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Neuropsychologia 81 (2016) 207–218
Contents lists available at ScienceDirect
Neuropsychologia
http://d0028-39
n CorrE-m1 Eq
journal homepage: www.elsevier.com/locate/neuropsychologia
Neural correlates of processing “self-conscious” vs. “basic”
emotions
Michael Gilead a,n,1, Maayan Katzir b,1, Tal Eyal b, Nira
Liberman a
a School of Psychological Sciences, Tel-Aviv University,
Ramat-Aviv, Tel-Aviv 69978, Israelb Department of Psychology,
Ben-Gurion University of the Negev, Beer-Sheva 84105, Israel
a r t i c l e i n f o
Article history:Received 17 June 2015Received in revised form9
December 2015Accepted 14 December 2015Available online 18 December
2015
Keywords:SelfEmotionmPFCdACCdlPFCSelf-controlPrideGuilt:
AngerJoy
x.doi.org/10.1016/j.neuropsychologia.2015.12.032/& 2015
Elsevier Ltd. All rights reserved.
esponding author.ail address: [email protected] (M.
Gilual contribution.
a b s t r a c t
Self-conscious emotions are prevalent in our daily lives and
play an important role in both normal andpathological behavior.
Despite their immense significance, the neural substrates that are
involved in theprocessing of such emotions are surprisingly
under-studied. In light of this, we conducted an fMRI studyin which
participants thought of various personal events which elicited
feelings of negative and positiveself-conscious (i.e., guilt,
pride) or basic (i.e., anger, joy) emotions. We performed a
conjunction analysisto investigate the neural correlates associated
with processing events that are related to self-consciousvs. basic
emotions, irrespective of valence. The results show that processing
self-conscious emotionsresulted in activation within frontal areas
associated with self-processing and self-control, namely, themPFC
extending to the dACC, and within the lateral-dorsal prefrontal
cortex. Processing basic emotionsresulted in activation throughout
relatively phylogenetically-ancient regions of the cortex, namely
invisual and tactile processing areas and in the insular cortex.
Furthermore, self-conscious emotions dif-ferentially activated the
mPFC such that the negative self-conscious emotion (guilt) was
associated with amore dorsal activation, and the positive
self-conscious emotion (pride) was associated with a moreventral
activation. We discuss how these results shed light on the nature
of mental representations andneural systems involved in
self-reflective and affective processing.
& 2015 Elsevier Ltd. All rights reserved.
1. Introduction
The self-conscious emotions acknowledged in
contemporarypsychological theories are guilt, shame, embarrassment,
and pride(Tangney, 2003; Tracy and Robins, 2004). These emotions
areprevalent in daily life (Hofmann et al., 2013), and are involved
invarious psychopathologies. For example, exaggerated feelings
ofguilt play a dominant role in depression (e.g., O’Connor et
al.,2002), shame is crucially involved in social-avoidance (e.g.,
Lutwakand Ferrari, 1997), and pride is a dominant emotion in
narcissism(Tracy et al., 2009). In light of this, furthering our
understanding ofthe psychological and biological mechanisms that
play a role in theprocessing of self-conscious emotions is an
immensely importantand clinically significant task.
Despite the involvement of self-conscious emotions in
pathologicaland normal behavior, only a handful of neuroimaging
studies havedirectly addressed this important topic. The goal of
the current studywas, accordingly, to investigate the neural
correlates of processingpositive and negative self-conscious vs.
non-self-conscious emotions,sometimes referred to as “basic
emotions” (Tracy and Robins, 2007).
09
ead).
How might the neural substrates that are involved in proces-sing
self-conscious emotions differ from those involved in pro-cessing
non-self-conscious emotions? To begin addressing thisquestion we
will turn to carefully examine the psychologicalprocesses that
distinguish basic and self-conscious emotions, andthen review
neural literature that addressed related questions.
1.1. Psychological differences between self-conscious and
basicemotions
Self-conscious emotions differ from non-self-conscious
(i.e.,basic) emotions in several respects. In what follows we
discuss twoof the main differences. The most straight-forward
distinctionbetween self-conscious and basic emotions pertains to
the objectthat is at the center of the emotional appraisal: for
self-consciousemotions, this object is the self, for basic emotions
it is not. Self-conscious emotions involve self-awareness,
self-evaluation, and aconsideration of how the self is being
evaluated by others, to agreater extent than basic emotions
(Baldwin and Baccus, 2004;Leary, 2007; Tracy and Robins, 2004;
Tangney, 2003). In lightof this, one might predict that processing
self-consciousemotions will activate neural substrates involved in
self-reflection(e.g., Kelley et al., 2002) and mentalizing (e.g.,
Mitchell et al.,2005; Denny et al., 2012), namely, the medial
Pre-Frontal Cortex(mPFC).
www.sciencedirect.com/science/journal/00283932www.elsevier.com/locate/neuropsychologiahttp://dx.doi.org/10.1016/j.neuropsychologia.2015.12.009http://dx.doi.org/10.1016/j.neuropsychologia.2015.12.009http://dx.doi.org/10.1016/j.neuropsychologia.2015.12.009http://crossmark.crossref.org/dialog/?doi=10.1016/j.neuropsychologia.2015.12.009&domain=pdfhttp://crossmark.crossref.org/dialog/?doi=10.1016/j.neuropsychologia.2015.12.009&domain=pdfhttp://crossmark.crossref.org/dialog/?doi=10.1016/j.neuropsychologia.2015.12.009&domain=pdfmailto:[email protected]://dx.doi.org/10.1016/j.neuropsychologia.2015.12.009
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M. Gilead et al. / Neuropsychologia 81 (2016) 207–218208
The second main distinction pertains to the goals associatedwith
the emotion; self-conscious emotions are associated withhigh-order
goals, and play an important role in adaptive socialbehavior such
as moral behavior (Tangney et al., 2007), coopera-tion (Dorfman et
al., 2014), and self-regulation (Eyal and Fishbach,2010;
Giner-Sorolla, 2001). To illustrate the latter,
self-consciousemotions are believed to direct behavior towards
long-term goals,and hence, play an important role in regulating
self-control con-flicts, i.e., conflicts between a long-term goal
that offers large yetdelayed benefits and a short-term temptation
that offers smalleryet immediate benefits (Williams and DeSteno,
2008; Zemack-Rugar et al., 2007). Consistent with this line of
theorizing, recentresearch has shown that self-conscious emotions
such as pride andguilt are associated with adherence to long-term
goals and theexertion of self-control, whereas emotions such as
joy, excitement,sadness and frustration are associated with pursuit
of short-termgoals and succumbing to temptations (Eyal and
Fishbach, 2010;Giner-Sorolla, 2001; Hofmann and Fisher, 2012;
Mukhopadhyayand Johar, 2007; Williams and DeSteno, 2008;
Zemack-Rugar et al.,2007). For example, in recent studies, we have
found that com-pared to a positive emotion that is not
self-conscious (i.e., joy),priming pride improved self-control
(Katzir et al., 2010, see alsoKatzir et al., 2015). Considering
that the brain areas commonlyassociated with conflict resolution
are the dorsal Anterior Cingu-late Cortex (dACC) and lateral-dorsal
prefrontal cortex (e.g., Bot-vinick et al., 2004; Cole and
Schneider, 2007; Hare et al., 2009;MacDonald et al., 2000; Ochsner
et al., 2012; Kerns et al., 2004),the processing of self-conscious
(vs. basic) emotions may be ex-pected to recruit these regions.
1.2 Past research on neural substrates of self-conscious
emotions
Several past studies investigated the neural substrates of a
spe-cific self-conscious emotion. These studies often (but not
always)reported activations in the medial prefrontal cortex: Shin
et al.(2000) compared the processing of guilt-evoking vs. neutral
sce-narios and found activation within the anterior temporal
poles,anterior cingulate gyrus, and left anterior insular
cortex/inferiorfrontal gyrus. Moll et al. (2007) conducted a study
on moral emo-tions that included guilt and anger, but did not
report the results ofa comparison between the two. Kedia et al.
(2008) conducted astudy on moral emotions in which they compared
guilt-evokingtrials to anger evoking trials and to
compassion-evoking trials. Theirresults showed that emotional (vs.
non-emotional) trials activatedthe dorsal mPFC, insula, amygdala,
and the temporo-parietal junc-tion—but there were no significant
differences in activation be-tween the different emotion
categories. Basile et al. (2011) com-pared guilt to sadness and
anger and found greater activationwithin the mPFC and the cingulate
gyrus in the guilt condition.Somerville et al. (2013) induced
self-evaluation by making partici-pants believe they are being
watched by a peer through a camera,and discovered heightened
engagement of the mPFC in adolescence(relative to childhood) that
persists into adulthood. Finally, Fourieet al. (2014) induced
feelings of guilt in the context of race relations,and found
activations in the ACC, anterior insula, the mPFC, pos-terior
cingulate cortex, and the precuneus.
Studies that focused on the neural substrates of a single
posi-tive self-conscious emotion did not observe activations in
themedial prefrontal cortex. Takahashi et al. (2008) compared
theneural correlates of processing pride as compared to joy,
anddiscovered that pride was associated with activation of two
re-gions involved in mentalizing tasks (the superior temporal
sulcusand temporal poles) but not in the mPFC. Another study
(Simon-Thomas et al., 2012) contrasted the processing of pride
withcompassion, and discovered activation in the posterior
cingulatecortex.
The studies outlined above investigated a single
self-consciousemotion, and contrasted it with non-emotional states,
or basicemotions. Thus, these studies were not in a position to tap
into thecorrelates of self-conscious emotions over and above
emotion-specific content. However, five previous studies looked at
neuralcorrelates that are associated with more than one
self-consciousemotion. Once again, these studies often reported
activations inthe mPFC: Wagner et al. (2011) investigated the
unique neuralcorrelates of guilt processing as compared with two
closely relatedemotions, sadness and shame. They found that guilt
was asso-ciated with greater activation of the mPFC as compared to
sadnessand shame. Takahashi et al. (2004) compared feelings of
guilt andembarrassment to a non-emotional control; once again, they
dis-covered activation in the mPFC as well as in the superior
temporalsulcus. Michl et al. (2014) replicated the paradigm from
Takahashiet al. (2004) but did not find mPFC activation for the
guiltcondition. However, Burnett et al. (2009) compared guilt
andembarrassment to basic emotions (disgust and fear) and
replicatedTakahashi et al.'s (2004) findings.
Importantly, all of these studies examined only negative
emo-tions. In light of that, further research is still warranted in
order tofully capture the construct of self-conscious emotions, as
it alsopertains to positive self-evaluation, namely, a sense of
pride. Theimportance of this point is further highlighted by the
fact thatwhereas negative self-conscious emotions often activated
themPFC (e.g., Takahashi et al., 2004; Burnett et al. 2009;
Wagneret al., 2011), the two studies that examined the processing
of pridedid not report activations within this region (Takahashi et
al.,2008; Simon-Thomas et al., 2012).
Two previous studies included within their design a positive
anda negative self-conscious emotion. However, they did not
compareself-conscious to non-self-conscious emotions: Roth et al.
(2014)compared the processing of a negative self-conscious
emotion(shame/guilt) to a positive self-conscious emotion (pride)
and anon-emotional control. Comparing self-conscious emotions vs.
noemotion (i.e., a condition in which participants waited for a
dis-tracting picture to appear) resulted in widespread activation
acrossthe cortex. Comparing pride to guilt and shame resulted in
greateractivation in widespread regions including the left superior
frontalgyrus, left ventral mPFC, middle and posterior cingulate
cortex, theinferior temporal gyrus, inferior parietal gyrus, the
left caudatebody and left lateral thalamus. The reverse contrast
(guilt andshame vs. pride) did not result in significant
differences. In Roth etal.'s study, the focus of investigation was
the processing of guilt ascompared to pride; in light of this, it
did not include basic-emotionconditions to allow researchers to
distinguish between self-con-scious and non-self-conscious
affective processing. Zahn et al.(2009) conducted a study on
social-moral emotions wherein theycompared the processing of events
that elicit the social emotions ofguilt, pride, gratitude and
anger. Their results showed that partici-pants’ reports of feeling
pride during the task were correlated withactivity in the septum,
and feelings of guilt were correlated withactivity in the subgenual
ACC. When the researchers looked at theneural activity during pride
trials (vs. guilt, gratitude and anger)they found activation in the
vmPFC, the ventral tegmental area, andthe parahippocampal gyrus;
there were no significantly strongeractivations for guilt (compared
with pride, gratitude, and anger).Similarly to Roth et al.'s (2014)
study, in this study as well, the focusof investigation was not
self-conscious emotions, therefore, it didnot directly address the
question of whether there are neural sub-strates that subserve
self-conscious (but not basic) emotions, irre-spective of the
dimension of valence.
Thus, at the current stage of investigation, further work
isneeded in order to uncover the neural substrates that
subserveself-conscious emotions—above and beyond emotion-specific
andvalence-specific processes.
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M. Gilead et al. / Neuropsychologia 81 (2016) 207–218 209
2. The current study
In order to investigate the neural correlates of
processingevents that elicit positive and negative self-conscious
emotions,we chose four prototypes of self-conscious and basic
emotions:pride (self-conscious, positive), guilt (self-conscious,
negative), joy(basic, positive) and anger (basic, negative). Based
on theoreticaland empirical distinctions between self-conscious and
basicemotions, we generated the following predictions and
researchquestions:
(i) Given past work (Eyal and Fishbach, 2010;
Giner-Sorolla,2001; Hofmann and Fisher, 2012; Katzir et al., 2010;
Shimoni et al.,2016; Williams and DeSteno, 2008; Zemack-Rugar et
al., 2007) onthe role of self-conscious emotions in self-control,
we predictedthat processing self-conscious emotions should activate
areas relatedto self-control, namely, the ACC and lateral-dorsal
prefrontal cortex(e.g., Botvinick et al., 2004; Cole and Schneider,
2007; Hare et al.,2009; MacDonald et al., 2000; Ochsner et al.,
2012; Kerns et al.,2004).
(ii) Given the inconclusive neural findings reviewed above,
wewere specifically interested to see whether both positive and
ne-gative self-conscious emotions activate the region previously
im-plicated in self-referential processing, namely, the mPFC (e.g.,
Dennyet al., 2012; Mitchell et al., 2006).
Furthermore, we were interested in two exploratory hy-potheses
concerning activation within the mPFC: extant evidenceshows that
the mPFC is divided into a dorsal region which is morestrongly
activated in reflection upon others, and a ventral regionwhich is
more strongly associated with self-reflection (Dennyet al., 2012;
Mitchell et al., 2006). Consequently, as a secondarygoal of the
current study, we wanted to see whether the specificself-conscious
emotions—i.e., guilt and pride—differentially relyupon these two
sub-regions. Additionally, past research showsthat the dorsal mPFC
is involved both in thinking of future eventsand recollection of
the past (e.g., Addis et al., 2007) whereas theventral mPFC is more
active when participants imagine (vs. re-member) events (Addis et
al., 2009). In light of this, as a secondarygoal of the current
study, we were interested in whether activitywithin this region
differs when processing emotional events whichoccurred in the past,
as opposed to events that are imagined tooccur in the future.
2.1. Method
2.1.1. ParticipantsTwenty right-handed participants (14 women,
average age 25.9
years, range 21–33 years) from Tel-Aviv University participated
inthe experiment. They were all native Hebrew speakers, none had
ahistory of neurological or psychiatric disorders, and all had
normalor corrected-to normal vision. One participant was excluded
fromthe final analysis due to excessive motion. They gave
writtenconsent prior to taking part in the experiment. The study
was
Table 1Pre-test data – Average (across stimuli) of number of
participants (out of 38) who repor
Stimulus classification Reported emotion
Guilt Pride
Guilt 24.5 (6.86) 0.5 (0.73)Pride 0.12 (0.34) 24.81 (7.07)Anger
0.125 (0.34) 0.18 (0.54)Joy 0.18 (0.40) 0.12 (0.34)
approved by the Institutional Review Board of the
SouraskyMedical Center, Tel-Aviv.
2.1.2. MaterialsWe created the stimuli based upon a pre-test.
First, based on
face validity, we compiled a list of 96 events (24 per emotion
ca-tegory) which we suspected might evoke a sense of guilt,
pride,anger, and joy. Each event was described in general terms and
afew examples of specific possible instantiations followed. For
ex-ample: “Neglecting to speak to someone close for a long time
(afriend; parent; sibling)”.
These scenarios were rated by thirty-eight participants (33
females,2 unknown, average age 23.11 years, range 19–26 years) who
an-swered two questions concerning each item: (i) whether it is
likelythat they would find themselves in the circumstance described
in thecourse of the next five years (by making a “yes”/“no”
response); (ii)whether such an event is typically associated with
feelings of guilt,pride, anger, joy or some other emotion (by
selecting one of the fiveoptions). Based upon participants’
responses we selected 16 events foreach emotion that were likely to
occur, and were typically associatedwith a specific emotion.
Specifically, for each emotion category, weselected stimuli such
that across all items in the category: (i) morethan 90% of
participants said it is likely that they would find them-selves in
these situations; (ii) at least 60% of participants categorizedthe
stimuli according to our designated classification.
Chi-squared tests indicated that participants classified
guiltstimuli as being associated with guilt more often than not,
χ2
(1,38)¼4.90, p¼ .026; pride stimuli as being associated with
pridemore often than not, χ2 (1,38)¼5.35, p¼ .020; anger stimuli
asbeing associated with anger more often than not, χ2
(1,38)¼5.35,p¼ .020; joy stimuli as being associated with joy more
often thannot, χ2 (1,38)¼26.34, po .001 (Table 1).
The first pre-test was meant to help generate a set of
eventsthat are common experiences for participants, and that
partici-pants label as being associated with one of the four
emotions. Inorder to make sure that the behavioral procedure
(described be-low) elicits the corresponding emotion, we conducted
a secondpre-test. Twenty-four participants (16 females, average age
25.00years, range 21–30 years) performed the experimental
procedurewith the exceptions that (1) they performed it
individually in frontof a computer rather than inside the scanner,
and therefore theirneural activation was not recorded, and (2)
following each block ofthe experimental procedure (i.e., a series
of 4 questions from thesame emotion, see below) they provided four
responses to indicatethe degree to which they feel the four
emotions (anger, guilt, joy,and pride) on a 5 point scale (1-low,
5-high).
The results indicated that the experimental procedure
suc-cessfully elicited the target emotions. Specifically,
participants re-ported feeling more intense pride following pride
blocks than joyblocks, F(1,23)¼34.00, po .001, ηp2¼ .60, and more
intense joyfollowing joy blocks than pride blocks, F(1,23)¼15.92,
po .001,ηp
2¼ .41. They also reported feeling more intense anger
following
ted that the stimulus elicits a given emotion. SDs in
parentheses.
Anger Joy Other
1.18 (1.37) 0.56 (0.81) 11.25 (6.78)0.18 (0.54) 8.56 (6.61) 4.31
(3.77)
24.81 (6.63) 0.18 (0.40) 12.68 (6.52)0.06 (0.25) 32.87 (4.22)
3.94 (4.75)
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Table 2Pre-test data – Average (across block type) of intensity
ratings of each emotionfollowing the 4 types of emotion blocks
(anger, guilt, joy, and pride). SEs inparentheses.
Block type Intensity of affect
Guilt Anger Pride Joy
Guilt 2.98 (0.35) 2.23 (0.35) 2.49 (0.26) 2.77 (0.27)Anger 1.71
(0.24) 2.88 (0.39) 2.36 (0.27) 2.77 (0.24)Pride 1.57 (0.22) 1.69
(0.26) 3.97 (0.31) 3.95 (0.30)Joy 1.53 (0.21) 1.58 (0.24) 3.26
(0.32) 4.42 (0.33)
M. Gilead et al. / Neuropsychologia 81 (2016) 207–218210
anger blocks than guilt blocks, F(1,23) ¼16.46, po .001, ηp2¼
.42,and more intense guilt following guilt blocks than anger
blocks, F(1,23)¼52.08, po .001, ηp2¼ .69 (see Table 2).
We also wanted to make sure that differences between
self-conscious and basic emotions do not reflect differences in
thedegree of social interaction in the chosen eliciting events. To
thatend, two independent judges coded the degree to which each
ofthe chosen 16 events entailed a social interaction on a 0–2
scale,(0-“no social interaction”; 1-“might involve social
interaction”; 2-“definitely involves social interaction”).
Inter-rater reliability washigh (r¼ .87). The emotion category
involving the least amount ofsocial interaction was guilt (M¼1.18),
followed by joy (M¼1.37),pride (M¼1.50), with anger being the most
socially-interactiveemotion (M¼1.75). Thus, on average, ratings of
social interactionfor self-conscious emotions (M¼1.34) were not
higher than thoseof basic emotions (M¼1.56).
2.1.3. Behavioral procedureParticipants were carefully
instructed and trained on the task
prior to entering the scanner. The training was repeated
verbatiminside the scanner. The items used for the training session
weretaken from a different pool of sentences than the main task.
Par-ticipants answered the emotion-eliciting questions by pressing
akey on a response box with their index and middle left hand
fin-gers. Stimuli were presented with Presentation version
14.9(Neurobehavioral Systems, CA, USA). Each question was
presentedon screen for 6000 milliseconds.
The experiment consisted of two consecutive sessions,
eachlasting 9 min and 14 s. We presented the stimuli in a
blockeddesign. Each block contained a series of 4 questions from
the sametemporal perspective and emotion, and was succeeded by a
10seconds fixation (Fig. 1). Each session contained 2 blocks from
eachof the 8 conditions (created by crossing the 4 emotions with
the2 temporal perspectives). We fully-randomized the order of
ex-perimental blocks, and of stimuli in each block. In total,
each
Fig. 1. An example block (Self-conscious, Negative, Past).
Example items are translatedconsisted of 8 block types (created by
crossing the variables: Past/Future � Self-conscmeant to elicit a
specific emotion. Each question was presented for 6 s.
participant answered all 128 questions (16 questions from each
ofthe 8 block types).
In the actual experiment, we made participants recollect
orimagine the emotion-eliciting events by asking them whether it
ispossible that each event would happen to them sometime in
thenext-five years and whether they have experienced an event
likethis sometime in the preceding five years. Participants were
cuedbefore each block with the appropriate temporal perspective
(fu-ture/past). On each experimental trial, participants saw the
emo-tion eliciting statement (e.g., “Neglecting to speak to
someoneclose for a long time [a friend; parent; sibling]). On each
experi-mental trial, participants responded whether the event
happenedto them or is likely to occur by choosing “yes” or “no”
buttons. Thelist of stimuli contained 128 questions: 16 events per
emotioncategory�4 emotion types: guilt, pride, anger and joy)�2
tem-poral perspectives (past vs. future). The complete list of
stimuli isprovided in Appendix A.
2.1.4. Imaging procedureWhole-brain T2*-weighted EPI functional
images were acquired
with a GE 3-T Signa Horizon LX 9.1 echo speed scanner
(Milwaukee,WI). 264 volumes were acquired (TR¼2000 ms, 200 mm FOV,
64�64matrix, TE¼35, 35 pure axial slices, 3.15�3.15�3.5 mm3 voxel
size,no gap). Slices were collected in an interleaved order. At the
beginningof each scanning session, 5 additional volumes were
acquired, to allowfor T1* equilibration (they were not included in
the analysis). Beforethe experiment, high-resolution anatomical
images (SPGR; 1 mm sa-gittal slices) were obtained. Head motion was
minimized by usingcushions arranged around each participant’s head,
and by explicitlyguiding the participants prior to entering the
scanner. Imaging datawere preprocessed and analyzed using SPM5
(Wellcome Departmentof Cognitive Neurology, London). A slice-timing
correction to the firstslice was performed followed by realignment
of the images to the firstimage. Next, data were spatially
normalized to an EPI template basedupon the MNI305 stereotactic
space. The images were then resampledinto 2-mm cubic voxels, and
finally smoothed with an 8-mm FWHMisotropic Gaussian kernel. The
general linear model was used for sta-tistical analyses. Eight
regressors (one for each stimulus condition)were used to model the
effects of interest; each consisted of a boxcarfunction convolved
with a standard hemodynamic response function.Two additional
regressors were included in the model to account
forsession-specific low-frequency effects. Based on our design
para-meters, SPM’s optimal high-pass filter cutoff was determined
usingDesign Magic high-pass filter optimization tool (developed by
MatthijsVink; http://www.ni-utrecht.nl/downloads/d_magic) and was
set at272. We computed the second-level analyses (in which
participantswere treated as random effects) using one-sample
t-tests. Significantregions of activationwere identified using a
threshold of po.001 witha cluster size threshold of 74 voxels,
corresponding to a threshold
from Hebrew (complete list of stimuli available in Appendix A).
The experimentious/Basic � Positive/Negative). Each block contained
four questions all of which
http://www.ni-utrecht.nl/downloads/d_magic
-
Table 3Behavioral Results. RTs (SDs in parentheses) in
milliseconds by Valence of emotion,Emotion Type and Temporal
Perspective.
Negative Positive
Self-conscious(Guilt)
Basic (Anger) Self-conscious(Pride)
Basic (Joy)
Future 2635 (598) 2499 (556) 2347 (499) 2130 (475)Past 2571
(662) 2550 (509) 2277 (464) 2197 (578)
M. Gilead et al. / Neuropsychologia 81 (2016) 207–218 211
of po.05, corrected for multiple comparison, as assessed
throughMonte Carlo simulations implemented in Matlab (Slotnick et
al., 2003).We ran 1000 iterations of the simulation using the
pre-definedparameters of our design, and the smoothness parameter
as estimatedin SPM.
In order to investigate the neural correlates of
self-consciousand basic emotions, regardless of emotion-specific
content, wesearched for regions that were activated for both
positive andnegative emotions. The Self-conscious4Basic emotions
conjunc-tion analysis (Guilt4Anger)∩(Pride4 Joy) was implemented
byrunning the contrast of Guilt4Anger (at a threshold of
0.031)inclusively masked with the contrast of Pride4 Joy (at a
thresholdof 0.031). Similarly, the Basic4Self-conscious conjunction
analysiswas implemented by running the contrast of Anger4Guilt
in-clusively masked with the contrast of Joy4Pride, using the
samethresholds. Since both contrasts are orthogonal, with a cluster
sizeof 74 voxels, this analysis tests against the conjunction null
atpo .05, corrected.
2.2. Results
2.2.1. Behavioral resultsParticipants indicated that they have
experienced or are likely
to experience the emotion-eliciting events on 93.82% of
trials.There were no significant differences in ratio of yes and no
re-sponses between the different emotions. There was no
significantdifference in response latencies for negative
self-conscious (guilt)and negative basic (anger) emotion questions,
F(1,18)¼1.18,p¼ .202. Participants responded more quickly to the
positive-va-lence basic emotion questions (joy; M¼2164 ms, SD¼509
ms)than to the positive self-conscious emotion questions
(pride;M¼2312 ms, SD¼453 ms), F(1,18)¼6.94, p¼ .016,2 ηp2¼ .30.
Parti-cipants also generally responded more quickly to
positive(M¼2238 ms, SD¼466 ms) compared to negative emotion
ques-tions (M¼2564 ms, SD¼544 ms), F(1,18)¼27.00, po .001, ηp2¼
.61.Furthermore, there was a significant interaction between
TemporalPerspective and Emotion Type, F(1,18)¼5.41, p¼ .030 (See
Table 3),such that participants responded more slowly to
self-consciousemotions than to basic emotions when thinking of the
future, F(1,18)¼10.54, p¼ .004, ηp2¼ .40, but not when thinking of
the past, F(1, 18)¼1.96, p ¼ .177. There were no other significant
effects.
2.2.2. Imaging data2.2.2.1. Temporal perspective. Temporal
perspective did not have asignificant main effect, nor did it
interact with any of the otherindependent variables. Therefore, we
collapsed all of our analysesacross this variable.
2.2.2.2. Self-conscious emotions (Guilt4Anger)∩(Pride4 Joy).
Aspredicted, processing self-conscious emotions activated
frontalregions associated with cognitive control: a region within
themPFC extending to the dACC, and within a lateral-dorsal
prefrontalregion (see Fig. 2 and Table 4).
2.2.2.3. Basic emotions (Anger4Guilt)∩(Joy4Pride).
Processingbasic emotions activated regions associated with embodied
ex-perience, namely, regions associated with visual-spatial
percep-tion and imagery (left superior occipital gyrus and
bilateral
2 The correlates of self-conscious and basic emotions were
identified usingconjunction analyses which pin-pointed significant
differences in neural activationthat were evident for both the
positive and negative emotions. Because there wereno differences in
response latencies for anger and guilt trials, the difference
inresponse latencies between joy and pride cannot account for our
findings regardingthe neural correlates of self-conscious and basic
emotions.
parahippocampal gyrus) and somatosensation (right
postcentralgyrus). Furthermore, basic emotions activated a region
of the leftinsula, which was found in previous studies to be
activated inemotional processing and interoception (e.g., Stein et
al., 2007;Zaki et al., 2012) (see Fig. 2 and Table 4).
2.2.2.4. Guilt4Other emotions. Guilt-evoking questions were
as-sociated with widespread activation throughout the cortex
(mostprominently in the dorsomedial and lateral PFC) in the right
lateralorbitofrontal cortex and in subcortical regions (most
prominentlyin the caudate nucleus and thalamus) (see Fig. 3 and
Table 4).
2.2.2.5. Pride4Other emotions. Pride-evoking questions were
as-sociated with activation of regions involved in
self-referentialprocessing and reward, namely, the ventromedial
prefrontal cortexextending to the orbitofrontal cortex (see Fig. 3
and Table 4).
2.2.2.6. Anger4Other emotions. Anger-evoking questions
wereassociated with activation of the left temporo-parietal
junctionand superior temporal sulcus (see Fig. 3 and Table 4).
2.2.2.7. Joy4Other emotions. Joy-evoking questions were
asso-ciated with activation of the bilateral superior occipital
gyrus (seeFig. 3 and Table 4).
2.2.2.8. Interpersonal4Non-interpersonal emotions. Processing
thesocial emotions (i.e., anger, guilt, & pride) compared to
non- socialemotions (i.e., joy) recruited regions involved in
social-cognition,namely the medial prefrontal cortex, precuneus and
temporo-parietal junction. Further activations included the
cerebellum andleft lateral prefrontal cortex (see Table 4).
2.2.2.9. Valence. Similarly to previous behavioral and neural
stu-dies which have shown an asymmetry between positive and
ne-gative valence (Baumeister et al., 2001), we observed robustand
widespread activations throughout cortical and sub-corticalregions
associated with negative emotions, whereas positiveemotions were
not associated with significant activation (seeTable 4).
2.2.2.10. ROI analysis. We defined as regions of interest the
twoclusters associated with processing self-conscious emotions
(themPFC extending to the ACC, and the lateral-dorsal
prefrontalcluster), and the five regions associated with processing
basic-emotions. We extracted parameter estimates from each ROI on
asubject-by-subject basis using MarsBaR v.042 (Brett et al.,
2002).We conducted a Valence (Negative, Positive) � Type
(Self-con-scious, Basic) repeated measured ANOVA on the mean beta
valuesof each of the seven ROIs. None of the regions displayed an
in-teraction of Valence and Type, max F(1,18)¼1.26, p¼ .276. In
themPFC, there was greater activity for self-conscious (vs.
basic)emotions, F(1,18)¼15.32, p¼ .001, and for negative (vs.
positive)valence, F(1,18)¼18.96, po .001; In the left middle
frontal gyrus,there was greater activity for self-conscious (vs.
basic) emotions, F
-
Fig. 2. Sagittal view of the brain showing neural activity
associated with self-conscious and basic emotions. Activations are
shown at a threshold of po .05 (corrected). Foractivation foci see
Table 4.
M. Gilead et al. / Neuropsychologia 81 (2016) 207–218212
(1,18)¼16.88, po .001, and for negative (vs. positive) valence,
F(1,18)¼38.13, po .001.
In the left insula, there was greater activity for basic (vs.
self-con-scious) emotions, F(1,18)¼12.65, p¼ .002, and for positive
(vs. nega-tive) valence, F(1,18)¼5.31, p¼ .034; In the left
parahippocampal gyrusthere was greater activity for basic (vs.
self-conscious) emotions, F(1,18)¼15.62, po.001; In the right
parahippocampal gyrus there wasgreater activity for basic (vs.
self-conscious) emotions, F(1,18)¼13.04,p¼ .002; In the left
superior occipital gyrus, there was greater activityfor basic (vs.
self-conscious) emotions, F(1,18)¼10.40, p¼ .005; In theright
postcentral gyrus there was greater activity for basic (vs.
self-conscious) emotions, F(1,18)¼11.79, p¼ .003. There were no
othersignificant differences.
3. Discussion
In the present study participants thought of various
personalevents which elicited feelings of self-conscious or basic
positive or
negative emotions. Self-conscious emotions were associated
withactivation within the mPFC extending to the dACC and within
thelateral-dorsal prefrontal cortex. We also observed that
self-con-scious emotions differentially activated regions within
the mPFCsuch that the negative self-conscious emotion (guilt) was
asso-ciated with a dorsal activation, and the positive
self-consciousemotion (pride) was associated with activation of the
ventralmPFC. The cluster co-activated by both types of
self-consciousemotions was located somewhere in between the ventral
anddorsal sub-regions. We did not find significant activations
asso-ciated with temporal perspective. Overall, we believe that
theseresults might shed light on the neural basis and phenomenology
ofself-reflective and affective processing. We now turn to
discusseach of our main findings.
3.1. Self-conscious emotions and self-regulation
As predicted, we found that processing of self-conscious
emo-tion, regardless of their valence, activated the dACC and
the
-
Table 4Regions identified in the whole brain analysis. All
activations are below a threshold of po .001 and a cluster extent
of 74 voxels (FWE-corrected, po .05).
Contrast Region Coordinates Significance level Voxels
x y z Z-score
Self-consious4BasicFrontal Medial Prefrontal Cortex �8 46 36
3.74 325
L Middle Frontal Gyrus �38 14 46 3.02 109Basic4Self-consious
Temporal L Parahippocampal Gyrus �28 �38 �24 3.63 434R
Parahippocampal Gyrus 14 �50 6 2.46 146
Occipital L Superior Occipital gyrus �46 �80 32 3.08 199Parietal
R Postcentral Gyrus 64 �16 30 3.01 279Insula L Insula �50 �6 �8
2.72 89
Guilt4OtherSub-lobar Caudate 8 8 8 5.24 3545Frontal R Inferior
Frontal Gyrus 58 26 14 5.14 769
Dorsomedial Prefrontal Cortex �6 16 68 4.91 2602L Inferior
Frontal Gyrus �52 20 8 4.78 2278R Inferior Frontal Gyrus 46 38 �14
4.6 220
Temporal L Fusiform Gyrus �48 �38 �8 4.41 207Parietal L
Precuneus �42 �64 42 4.04 782Occipital R Lingual Gyrus 14 �86 2 3.5
113Cerebellum L Inferior Semi-Lunar Lobule �24 �80 �38 4.54 209
R Uvula 22 �80 �24 3.8 238Pride4Other
Frontal Ventromedial Prefrontal Cortex �2 62 24 4.67
278Anger4Other
Temporal L Temporo-Parietal Junction �40 �60 26 4.38 792L
Superior Temoral Sulcus �52 �6 �28 4.3 250
Joy4OtherOccipital R Superior Occipital Gyrus 40 �84 32 4.69
215
L Superior Occipital Gyrus �36 �86 36 3.71
125Social4Non-social
Frontal Dorsomedial Prefrontal Cortex �8 52 28 4.98 2894L
Inferior Frontal Gyrus �50 26 �10 3.98 611L Middle Frontal Gyrus
�38 10 50 3.95 572L Middle Frontal Gyrus �34 52 8 3.87 87
Temporal L Temporo-Parietal Junction �50 �64 18 4.55
1422Parietal Precuneus �2 �62 38 4.25 733Cerebellum L Inferior
Semi-Lunar Lobule �22 �80 �44 3.87 85
R Inferior Semi-Lunar Lobule 26 �80 �42 3.82
93Negative4Postive
Frontal L Middle Frontal Gyrus �42 10 52 5.27 3354Superior
Frontal Gyrus 2 40 52 5.21 3970
R Inferior Frontal Gyrus 42 30 �18 4.99 691Occipital L Middle
Temporal Gyrus �50 �64 20 5.17 2951Temporal L Inferior Temporal
Gyrus �52 �8 �28 4.76 283Cerebellum L Pyramis �20 �86 �32 4.82
4571
Positive4NegativeNon identified
M. Gilead et al. / Neuropsychologia 81 (2016) 207–218 213
lateral-dorsal prefrontal cortex. Much research shows that
thedACC and the lateral-dorsal prefrontal cortex are involved in
ef-fortful cognitive control (e.g., Cole and Schneider, 2007; Hare
et al.,2009; Ochsner et al., 2012). Specifically, it has been
suggested thatthe dACC is involved in monitoring cognitive conflict
(e.g., Botvi-nick et al., 2004) and that lateral-dorsal regions are
involved inimplementation of control over conflict (MacDonald et
al., 2000).
The involvement of cognitive-control regions in the processingof
self-conscious emotions is consistent with the view according
towhich the function of self-consciousness is to allow flexible
andcomplex control of behavior (e.g., Anderson, 1983). More
specifi-cally, self-conscious emotions can help us manage conflicts
be-tween phylogenetically-ancient hedonic value (the taste of
burgerand fries), and phylogenetically-novel, socially-constructed
values(being thin, not hurting animals). Supporting this idea,
recentbehavioral work suggests that both negative (Zemack-Rugar et
al.,2007) and positive (Eyal and Fishbach, 2010; Katzir et al.,
2010;
Shimoni et al., 2016; Williams and DeSteno, 2008)
self-consciousemotions lead people to exert greater degrees of
self-control. Forexample, in recent studies, we (Katzir et al.,
2010, 2015) haveshown that thinking of pride- (vs. joy-) evoking
events facilitatesself-control, as evident in (1) more successful
inhibition on ananti-saccade task (performance on which is known to
rely on thedACC and lateral-dorsal prefrontal cortex, e.g., Klein
et al., 2007;Pierrot-Deseilligny et al., 2003), and (2) reduced
congruency effecton a switching task (reduced congruency effect
indicates betterconflict resolution which is known to rely on the
dACC and lateral-dorsal prefrontal cortex, Kerns et al., 2004).
Thus, the current workjoins this previous behavioral evidence in
highlighting the role ofself-conscious emotions in heightened
self-control.
The finding that self-conscious emotions, irrespective of
va-lence, activate medial frontal regions is also consistent with
extantneuropsychological literature that examined the behavior of
in-dividuals with medial frontal cortex lesions (e.g., Sturm et
al.,
-
Fig. 3. Sagittal view of the brain showing neural activity
associated with specific emotions. Activations are shown at a
threshold of po .05 (corrected). For activation foci seeTable
4.
M. Gilead et al. / Neuropsychologia 81 (2016) 207–218214
2013; Beer et al., 2003; 2006; Moll et al., 2011; Krajbich et
al.,2009). This work documented brain-lesioned individuals'
reducedaffect in response to guilt-, shame-, and
embarrassment-evokingsituations. The current findings, which
implicate medial frontalregions also in positive self-conscious
affect, raise the possibilitythat frontal-lesioned individuals
should also be deficient in theirability to feel a sense of pride,
even when their behavior justlywarrants it. This prediction, to the
best of our knowledge, has notbeen directly tested, and should be
investigated in future work.
3.2. Self-conscious emotions and self-related processing
Our results show that affective self-processing was
associatedwith activation of the mPFC. This region was previously
shown tobe involved in the cognitive aspects of self-reflective
processing,namely, in processing knowledge about the self (e.g.,
Kelley et al.,2002) and in processing the mental states of the self
and others(Mitchell et al., 2005). Past research has consistently
shown thatnegative self-conscious emotions activate the mPFC
(e.g.,
Takahashi et al., 2004; Burnett et al., 2009; Wagner et al.,
2011).The current study joins a study by Zahn et al. (2009) in
showingthat feelings of pride activate the mPFC. However two
previousstudies that examined the processing of pride did not
report ac-tivations within this region (Takahashi et al., 2008;
Simon-Thomaset al., 2012).
It is likely that methodological differences between the
currentstudy and the studies by Takahashi et al. (2008) and
Simon-Tho-mas et al. (2012) produced these diverging findings. In
Takahashiet al.'s study participants read sentences describing
pride-relatedevents (e.g., “I graduated from the most prestigious
university); inSimon-Thomas et al.'s study participants observed
pride-relatedimages (e.g., images of graduation). It is possible
that the stimuli inthese studies were not sufficiently
self-relevant, and therefore didnot produce sufficiently strong
feelings of pride. This possibilityshould be studied in future
research comparing different ap-proaches for the elicitation of
self-conscious emotions.
Extensive findings show that within the mPFC, a
meaningfuldistinction may be made between the dorsal region, which
is more
-
M. Gilead et al. / Neuropsychologia 81 (2016) 207–218 215
strongly activated when thinking of other people, and the
ventralregion, which is associated with thinking about the self and
closeothers (e.g., Denny et al., 2012; Mitchell et al., 2006).
Interestingly,our results also exhibited a distinction between the
ventral anddorsal mPFC, wherein guilt was associated with more
dorsal ac-tivation, and pride was associated with ventral
activation.
The association between the ventral mPFC and pride is
con-sistent with recent work (Chavez and Heatherton, 2015)
showingthat the individuals' level of self-esteem (which can be
seen as asustained feeling of pride) is correlated with structural
integrityand functional connectivity between the ventral mPFC and
sub-cortical regions involved in processing rewards (i.e., the
ventralstriatum). These findings, alongside with the current
research,may suggest that the differential involvement of the
ventral mPFCin self- (vs. other-) focused processing could be
related to its roleas part of a network of regions that subserve
positive-affectiveprocessing (e.g., Brown et al., 2011; Liu et al.,
2011).
However, it remains possible that guilt and pride
differentiallyactivate the ventral and dorsal mPFC because they
differ in thedegree of self- and other-related processing. Guilt is
often relatedto moral considerations; in such cases it likely
entails the re-presentation of both a moral agent (the self) and a
moral patient(the other; Gray et al., 2012; e.g., I feel guilty
because I hurt them).Although feelings of pride often involve the
perspective of others(e.g., I bet they are proud of me), it is also
common to feel prideirrespective of the consideration of others
(e.g., I am proud ofmyself for accomplishing this). Thus, future
research should at-tempt to carefully disentangle the involvement
of representationsof self and others in feelings of guilt and
pride, and thereby mayhelp shed light on the nature of the
functional distinction betweenthe ventral and dorsal mPFC.
An additional observed distinction between the dorsal and
ventralmPFC is that the dorsal mPFC is involved both in thinking of
futureevents and recollection of the past (e.g., Addis et al.,
2007) whereas theventral mPFC is more active when participants
imagine (vs. re-member) events (e.g., Addis et al., 2009). Contrary
to our expectation,in the current investigation we did not find
significant differences inactivation for future and past events in
the mPFC (or within any otherbrain region). Although surprising,
this null finding is consistent withmuch previous research
suggesting that the neural systems whichallow us to imagine future
worlds and to recollect the past sub-stantially overlap (e.g.,
Schacter et al., 2007; Buckner and Carroll,2007); thus, it is
likely that any differences between future- and past-oriented
processing may have been too subtle to be detected in thecurrent
paradigm.
Finally, much literature shows that many psychopathologiesare
associated with dis-regulated activity of the medial
prefrontalcortex. For example, research shows that the vmPFC is
hypoacti-vated in post-traumatic stress disorder (Shin and
Liberzon, 2010)and hyperactivated in obsessive-compulsive disorder
(e.g., Sturmet al., 2013). In contrast, the dmPFC was found to be
hypoactivatedduring reward-seeking among heavy drinkers (e.g., Bogg
et al.,2012). The finding whereby the dmPFC is associated with
guiltwhereas the vmPFC is associated with pride may help in
devel-oping a more refined model of the link between dysregulated
self-conscious affective processing, psychopathology, and mPFC
dis-regulation. For example, our findings may suggest that
interven-tions that apply repeated transcranial magnetic (rTMS) in
order totreat depressive and manic states (e.g., Pallanti et al.,
2014), maybenefit from targeting the dmPFC and vmPFC,
respectively.
3.3. Self-conscious vs. social emotions
Self-conscious emotions are believed to have developed as
anadaptation to the complex social organization of the ancient
humansocieties (Dunbar, 1998), and are central in motivating
socially
appropriate and desired behaviors (e.g., Tangney et al., 2007).
In lightof this, it is unsurprising that self-conscious emotions
are also ne-cessarily “social emotions”. Many aspects of the self
are socially con-structed, in that they pertain to a valuation that
is dependent uponsocial norms (e.g., “I am smart/stupid relative to
my peers”, “I am amoral/immoral person”). Indeed, much research
shows that there issubstantial overlap between the brain regions
involved in processingsocial information and self-relevant
information (e.g., Mitchell et al.,2005). This convergence was also
evident in our study, wherein self-conscious emotions, once again,
activated the mPFC.
However, it is important to note that self-conscious emotionsand
social-emotions are not one and the same. An emotion can
beinherently inter-personal without requiring any self-focus or
self-reflection; for example, anger is clearly an inter-personal
emotionin that it is typically directed towards other people, yet
it does notrequire much self-reflection. Indeed, contrasting the
widely ac-knowledged “inter-personal emotions” (i.e., guilt, pride,
anger)with a non- inter-personal emotion (joy) revealed
activationwithin a network of regions involved in social-cognition,
whichincludes the precuneus, the posterior superior temporal
sulcus(pSTS) extending to the temporo-parietal junction and to
thetemporal pole, as well as activation within the mPFC (Van
Over-walle and Baetens, 2009).
These findings are consistent with those of several other
stu-dies investigating the correlates of inter-personal and
non-inter-personal emotions (e.g., Britton et al., 2006; Frewen et
al., 2011).Furthermore, they are somewhat consistent with two
previousstudies which investigated self-conscious emotion
processing (e.g.,Guilt – Takahashi et al., 2004; Pride – Takahashi
et al., 2008). Ourfindings further suggest that the superior
temporal sulcus andprecuneus activation evident in previous studies
on guilt and pridemight be related to the social interaction that
is inherent to theseemotions rather than the “self-referential”
aspects of self-con-scious emotions. In fact, our results show that
pSTS activation wasmost markedly associated with emotions of anger,
which is aninter-personal, but not a self-conscious emotion.
3.4. Basic emotions and embodied experience
It is argued that self-conscious and basic emotions are likely
todiffer with regards to the abstractness of mental
representationsthat they typically rely upon (Karsh and Eyal, 2015;
Agerströmet al., 2012). The self-construct is an entity that has
concrete-physical properties (e.g., “I have blonde hair”), as well
as im-material properties (e.g., “I am an honest person”; “I am
greedy”)that rely on an abstract-symbolic representational medium.
Incontrast, basic emotions are believed by some to exist in
animalsand infants, and thus may precede the emergence of
abstract-symbolic thought (e.g., Izard et al., 1995). Consequently,
basicemotions may rely to a greater extent upon phylogenetically
andontogenetically earlier-developed, modality-specific systems,
thatdo not necessarily employ abstract mental
representations,whereas self-conscious emotions are likely to rely
on the phylo-genetically-novel prefrontal cortex, which is involved
in the pro-cessing of more abstract mental representations (e.g.,
Badre et al.,2010; Mian et al., 2014; Straube et al., 2013; Cole et
al., 2011; Tanjiet al., 2007; Wang et al., 2010; But see, for
example, Barrett, 2006,for an approach according to which all human
emotion relies onabstract mental representation).
Our results show that processing basic emotions (i.e., anger
andjoy) compared with self-conscious emotions, resulted in
activationthroughout relatively phylogenetically and
ontogenetically earlier-developed regions of the cortex, namely in
visual and tactileprocessing areas (i.e., occipital cortex,
somatosensory cortex, andparahippocampal gyrus) and in the insular
cortex. Notably,whereas self-conscious emotions recruited the
frontal cortex, no
-
M. Gilead et al. / Neuropsychologia 81 (2016) 207–218216
frontal activations were evident for basic-emotion processing
(ascompared to self-conscious emotions). These findings are
con-sistent with the idea according to which self-conscious
emotionsare a relatively recent adaptation (Demoulin et al., 2004),
whereasbasic emotions originate early in our evolutionary ancestry
(Ek-man, 1992).
Furthermore, these findings are consistent with somatic
(e.g.,the James-Lange approach, e.g., Lang, 1994) and embodied
(e.g.,Niedenthal, 2007) theories of emotion which stress the
im-portance of bodily states in emotional experience. However,
theyalso imply that the association between somatic and
emotionalstates is particularly relevant to basic emotions, and
that self-conscious emotions may be more abstract and dis-embodied
innature.
3.5. Concluding comments
Research in recent years has begun to delineate the
neuralsystems that are involved in the processing of self-related
in-formation (e.g., Mitchell et al., 2006) and the processing of
affec-tively laden stimuli (e.g., Lindquist et al., 2012). However,
researchthat attempts to understand the neural systems involved in
theprocessing of self-conscious emotions has been surprisingly
scarce.By situating our investigation within a broader theory of
thefunctionality of self-conscious emotions, the current study
ad-dressed a major gap in the research into self-conscious
emotions,and provided an important piece of evidence for this
cumulativeendeavor. Of course, much more research into the
affective com-ponent of self-processing is clearly warranted, as it
may shed lighton the age-old mystery of self-reflection, as well as
provide a morerefined framework for understanding and treatment
ofpsychopathologies.
Acknowledgments
This work was supported by Grants from The Israel
ScienceFoundation to Nira Liberman (Grant no. 92/12) and to Tal
Eyal(Grant no. 923/09), by the I-CORE Program of the Planning
andBudgeting Committee and The Israel Science Foundation (Grantno.
51/11) and by a research Grant from the Israeli FoundationTrustees
to Maayan Katzir (Fund for Doctoral Students no. 30).Michael Gilead
is financially supported by fellowships from theFulbright and
Rothschild Foundations.
Appendix A: Stimuli
The Appendix presents the 16 stimuli used for each
emotion.Although each stimulus appeared both in the future
condition andin the past condition, for brevity, we present in this
appendix thestimuli either in their past form or in their future
form.
-
M. Gilead et al. / Neuropsychologia 81 (2016) 207–218 217
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Neural correlates of processing “self-conscious” vs. “basic”
emotionsIntroductionPsychological differences between
self-conscious and basic emotions1.2 Past research on neural
substrates of self-conscious emotions
The current studyMethodParticipantsMaterialsBehavioral
procedureImaging procedure
ResultsBehavioral resultsImaging dataTemporal
perspectiveSelf-conscious emotions (GuiltgtAnger)∩(PridegtJoy)Basic
emotions (AngergtGuilt)∩(JoygtPride)GuiltgtOther
emotionsPridegtOther emotionsAngergtOther emotionsJoygtOther
emotionsInterpersonalgtNon-interpersonal emotionsValenceROI
analysis
DiscussionSelf-conscious emotions and
self-regulationSelf-conscious emotions and self-related
processingSelf-conscious vs. social emotionsBasic emotions and
embodied experienceConcluding comments
AcknowledgmentsAppendix A: StimuliReferences