Step

Fig. 2. (a) Adaptive CS was significantly related to happiness, but did not show an association with trait anxiety or depression. (b) Maladaptive CS was positively related to trait anxiety and depression, but negatively related to happiness.

4.4. Amygdala-seed FC during passive response to negative emotion mediated the association between maladaptive CS and negative emotional reactivity

Activation maps demonstrated increased activation in the right parahippocampal gyrus extending to amygdala, bilateral hippocampus/ thalamus, middle occipital gyrus (MOG), posterior cingulate cortex (PCC) and superior frontal gyrus (SFG) and decreased activation in the bilateral superior temporal gyrus and medial frontal gyrus in the

Table 2

Hierarchical regression analysis predicting negative emotional reactivity ratings as a function of age, gender, trait anxiety, depression, happiness and maladaptive CS.

variability

Step 1

Intercept Age Gender Adjusted R2

Step2

Intercept Age Gender Trait anxiety Depression Happiness Adjusted R2

Step3

Intercept Age Gender Trait anxiety Depression Happiness

Maladaptive CS

Adjusted R2 Estimate

− 1.691 0.151 0.488 0.125

0.780 0.133 0.493 − 0.026 0.006 − 0.010 0.149

1.445 0.117 0.406 − 0.031 − 0.005 − 0.008

0.608

0.228 SE

0.923 0.047 0.140

1.298 0.047 0.139 0.013 0.014 0.005

1.247 0.045 0.134 0.013 0.014 0.004

0.150

p-Value

0.069 0.002 0.001

0.549 0.005 0.001 0.051 0.650 0.035

0.248 0.011 0.003 0.014 0.694 0.062

<0.001

WatchNeg > WatchNeu contrast. (Supplementary Tables S2, Figure S1). Within the amygdala ROI analysis in the WatchNeg > WatchNeu contrast, there was increased activity in the right amygdala (tmax = 9.77, k = 75, peak MNI coordinate: x = 24, y = − 3, z = − 21 at p < 0.05, FWE-SVC corrected) and left amygdala (tmax = 9.39, k = 64, peak MNI coordinate: x = − 21, y = − 6, z = − 18 at p < 0.05, FWE-SVC corrected). The averaged extracted betas for the bilateral amygdala were positively linked to negative ratings in the WatchNeg condition (r = − 0.208, p = 1.30 × 10− 2). Results from multiple regression analysis showed no effects of maladaptive CS on brain activation changes.

We tested whether maladaptive CS use was related to amygdala-seed FC during viewing of the negative pictures. The seeds region consisted of all voxels in an 8 mm radius around the peak of the activation of bilateral amygdala in the contrast of WatchNeg > WatchNeu. After controlling for the effects of age and sex, multiple regression analysis revealed that maladaptive CS was negatively correlated with FC between the right amygdala and areas in the right middle frontal gyrus (MFG, t = − 4.38, k = × 210, peak MNI coordinate: x = 42, y = − 4, z = 50, mass p-FWE = 7.53 10− 3). Similarly, maladaptive CS use was linked to lower FC between the left amygdala and areas in the right MFG (t = − 4.64, k = 114, peak MNI coordinate: x = 40, y = 2, z = 62, mass p-FWE = 3.13 × 10− 3 , Fig. 3. a1). Next, we tested whether amygdala-MFG FC mediated the relation between maladaptive CS and negative emotional reactivity indicated by reduced ratings during WatchNeg > WatchNeu, by computing 5000 bootstrap resamples. Bilateral amygdala-MFG connectivity betas were extracted from all voxels in the gPPI analysis and averaged for each participant. Mediation analysis showed that maladaptive CS scores were negatively related to the FC strength of the amygdala and MFG (r = − 0.35, p < 0.001) but positively associated with subjective ratings of WatchNeg > WatchNeu (r = 0.23, p = 0.004). The mediation analysis showed a significant indirect effect of amygdala-MFG connectivity on the relation between maladaptive CS and negative emotional reactivity. The bootstrapped indirect effect was 0.08, with the 95% confidence interval ranging from 0.01 to 0.17 (Fig. 3. a2). Altogether these tests supported the hypotheses that maladaptive CS is associated with

Table 3

ROIs used for gPPI analyses.

Side

WatchNeg > WatchNeu

Amygdala L R

RegulateNeg > WatchNeg

Superior Frontal Gyrus L R

Inferior Frontal Gyrus L R

Middle Frontal Gyrus L R

Middle Temporal Gyrus L R

Inferior Parietal Gyrus/ Supramarginal L

Gyrus R

Medial orbitofrontal cortex L R ROIs: region of interest, BA: brodmann area.

MNI Coordinates x y z

− 21 24

− 9 21 − 45 33 − 42 45 − 54 54 − 51

57 − 6 6 − 6 − 3

18 54 36 21 15 24 –33 − 27 − 57

− 54 52 57 − 18 − 21

63 30 − 6 − 12 48 42 − 6 − 9 48

45 − 19 − 16 BA

BA6 BA10 BA6 BA47 BA6 BA8 BA21 BA20 BA39

BA40 BA11 BA11

amygdala-PFC connectivity during negative emotional reactivity, and this functional connectivity mediated the association between maladaptive CS and negative reactivity.

4.5. Maladaptive CS is associated with lower IFG-amygdala FC during negative emotion regulation

Analysis of overall activation patterns during downregulation of negative feelings showed increased activation in regions of the bilateral tion of negative emotion. Results revealed that FC was positively asso-

SFG, rostral inferior frontal gyrus (IFG), MFG, middle temporal gyrus regions of the parahippocampal gyrus (P < 0.05, FWE-corrected). (Supplementary Tables S3, Figure S3). There were no significant results from our multiple regression analysis, which indicated no effects of maladaptive CS and adaptive CS on brain activation changes during RegulateNeg > WatchNeg contrast. To examine the potential links between CS use and FC during emotion regulation, we implemented PPI analyses with the prefrontal lobe regions. PPI analysis results revealed that maladaptive CS scores were related to lower FC between the left IFG and areas in the right subcortical region (t = − 4.24, k = 170, peak MNI coordinate = 30, − 16, − 24 at mass p-FWE = 1.01 × 10− 2), such as the amygdala, hippocampus and parahippocampal gyrus. The association between extracted values of FC strength for the IFG-amygdala and maladaptive CS is visualized in Fig. 3. b.

4.6. Adaptive CS is associated with higher OFC-subcortical FC during negative emotion regulation

Based on the same ROI seeds, we also investigated whether adaptive CS use was linked to stronger FC between the prefrontal cortex and subcortical regions, which might facilitate increased top-down regulaand inferior parietal gyrus (IPG), as well as the decreased activity in

ciated with adaptive CS for the left OFC seed in the lateral and medial PFC, including for connectivity to the left ACC, paracingulate gyrus (t = 5.85, k = 1339, peak MNI coordinate: x = − 6, y = 30, z = 24 at mass p-FWE = 2.09 × 10− 5), a region in the right MCC, precentral gyrus and SMA (t = 5.02, k = 355, peak MNI coordinate: x = 6, y = − 12, z = 36 at mass p-FWE = 6.04 × 10− 4), and a region of the right putamen and caudate (t = 4.32, k = 327, peak MNI coordinate: x = 22, y = 6, z = 18 at mass p-FWE = 8.08 × 10− 4). The associations between adaptive CS and extracted FC strength of the left OFC-ACC/SFG, OFC-PCC/SMA and OFC-putamen/caudate connectivity are visualized in Fig. 4 a.

Fig. 3. (a1) Maladaptive CS was negatively correlated with FC between the bilateral amygdala and right MFG during passive negative stimulus watching (P < 0.05, whole-brain FWE-corrected), and (a2) The extracted values of FC strength of amygdala-MFG mediated the association between maladaptive CS and reduced ratings during WatchNeg > WatchNeu. (b1) During down regulation processing, maladaptive CS use was negatively correlated with FC between the IFG and right amygdala (voxel-wise threshold level of p < 0.001 uncorrected and a cluster-level threshold of p < 0.05 FEW). CS = coping style, FWE = family-wise error, MFG = middle frontal gyrus, IFG = inferior frontal gyrus.

5. Discussion

The present study provides insight into whether a two-factor model of coping styles was associated with individual differences in trait negative emotion, reactivity and downregulation of negative emotions, as well as underlying patterns of neural activity and connectivity. Variation in the specific coping strategies that individuals reported using in daily life was highly related to one another, loading onto latent factors for maladaptive and adaptive CS. This two-factor model was further elaborated on by results that maladaptive CS were positively associated with individual anxiety and depression, while adaptive CS was only positively related to individual happiness, which was in line with previous studies (Hampel & Petermann, 2006; Mahmoud et al., 2012). As expected, our results showed that maladaptive CS (rumination, catastrophizing, worry, punishment and other blame) was associated with stronger negative emotional reactivity and decreased connectivity between the bilateral amygdala and MFG in response to unpleasant emotional stimuli. However, we did not find any associations between adaptive CS and regulatory ability at the behavioral level. These results partly supported our hypothesis that maladaptive CS has a significant influence on trait emotional experiences and state negative emotional reactivity. However, consistent effects of adaptive CS on emotional health were not found.

During the regulation stage of our fMRI task, maladaptive CS was negatively associated with connectivity between the left IFC and amygdala. On the other hand, adaptive CS (reappraisal, refocus on planning, positive reappraisal, putting into perspective and acceptance) was correlated with the FC strength of the OFC-ACC/SFG, OFC-MCC and OFC- putamen/caudate connections during the explicit emotional regulation by reappraisal. However, neither maladaptive nor adaptive CS showed any association with regulatory ability. Together these results partly supported our ideas that maladaptive and adaptive coping styles influence individual state negative emotion regulation and corresponding neural correlates.

5.1. Maladaptive CS and adaptive CS

Adaptive and maladaptive coping styles are not necessarily reciprocal or opposite one another (Aldao & Nolen-Hoeksema, 2012). Our results showed a significant positive correlation between adaptive and maladaptive CS, which can be better understood in light of previous research that suggested some people frequently use both adaptive and maladaptive strategies in daily life (Dixon-Gordon et al., 2015). For instance, individuals who are prone to experience stronger negative emotions generally may be more likely to engage in either maladaptive regulation strategies or adaptive CS, due to the greater total quantity (and intensity) of emotions that present for the possibility of regulation. Our results suggested that maladaptive CS have stronger associations with trait negative emotionality (e.g. trait anxiety and depression) while adaptive CS was only related with positive emotion (happiness) in this study. A potential explanation is that the effects of maladaptive CS on psychopathology are maladaptive most of the time, but the effects of adaptive CS may be more context-dependent (Aldao et al., 2010, 2014). Therefore, our results suggested that maladaptive CS play a critical role in negative emotional experience, while associations with adaptive CS use were less clear.

5.2. Maladaptive CS and negative emotional reactivity

Maladaptive CS, including high use of rumination, worry,

Fig. 4. Adaptive CS related to down regulation of negative emotions. Adaptive CS positively correlated with FC between the left OFC and left ACC, right MCC and right putamen/caudate during RegulateNeg > WatchNeg condition (voxel-wise threshold level of p < 0.001 uncorrected and a cluster-level threshold of p < 0.05 FEW). OFC = orbitofrontal cortex, ACC = anterior cingulate cortex, MCC = middle cingulate cortex.

catastrophizing, self-punishment and others-blame, have been suggested to be engaged with an attentional bias oriented toward negative feelings (Morrison & O’Connor, 2008; Romens & Pollak, 2012), exaggerated responses to negative stimuli (Rauch et al., 2000; Ray et al., 2009), and aggressive and negative views about the self (Salmivalli, 2001; Van Buren & Cooley, 2002). We did not find that maladaptive CS were related to higher levels of amygdala activity. When using the bilateral amygdala regions that activated during the WatchNeg condition as ROIs for gPPI analysis, we found that bilateral amygdala-MFG functional connectivity was associated with maladaptive CS. This FC mediated the association of maladaptive CS with negative emotion reactivity. Brain activity in the amygdala has been suggested to be recruited in negative stimulus response (Hamann & Mao, 2002; Murray, 2007), while brain activity in the region of the MFG has been suggested as a correlate of emotion reappraisal in daily life (Grecucci et al., 2013). Additionally, passive negative stimulus processing is usually synchronized with self regulation, reflected in recruitment of both the amygdala and ventral regions of the prefrontal cortex (M. L. Phillips et al., 2008; Wagner & Heatherton, 2013). Inferentially, decreased amygdala-PFC functional connectivity might play a role in automatic regulation during passive negative stimulus processing. Thus, assuming that maladaptive CS itself might be associated with more negative meaning generation and may involve inhibitory effects of spontaneous regulation (Moritz et al., 2016), individuals with a high tendency toward maladaptive CS might have more intense reactivity to any given negative stimulus.

5.3. CS and negative emotion downregulation

The activated regions in our reappraisal-strategy regulation task included the left and right ventrolateral MFG, lateral SFG, rostral IFG, MTG and the IPG, and suggested the engagement of selective attention, inhibition, cognitive control and conflict monitoring (Morawetz, Bode, Baudewig, et al., 2017; Motzkin et al., 2015), likely resulting in successful facilitation of emotion regulation. Following the assumption that maladaptive CS—which is characterized by lacking emotional control and by disengagement with negative stimuli—might be related to less emotion regulation success (Engen & Anderson, 2018), adaptive CS might be associated with greater emotion regulation success, since it might include forms of antecedent regulation with self-control-related valuation processes (Hare et al., 2009). We did not find that maladaptive and adaptive CS were linked to regulatory ability at the behavioral level but did find evidence for a relation at the neural level. In line with previous research (Kanske et al., 2011), the discrepant associations of CS with neural and behavioral measures might be explained by differing measurement sensitivities of functional connectivity and subjective ratings. The distinctive functional connectivity patterns linked to maladaptive CS and adaptive CS in the current study help reveal a clearer picture of how habitual coping styles might influence emotion regulation and associated neural systems.

We observed that in those with higher levels of maladaptive CS, left IFG/vlPFC, as a top-down-control-related PFC region, showed lower functional connectivity to subcortical regions, including the amygdala, hippocampus and parahippocampal gyrus. The left IFG plays an important role in selective attention and inhibition (H¨ ansel & von K¨ anel, 2008), cognitive control (Kohn et al., 2015; Ochsner et al., 2012) and inner speech (Messina et al., 2016) during the regulation process. From the cognitive control of emotion perspective (Silvers & Guassi Moreira, 2019), maladaptive CS could be interpreted as a potential inefficiency to engage prefrontal top-down circuitry upon the amygdala to modulate activity. Successful reappraisal involves both inhibiting the negative appraisal and generating a positive appraisal for a given emotional stimulus. A previous study suggested that co-activation of the vlPFC and amygdala might lead to successful reappraisal (Wager et al., 2008). Consistent with this view, the negative association with IFG-amygdala functional connectivity in the current study indicated that maladaptive CS might facilitate the generation of more negative appraisals during reappraisal of stimuli. However, previous findings have suggested that inverse amygdala-PFC connectivity was related to better regulation success using suppression (Lee et al., 2012). Similarly, inverse associations between amygdala and ventromedial PFC (vmPFC) were related to better regulation by using distraction in older adults (Urry et al., 2006). Since attentional deployment, cognitive change and response modulation are different stages of emotion regulation, the corresponding neural correlates and functional connectivity might be variable across these strategies (Morawetz, Bode, Derntl, et al., 2017). Additionally, different PFC functional connectivity patterns have been found between younger adults and older adults (Allard & Kensinger, 2014). Altogether, the inconsistent connectivity patterns associated with successful regulation across studies should be further explored in relation to the different strategies used in the experimental paradigms and with respect to the age of samples.

Adaptive CS might involve the suppression of negative stimuli- elicited retrieval, which is supported by the attentional control mechanism recruited by functional connectivity in PFC regions and the ACC (Depue et al., 2016; Guo et al., 2018). Our results suggested that dependent on levels of adaptive CS, the OFC showed greater functional connectivity with ACC, MCC, and putamen. The OFC seed in our study is considered anatomically synonymous with the vmPFC (Blair, 2007). Previous research suggested that appraisal and expression of negative emotion might recruit activity of dorsal-caudal regions in the ACC and mPFC, while ventral-rostral portions of the ACC and mPFC seem to have a direct regulatory role on emotional responses involving subcortical regions (Etkin et al., 2011; Li, Biswal, et al., 2019, Li, Duan, et al., 2019). A number of studies have revealed the important role of connectivity between the ACC/MCC and vmPFC in detecting emotion conflict and modulating activity in the amygdala (Banks et al., 2007; Egner et al., 2008; Etkin et al., 2006). This conflict-driven regulatory mechanism likely facilitates positive appraisal, which competes with the initial negative emotional appraisal during the valuation process. The MCC and putamen is involved in monitoring reward response behavior (Haber, 2017; Haruno & Kawato, 2006) and punishment avoidance (Vogt, 2009), and has been suggested to play a crucial role in facilitating intentional motor control over emotional responses (Felix et al., 2013). Adaptive strategies, such as reappraisal, likely rely upon conscious control over affect-related attention and involve recruitment of a top- down dorsal attention system from the dlPFC to ACC/MCC (Kohn et al., 2015; Ligeza et al., 2016). In line with this idea, we speculate that adaptive CS might include trait-like tendencies that involve attentional deployment and reward monitoring during emotion regulation.

5.4. Limitations

Some limitations in the present study should be considered in future research. First, the measurement of CS was calculated by self-report endorsement of various strategies or coping styles. We focused on the integrated correlation of coping style with individual reactivity and downregulation of negative emotions. However, this way of measuring coping styles might be limited to capturing coping strategies people usually engage in during daily life, rather than during any particular negative experience. Second, certain results of the current study warrent further investigation, such as the lack of a behavioral correlation between CS and regulatory ability and the neurofunctional correlates of coping styles during emotion regulation. Third, more elaborate experimental designs need to be used to investigate the specific cognitive processes underlying the effects of maladaptive CS on emotion regulation. For instance, whether maladaptive CS actively makes negative emotions more intense or whether it simply disrupts the use of adaptive CS strategies and subsequent regulatory success. Fourth, future research should draw attention to the interactions between different coping strategies and investigate how people develop different styles of coping through the process of interaction with their living environments. To gain a better understanding of the associations between the different

coping strategies used in daily life and the impact that context can have on CS, experience sampling methods could be used to capture variation in coping strategy engagement at multiple time points, as well as the influence of contextual information on strategy choices, such as the intensity or controllability of triggering events (Aldao et al., 2015). Considering these contextual factors would help to more fully understand the inherent adaptiveness of certain coping styles.

6. Conclusion

In the present study, we found that maladaptive CS plays a critical role in emotional well-being, showing stronger associations with trait and state negative emotions, when compared to adaptive CS. Delineating the associations between CS and emotional well-being could help us better understand protective and risk factors for mental health and provide effective intervention. In addition, our current work provides support for theories that emphasize the importance of frontal- subcortical interactions in successfully down-regulating negative emotions and show that these patterns of neural connectivity co-vary with individual differences in everyday use of coping strategies. Overall, this research helps to bridge the gap between our understanding of emotion downregulation in daily life and in laboratorial experiments.

Funding

This research was supported by the National Natural Science Foundation of China (31470981; 31571137; 31500885; 31600878; 31771231), Natural Science Foundation of Chongqing (cstc2015jcyjA10106). Scott Blain was supported by the National Science Foundation Graduate Research Fellowship Program (1348264).

CRediT authorship contribution statement

Xiaoqin Wang: Conceptualization, Investigation, Data curation, Formal analysis, Visualization, Writing - original draft. Scott D. Blain: Conceptualization, Methodology, Writing - review & editing. Dongtao Wei: Conceptualization. Wenjing Yang: Conceptualization, Writing - review & editing. Junyi Yang: Data curation. Kaixiang Zhuang: Methodology. Li He: Methodology. Colin G. DeYoung: Conceptualization, Methodology, Writing - review & editing. Jiang Qiu: Conceptualization, Investigation, Resources, Project administration.

Declaration of Competing Interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

Appendix A. Supplementary data

Supplementary data to this article can be found online at https://doi. org/10.1016/j.bandc.2020.105631.

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