the MPFC, the lateral PFC, the
posterior cingulate cortex/precuneus, and the superior
temporal sulcus/gyrus. For all regions, t tests confirmed
that both self and other blocks showed significantly
greater activation than in–out blocks (all t > 2.4, p <
.05). In addition, self and other blocks showed equiva-
lent levels of activation (all t <1,p = ns) for all regions
Ochsner et al. 3
except for the lateral prefrontal region for which there a
marginally significant difference [t(12) = 2.15, p < .06].
Regions specifically associated with either the evalua-
tion of one’s own, or an other person’s, emotional
experience were identified by directly contrasting self
and other blocks. As shown in Figure 3, self blocks
selectively activated two clusters in the MPFC as well
as in the right middle temporal gyrus, whereas other
blocks selec tively activated the left ventral lateral pre-
frontal cortex and the cuneus in the medial occipital
lobe (Table 4). Plan to comparison t tests on measures of
block average percent signal change confirmed that self
blocks selectively recruited two regions of the MPFC
(t > 2.3, p < .05) and that other blocks selectively
recruited the lateral PFC and the occipital cortex (t >
2.4, p < .05).
DISCUSSION
This is the first study to directly compare neural systems
involved in attributing emotional experiences to oneself
and to other individuals. With equal speed participants
discerned equal proportions of pleasant, unpleasant,
and neutral affective states in themselves and in individ-
uals depicted in photographic scenes.
Common Processes Mediating Attribution
of Emotion to Self and Other
In comparison to baseline judgments of the spatial
characteristics of images, neural correlates of self- and
other-focused attributions were similar in three impor-
tant ways. First, self- and other-oriented judgments
commonly recruited regions of the MPFC that previously
have been implicated both in TOM (Gallagher & Frith,
2003; Frith & Frith, 1999) and in the representation of
meta-states of self-awareness, thought to be necessary
for r eflecting upon (Lane & McRae, in press) and
regulating (Ochsner, Ray, Gabrieli, & Gross, in press;
Ochsner & Gross, 2004) one’s own emotional state.
Common recruitment of this region by self and other
emotion perception supports the hypothesis that a
central sys tem supports intentional attributions about
one’s own and other individual’s internal states (Gal-
lagher & Frith, 2003; Premack & Woodruff, 1978; cf.
Vogeley et al., 2001) and may suggest that judgments of
the internal feelings of others are guided by an under-
Figure 1. Behavioral data
for self, other, and in–out
judgments. (A) Proportions of
affect judgments for self and
other blocks, and inside versus
outside judgments for in–out
blocks. (B) Group averaged
response times for judgments
made for self, other, and in–out
blocks.
4 Journal of Cognitive Neuroscience Volume 16, Number 10
standing of our own feelings in response to the events
we see them experiencing.
Second, both self and other judgments recruited a
region of the left inferior lateral prefrontal cortex (BA
45) thought to be involved in mediating competition, or
resolving interference, between competing associations
in verbal working memory (Bunge, Ochsner, Desmond,
Glover, & Gabrieli, 2001; Jonides, Smith, Marshuetz,
Koeppe, & Reuter-Lorenz, 1998). Joint activation of the
MPFC and a very similar region of the left inferior lateral
PFC has been observed during both the appraisal of
aversive stimuli as negative as well as the appraisal of
neutral stimuli as negative (Ochsner & Gross, 2004). In
combination with the present findings, this suggests that
the MPFC and the inferior lateral PFC might work in
concert to media te interference between , and select the
appropriate, semantic description of emotional states.
Third, both self- and other-oriented judgments re-
cruited regions spanning the junction of the posterior
cingulate cortex and the precuneus, as well as regions of
the superior temporal sulcus/gyrus. The posterior cin-
gulate has been associated with evaluating the affective
valence of external stimuli (Maddock et al., 2003; Mad-
dock & Buonoco re, 1997), and has been activated when
Table 1. Group Activations for Self > In–Out Contrast
Coordinates
Region of Activation Brodmann x y z Z score Volume (mm
3
)
Superior frontal gyrus 9 6 52 32 3.59 1072
Superior FG 9 0 56 36 3.53 (L)
Superior FG L8 12 58 44 3.37 (L)
Superior frontal gyrus L8 12 36 52 3.89 552
Superior FG L9 0 56 36 3.53 (L)
Superior FG L9 12 58 44 3.37 (L)
Superior frontal gyrus L10 26 58 30 3.14 256
Superior FG L10 20 66 34 3.07 (L)
Superior FG L10 28 50 32 2.84 (L)
Superior frontal gyrus R8 16 34 52 3.45 168
Medial frontal gyrus L10 10 56 14 3.95 2784
Medial FG 9/10 2 56 24 3.79 (L)
Medial FG 10 2 56 8 3.60 (L)
Anterior cingulate 24 4 26 28 3.16 216
Anterior cingulate 24 2 18 40 4.00 376
Posterior cingulate/precuneus L31 8 48 34 3.09 648
Posterior cingulate L31 4 56 22 3.09 (L)
Posterior cingulate L31 12 54 26 3.06 (L)
Superior temporal gyrus L38 52 16 10 4.75 1672
Superior TG L22 52 12 2 3.61 (L)
Inferior FG L45 52 22 12 3.90 (L)
Middle temporal gyrus L21/22 60 34 2 3.74 544
Middle temporal gyrus R22 52 38 0 3.58 624
Middle TG R21/22 46 44 4 3.19 (L)
Inferior parietal lobe L39 54 70 46 3.15 200
Caudate body L 2 8 8 3.08 232
Note: Local maxima for clusters are denoted with (L). R and L hemispheres are not designated for maxima within 6 mm of midline. Coordinates are
in MNI space.
Ochsner et al. 5
participants judged the morality of their own or some-
one else’s behavior (Greene, Sommerville, Nystrom,
Darley, & Cohen, 2001). The precuneus has been asso-
ciated with adopting a first-person as compared to a
third-person spatial perspective (Vogeley & Fink, 2003),
which may underlie recruitment of this region during
Table 2. Group Activations for Other > In–Out Contrast
Coordinates
Region of Activation Brodmann x y z Z score Volume (mm
3
)
Medial frontal gyrus 10 2 56 10 3.03 264
Medial FG 9 2 56 22 2.94 (L)
Medial FG 10 6 54 26 2.89 (L)
Superior frontal gyrus 9 6 52 32 3.26 352
Superior FG 9 2 56 40 3.09 (L)
Superior temporal gyrus R22 44 42 6 3.54 488
Middle TG R22 52 38 2 3.24 (L)
Middle occipital/temporal R19/37/39 56 70 6 3.75 256
Precuneus L31 10 58 32 4.12 848
8 52 24 3.22 (L)
18 56 30 2.75 (L)
Parahippocampal gyrus R 28 26 10 3.49 160
Note: Local maxima for clusters are denoted with (L). R and L hemispheres are not designated for maxima within 6 mm of midline. Coordinates are
in MNI space.
Table 3. Group Activations Commonly Activated by Self and Other Judgments (Revealed by Other > In–Out Contrast Masked by
Self > In–Out Contrast)
Coordinates
Region of Activation Brodmann x y z Z score Volume (mm
3
)
Medial frontal gyrus 9 2 56 40 3.17 440
Medial FG 9 4 52 32 2.92 (L)
Medial frontal gyrus 10 2 58 12 2.90 256
Medial FG 9 6 54 26 2.62 (L)
Medial FG 9 2 54 24 2.56 (L)
Superior frontal gyrus 8 18 38 54 2.86 232
Superior FG 8 10 38 56 2.83 (L)
Superior frontal gyrus 10 26 56 30 2.88 160
Inferior frontal gyrus 45 52 22 12 3.10 192
Precuneus 7 6 60 30 3.85 672
Precuneus 31/ 7 14 58 32 3.56 (L)
Posterior cingulate 31 4 54 24 3.00 (L)
Superior temporal sulcus 22 46 40 2 3.54 256
Superior temporal sulcus 22 60 42 2 2.68 128
Superior temporal gyrus 22 64 32 12 3.17 104
Note: Local maxima for clusters are denoted with (L). R and L hemispheres are not designated for maxima within 6 mm of midline. Coordinates are
in MNI space.
6 Journal of Cognitive Neuroscience Volume 16, Number 10
episodic memory retrieval (Kr ause et al., 1999; Fink et al.,
1996). Being able to toggle between first- and third-
person perspectives (‘‘I feel sad’’ vs. ‘‘I keep sighing, I
guess I feel sad ’’) may be essential for making attribu-
tions about our own or other’s feelings, actions, and
attributes. In keeping with this notion, the precuneus
has been recruited both by TOM attributions (Wicker
et al., 2003; Gallagher, Happe, et al., 2000), and when
judging the self-descriptiveness of trait words (Kelley
et al., 2002; Kircher, Brammer, et al., 2002; Lieberman,
Gaunt, Gi lbert, & Trope, 2002) or thoughts (Kjaer,
Nowak, & Lou, 2002). The STG may be involved in the
bottom-up registration of cues that imply intent ional
action (Gallagher & Frith, 2003; cf. Saxe & Kanwisher,
2003), as suggested by its activation during the percep-
tion of biological motion implied by point-light displays,
or actually produced by hands, bodies, mouths, and eyes
(for reviews, see Gallagher & Frith, 2003; Puce & Perrett,
2003; Allison, Puce, & McCarthy, 2000), and its activation
when participants analyze physical rather than mental
causes for described actions (Gallagher, Happe, et al.,
2000). Recruitment of these regions in addition to the
MPFC suggests that self and other perceptions of emo-
tion rely on a network of regions whose constituent
members encode affective, spatial , and nonverbal cues
relevant to app raising t he affective significance of a
stimulus to oneself or others. Just as post erior temporal
and lateral prefrontal cortical regions support bottom-
up and top-down processes during recognition of non-
social objects (Kosslyn et al., 1994), posterior cortical
regions may support bottom-up recognition o f inten-
tional behaviors, whereas the MPFC may support rea-
soned top-down attributions about the mental states
that guide them.
Distinct Processes Mediating Attribution of
Emotion to Self and Other
Direct comparison of the two judgment types provided
the strongest test of differential recruitment during self
and other emotion perception. Because self and other
judgments both involved the same type of affective
evaluation, differences observed when the two condi-
tions were contrasted directly should reflect differences
in the deployment of attention to, and elaboration of,
internal or external cues that differentially contribute to
each type of judgment. With this in mind, three key
findings were observed.
2
First, self judgments selectively activated the MPFC,
suggesting that distinct subregions within the MPFC are
involved in self-focused attention and as opposed to
attributional process generally applicable to understand-
ing one’s own and other individual’s emotional states.
3
This finding is consistent with studies showing greater
MPFC activation when judging whether trait adjectives
or phrases describe oneself as compared to another
familiar person (Lieberman, Jarcho, & Satpute, in press;
Kelley et al., 2002; Kjaer et al., 2002; Craik et al., 1999).
Second, other judgments selectively activated the left
inferior lateral prefrontal cortex, which is consistent
with the general role played by lateral prefrontal areas
in the maintenance and manipulation of information
Figure 2. Results of other > in–out group contrast masked by self > in–out group contrast revealing regions commonly activated above baseline
superimposed on canonical T1 anatomical images illustrating. Top row shows regions of the left inferior prefrontal cortex (A), the dorsal medial
prefrontal cortex (B and D), the posterior cingulated/precuneus (C ), and the superior temporal gyrus (E) recruited both by self and other
judgments more than in the perceptual baseline condition. Bottom row shows present signal change for each block type for each activated region
with error bars showing standard error of the mean. Activation in self and other conditions is statistically equivalent ( p > .05) for all regions and
approached significance only for the left inferior prefrontal cortex (A, p < .052) and the posterior cingulate/precuneus (C, p < .06).
FPO
Ochsner et al. 7
about the external world (Smith & Jonides, 1999)
4
as
well as the retrieva l of semantic/contextual knowledge
that could be used to interpret social targets (Wagner,
Pare-Blagoev, Clark, & Poldrack, 2001). The recruit-
ment of Broca’s area is interesting because this region
has been implicated in representing the goals of actions
that either are observed or executed (Heiser, Iacoboni,
Maeda, Marcus, & Mazziotta, 2003; Iacoboni, Woods,
et al., 1999). Notably , the medial/lateral prefrontal split
for self and other judgments parallels the finding of
MPFC activation for emotion regulation strategies that
reappraise the self-relevance of aversive scenes, as com-
pared to lateral PFC activation for strategies that reap-
praise the actions and outcomes of actors depicted in
those scenes (Ochsner, Ray, et al., in press). Third, re-
cruitment of middle temporal cortex for self judgments
may reflect this regions role in representing semantic
and linguistic content, including the self monitoring
(Hashimoto & Sakai, 2003) and retrieval of verbal infor-
mation that conveys emotional (Mitchell, Elliott, Barry,
Cruttenden, & Woo druff, 2003) or personal (Paller et al.,
2003) information, where as activation of the medial
occipital cortex for other judgments may reflect height-
ened attention to external visual inputs when evalu-
ating visual cues to another’s mental state (Culham &
Kanwisher, 2001).
5
Role of the MPFC in Emotion and Social Cognition
Taken together, the present findings suggest that dis-
tinct but highly overlapping neural systems support the
attribution of emotion to oneself and to others. A key
player in both networks was the MPFC. Portions of the
MPFC supported both the attribution of emotional
states to oneself and to other pictured individuals, which
supports the hypothesis that this region is the hub of a
system mediating inferences about one’s own and other
individual’s ment al states (Lane & McRae, in press;
Gallagher & Frith, 2003). In addition, distinct MPFC
regions were more activated for self judgments, whereas
lateral PFC regions were more activated for other judg-
ments. This finding suggests that distinct control sys-
tems are involved in atten ding to a nd e laborating
internally as compared to externally generated informa-
Figure 3. Results of group contrasts between self and other judgments revealing regions more activated during each type of judgment. Graphs
show percent signal change for representative ROIs on self, other, or in–out blocks with error bars showing standard error of the mean. (A) Medial
view of left hemisphere showing two clusters in the MPFC identified in the self > other contrast. (B) Axial images showing (left panel) activations of
the left lateral prefrontal cortex (intersecting Broca’s area) and (right panel) the occipital cortex identified in the other > self contrast.
FPO
8 Journal of Cognitive Neuroscience Volume 16, Number 10
tion (cf. Christoff, Ream, & Gabrieli, in press; Christoff,
Ream, Geddes, & Gabrieli, 2003), a distinction that also
has been observed in the context of emotion regulation
(Och sner, Ray, et al., in press) rather than emotion
attribution. That being said, there remain at least three
important and interrelated questions concerning the
specific functional roles played by the MPFC.
The first question concerns the precise characteriza-
tion of MPFC functions , and whether MPFC activation
reflects a special kind of process devoted to mental state
attributions, social cognition, and/or self-mo nitoring
more generally. This question arises because MPFC
activation has been observed in contexts that at first
blush do not appear to require mental state inferences
to be drawn, including inductive reasoning (Goel, Gold,
Kapur, & Houle, 1997), judging semantic coherence
(Ferstl & von Cramon, 2002) or word familiarity (Hen-
son, Rugg, Shallice, Josephs, & Dolan, 1999), and pro-
spective memory (Burgess, Scott, & Frith, 2003). Should
these results be taken to suggest that the MPFC carries
out some type of cognitive processing common to
social-cognitive and self-referential judgment, or should
the reverse conclusion be drawn, that apparently cogni-
tive tasks require social-cognitive and self-referential
processing?
6
On one hand, it is difficult to address this
question in the absence of single study comparisons of
tasks requiring mental state attributions with tas ks that
do not require them, in order to determine whether
similar or different regions of the MPFC are being
recruited in the two cases.
7
On the other hand, many
of the tasks that recruit dorsal (coarsely defined as z >0,
see Table 5, Figure 4) regions of the MPFC (see right-
most column of Table 5) could be described as requiring
a common form of metacognitive processing. Social
behaviors, personal characteristics, emotional experien-
ces, linguistic utterances, and inductive problems can all
be seen as examples of stimuli for which an attributed
meaning is a meta-level emergent property of multiply
interpretable inputs that, in and of themselves, do not
directly imply a single interpretation. From this perspec-
tive, the MPFC might be important for the metacognitive
ability to re-r epresen t affective, cognitive, and other
types of inputs in a self-generated symbolic (perhaps
linguistically describable, e.g., ‘‘I feel good,’’ or ‘‘He is
sad.’’) format (Christoff, Ream, & Gabrieli, in press;
Gallagher, Jack, Roepstorff, & Frith, 2002). By contrast,
ventral (for present purposes defined as z < 0) regions
of the MPFC have been strongly associated with repre-
senting the affective value of stimuli (e.g., Lane & McRae,
in press; Fellows & Farah, 2003; Bechara, Damasio, &
Damasio, 2000), in part because of connections with the
amygdala and autonomic centers more robust than
those present for the dorsal regions of the MPFC (Ongur
& Price, 2003; Ongur, Ferry, & Price, 2000). At present,
inspection of Table 5 and Figure 4 suggests that the
ventral MPFC is activat ed by, but not cons istently asso-
ciated with, specific types of self-referential or social-
cognitive judgments. To date, studies have not been
specifically designed to address the differential roles of
dorsal and ventral medial prefrontal regions in self-
referential and social-cognitive processing.
The second question concerns role specific com-
putations carried out by the MPFC in the present
experiment, which are illuminated by the precedin g
discussion. All of the activations in the present study fall
within the dorsal region of the MPFC, which is consis-
tent with the notion that the mental state attributional
processes isolated here are related to two self-referential
Table 4. Group Activations for Self > Other and Other > Self Contrasts
Coordinates
Region of Activation Brodmann x y z Z score Volume (mm
3
)
Self > Other Contrast
Superior frontal gyrus 9 2 58 38 3.17 200
Medial frontal gyrus 10 2 50 16 3.03 192
Middle temporal gyrus L21 62 34 6 3.04 160
Other > Self Contrast
Inferior frontal gyrus L44 58 6 18 3.19 248
Inferior FG L45 58 12 24 2.83 (L)
Medial occiptial gyrus/cuneus 17&18 2 80 6 3.66 456
Cuneus 18 2 80 16 3.10 (L)
Note: Local maxima for clusters are denoted with (L). R and L hemispheres are not designated for maxima within 6 mm of midline. Coordinates are
in MNI space.
Ochsner et al. 9
Table 5. Medial prefrontal activation coordinates for studies involving self-referential or social-cognitive judgments (shown in Figure 4)
Identifier Study x y z Target Judgment Type Contrast Region Figure
Blue circle Ochsner et al., this volume 6 52 32 self affective evaluation self appraisal of emotion vs. in–out perceptual
judgment
dMPFC 4A
Blue circle Ochsner et al., this volume 12 36 52 self affective evaluation self appraisal of emotion vs. in–out perceptual
judgment
dMPFC 4A
Blue circle POchsner et al., this volume 10 56 14 self affective evaluation self appraisal of emotion vs. in–out perceptual
judgment
dMPFC 4A
Blue circle Ochsner et al, this volume 4 26 28 self affective evaluation self appraisal of emotion vs. in–out perceptual
judgment
dMPFC 4A
Blue circle Ochsner et al., this volume 2 58 38 self affective evaluation self appraisal of emotion vs. other appraisal of
emotion
dMPFC 4A
Blue circle Ochsner et al., this volume 2 50 16 self affective evaluation self appraisal of emotion vs. other appraisal of
emotion
dMPFC 4A
Yellow sq. Ochsner et al., this volume 2 56 40 self and social affective evaluation both self and other appraisals vs. in–out dMPFC 4A
Yellow sq. Ochsner et al., this volume 2 58 12 self and social affective evaluation both self and other appraisals vs. in–out dMPFC 4A
Yellow sq. Ochsner et al., this volume 18 38 54 self and social affective evaluation both self and other appraisals vs. in–out dMPFC 4A
Blue circle Ochsner et al., this volume 2 56 10 social affective evaluation other appraisal of emotion vs. in–out perceptual
judgment
dMPFC 4A
Blue circle Ochsner et al., this volume 6 52 32 social affective evaluation other appraisal of emotion vs. in–out perceptual
judgment
dMPFC 4A
a Gusnard et al., 2001 11 30 44 self affective evaluation appraise own feeling vs. in–out judgment dMPFC 4A
a Gusnard et al., 2001 7 45 25 self affective evaluation appraise own feeling vs. in–out judgment dMPFC 4A
a Gusnard et al., 2001 3 53 24 self affective evaluation appraise own feeling vs. in–out judgment dMPFC 4A
a Gusnard et al., 2001 11 23 52 self affective evaluation appraise own feeling vs. in–out judgment dMPFC 4A
a Gusnard et al., 2001 9 39 42 self affective evaluation appraise own feeling vs. in–out judgment dMPFC 4A
a Gusnard et al., 2001 5 3 48 self affective evaluation own feeling vs. fixation baseline ACC 4A
b Lane et al., 1997 0 50 16 self affective evaluation appraise own feeling vs. in–out judgment dMPFC 4A
b Lane et al., 1997 8108 self affective evaluation appraise own feeling vs. in–out judgment vMPFC 4A
c Paradiso et al., 1999 0 29 35 self affective evaluation rate valence of response for pleasant vs.
unpleasant photos
dMPFC 4A
c Paradiso et al., 1999 7 28 35 self affective evaluation rate valence of response for pleasant
vs. unpleasant photos
dMPFC 4A
10 Journal of Cognitive Neuroscience Volume 16, Number 10
d Paulus & Frank, 2003 14 40 1 self affective evaluation judge own preference vs. visual discrimination ACC 4A
d Paulus & Frank, 2003 9 50 4 self affective evaluation judge own preference vs. visual discrimination vMPFC 4A
e Phan et al., 2003 3 54 27 self affective evaluation rate own arousal for aversive images dMPFC 4A
e Phan et al., 2003 3 54 33 self affective evaluation rate own arousal for aversive images dMPFC 4A
e Phan et al., 2003 12 60 27 self affective evaluation rate own arousal for aversive images dMPFC 4A
e Phan et al., 2003 21 51 33 self affective evaluation rate own arousal for aversive images dMPFC 4A
e Phan et al., 2003 3 45 39 self affective evaluation rate own arousal for aversive images dMPFC 4A
e Phan et al., 2003 3 51 27 self affective evaluation rate own arousal for aversive images dMPFC 4A
e Phan et al., 2003 3 39 30 self affective evaluation rate own arousal for aversive images dMPFC 4A
f Porro et al., 1998 7 57 0 self affective evaluation correlation with self-rated intensity of painful
stimulus
vMPFC 4A
g Simpson et al., 2001 1 41 8 self affective evaluation decreases relative to fixation predict preshock
anxiety
vMPFC 4A
g Simpson et al., 2001 1178 self affective evaluation decreases relative to fixation predict preshock
anxiety
vMPFC 4A
h Tabert et al., 2001 2 47 24 self affective evaluation judge most unpleasant of 3 negative words vs.
judge most neutral of 3 neutral words
dMPFC 4A
h Tabert et al., 2001 7212 self affective evaluation judge most unpleasant of 3 negative words vs.
judge most neutral of 3 neutral words
vMPFC 4A
h Tabert et al., 2001 13 55 24 self affective evaluation judge most unpleasant of 3 negative words
vs. judge most neutral of 3 neutral words
dMPFC 4A
i Taylor et al., 2003 1 26 29 self affective evaluation rate arousal to aversive vs. non aversive photos ACC 4A
i Taylor et al., 2003 1 26 29 self affective evaluation rate arousal to aversive vs. non aversive photos ACC 4A
j Zysett et al., 2002 6 55 13 self affective evaluation judge preferences vs. retrieve semantic
information
dMPFC 4A
k Zysett et al., 2003 5 49 16 self affective evaluation judge preferences vs. retrieve semantic
information
dMPFC 4A
k Zysett et al., 2003 25 45 28 self affective evaluation judge preferences vs. retrieve semantic
information
dMPFC 4A
k Zysett et al., 2003 11 42 2 self affective evaluation judge preferences vs. retrieve semantic
information
vMPFC 4A
k Zysett et al., 2003 13 62 1 self affective evaluation judge preferences vs. retrieve semantic
information
vMPFC 4A
(continued on next page)
Ochsner et al. 11
Table 5. (continued )
Identifier Study x y z Target Judgment Type Contrast Region Figure
l Kampe et al., 2003 8 60 22 self recognize identity hearing own name vs. other name dMPFC 4A
l Kampe et al., 2003 0 20 58 self recognize identity hearing own name vs. other name dMPFC 4A
l Kampe et al., 2003 6 60 20 self recognize identity hear own name and perceive gaze direction
vs. non self stimuli
dMPFC 4A
m Kircher et al., 2000 0 6 37 self recognize identity view own face vs. unknown face ACC 4A
m Kircher et al., 2000 3 36 4 self recognize identity view own face vs. unknown face ACC 4A
m Kircher et al., 2000 6 42 2 self recognize identity view own face vs. unknown face ACC 4A
n Nakamura et al., 2001 14 54 12 self recognize identity self voice recognition vs. vowel recognition dMPFC 4A
o Sugiura et al., 2000 7 44 0 self recognize identity passive viewing of own face ACC 4A
o Sugiura et al., 2000 7 32 25 self recognize identity recognition judgments of own face ACC 4A
o Sugiura et al., 2000 7 31 26 self recognize identity recognition judgments of own face vs. passive
view of own face
ACC 4A
o Sugiura et al., 2000 25 36 16 self recognize identity recognition judgments of own face vs. passive
view of own face
ACC 4A
p Greene et al., 2001 1 52 17 self perspective-taking judge morality for personal vs. nonpersonal
moral dilemmas
dMPFC 4A
q Ruby & Decety, 2003 24 50 6 self perspective-taking 1st person vs. 3rd person conceptual
perspective taking
vMPFC 4A
q Ruby & Decety, 2003 46812 self perspective-taking 1st person vs. 3rd person conceptual perspective
taking
dMPFC 4A
r Vogeley et al., 2001 6 54 4 self perspective-taking judging own intentions for imagined actions ACC 4A
r Vogeley et al., 2001 12 50 4 self perspective-taking judging own intentions for imagined actions ACC 4A
s Craik et al., 1999 6 56 8 self self-descriptiveness judge self relevance of words dMPFC 4A
s Craik et al., 1999 6 40 28 self self-descriptiveness judge self relevance of words dMPFC 4A
t Fossati & Hevenor, 2003 16 40 27 self self-descriptiveness self referential judgment vs. letter recognition
control
dMPFC 4A
t Fossati & Hevenor, 2003 10 49 16 self self-descriptiveness self referential judgment vs. letter recognition
control
dMPFC 4A
u Johnson et al., 2002 0 54 8 self self-descriptiveness yes/no to self reflective vs. semantic questions dMPFC 4A
v Kelley et al., 2002 10 52 2 self self-descriptiveness self-relevant judgments vs. other relevant
judgments
dMPFC 4A
12 Journal of Cognitive Neuroscience Volume 16, Number 10
w Kircher et al., 2002 12 22 31 self self-descriptiveness self descriptive vs. non self-descriptive
judgments
ACC 4A
x Lieberman et al., in press 45812 self self-descriptiveness self descriptiveness Js for high experience vs.
low experience domains
vMPFC 4A
x Lieberman et al., in press 10 6 54 self self-descriptiveness self descriptiveness Js for high vs. low experience
domains P’s nonschematic for trait
dMPFC 4A
x Lieberman et al., in press 12 52 32 self self-descriptiveness self descriptiveness Js for high vs. low experience
domains P’s nonschematic for trait
dMPFC 4A
x Lieberman et al., in press 20 52 10 self self-descriptiveness self descriptiveness Js for low vs. high experience
domains P’s nonschematic for trait
vMPFC 4A
x Lieberman et al., in press 22 30 16 self self-descriptiveness self descriptiveness Js for low vs. high experience
domains P’s nonschematic for trait
vMPFC 4A
x Lieberman et al., in press 14 30 48 self self-descriptiveness self descriptiveness Js for high vs. low experience
domains P’s schematic for trait
dMPFC 4A
x Lieberman et al., in press 65410 self self-descriptiveness self descriptiveness Js for high vs. low experience
domains (P’s schematic for trait)
vMPFC 4A
y Macrae et al., 2004 24 58 1 self self-descriptiveness predicts subsequent memory after self-relevance
judgment
dMPFC 4A
y Macrae et al., 2004 0 50 8 self self-descriptiveness predicts subsequent memory after self-relevance
judgment
dMPFC 4A
y Macrae et al., 2004 9 50 8 self self-descriptiveness judge self relevance vs. perceptual baseline dMPFC 4A
aa Cato et al., 2004 22 42 31 self self-generation generate vs. repeat negative as opposed to
neutral words
dMPFC 4A
aa Cato et al., 2004 3 60 12 self self-generation generate vs. repeat positive as opposed to
neutral words
dMPFC 4A
aa Cato et al., 2004 4 60 29 self self-generation negative vs. neutral word generation dMPFC 4A
bb Crosson et al., 1999 7 60 28 self self-generation self generate vs. repeat emotion words dMPFC 4A
bb Crosson et al., 1999 4 17 47 self self-generation self generate vs. repeat emotion words ACC 4A
bb Crosson et al., 1999 7 22 43 self self-generation self generate vs. repeat neutral words ACC 4A
cc Ochsner et al., in press 10 18 62 self self-generation decrease negative affect vs. reappraisal vs. look
at negative image
dMPFC 4A
cc Ochsner et al., in press 16 46 42 self self-generation decrease negative affect vs. reappraisal vs. look
at negative image
dMPFC 4A
(continued on next page)
Ochsner et al. 13
Table 5. (continued )
Identifier Study x y z Target Judgment Type Contrast Region Figure
cc Ochsner et al., in press 8 46 48 self self-generation decrease negative affect vs. reappraisal vs. look
at negative image
dMPFC 4A
cc Ochsner et al., in press 10 .50 34 self self-generation increase vs. decrease negative emotion via
reappraisal
dMPFC 4A
cc Ochsner et al., in press 4 64 32 self self-generation increase vs. decrease negative emotion via
reappraisal
dMPFC 4A
cc Ochsner et al., in press 4 68 24 self self-generation increase vs. decrease negative emotion via
reappraisal
dMPFC 4A
cc Ochsner et al., in press 10 2 66 self self-generation increase negative affect vs. reappraisal vs. look at
negative image
ACC 4A
cc Ochsner et al., in press 6 48 40 self self-generation increase negative affect vs. reappraisal vs. look at
negative image
dMPFC 4A
cc Ochsner et al., in press 18 10 44 self self-generation increase negative affect vs. reappraisal vs. look at
negative image
dMPFC 4A
dd Partiot et al., 1995 12 38 36 self self-generation imagine events/feelings during preparation for
moms funeral
dMPFC 4A
dd Partiot et al., 1995 2384 self self-generation imagine events/feelings during preparation for
moms funeral
vMPFC 4A
dd Partiot et al., 1995 18 42 32 self self-generation imagine events/feelings during preparation for
moms funeral
dMPFC 4A
dd Partiot et al., 1995 10 46 24 self self-generation imagine events/feelings during preparation for
moms funeral
dMPFC 4A
ee Pietrini et al., 2000 2 6 44 self self-generation increase for imagined aggressive vs. imagined
neutral behavior
ACC 4A
ee Pietrini et al., 2000 43212 self self-generation decrease for imagined aggressive vs. imagined
neutral behavior
vMPFC 4A
ee Pietrini et al., 2000 2 60 12 self self-generation decrease for imagined aggressive vs. imagined
neutral behavior
dMPFC 4A
ee Pietrini et al., 2000 16 60 20 self self-generation decrease for imagined aggressive vs. imagined
neutral behavior
dMPFC 4A
ee Pietrini et al., 2000 4 58 8 self self-generation decrease for imagined restrained aggression
vs. imagined neutral behavior
dMPFC 4A
ff McGuire et al., 1996 8 38 24 self self-generation correlates with self generated stimulus
independent thoughts
dMPFC 4A
14 Journal of Cognitive Neuroscience Volume 16, Number 10
ff McGuire et al., 1996 4 28 36 self self-generation correlates with self generated stimulus
independent thoughts
dMPFC 4A
ff McGuire et al., 1996 10 48 0 self self-generation correlates with self generated stimulus
independent thoughts
dMPFC 4A
ff McGuire et al., 1996 4 44 8 self self-generation correlates with self generated stimulus
independent thoughts
dMPFC 4A
1 Berthoz et al., 2002 6 14 60 social good/bad evaluation reaction to transgression of norms vs. normal
stories
dMPFC 4B
1 Berthoz et al., 2002 6 36 54 social good/bad evaluation reaction to transgression of norms vs. normal
stories
dMPFC 4B
1 Berthoz et al., 2002 6 32 54 social good/bad evaluation reaction to transgression of norms vs. normal
stories
dMPFC 4B
1 Berthoz et al., 2002 2 36 52 social good/bad evaluation reaction to embarrassing stories vs. normal
stories
dMPFC 4B
1 Berthoz et al., 2002 8 52 18 social good/bad evaluation reaction to transgression of norms vs. normal
stories
dMPFC 4B
1 Berthoz et al., 2002 10 54 24 social good/bad evaluation reaction to transgression of norms vs. normal
stories
dMPFC 4B
1 Berthoz et al., 2002 4 54 36 social good/bad evaluation reaction to embarrassing stories vs. normal
stories
dMPFC 4B
1 Berthoz et al., 2002 12 58 8 social good/bad evaluation reaction to transgression of norms vs. normal
stories
dMPFC 4B
2 Cunningham et al., 2003 12 40 20 social good/bad evaluation good/bad vs. relative age judgment of target
person
ACC 4B
3 Farrow et al., 2001 2 49 19 social good/bad evaluation judge other’s emotions vs. general
knowledge inference
vMPFC 4B
3 Farrow et al., 2001 14 60 26 social good/bad evaluation judge other’s emotions vs. general knowledge
inference
dMPFC 4B
3 Farrow et al., 2001 12 56 34 social good/bad evaluation judge forgivability of crime vs. general knowledge
inference
dMPFC 4B
3 Farrow et al., 2001 4 65 17 social good/bad evaluation judge forgivability of crime vs. judge other’s
emotions
dMPFC 4B
3 Farrow et al., 2001 14 59 31 social good/bad evaluation judge forgivability of crime vs. judge other’s
emotions
dMPFC 4B
(continued on next page)
Ochsner et al. 15
Table 5. (continued )
Identifier Study x y z Target Judgment Type Contrast Region Figure
3 Farrow et al., 2001 2 43 44 social good/bad evaluation judge forgivability of crime vs. judge other’s
emotions
dMPFC 4B
4 Teasdale et al., 1999 3 3 42 social good/bad evaluation understand and interpret negative captions for
photos
ACC 4B
4 Teasdale et al., 1999 6 39 20 social good/bad evaluation understand and interpret negative captions for
photos
ACC 4B
4 Teasdale et al., 1999 3 31 28 social good/bad evaluation understand and interpret negative captions for
photos
ACC 4B
4 Teasdale et al., 1999 3 3 42 social good/bad evaluation understand and interpret positive vs. negative
captions for photos
ACC 4B
4 Teasdale et al., 1999 0 42 15 social good/bad evaluation understand and interpret positive vs. negative
captions for photos
ACC 4B
5 Wicker et al., 2003 4 45 34 social good/bad evaluation judge intentions from clips of moving gaze
followed by emotionally expressive eyes
dMPFC 4B
5 Wicker et al., 2003 1 51 18 social good/bad evaluation judge intentions from clips of moving gaze
followed by emotionally expressive eyes
dMPFC 4B
5 Wicker et al., 2003 3 27 37 social good/bad evaluation judge intentions from clips of moving gaze
followed by emotionally expressive eyes
ACC 4B
5 Wicker et al., 2003 1 37 10 social good/bad evaluation judge intentions from clips of moving gaze
followed by emotionally expressive eyes
ACC 4B
5 Wicker et al., 2003 1 34 18 social good/bad evaluation judge intentions from clips of moving gaze
followed by emotionally expressive eyes
vMPFC 4B
6 Winston et al., 2003 16 42 8 social good/bad evaluation judge trustworthiness vs. gender of face vMPFC 4B
7 Calder et al., 2002 2 44 36 social gaze perception view increasing proportion of faces with averted
gaze
dMPFC 4B
7 Calder et al., 2002 26 44 8 social gaze perception view increasing proportion of faces with averted
gaze
vMPFC 4B
8 Hooker et al., 2003 4 23 44 social gaze perception judge gaze direction vs. arrow direction dMPFC 4B
9 Kampe et al., 2003 8 50 14 social gaze perception view faces with directed vs. averted gaze dMPFC 4B
10 Platek et al., 2003 5 34 57 social gaze perception mind in eyes task: judge, ‘‘what are those eyes
thinking?’’
dMPFC 4B
11 Goel et al., 1995 12 38 32 social mental state inference judging another person’s knowledge of objects dMPFC 4B
11 Goel et al., 1995 6 46 28 social mental state inference judging another person’s knowledge of objects dMPFC 4B
16 Journal of Cognitive Neuroscience Volume 16, Number 10
12 Heekeren et al., 2003 8 45 22 social mental state inference moral vs. semantic judgment vMPFC 4B
12 Heekeren et al., 2003 1 55 2 social mental state inference moral vs. semantic judgment vMPFC 4B
12 Heekeren et al., 2003 6 61 27 social mental state inference moral vs. semantic judgment vMPFC 4B
13 Iacoboni et al., 2004 2 52 26 social mental state inference viewing social, interactive vs. solitary figure films dMPFC 4B
14 Mitchell et al., 2002 0 54 21 social mental state inference judge applicability of terms for describing people
vs. objects
dMPFC 4B
14 Mitchell et al., 2002 3 39 0 social mental state inference judge applicability of terms for describing people
vs. objects
vMPFC 4B
14 Mitchell et al., 2002 12 36 0 social mental state inference judge applicability of terms for describing people
vs. objects
dMPFC 4B
15 Mitchell et al., 2004 12 51 36 social mental state inference form impression of pictured person vs. judge
sequence of photo presentations
dMPFC 4B
15 Mitchell et al., 2004 6 48 48 social mental state inference form impression of pictured person vs. judge
sequence of photo presentations
dMPFC 4B
15 Mitchell et al., 2004 6 51 39 social mental state inference form impression of pictured person vs. judge
sequence of photo presentations
dMPFC 4B
15 Mitchell et al., 2004 9 33 57 social mental state inference form impression of pictured person vs.
judge sequence of photo presentations
dMPFC 4B
15 Mitchell et al., 2004 0 45 36 social mental state inference form impression of pictured person vs. judge
sequence of photo presentations
dMPFC 4B
15 Mitchell et al., 2004 6 57 33 social mental state inference form impression of pictured person vs. judge
sequence of photo presentations
dMPFC 4B
15 Mitchell et al., 2004 12 36 57 social mental state inference form impression of pictured person vs. judge
sequence of photo presentations
dMPFC 4B
15 Mitchell et al., 2004 9 57 27 social mental state inference form impression of pictured person vs. judge
sequence of photo presentations
dMPFC 4B
15 Mitchell et al., 2004 6 51 45 social mental state inference form impression of pictured person vs. judge
sequence of photo presentations
dMPFC 4B
15 Mitchell et al., 2004 0 39 51 social mental state inference form impression of pictured person vs. judge
sequence of photo presentations
dMPFC 4B
15 Mitchell et al., 2004 9 63 21 social mental state inference form impression of pictured person vs. judge
sequence of photo presentations
dMPFC 4B
15 Mitchell et al., 2004 12 21 60 social mental state inference form impression of pictured person vs. judge
sequence of photo presentations
dMPFC 4B
(continued on next page)
Ochsner et al. 17
Table 5. (continued )
Identifier Study x y z Target Judgment Type Contrast Region Figure
15 Mitchell et al., 2004 15 24 57 social mental state inference form impression of pictured person vs. judge
sequence of photo presentations
dMPFC 4B
16 Wood et al., 2003 16 30 50 social mental state inference categorize social phrases dMPFC 4B
16 Wood et al., 2003 12 41 42 social mental state inference categorize social phrases dMPFC 4B
16 Wood et al., 2003 8 48 31 social mental state inference categorize social phrases dMPFC 4B
16 Wood et al., 2003 12 34 50 social mental state inference categorize social words dMPFC 4B
16 Wood et al., 2003 4 45 38 social mental state inference categorize social words dMPFC 4B
17 Brunet et al., 2000 16 44 20 social view moving shapes view clips that evoke intentional vs. physical
causality inferences
dMPFC 4B
17 Brunet et al., 2000 8 32 4 social view moving shapes view clips that evoke intentional vs. physical
causality inferences
vMPFC 4B
17 Brunet et al., 2000 8 36 0 social view moving shapes view clips that evoke intentional vs. physical
causality inferences
vMPFC 4B
17 Brunet et al., 2000 4 4 38 social view moving shapes view clips that evoke intentional vs. physical
causality inferences
vMPFC 4B
17 Brunet et al., 2000 8 34 2 social view moving shapes view clips that evoke intentional vs. physical
causality inferences
ACC 4B
17 Brunet et al., 2000 8 36 0 social view moving shapes view clips that evoke intentional vs. physical
causality inferences
vMPFC 4B
17 Brunet et al., 2000 4 56 44 social view moving shapes view clips that evoke intentional vs. physical
causality inferences
dMPFC 4B
17 Brunet et al., 2000 22 38 20 social view moving shapes view physical causality clips with characters
vs. without
vMPFC 4B
18 Castelli et al., 2000 4 60 32 social view moving shapes observe complex intentional vs. nonintentional
control movements
dMPFC 4B
18 Castelli et al., 2000 6 58 32 social view moving shapes observe complex intentional vs. nonintentional
control movements
dMPFC 4B
19 Martin & Weisberg, 2003 3 52 11 social view moving shapes view clips of social vs. mechanical movements vMPFC 4B
20 Ruby & Decety, 2001 14 72 10 social perspective-taking 3rd vs. 1st person spatial perspective taking dMPFC 4B
20 Ruby & Decety, 2001 28 50 8 social perspective-taking 3rd vs. 1st person spatial perspective taking vMPFC 4B
21 Ruby & Decety, 2003 0 20 70 social perspective-taking 3rd vs. 1st person conceptual perspective taking dMPFC 4B
18 Journal of Cognitive Neuroscience Volume 16, Number 10
21 Ruby & Decety, 2003 10 24 56 social perspective-taking 3rd vs. 1st person conceptual perspective taking dMPFC 4B
21 Ruby & Decety, 2003 8 40 52 social perspective-taking 3rd vs. 1st person conceptual perspective taking dMPFC 4B
21 Ruby & Decety, 2003 24 48 42 social perspective-taking 3rd vs. 1st person conceptual perspective taking dMPFC 4B
22 Baron-Cohen et al., 1999 9 50 20 social TOM infer state of mind vs. identify gender dMPFC 4B
22 Baron-Cohen et al., 1999 6 6 53 social TOM infer state of mind vs. identify gender dMPFC 4B
22 Baron-Cohen et al., 1999 0 47 9 social TOM infer state of mind vs. identify gender dMPFC 4B
23 Fletcher et al., 1995 0 38 24 social TOM read theory of mind vs. physical stories ACC 4B
23 Fletcher et al., 1995 12 42 40 social TOM read theory of mind stories vs. unlinked
sentences
dMPFC 4B
23 Fletcher et al., 1995 12 36 36 social TOM read theory of mind stories vs. unlinked
sentences
dMPFC 4B
24 Gallagher et al., 2000 8 50 10 social TOM read TOM vs. non TOM stories dMPFC 4B
24 Gallagher et al., 2000 10 48 12 social TOM view TOM vs. non TOM stories & cartoons dMPFC 4B
25 Happe et al., 1996 10 44 16 social TOM read TOM vs. non TOM stories dMPFC 4B
26 Saxe & Kanwisher, 2003 6 57 18 social TOM read false belief vs. false photograph stories dMPFC 4B
27 Vogeley et al., 2001 6 56 2 social TOM judge intentions of others vs. self described
in vignettes
ACC 4B
27 Vogeley et al., 2001 4 28 30 social TOM judge intentions of others vs. self described
in vignettes
ACC 4B
28 Gallagher et al., 2002 8 54 12 social interactive rock paper scissors game vs. mentalizing human
as opposed to rule-following computer
ACC 4B
28 Gallagher et al., 2002 10 50 30 social interactive rock paper scissors game vs. mentalizing human
as opposed to rule-following computer
dMPFC 4B
28 Gallagher et al., 2002 2 46 14 social interactive rock paper scissors game vs. mentalizing human
as opposed to random computer
dMPFC 4B
29 Rilling et al., 2002 3 51 6 social interactive prisoner’s dilemma vs. human; mutual
cooperation vs. all other dyadic choices
ACC 4B
29 Rilling et al., 2002 3 48 12 social interactive prisoner’s dilemma vs. human; mutual
cooperation vs. all other dyadic choices
vMPFC 4B
29 Rilling et al., 2002 6 51 18 social interactive prisoner’s dilemma vs. human; mutual
coop/mutual defect vs. all other dyadic choices
vMPFC 4B
(continued on next page)
Ochsner et al. 19
and social-cognitive processes not specifically associa ted
with emotion per se. A within-study comparison of
emotional and nonemotional self- and other-cued judg-
ments would be necessary to derive firm conclusions in
this regard.
Finally, a third important question concerns the func-
tional importance attached to the relative ubiquity of
MPFC activation across a variety of self-referential and
social-cognitive judgments (see Table 5 and Figure 4),
including conditions in which participants are allowed to
simply ‘‘rest’’ while inside the s canner and are not
explicitly directed to engage in either type of processing
(Gusnard & Raichle, 2001; Gusnard et al., 2001;
McGuire, Paulesu, Frackowiak, & Friat, 1996). Compar-
atively greater MPFC activation has been found during
rest when participants are free to a think about whatever
they wish as compared to tasks that require specific
forms of executive control (Gusnard & Raichle, 2001),
self-referential (e.g., Kelley et al., 2002), or social cogni-
tive (e.g., Mitchell, Macrae, Schooler, Rowe, & Milne,
2002) processing. This pattern of heightened MPFC
activation and metabolic stability at rest has been taken
as support for the hypothesis that the MPFC carries out
a default- state monitor ing function which involves mon-
itoring of the internal milieu and represents a physio-
logic baseline for brain imaging studies (Gusnard &
Raichle, 2001). This hypothesis generally is consistent
with the available data, and intriguingl y suggests that
self-referential and/or social-cognitive processing may be
a natural part of stream of consciousness thought
(Mitchell, Macrae, Schooler, et al., 2002; Christoff, Ream,
& Gabrieli, in press). Because the present study did not
include a resting condition, relative activations or de-
activations relative to a putative default state unfortu-
nately cannot be evaluated. It should be noted,
however, that comparisons with a resting baseline state
would only have clear psychological meaning with re-
spect to whatever psychological processes are engaged
during that state. Because resting baselines do not
involve a directed task, the precise nature of psycho-
logical processes may v ary. This means that differ-
ences in activation with respect to that state may not
be clearly interpretable in terms of specific underlying
psychological processes (cf. Stark & Squire, 2001), and
could explain why deactivations for self-ref erential or
social-cognitive processing are not always found in com-
parison to the resting state (e.g., Iacoboni, Lieberman,
et al., 2004; Mitchell, Macrae, & Banaji, 2004; Zysset,
Huber, Ferstl, & von Cramon, 2002).
METHODS
Participants
Thirteen participants (7 women, M age = 29.5 years)
were recruited in compliance with the human subjects
regulations of Stanford University.
Behavioral Paradigm
Participants viewed three types of mixed blocks of
positive, negative, and neutral photos selected from
the International Affective Picture System (Lang, Green-
wald, Bradley, & Hamm, 1993). Prior to the onset of
each block, one of three instructional cues was pre-
sented in the center of the screen for 4 sec. On ‘‘self’’
blocks, participants were instructed to judge whether
they felt pleasant, unpleasant, or neutral in response to
each photo. On ‘‘other’’ blocks, p articipants judged
whether the central figure for each photo felt pleasant,
unpleasant, or neutral. On ‘‘in–out’’ blocks, participants
judged whether each photo had been taken inside,
outside, or whether it could not be determined (i.e.,
they were unsure) in which location the photo had been
taken. Each block was comprised of a series of six trials.
On each trial, a photo was presented for 2 sec followed
by a 3-point rating scale for 1.5 sec and a 500-msec
intertrial interval. The rating scale displayed appropriate
response options for self, other, and in–out blocks, and
served to guide judgments that participants made using
three fingers of their right hand on a four-button
response box. To insure th at participants would be
experiencing pleasant or unpleasant affect throughout
the task, each block contai ned either two pleasant and
three unpleasant or three pleasant and two unpleasant
Note: Articles are organized alphabetically by judgment target and judgment type. Column labels indicate: Identifier = identifier for activation in
Figure 1; Study = study listed in references; x, y, z = coordinates given in published study, with +x = right and x = left, +y = rostral/anterior and
y = posterior, +z = dorsal and z = ventral relative to origin; Target = judgment target, ‘‘self’’ in case of self-referential judgments, ‘‘social’’ in
case of social-cognitive judgments involving inferences about the mental or emotional states of another person; Contrast = comparison producing
activation focus; Region = region of the medial frontal cortex activated, with dMPFC indicating dorsal medial prefrontal cortex with z coordinate >
0 and falling within BAs 8, 9, 10, or dorsal portions of 32, vMPFC indicating ventromedial prefrontal cortex with z < 0 and falling within BAs 10, 11,
14, 25, and ventral portions of 32, and ACC indicating the anterior cingulate cortex, which includes BAs 24 and 32 (Ongur & Price, 2003; Ongur et al.,
2000). Judgment types are as follows: affective evaluation = assessing valence or arousal of own emotional response, or judging personal preference
for a stimulus; recognize identity = view one’s own face or hear one’s own name; perspective taking = imagining first- or third-person spatial (what
do I/they see?) versus conceptual (what do I/they think/feel/believe?) perspective; self-generation = generate words/thoughts; mental state
inference = judge states or traits of depicted individuals; view moving shapes = view video clips of abstract moving shapes whose motions tend to
elicit either attributions of intentionality, or require some other type of nonmental inference to describe/understand; TOM = view and/or judge
vignettes explicitly designed to require theory of mind inferences in order to comprehend them; interactive = participants believe they are playing a
game in real time with another participant. Abbreviations: sq. = square, Js = judgments, TOM = theory of mind judgments, coop = cooperation,
pleas = pleasant, P’s = participants.
20 Journal of Cognitive Neuroscience Volume 16, Number 10
images, and one neutral image as determined by nor-
mative ratings (Lang et al., 1993). Valence and arousal
ratings for pleasant, unpleasant, and neutral images
were equated across three stimulus sets that were
counterbalanced across judgmen t types. Valenced and
neutral images were randomly intermixed within blocks,
and six blocks of each instruction type were presented in
pseudorandom order within a single 8 min 24 sec scan.
Stimulus presentation and response collection were
controlled by the program Psyscope 1.2.5 running on a
Macintosh G3 Computer. Stimuli were back projected
onto a screen mounted on a custom head coil that
limited head motion using a bitebar.
MRI Data Acquisition
Whole-brain fMRI data (32 axial slices, 3.5 mm thick)
were collected at 3T (GE Signa LX Horiz on Echos peed
scanner) with a T2*-sensitive gradient-echo spiral-out
pulse sequence (30 msec TE, 2000 msec TR, 2 inter-
Figure 4. Medial prefrontal activation coordinates for studies involving self-referential or social cognitive judgments shown in Table 5. (A) Left
and right medial views of activations across a series of studies related to judgments of self-reference. Identifier letters correspond to specific
activation coordinates for studies listed in the top half of Table 5. Blue circles and yellow squares indicate activations from the present study related
to self appraisals of emotion and both self and other appraisals of emotion, respectively. (B) Left and right medial views of activations across a
series of studies related to social cognitive judgements. Identifier numbers correspond to specific activation coordinates for studies listed in
the bottom half of Table 5. Blue circles and yellow squares indicate activations from the present study related to other appraisals of emotion
and both self and other appraisals of emotion, respectively. (C) Figure key indicating relative locations of Brodmann’s areas located in the
medial frontal cortex (Ongur & Price, 2003; Ongur et al, 2000). For descriptive purposes, dorsal medial prefrontal (MPFC) regions have
a z-coordinate > 0 and fall within BAs 6, 8, 9, or 10, ventromedial prefrontal regions (vMPFC ) have z < 0 and fall within BAs 10, 11, 14, 25,
and ventral portions of 32, and ACC indicating the anterior cingulate cortex, which includes BAs 24 and 32. For descriptive information
pertaining to specific studies, see Table 5.
FPO
Ochsner et al. 21
leaves, 60 8 flip angle, 24 cm field of view , 64 by 64 data
acquisition matrix). T2-weighted flow-compensated
spin-echo scans were acquired for anatomical reference
using the same slice prescription (2000 msec TR, 85 msec
TE), and high-order shimming was performed before
functional scans (Glover, 1999).
MRI Data Analysis
Functional images were slice time and motion-corrected
using SPM99 (Wellcome Department of Cognitive Neu-
rology). Anatomical images next were coregistered to
the mean func tional image, and normalized to a stan-
dard template brain. Functional images were then nor-
malized using those parameters, interpolated to 2 2
2 mm voxels, and smoothed with a gaussian filter (6 mm
full width half maximum). To remove drifts within
sessions, a high-pass filter with a cutoff period two times
the block length was applied.
A mixed design was used to model first-level fixed-
effects for each participant. The 4-sec instructional cue
proceeding each block was modeled with a canonical
hemodynamic response function; the 24-sec photo
blocks were modeled as a boxcar regressor convolved
with the canonical hemodynamic response. A general
linear model analysis was used in SPM99 to create
contrast images for each participant summarizing differ-
ences between block types. These images were used to
create second-level group average SPM{t} maps that
were thresholded at p < .005, uncorrected for multiple
comparisons, with an extent threshold of 20 voxels.
These parameters correspond to an overall alpha level
of p < .05, corrected for multiple comparisons a s
calculated by the Monte Carlo simulation method of
Forman et al. (1995) implemented in AFNI, and has been
employed in numero us prior studies (e.g., Wood, Ro-
mero, Makale, & Grafman, 2003; Konishi, Nakajima,
Uchida, Kikyo, et al., 1999; Poldrack et al., 1999; Wagner,
1999; Konishi, Nakajima, Uchida, Sekih ara, & Miyashita,
1998). Maxima are reported in ICBM152 coordinates as
in SPM99. To formally identify regions active for both the
self > in–out and other > in–out contrasts, the t-map
for the first contrast was used as an inclusive mask for
the second contrast, with each voxel-level thresholded
at p < .01. Using the Fisher method for combining
p values, this analysis yields regions active with prob-
ability p < .001 across both tasks. For functionally
defined regions shown at the group level to be in-
volved in attributing emotion to self, to other, or to
both, measures of mean percent signal change for a
given type of instruction block (relative to the mean
activation of that region across the entire study) were
extracted from the peak activated voxel, that is, appar-
ent deactivations relative to the zero line are not
deactivations per se, but simply reflect lesser activation
with respect to the mean level of activation in that
given region of interest
8
.
UNCITED REFERENCES
Kelley & Berridge, 2002
Kircher et al., 2001
Acknowledgments
We thank Elaine Robertson and Hedy Kober for assistance in
preparation of the manuscript, Tor Wager for assistance with
Figure 4, and we acknowledge support by a grant from the
John and Dodie Rosekranz Endowment (SCM), grant BCS-
93679 from the National Science F oundation (KNO), a
NARSAD Young Investigator Grant (KNO), and grant RR
09784 from the NIH (GHG).
Reprint requests should be sent to Kevin Ochsner, Department
of Psychology, Columbia University, 369 Schermerhorn Hall,
1190 Amsterdam Avenue, New York, NY 10027, or via e-mail:
ochsner@psych.columbia.edu.
Notes
1. Although not of primary interest here, it is worth noting
that neither the contrast of in–out > self nor in–out > other
blocks showed activation in the medial or lateral prefrontal
regions. The in–out > self contrast showed bilateral acti-
vation of the lateral superior parietal cortex and the cuneus
consistent with the spatial nature of the task (Culham &
Kanwisher, 2001), as well as activation of the primary visual
cortex, consistent with the findings reported in Figure 3 and
Table 4, indicating relatively lesser engagement of early visual
processing during the internally queued self blocks. The in–
out > other contrast showed activation of the left parietal
cortex and the cuneus, also consistent with a greater reliance
on visual–spatial processing in the in–out cued blocks.
Notably, greater activation of the medial prefrontal cortex
was not found during the in–out cued blocks (even when
thresholds were lowered to .1), which suggests that at least
under the conditions of the present experiment, medial
prefrontal recruitment reflects greater activation with respect
to our perceptual baseline, although as not ed i n the
discussion, the design of the present study does not permit
comparison of activations in the self and other blocks to
resting baseline because such a baseline was not included in
the present study.
2. It is important to note that alternative accounts of the
present findings in terms of differential response difficulty or
arousal are unlikely both for empirical and theoretical reasons.
The similar speed and frequency with which self and other
judgments were made makes it u nlikely that differences
between the brain regions recruited by each judgment type
arise either from differential difficulty in making them, or from
differential arousal (which would be expected to motivate
more rapid responding in one condition as compared to the
other). Similarly, although it could be argued that greater
activation for either self or other judgments reflects greater
arousal in one condition, this seems unlikely because
activations of arousal-related limbic structures have been
found to decrease when participants rate emotional responses
to the same sorts of aversive photos used here (Taylor, Phan,
Decker, & Liberzon, 2003).
3. Additional differences between self and other judgments
were revealed in contrasts of each judgment with their
common perceptual baseline. These differences may not be
as large or as reliable as those revealed by directly contrasting
self and other judgments to one another, as described above,
22 Journal of Cognitive Neuroscience Volume 16, Number 10
but may nevertheless provide information about the differ-
ential recruitment of systems that may be involved in attention
to and encoding of internal as opposed to external cues
relevant to emotional attribut ions. Fo r self j udgments,
activation was observed in dorsal anterior cingulate regions
associated with attention to and monitoring of internal
responses (Botvinick, Braver, Barch, Carter, & Cohen, 2001)
including self-generated changes in emotion (Ochsner, Ray,
et al., in press; Ochsner, Bunge, Gross, & Gabrieli, 2002) as
well as inferior parietal regions involved in selective attention
and representing the body in space (Culham & Kanwisher,
2001). For other judgments, activation was observed in a
parahippocampal region implicated in encoding visual and
spatial cues that designate specific places (Epstein & Kan-
wisher, 1998).
4. Additional differences between self and other emotion
perception concerned the late rality of activated regions.
Whereas self judgments recruited primarily the left frontal,
temporal, parietal, and cingulate regions, other judgments
recruited the left frontal and cingulate regions in combination
with the right temporal and p arahippocampal regions.
Although the reasons for these laterality findings are not
immediately clear, they might relate to preferential recruit-
ment of left lateralized systems for either self-focused
processing (Kelley et al., 2002; Turk et al., 2002) or mentally
manipulating verbal information (Smith, Jonides, Marshuetz, &
Koeppe, 1998) as may occur when reasoning about mental
states, and to right lateralized systems specialized for encoding
visual spatial and nonverbal cues (Kosslyn & Koenig, 1992)
recruited when directing attention outwards to judge the
emotions experienced by pictured persons.
5. It is possible that when processing visual cues necessary to
draw inferences about the internal emotional states of others’
results in an interaction between bottom-up and top-down
processing that enhances attention, and visual cortical
activation, in a way that does not occur when one is making
a simple perceptual judgment about the visual properties of
the stimulus during the in–out blocks. This pattern of
activation was not expected a priory, however, and awaits
replication in future studies.
6. In the linguistic coherence study of Ferstl and von Cramon
(2002), for example, participants may have spontaneously
experienced self-reflective mental states when determining
whether sentences like, ‘‘Sometimes a truck drives by the
house,’’ cohere with sentences like, ‘‘That’s when the dishes
start to rattle.’’ Hearing these sentences may evoke mental
images in which they watch the dishes rattle in a kitchen while
another individual drives a truck outside, which participants
use to draw inferences about what they would believe or infer
when experiencing this event (cf. Frith & Frith, 1999).
7. It should be noted, however, that within-study compar-
isons of all potentially relevant conditions is plainly impractical,
and that meta-analytic procedures that statistically evaluate
similarity of activation foci across studies may be able to
identify distinct functional subregions within the MPFC (Kober,
Wager, & Ochsner, 2004). Another tack that can be taken
towards understanding MPFC function is to consider when it is
not recruited in the context of specific kinds of self versus
other processing. For example, activation of the MPFC has
been found when participants either imitated the gestures of
another person, or when they watched that person imitate
their gestures (Decety, Chaminade, Grezes, & Meltzoff, 2002),
and also when participants either reasoned about their own
behavioral intentions or those of a described individual
(Vogeley et al., 2001). These results join similar findings from
the present study in supporting a general role for the MPFC in
mental state attributions. However, those two studies both
found activation of parietal regions uniquely associated with
either self- and/or other-focused processing, which were not
observed in the present study. Although the reasons for these
discrepancies are not clear, one possibility is that the particular
systems important for distinguishing oneself and others will
vary depending upon the dimension of similarity in question.
Parietal systems have been implicated in the agentic control of
action (Farrer et al., 2003; Ramachandran, 1998), which may be
important during motor imitation and when drawing infer-
ences about the intentions of actors from their physical actions
described in vignettes. By contrast, emotional attributions (at
least in the present context) may depend more heavily on
decoding the meanings of interoceptive and exteroceptive
cues, as suggested by judgment-specific patterns of medial and
lateral prefrontal activation.
8. Two notes are important with respect to interpretation of
our measure of percent signal change. The first is that peak
voxel activations were selected rather then average cluster
activations because prior research has shown peak voxel
activity to be more strongly correlated with electrophysiolog-
ical measures of activation than is cluster average activity
(Arturs &amp; Boniface, 2003). Arturs and Boniface (2003)
examined the relationship between BOLD activity in the
somatosensory cortex and somatosensory evoked potentials
elicited by median nerve stimulation. They found that peak
voxel rather than cluster average measures of percent signal
change most strongly correlated with somatosensory evoked
potentials, suggesting that peak voxel activity might best
correlate with the neural generators of the BOLD response.
The second is that because percent signal change is calculated
with respect to the mean level of activity within a given region,
apparent deactivations with respect to the zero line do not in
fact reflect deactivations of the sort described by Gusnard et al.
(2001) with respect to a resting baseline. Rather, variations of
percent signal change here reflect deviations with respect to
an average level of activation, and are properly interpreted as
relative differences between conditions rather than activations
or deactivations with respect to a physiologic baseline
measure. This measure of computing percent signal change
against the mean of activity for a region across time and
across task conditions reflects a desire to compute a
‘‘descriptive’’ index of relative activation across task con-
ditions that is not influenced by factors that could impact the
global mean, and is not influenced by uncontrolled variability
in the psychological processes engaged by participants, as
would be the case if percent signal change were computed
with respect to a resting baseline that does not control
psychological processing.
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