(PDF) Cognitive Framing Modulates Emotional Processing through Dorsolateral Prefrontal Cortex and Ventrolateral Prefrontal Corte

| | Received: 31 March 2020    Revised: 22 June 2020    Accepted: 2 July 2020 DOI: 10.1002/brb3.1761 ORIGINAL RESEARCH Cognitive framing modulates emotional processing through dorsolateral prefrontal cortex and ventrolateral prefrontal cortex networks: A functional magnetic resonance imaging study Ulrich Kirk1  | Lau Lilleholt2 | David Freedberg3,4 1 Department of Psychology, University of Southern Denmark, Odense, Denmark Abstract 2 Department of Psychology, University of Introduction: In this study, we show new evidence for the role of ventrolateral pre- Copenhagen, Copenhagen, Denmark frontal cortex-dorsolateral prefrontal cortex (VLPFC-DLPFC) networks in the cogni- 3 Department of Art History and Archaeology, Columbia University, New tive framing of emotional processing. York, NY, USA Method: We displayed neutral and aversive images described as having been sourced 4 Italian Academy for Advanced Studies, from artistic material to one cohort of subjects (i.e., the art-frame group; n  =  19), Columbia University, New York, NY, USA while identical images, this time identified as having been sourced from documentary Correspondence material (i.e., the doc-frame group; n = 20) were shown to a separate cohort. Ulrich Kirk, Department of Psychology, University of Southern Denmark, Odense, Results: Using functional magnetic resonance imaging (fMRI), we employed a lin- Denmark. ear parametric model showing that relative to the doc-frame group the art-frame Email:

group exhibited a modulation of amygdala activity in response to aversive images. Funding information The attenuated amygdala activity in the art-frame group supported our hypothesis Danish Agency for Science, Technology and Innovation (UK); Lundbeckfonden (UK) that reduced amygdala activity was driven by top-down DLPFC inhibition of limbic responses. A psychophysiological interaction (PPI) analysis demonstrated that VLPFC activity correlated with amygdala activity in the art-frame group, but not in the doc- frame group for the contrast [Aversive > Neutral]. Conclusion: The role of the VLPFC in cognitive control suggests the hypothesis that it alongside DLPFC insulates against embodied emotional responses by inhibiting au- tomatic affective responses. KEYWORDS amygdala, dorsolateral prefrontal cortex, emotion, framing, functional magnetic resonance imaging, ventrolateral prefrontal cortex 1 |  I NTRO D U C TI O N interesting, gratifying, and worth living. For decades' researchers have sought to understand how we process emotional stimuli and No single aspect of our mental life is more profound than our abil- regulate emotional experiences, an endeavor that has provided ity to experience emotions. Emotions are what makes our lives us with numerous valuable insights. Furthermore, studies have The peer review history for this article is available at https://publo​ns.com/publo​n/10.1002/brb3.1761. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. © 2020 The Authors. Brain and Behavior published by Wiley Periodicals LLC. Brain and Behavior. 2020;00:e01761.  | wileyonlinelibrary.com/journal/brb3     1 of 8 https://doi.org/10.1002/brb3.1761 | 2 of 8       KIRK et al. investigated how contextual factors influences the way in which underpinnings of cognitive reappraisal are associated with increased emotional stimuli are processed (Aldao, 2013; Aldao & Tull, 2015; activation in the lateral and medial prefrontal regions and decreased Gross, 2015). activation in the amygdala and medial orbitofrontal cortex. Similarly, In one of the studies examining this question, Gerger, Leder, and both Urry et al. (2006) and Delgado, Nearing, LeDoux, and Phelps Kremer (2014) illustrated that framing emotional stimuli as art pro- (2008) showed that cognitive reappraisal influences amygdala ac- foundly alters people's emotional experience. More specifically, the tivity through connections to regions of the ventromedial PFC. results from this study indicated that framing highly aversive pic- Furthermore, Kanske, Heissler, Schönfelder, Bongers, and Wessa tures as art decreases the intensity of emotional experiences and (2011) found that both cognitive reappraisal and distraction strat- allow people to appraise aversive stimuli more positively. Although egies rely on control areas in the medial and dorsolateral prefrontal Gerger et al. (2014) provide evidence that framing aversive stimuli and inferior parietal cortex, and leads to decreased activity in the as art alters the way in which people appraise and experience aver- bilateral amygdala. sive stimuli, very little is known about the neurobiological under- In almost all studies on emotion processing and regulation, sub- pinnings responsible for this apparent framing effect. The purpose jects are either specifically instructed to employ a reappraisal strat- of the present study is therefore to investigate how highly aversive egy or utilize attentional control in order to effectively alter the pictures framed as art are processed compared to identical pictures emotional impact of a stimulus, or to use a “natural” mode of perceiv- framed as documentary photographs by using functional magnetic ing the emotionality of the stimulus, without any specific instruction. resonance imaging (fMRI). As opposed to this, the present study extends this line of research It is assumed that framing stimuli as art provides the cognitive by investigating whether aversive pictures framed as art promote system with cues on how to process and respond to objects with elevated levels of neural activity in regions responsible for top-down artistic qualities 4. For instance, Leder and colleague (Leder, Belke, appraisal processes. Using aversive and neutral images taken from Oeberst, & Augustin, 2004) argue that framing stimuli as art give rise IAPS database (Lang, Bradley, & Cuthbert, 1997), we told one group to certain expectations such as the belief that one will have a grati- of subjects that they were viewing images displayed in museums of fying and positive emotional experience. Furthermore, scholars have contemporary art, while the other group was told that the images argued that framing stimuli as art enables a specific style of percep- were sourced from documentary material and thus depicted real-life tion which allow people to adopt a more distanced perspective on events. Thus, in order to probe the cognitive modulation of the emo- what is depicted and appraise the aesthetic qualities of the artwork tional experience, we changed the cognitive frame in which the same (Cupchik, 2002). In support of this view, multiple studies have shown highly arousing stimuli were presented. that people engage in more elaborative processing (Nadal, Munar, Following the literature on emotion processing and regulation Capo, Rossello, & Cela-Conde, 2008) and attend more to stylistic and (Delgado et  al.,  2008; Kanske et  al.,  2011; Ochsner et  al.,  2002; formal properties of visual stimuli, when percepts are placed in an Urry et al., 2006), we hypothesized that amygdala activity in re- art context (Cupchik, Vartanian, Crawley, & Mikulis, 2009; Jacobsen, sponding to aversive images would be attenuated in the case of Schubotz, Höfel, & Cramon, 2006; Kirk, Skov, Hulme, Christensen, & subjects belonging to the art-frame group, while in the case of the Zeki, 2009). Lastly, placing visual stimuli in an art context signals that doc-frame group such activity would be comparatively elevated. the visual content is just a figment of the artist's imagination and can Furthermore, in line with previous research (Delgado et  al.,  2008; be considered as fictional (Gerger et al., 2014). Kanske et al., 2011; Ochsner et al., 2002; Urry et al., 2006), we hy- Previous research indicates that appraising an aversive stim- pothesized that the emotional response of subjects in the art-frame ulus as fictional rather than realistic profoundly reduces its emo- group would be modulated by increased activity in regions associ- tional impact (Mocaiber et al., 2010, 2011). Furthermore, it has ated with top-down appraisal processes, such as dorsolateral pre- been shown that adopting a psychological distance from aversive frontal cortex (DLPFC). Finally, given that DLPFC and ventrolateral stimuli reduces signs of arousal and bodily emotional expressions prefrontal cortex (VLPFC) have been shown to be interconnected (Gross, 1998; Lazarus & Alfert, 1964; Speisman, Lazarus, Mordkoff, both in nonhuman primates (Barbas, 2000; Barbas & Pandya, 1989; & Davison, 1964). Hence, evidence suggest that placing stimuli in an Yeterian, Pandya, Tomaiuolo, & Petrides, 2012) and humans (Goulas, artistic context fosters appraisal processes that allows the observer Uylings, & Stiers, 2012), we further hypothesized that VLPFC along- to suppress his or her initial emotional reaction and judge aversive side DLPFC would be associated with decreased amygdala activity in stimuli more positively (Gerger et al., 2014). Consequently, it seems the art but not the doc-frame group. plausible that framing aversive stimuli as art leads to increased neu- ral activity in brain regions associated with top-down appraisal pro- cesses, thereby allowing the observer to perceive aversive stimuli 2 | M ATE R I A L A N D M E TH O DS differently and have a positive emotional experience. Neuroimaging studies have shown that several brain regions 2.1 | Subjects play an important role in top-down appraisal processes, such as cognitive reappraisal and attentional control. For instance, Ochsner, To test our hypotheses, we enrolled thirty-nine subjects in an fMRI Bunge, Gross, and Gabrieli (2002) found that the neurobiological paradigm and divided them into two groups: an art-frame group and KIRK et al. |       3 of 8 a doc-frame group. The art-frame group consisted of 19 subjects and 7 = extremely frightening). We used an event-related fMRI design. the doc-frame group of 20 subjects. The art-frame group included On each trial, an image was presented for 5 s, followed by an in- eight women and 11 men (mean age 22.7), while the doc-frame ter-trial interval for 4–14 s. The images were presented at a screen group included nine women and 11 men (mean age 24.2). The two resolution of 1,024  ×  768 pixels and centered in a 500  ×  500- groups did not differ in terms of mean age or gender distribution and pixel resolution surrounded by a black background. Stimuli were randomly assigned to either the art or the doc-frame group. were presented, and responses collected using NEMO (Human None of the subjects were educated in the arts. All subjects had nor- Neuroimaging Lab, Virginia Tech Carilion Research Institute). The mal or corrected-to-normal vision, and none had a history of neu- stimuli were back-projected via an LCD projector onto a transpar- rological or psychiatric disorders. All procedures and experiments ent screen positioned over the subjects' head and viewed through reported involving human subjects were approved and conducted in a tilted mirror fixed to the head coil. accordance with the Institutional Review Board of Virginia Tech. All volunteers in the study participated in the experimental tasks after giving informed consent. 2.3 | fMRI data acquisition The anatomical and functional imaging was performed using iden- 2.2 | fMRI task tical 3 Tesla Siemens Trio scanners. High-resolution T1 weighted scans were acquired using an MPRAGE sequence (Siemens). 2.2.1 | Image-viewing paradigm Functional imaging used an EPI sequence with a repetition time (TR) of 2,000 ms, echo time (TE) = 30 ms, flip angle = 90°, 220 mm All subjects were scanned in a passive task involving viewing neu- field of view (FOV), 64 × 64 matrix. Functional slices were oriented tral and aversive IAPS images. Prior to scanning, subjects in the 30° superior-caudal to the plane through the anterior and posterior two groups were given a different set of instructions. The art- commissures in order to reduce signal dropout due to magnetic field frame group received the following instructions: “Today you will in-homogeneities (Deichmann, Gottfried, Hutton, & Turner,  2003). be viewing images of artwork while you undergo an MRI scan. Each functional image was acquired in an interleaved way, compris- Some of these images have a very strong emotional effect. Inside ing 34 4 mm axial slices for measurement of the blood oxygenation the scanner you will be presented with 80 photographs that have level-dependent (BOLD) effect (Ogawa, Lee, Kay, & Tank,  1990), been exhibited in contemporary art museums as works of art.” yielding 3.4 mm × 3.4 mm × 4.0 mm voxels. Subjects in the doc-frame group received the following instruc- tions: “Today you will be viewing photographs while you undergo an MRI scan. Some of these images have a very strong emotional 2.4 | fMRI data analysis effect. Inside the scanner you will be presented with 80 photo- graphs that depict real-life events.” Image preprocessing and data analysis were performed using SPM8 In the scanner, subjects were presented with 80 IAPS images (Wellcome Trust Centre for Neuroimaging). The preprocessing pro- selected from the IAPS database (Lang et al., 1997). Specifically, cedures have been described in our previous work (Kirk, Pagnoni, subjects were presented with images belonging to two emotional Hétu, & Montague, 2019). Briefly, preprocessing steps included mo- categories, namely 40 aversive images (valence: mean 2.9, range tion correction, co-registration, slice timing, normalization, and spa- 1.7–3.9; arousal mean 5.5, range 3.5–7.4) and 40 neutral images tial smoothing. For the analysis, a general linear model (GLM) was (valence: mean 5.1, range 4.23–5.9; arousal mean 3.1, range 1.7– applied to the fMRI time series where image onset was modeled 5.2). The procedure was presented in a pseudorandomized fashion, as single impulse response functions including image duration and and picture order was counterbalanced across subjects. During the then convolved with the canonical hemodynamic response function scanning session, subjects were instructed to passively view the (HRF). images. Postscanning, subjects were asked to complete an un- A parametric regression analysis was used (Büchel, Holmes, expected task of behavioral rating of the images, while making a Rees, & Friston, 1998; Phan et al., 2004) that allowed to model lin- self-paced subjective aversion rating using a Likert scale (1–7). It ear 1st order hemodynamic responses using orthogonalized polyno- is important to emphasize that subjects were not instructed about mial expansions. This was performed for each of the two conditions this subsequent behavioral rating-task prior to the scanning ses- (aversive and neutral images) using subject-specific behavioral rat- sion. The same images were presented to both groups with the ings for each image. Residual effects of head motion were corrected only difference being the initial framing of the images presented for by including the six estimated motion parameters for each sub- as either art or documentary. In the behavioral task, the images ject as regressors of no interest. The procedures for first-level and were displayed in a randomized order compared with the scanning second-level, random effects (RFX) analysis have been described session. Prior to the behavioral task, subjects were specifically in- in our previous work (Kirk et al., 2009). The statistical results given structed to rate how frightening each images were perceived in were based on a single-voxel t-statistics corresponding to p  <  .05 the moment on a scale ranging from 1 to 7 (1 = not frightening; corrected for multiple comparisons using the false discovery rate | 4 of 8       KIRK et al. statistic (FDR) with an extent threshold of >10 voxels. The co-or- dinates of all activations are reported in MNI space. Data were dis- played using the xjView toolbox. For the functional connectivity analysis, we implemented psycho- physiological interaction analysis (PPI) (Friston et al., 1997). The PPI assess changes in functional connectivity between the seed region of the right amygdala and other brain regions whose activity covary with the amygdala. The PPI employed a regressor representing the decon- volved time series of neural activity within a 4-mm sphere centered in the right amygdala (x,y,z = 20 –2 –16), which constituted the phys- iological variable. A second regressor representing the psychological F I G U R E 1   Mean behavioral responses. In the scanner, variable, specifically the contrast [Aversive > Neutral]. Finally, a third participants were presented with the IAPS images that were regressor representing the cross product of the previous two (the PPI displayed for 5 s each during a passive scanning session. In a term). The model also included motion parameters as regressors of subsequent behavioral session, participants provided self-paced fear ratings on a Likert-type scale for each image. Participants were no interest. The PPI was carried out in each subject and entered into not informed about the behavioral task until after the scanning random effects analysis separately for each of the two groups. session. The mean rating and standard error (SE) for the art-group aversive images were 0.31 (0.19) and neutral images −0.94 (0.20). The mean rating and SE for the doc-group aversive images were 3 |   R E S U LT S 0.48 (0.14) and neutral images −1.31 (0.20). Statistical analysis showed no significant difference between the two groups 3.1 | Behavioral results that the aversive images in the art-frame group drove the reactivity We were interested in seeing if there were differences in behav- compared to the aversive images in the doc-frame group (Figure 2, ioral ratings for aversive and neutral images across the art and bottom). There were no differences across the groups with regard doc-frame groups. Note that the behavioral ratings of the images to the neutral images. These results suggest that the art frame does were collected post hoc and thus not during the scanning run indeed recruit neural activity in regions associated with top-down (see Methods). To further examine if there were any differences appraisal and cognitive control such as the DLPFC. between the two groups a mixed ANOVA was used to inspect For the opposite contrast [doc aversive > art aversive], we found group (art-frame, doc-frame) by image type (aversive, neutral). that the reactivity to fearful images in the right amygdala (x,y,z = 20 There was no significant interaction between group and rating (F –2 –16; p < .05, FDR-corrected; voxels = 43) exhibited a steeper slope (1,37) = 2.01; p = .165). Similarly, there was not a significant main related to the parametric analysis in the doc-frame group (Figure 3, effect for group (F (1,37) = 0.26; p = .615). However, there was a top). Furthermore, using average β-estimates extracted from the substantial main effect for image type (F (1,37) = 65.63, p < .01, right amygdala showed that the activation was driven by modulated 𝜂 2p  =  0.639) with both groups rating the aversive images more reactivity in this region in the doc-frame group (Figure 3, bottom). frightening than the neutral images (Figure 1). We then assessed functional connectivity in the amygdala region of interest across the entire time series for the contrast [aversive > neu- tral images] separately for the two groups (Figure  4). An analysis of 3.2 | fMRI results functional connectivity implemented as an psychophysiological in- teraction analysis revealed that the art-frame group demonstrated We aimed to identify neural regions that exhibited a modulation a strong positive coupling with one region, in a whole brain analysis, across the two groups when presented with aversive images. We namely the right ventrolateral prefrontal cortex (VLPFC) (x,y,z = 48 38 used a parametric regression model that identifies areas of the –6; p < .05, FDR-corrected). This result suggests that in the art-frame brain that identifies areas of the brain whose activation amplitude group, viewing aversive images compared to neutral ones is associated scales linearly with subjective ratings. Note that we only report the with a stronger amygdala-VLPFC functional connectivity. linear effects (1st order polynomial expansions) and not the simple In contrast, the doc-frame group showed a positive coupling with average effects (i.e., zero order polynomial expansions) in that the the visual cortex, specifically the lingual gyrus BA 18 (x,y,z = 3 –71 1; latter did not yield significant activations even when lowering the p < .05, FDR-corrected). Such a strong correlation between a visual threshold (p  <  .001, uncorrected). When computing the contrast cortical region and the amygdala in processing aversive images may [art aversive >  doc aversive], we found that the reactivity to fear- preclude a mode of disengagement of the aversive material depicted ful aversive images in the left DLPFC (x,y,z  =  −40 34 44; p  <  .05, in the doc-frame group. FDR-corrected; voxels = 35) exhibited a steeper slope related to the Finally, we formally assessed the differences between the two parametric analysis in the art-frame group (Figure 2, top). We sub- PPI contrasts, albeit the did you yield significant voxels, even when sequently extracted β-estimates from the DLPFC region and found lowering the threshold (p < .001, uncorrected). KIRK et al. |       5 of 8 F I G U R E 2   Neural activity in DLPFC in the main effect [art aversive > doc aversive]. Top: DLPFC display linear increase with fear responses for aversive images in the art-frame group compared with the doc-frame group. Bottom: Average beta values extracted for each group in the left DLPFC display better fit in terms of beta values for aversive images in the art-frame group relative to the same images presented to the doc-frame group. There are no differences between groups for neutral images. Error bars indicate SE F I G U R E 3   Neural activity in amygdala in the main effect [doc aversive > art aversive]. Top: Right amygdala display linear increase with fear responses for aversive images in the doc-frame group compared with the art-frame group. Bottom: Average beta values extracted for each group in the right amygdala display higher beta values for aversive images in the doc-frame group relative to the same images presented to the art-frame group. There are no differences between groups for neutral images. Error bars indicate SE 4 | D I S CU S S I O N framed as documentary by using fMRI. We hypothesized that the amygdala would reflect a steeper slope in the subjects belonging The purpose of this study was to investigate how highly aversive to the art-frame group compared to the doc-frame group. In line pictures framed as art are processed compared to identical pictures with this, we further hypothesized that amygdala activity would | 6 of 8       KIRK et al. F I G U R E 4   Group-specific changes in effective connectivity for [aversive > neutral images]. Psychophysiological interaction analysis (PPI) displaying increased coupling between the right amygdala seed region and the right VLPFC in the art-frame group. By contrast, the doc- frame group displayed greater coupling between the amygdala seed region and the lingual gyrus in the visual cortex exhibit stronger coupling by VLPFC alongside DLPFC in the case of such as fear, allowing for a more distanced and reflective perspec- the art but not the doc-frame group. In support of our hypotheses, tive. In a similar vein, Scherer argues that artistic stimuli do not fos- the fMRI data showed that amygdala activity was down-regulated ter what he terms utilitarian emotions but rather what he refers to while DLPFC was up-regulated in subjects belonging to the art- as aesthetic emotions (Scherer,  2004, 2005). Utilitarian emotions frame group. Conversely, for subjects belonging to the doc-frame correspond to everyday emotions such as fear, joy anger, disgust, group amygdala activity exhibited a steeper slope in the parametric and sadness. These emotions are utilitarian in the sense that they analysis. Moreover, framing aversive images as art was found to facilitate adaptive response (e.g., fight, flight) to events that have im- be associated with functional connectivity between amygdala and portant consequence for our survival and personal well-being. On VLPFC, whereas the doc-frame led to increased connectivity be- the contrary, aesthetic emotions do not have an adaptive function tween amygdala and visual cortex. Taken together, these findings and are not shaped by the appraisal of the artwork's ability to satisfy indicate that images framed as art lead to reactivity in regions as- physical needs or further current goals. Rather, aesthetic emotions, sociated with top-down appraisal and cognitive control compared such as fascination, admiration, and rapture, are produced by the to images framed as documentary photographs. Consequently, the appreciation of the intrinsic qualities of the artwork. In line with findings from this study suggest that framing highly aversive im- Scherer's distinction between utilitarian and aesthetic emotions, it ages as art, result in stronger VLFPC coupling of amygdalic activity. can thus be argued that activation of VLPFC-DLPFC inhibits adap- However, it should be noted that the formal comparison between tive utilitarian emotions, allowing for a more distanced and reflective the PPI contrast for the two groups did not yield significant voxels, perspective in which aesthetic emotions can arise. which may suggest insufficient statistical power. In agreement with previous neuroimaging studies, the fMRI Although the fMRI data suggest that framing aversive visual data suggest that DLPFC is partially responsible for top-down ap- stimuli as art recruit neural activity in regions associated with top- praisal processes (Delgado et al., 2008; Kanske et al., 2011; Urry down appraisal processes, this was not reflected in the behavioral et al., 2006). Specifically, the fact that reactivity in DLPFC was rating task. Specifically, no significant difference was observed be- found to be associated with amygdala reactivity strongly indi- tween behavioral ratings of aversive IAPS images across the art and cates that these brain regions are involved in top-down appraisal doc-frame groups. One possible explanation for this result is the processes aimed at regulating automatic emotional responses well-known fact that small sample sizes yield low statistical power (Delgado et  al.,  2008; Kanske et  al.,  2011; Ochsner et al., 2002; (Button et al., 2013). Urry et al., 2006). Interestingly, the PPI analysis showed that sub- The fMRI data presented here are consistent with previous re- jects belonging to the art-frame, but the not doc-frame group search which suggest that placing visual stimuli in an artistic con- demonstrated a significant positive coupling between VLPFC and text fosters top-down appraisal processes aimed at inhibiting innate amygdala. This is particularly interesting as it suggests that VLPFC emotional responses. In extension, this result lends additional sup- plays an important role in top-down appraisal processes by inhib- port to the empirical observation that appraising aversive stimuli as iting automatic affective responses. fictional rather than realistic reduces its emotional impact (Mocaiber Although most appraisal theorists agree that context plays an et al., 2010, 2011). Accordingly, framing aversive images as art seems important role in emotion regulation and appraisal processes, very to provide the cognitive system with cues that fosters a specific style few studies have investigated the effects of contextual framing of perception allowing the observer to adopt a more distanced and (Aldao, 2013; Aldao & Tull, 2015; Gross, 2015). The results presented reflective perspective of what is depicted. A possible explanation for here are thus important: First because they indicate that contextual this finding is that artistic stimuli, as opposed to real-world stimuli, factors influence how people spontaneously process emotional do not pose any risk for our survival and personal well-being, due stimuli; second because they show that people automatically engage to its fictitious nature (Gerger et al., 2014). Hence, framing aversive in more effective emotion regulation strategies when highly aversive stimuli as art eliminates the need for an adaptive emotional response percepts are placed in an art context. KIRK et al. |       7 of 8 5 | CO N C LU S I O N Cupchik, G. C., Vartanian, O., Crawley, A., & Mikulis, D. J. (2009). Viewing artworks: Contributions of cognitive control and percep- tual facilitation to aesthetic experience. Brain and Cognition, 70, Building on the experimental framework put forward here future 84–91. studies should strive to investigate how different contextual frames Deichmann, R., Gottfried, J., Hutton, C., & Turner, R. (2003). Optimized influence how people process highly emotional stimuli and indeed EPI for fMRI studies of the orbitofrontal cortex. NeuroImage, 19, 430–441. https://doi.org/10.1016/S1053​-8119(03)00073​-9 asking about subjects emotional experience (Kron, Goldstein, Lee, Delgado, M. R., Nearing, K. I., LeDoux, J. E., & Phelps, E. A. (2008). Neural Gardhouse, & Anderson,  2013; Kron, Pilkiw, Banaei, Goldstein, & circuitry underlying the regulation of conditioned fear and its rela- Anderson, 2015). Investigating this is an important next step in the tion to extinction. Neuron, 59, 829–838. https://doi.org/10.1016/j. study of emotions which would provide us with a more detailed un- neuron.2008.06.029 derstanding of the relationship between context and emotion pro- Friston, K. J., Buechel, C., Fink, G. R., Morris, J., Rolls, E., & Dolan, R. J. (1997). Psychophysiological and modulatory interactions in neu- cessing and regulation. roimaging. NeuroImage, 6, 218–229. https://doi.org/10.1006/ nimg.1997.0291 AC K N OW L E D G M E N T S Gerger, G., Leder, H., & Kremer, A. (2014). Context effects on emo- This work was supported by an International Network Programme tional and aesthetic evaluations of artworks and IAPS pictures. Acta Psychologica, 151, 174–183. Grant from the Danish Agency for Science, Technology and Goulas, A., Uylings, H. B. M., & Stiers, P. (2012). Unravelling the intrinsic Innovation (UK), and a stipend for sabbatical leave from functional organization of the human lateral frontal cortex: A parcel- Lundbeckfonden (UK). lation scheme based on resting state fMRI. Journal of Neuroscience, 32, 10238–10252. Gross, J. J. (1998). Antecedent-and response-focused emotion regula- C O N FL I C T O F I N T E R E S T tion: Divergent consequences for experience, expression, and physi- The authors declare no conflict of interest. ology. Journal of Personality and Social Psychology, 74, 224. Gross, J. J. (2015). Emotion regulation: Current status and future pros- AU T H O R C O N T R I B U T I O N pects. Psychological Inquiry, 26, 1–26. https://doi.org/10.1080/10478​ 40X.2014.940781 UK and DF designed the study; UK and DF performed the research; Jacobsen, T., Schubotz, R. I., Höfel, L., & Cramon, D. Y. V. (2006). Brain UK and DF analyzed data; and UK, DF, and LL wrote the article. correlates of aesthetic judgment of beauty. NeuroImage, 29, 276–285. https://doi.org/10.1016/j.neuro​image.2005.07.010 DATA AVA I L A B I L I T Y S TAT E M E N T Kanske, P., Heissler, J., Schönfelder, S., Bongers, A., & Wessa, M. (2011). How to regulate emotion? Neural networks for reappraisal and dis- The results generated during the current study are available from traction. Cerebral Cortex, 21, 1379–1388. https://doi.org/10.1093/ the corresponding author on reasonable request. cerco​r/bhq216 Kirk, U., Pagnoni, G., Hétu, S., & Montague, R. (2019). Short-term mind- ORCID fulness practice attenuates reward prediction errors signals in the Ulrich Kirk  https://orcid.org/0000-0002-2121-2016 brain. Scientific Reports, 9, 6964. Kirk, U., Skov, M., Hulme, O., Christensen, M. S., & Zeki, S. (2009). Modulation of aesthetic value by semantic context: An fMRI study. 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