Neuropsychologia 81 (2016) 207–218

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Neuropsychologia

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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: michael.gilead@gmail.com (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).

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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.

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)

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.

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(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

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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

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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

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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.

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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