Pain empathy, the capacity to resonate with, relate to, and share others’ pain, is an essential part of human experience. Among other functions, it motivates helping and cooperative behaviors and aids in learning to avoid harmful situations. Vicarious pain can be triggered by observing or imagining another individual’s painful state and can be elicited by multiple types of social cues, particularly the observation of an inflicted physical injury or a facial expression of pain (Decety and Ickes, 2009; Jauniaux et al., 2019; Vachon-Presseau et al., 2012; Yesudas and Lee, 2015). While stimuli depicting the noxious stimulation of body limbs [i.e. observation of noxious stimulation (NS) induced vicarious pain (NS vicarious pain)] provide objective cues about the sensory component of the observed pain, the observation of facial expressions of pain [i.e. facial expressions induced vicarious pain (FE vicarious pain)] is considered more subjective and indirect as the pain experience of the expresser needs to be interpreted by the observer (Hadjistavropoulos et al., 2011; Vachon-Presseau et al., 2012). Functional magnetic resonance imaging (fMRI) studies employing corresponding pictorial stimuli have identified distinct and common neural substrates of pain empathy across vicarious pain induction procedures (Jauniaux et al., 2019). For example, Vachon-Presseau et al., 2012 demonstrated that NS vicarious pain increased activity in core regions of the mirror neuron system, specifically inferior frontal and posterior regions engaged in coding sensory-somatic information (Rizzolatti and Craighero, 2004) while the presentation of a facial expression of pain led to stronger increases in the medial prefrontal cortex and precuneus which have been associated with social cognitive processes such as mentalizing and theory of mind (Amft et al., 2015; Gallo et al., 2018; Mitchell, 2009; Northoff et al., 2006). Despite the different psychological domains engaged in the pain empathic responses induced by NS and FE both elicit vicarious pain experience (Timmers et al., 2018), encompassing pain-specific processes such as recognizing and understanding the painful state of the other person and affective sharing of pain but also non-specific processes that are shared between pain and other non-painful experiences such as arousal and negative affect (Zaki et al., 2016). In line with the shared underlying mental processes previous neuroimaging meta-analyses revealed that the observation of acute pain infliction and painful facial expressions commonly activate core empathy and nociceptive pain regions specifically the insular and cingulate cortices (Jauniaux et al., 2019; Lamm et al., 2011; Timmers et al., 2018). The overlapping activations have been suggested to reflect shared neural representations of vicarious pain (Jauniaux et al., 2019; Lamm et al., 2011; Timmers et al., 2018).

However, overlapping functional activations within these regions do not necessarily reflect shared underlying neural representations of a specific mental process (Zaki et al., 2016), given that (1) due to local spatial dependencies the main focus of traditional mass-univariate fMRI analytic approaches (i.e. conducting massive number of tests on brain voxels one at a time) is not on single-voxel activity, but on smoothed, regional differences in brain activity across multiple tasks or stimuli (Haynes, 2015) and (2) brain regions may contain multiple, distinct populations of neurons and averaging across those neuron populations yields nonspecific signals (Haxby et al., 2014; Zaki et al., 2016). For instance, electrophysiological and optogenetic studies have identified distinct neuronal populations in the anterior cingulate and insular cortices that activate during several functional domains, including pain- and empathy-related processes as well as attention, salience, social observation learning and reward expectancy (Allman et al., 2011; Chen, 2018; Kvitsiani et al., 2013; Sakaguchi et al., 2018; Shidara and Richmond, 2002; Shura et al., 2014; Sikes and Vogt, 1992). Studies employing mass-univariate fMRI analyses suggest that both regions are engaged by various experimental paradigms including not only experienced and observed pain, but also reward, arousal, salience and attention (Cauda et al., 2012; Shackman et al., 2011; Uddin, 2015; Wager et al., 2016; Yarkoni et al., 2011). Despite the overlapping fMRI activation in response to different experimental manipulations the underlying brain representations may be separable (Corradi-Dell'Acqua et al., 2016; Krishnan et al., 2016; Woo et al., 2014), emphasizing that more fine-grained analyses are required to determine process-specific shared or distinct neural representations (Zaki et al., 2016).

In an effort to overcome these limitations, recent studies have proposed several strategies to investigate the ‘shared representation’ question, including pharmacological (see e.g. Rütgen et al., 2015) and multivariate pattern analysis (MVPA) approaches. Compared to conventional analytic approaches, MVPA can be effective in extracting information at much finer spatial scales (e.g. below the intrinsic resolution determined by the voxel size by pooling together weak feature-selective signals in each voxel; Kamitani and Tong, 2005; Woo et al., 2017) and represents a more suitable approach to support or reject claims about neural mechanisms that are shared between mental processes (Chikazoe et al., 2014; Peelen and Downing, 2007; Zaki et al., 2016). In support of this view, using MVPA approaches researchers have demonstrated shared neural representations across mental processes (including self-experienced and observed pain) in both humans and animals (Carrillo et al., 2019; Corradi-Dell'Acqua et al., 2011; Corradi-Dell'Acqua et al., 2016). Moreover, a growing number of recent studies have demonstrated functional independence of overlapping univariate activation in these brain regions using MVPA (Krishnan et al., 2016; Peelen et al., 2006; Woo et al., 2014), including separable neural representations of physical and social rejection pain within the dorsal anterior cingulate cortex (Woo et al., 2014) and of modality-specific aversive experience in the anterior insular cortex (Krishnan et al., 2016).

Nevertheless, shared multivariate patterns do not necessarily imply process-specific common neural representations per se given that the shared neural representations could simply reflect common demands on basal processing domains such as attention or arousal (Corradi-Dell'Acqua et al., 2011; Corradi-Dell'Acqua et al., 2016; Krishnan et al., 2016). For instance, Corradi-Dell'Acqua et al., 2016 found shared neural patterns between vicarious and self-experienced pain in the left anterior insula and further demonstrated that the common local patterns were not specific to pain-related processing, but also represented disgust and unfairness suggesting modality-unspecific processing of aversive and arousing experiences.

This leads to the questions of (1) whether or not NS and FE-induced vicarious pain share pain-associated common neural representations, and further (2) whether a general (i.e. across NS and FE vicarious pain modalities) neural signature of vicarious pain, which is specific to the pain empathic response rather than capturing unspecific processes such as negative emotional experience or arousal, can be determined. More specifically, we examined the following three questions in this study: (i) whether NS and FE-induced vicarious pain-predictive signatures share spatially (correlation and distribution) and functionally (predictions of cross-modality vicarious pain versus corresponding non-painful control stimuli) similar neural representations, (ii) whether a general and specific vicarious pain-predictive neural signature, which should (1) generalize across different vicarious pain stimuli and (2) not be sensitive to predict unspecific negative affect or arousal induced by non-painful negative stimuli, and (3) be ‘activated’ by the direct experience of somatic pain as reflected by an accurate prediction of self-experienced somatic pain, can be determined.

To this end, we employed MVPA to fMRI data from an experiment during which participants were presented with stimuli depicting the infliction of noxious stimulation of body limbs (NS vicarious pain) or painful facial expressions (FE vicarious pain) as well as corresponding non-painful control stimuli (Figure 1A). Given that relative to their control stimuli both sets of painful stimuli were perceived as more painful in terms of recognized and shared pain as well as more arousing and negative (details see Results and Figure 1B), we additionally asked participants to undergo an emotion processing paradigm with non-painful high-arousal negative stimuli and low-arousal neutral stimuli from the International Affective Picture System (IAPS) (details see Materials and methods) to further test the specificity of the shared neural representations with respect to the vicarious experience of pain rather than emotional arousal or negative affect. To determine the association of the vicarious pain signature with direct pain experience, we included an independent fMRI dataset that collected ratings of self-experienced pain during thermal pain induction (details are provided in Materials and methods and in Wager et al., 2013; Woo et al., 2015).

Figure 1

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Given that small sample sizes may lead to a large cross-validation error which is the discrepancy between the prediction accuracy measured by cross-validation and the expected accuracy on new data (Varoquaux, 2018) and fMRI-based inferences on regions that are most predictive substantially benefit from larger samples (Chang et al., 2015) we included a comparably large sample of n = 238 individuals (details see Materials and methods and Li et al., 2018; Xu et al., 2020a).

Several studies have explored the neural underpinnings of vicarious pain in humans and suggested overlapping univariate fMRI activations in the anterior cingulate and insular cortices across different vicarious pain induction procedures (for meta-analyses see e.g. Jauniaux et al., 2019; Timmers et al., 2018). However, the conventional univariate approach lacks anatomical and functional specificity to test the question of whether vicarious pain across different modalities share common and process-specific neural representations (Iannetti et al., 2013; Krishnan et al., 2016; Woo et al., 2014; Zaki et al., 2016). Here, we employed a fine-grained MVPA approach which is sensitive and specific to particular types of mental processes including pain (Kragel et al., 2018; Peelen et al., 2006; Wager et al., 2013; Woo et al., 2017) to explore (1) whether shared neural representations of vicarious pain can be determined across different induction procedures (FE, NS) and (2) whether the shared neural representation is sensitive to pain-unspecific components of the vicarious pain response (arousal, negative affect) and related to the experience of somatic pain. We demonstrated that shared multivariate patterns encoding NS and FE vicarious pain can be determined at the whole-brain level and that across different analytic approaches the mid-insular cortex was consistently engaged across induction procedures. Furthermore, we demonstrated that these patterns were not sensitive to respond to the processing of non-painful high-arousal negative stimuli in the same sample, together with the findings showing that NS vicarious pain predicted FE vicarious pain (and vice versa) more accurately as compared with the predictions using a negative emotion decoder, suggesting that the common vicarious pain representations do not simply reflect shared unspecific processes of negative affect or arousal. Moreover, the shared vicarious pain representations predicted self-experienced thermal pain in an independent sample, suggesting an association between the neural expression and processes directly related to the experience of pain. Together these results provide evidence for a generalized neural representation of vicarious pain, particularly in the mid-insula, and demonstrated that the shared neural signature may specifically capture pain-associated aspects of the vicarious pain response rather unspecific processes such as aversive experience or arousal.

The idea that vicarious pain across different induction procedures share common neural representations has been supported by meta-analyses covering previous fMRI pain empathy studies that demonstrated overlapping activations in the insular and cingulate cortices (Jauniaux et al., 2019; Timmers et al., 2018). In line with these meta-analytic findings we found that these regions were consistently engaged during both NS and FE vicarious pain. However, the insular and anterior cingulate cortices are involved in a wide range of mental processes including representation of interoceptive and affective states as well as salience detection (Craig, 2009; Critchley et al., 2004; Timmers et al., 2018; Uddin, 2015), suggesting that the overlapping activity might be due to common underlying mental processes such as detecting and orienting attention toward salient stimuli or unspecific emotional arousal (Corradi-Dell'Acqua et al., 2011; Corradi-Dell'Acqua et al., 2016; Valentini and Koch, 2012).

To systematically test whether vicarious sharing of pain elicited by different social cues shares common neural representations, we developed and compared multivariate patterns that predicted NS and FE vicarious pain evoking stimuli respectively. While mass-univariate analysis results reflect the presence of intermingled neuronal populations related to stimulus-specific representations, MVPA investigates whether idiosyncratic spatial variations in the fMRI signal are shared or dissociated across different conditions and thus might be more suitable to determine process-specific representations in meso-scale neural circuits (Kamitani and Tong, 2005; Kriegeskorte et al., 2006; Peelen and Downing, 2007). Moreover, previous studies have suggested that whole-brain predictive models could better capture emotional processes compared to regional approaches, such as decoding of a single brain region or searchlight-based methods (Kragel et al., 2018; Woo et al., 2017). To this end, we first identified whole-brain fMRI patterns that accurately predicted NS and FE vicarious pain, respectively. We found that the NS and FE vicarious pain-predictive patterns were spatially correlated and both could classify within- and between-modality painful versus non-painful stimuli at the whole-brain level, suggesting that NS and FE vicarious pain share distributed processing across multiple systems and component processes. In line with previous studies demonstrating that while NS vicarious pain provides objective cues about the sensory component of the observed pain the FE vicarious lacks such information and is more subjective and indirect as the pain experience of the expresser need to be interpreted by the observer (Hadjistavropoulos et al., 2011; Vachon-Presseau et al., 2012), the decreased accuracies and effect sizes in the cross-modality predictions additionally suggest partly distinguishable neural representations of NS and FE vicarious pain possibly reflecting the engagement of different component processes.

In the context of previous studies suggesting that pain empathy deficits are mediated by regional-specific brain lesions and functional dysregulations (Leigh et al., 2013; Shamay-Tsoory et al., 2009; Xu et al., 2020b) the question for the contribution of specific brain regions arises. Thresholding the vicarious pain patterns (at FDR q < 0.05, two-tailed) allowed us to identify voxels that reliably contributed to the respective decoders and revealed that specifically the bilateral mid-insula provided important features to predict both NS and FE vicarious pain. Moreover, the mid-insula partial vicarious pain patterns were highly spatially correlated and both could significantly predict within- and between-modality vicarious pain-related experience. Consistent with this, searchlight-based classification analyses also demonstrated that mid-insula local patterns produced significant within- and between-modality predictions of vicarious pain. Our results are in line with a previous meta-analysis showing that the mid-insula responds specifically to empathy for pain across different task paradigms compared to empathy for non-pain negative affective states (Timmers et al., 2018), which together with the present findings suggests that the mid-insula represents a core neural substrate for vicarious pain.

Although multivariate predictive models can capture information at much finer spatial scales and consequently anatomical specificity (Kamitani and Tong, 2005; Woo et al., 2017), the question of the specific mental processes captured by our vicarious pain-predictive patterns remains unclear. Pain empathy is a multi-component process that includes the vicarious sharing of pain but may also evoke emotional arousal and negative affect, and these unspecific processes can be captured by the decoders. To determine the functional specificity of the neural representations, we applied the vicarious pain-predictive patterns to data from an emotion processing paradigm acquired in the same sample as well as to data from a thermal pain induction experiment in an independent sample and found that (1) the vicarious pain patterns performed only modest for discriminating high-arousal (non-painful) negative stimuli from low-arousal neutral stimuli and (2) the whole-brain and mid-insula patterns predicted levels of self-experienced thermal pain with high accuracies. Finally, we developed a general vicarious pain-predictive pattern across NS and FE vicarious pain induction procedures and demonstrated that it accurately predicted both NS and FE vicarious pain (accuracies > 87%) as well as thermal pain intensities (accuracies > 82%), yet classified non-painful negative versus neutral stimuli with comparably low accuracy (59%). In line with the prediction results, meta-analytic decoding analysis revealed that this general vicarious pain pattern was highly correlated with the domains of ‘painful’ and ‘pain’, but not with ‘arousal’, ‘valence’ or ‘negative’ (not shown in the top 100 relevant terms). Together these findings suggest a shared neural representation of vicarious pain and a high-specificity of the whole-brain and specifically the mid-insula patterns for the vicarious experience of pain. A previous study developed a vicarious pain signature (VPS) that was sensitive and specific to NS vicarious pain, but not sensitive to the intensity of self-experienced somatic pain (Krishnan et al., 2016). Examining similarities with our general vicarious pain-predictive pattern revealed only modest spatial correlations between the two patterns (r = 0.04). The different instructions employed in the experiments might have contributed to the low overlapping spatial distributions such that participants in the previous study were required to explicitly rate their emotional response to the stimuli, whereas we decided for an implicit processing (passive viewing) paradigm to match instructions across the vicarious pain and negative emotional processing paradigm and to additionally control for cognitive processes which can modulate empathic reactivity and painful experience as well as the specific neural networks engaged (Jauniaux et al., 2019; Urien and Wang, 2019). Moreover, we found that the present pattern could successfully predict pain experience during thermal heat stimulation while the VPS was not sensitive to self-experienced pain. The observed differences might be explained in terms of (1) the considerably larger sample size included in the present study and prediction accuracy (as reflected by prediction-outcome correlation) of self-experienced pain experience increased as a function of sample size used to develop the NS vicarious pain decoder (see additional analysis presented in Figure 7—figure supplement 2) and (2) differences between paradigms and instructions such that, for example, a recent meta-analysis of pain empathy studies showed that the mid-cingulate gyrus was more activated by explicit cognitive/evaluative paradigms while the right inferior frontal gyrus and anterior insula were more activated by implicit perceptual/affective paradigms (Timmers et al., 2018).

Our results highlighted the mid-insula as a key region sharing similar neural representations across NS and FE vicarious pain suggesting that it may contribute to the core vicarious pain experience that characterizes pain empathy. Consistent with the whole-brain results, the shared information in the mid-insula was specific to vicarious pain rather than negative affect or arousal. Previous non-human primate and human studies indicate that the posterior and mid-insula receive nociceptive information from thalamic nuclei (Craig et al., 1994; Craig et al., 2000) which are in turn conveyed to the anterior insula for progressive integration with higher level affective and interoceptive experience (Corradi-Dell'Acqua et al., 2011; Corradi-Dell'Acqua et al., 2016; Singer et al., 2009). Although overarching models of the neural basis of pain empathy and neuroimaging meta-analyses (Jauniaux et al., 2019; Timmers et al., 2018) emphasize the role of the anterior insula in pain empathy processing, accumulating evidence from studies examining shared and process-specific representations of vicarious pain suggest a specific role of the mid-insula in vicarious pain (Corradi-Dell'Acqua et al., 2011; Krishnan et al., 2016), whereas the (left) anterior insula also responded to negative stimuli in general (Corradi-Dell'Acqua et al., 2011) and across modalities (Corradi-Dell'Acqua et al., 2016). Importantly, the peak anterior insula coordinates identified in these previous studies did not overlap with our mid-insula mask or the mid-insula region that exhibited reliable predictive features in both NS and FE vicarious pain whole-brain patterns determined in the present study, suggesting a more specific role of the mid-insula in pain-related components of the vicarious pain response (see also recent meta-analysis by Timmers et al., 2018 demonstrating a specific role of the mid-insula in pain empthy). In support of our findings a previous study employed a similar whole-brain MVPA approach to predict NS vicarious pain induced by an evaluative paradigm also identified the bilateral mid-insula as reliable (q < 0.05, FDR corrected) predictive regions (Krishnan et al., 2016), further conforming the reliable contribution of this region in encoding vicarious pain. Studies examining the functional and anatomical organization of the insular cortex with intracerebral electrical stimulation have demonstrated that painful sensations can be elicited by stimulation of the middle but not the anterior insula (Afif et al., 2010). Together with the functional relevance of the mid-insula to predict objective and subjective pain experience in an independent sample and the contribution of this region to nociception as well as vicarious pain (Botvinick et al., 2005; Krishnan et al., 2016; Lamm et al., 2011; Timmers et al., 2018; Wager et al., 2013), our findings suggest that the shared representations in the mid-insula across vicarious pain induction procedures may specifically code the automatic pain sharing which resonates with embodies conceptualizations of vicarious pain (see e.g. Corradi-Dell'Acqua et al., 2011 for a convergent interpretation). However, consistent with previous evidence that (NS) vicarious pain representation is distributed across brain regions and single local regions exhibit considerably lower effect sizes compared to whole-brain predictive models (Krishnan et al., 2016), we found that the prediction effect sizes for the mid-insula were smaller than those observed in our whole brain analyses. These findings suggest that despite the key role of the mid-insula in vicarious pain experience this region is not sufficient to fully capture this process.

Consistent with previous studies suggesting that the anterior cingulate cortex represents a core brain region for emotional empathy in general and pain empathy in particular (Fan et al., 2011; Jauniaux et al., 2019; Timmers et al., 2018), we found overlapping deactivations in the rostral and ventral anterior cingulate cortex in mass-univariate analyses and shared patterns in the dorsal, rostral and ventral anterior cingulate cortex in searchlight-based prediction analyses between NS and FE vicarious pain. However, no overlapping reliable predictive voxels for whole-brain NS and FE pain-predictive patterns were found in cingulate regions suggesting a differential involvement of this region during FE and NS vicarious pain induction procedures. From a methodological perspective, these results may reflect that the whole-brain predictive model could provide a more specific neural description of a behavior or mental process (Kragel et al., 2018; Woo et al., 2017). In line with our findings, previous studies also showed that significant activation and searchlight-based prediction in local regions do not necessarily imply reliable predictive features in whole-brain predictive models (Krishnan et al., 2016; Woo et al., 2014). From a brain systems perspective, these findings may indicate that the anterior cingulate cortex is not specifically involved in vicarious pain elicited across induction procedures. Although the anterior cingulate cortex has been reliably identified in meta-analytic studies covering brain activation patterns during (pain) empathy induction procedures (see e.g. Jauniaux et al., 2019; Timmers et al., 2018), the anterior cingulate has also been associated with a number of basal processes, including arousal and salience, and activation in this region may reflect rather unspecific neural responses.

The present study has limitations that should be addressed in future studies. Compared to the homogeneous stimuli within the conditions of the vicarious pain and the self-experienced pain paradigm the stimuli displaying emotional evocative scenes from the IAPS database may have led to a higher inter-trial variance in the negative processing experiment. Although the inter-stimulus variance should not systematically differ between the experimental conditions employed to develop the corresponding decoder, we cannot fully exclude that this may have partly contributed to the low accuracies of the emotional processing decoder with respect to predicting self-experienced pain ratings. Moreover, the current study employed a passive observation paradigm and a recent meta-analysis revealed that vicarious pain induced by cognitive/evaluative and affective/perceptual paradigms elicited activations in overlapping yet also different brain regions (Timmers et al., 2018). Whether the present conclusions could generalize to more ‘active’ engagements in empathy (e.g. explicitly asking subjects to imagine that the injury occurring in the picture displayed was happening to them) remains to be determined.

In conclusion, by applying a novel whole-brain as well as local-region-based MVPA approaches in a large sample of healthy adults, our results provide the first neuroimaging evidence that NS and FE vicarious pain share common neural representations, especially in the mid-insula which may specifically encode the vicarious sharing of pain that specifically characterizes pain empathy. Moreover, we also provide a general vicarious pain-predictive pattern (across NS and FE vicarious pain stimuli), which may be employed in future studies to facilitate inferences about pain empathy across modalities as well as self-experienced pain. Our study offers a new approach to better understand pain empathy by exploring common neural representations and linking these shared representations to felt pain.

The functional MRI, numerical data as well as the Matlab scripts used to generate the figures have been deposited on the figshare repository under accession code 11994498 (https://figshare.com/articles/Vicarious_pain_dataset/11994498) Statistical and pattern weight maps are available on the Neurovault repository under collection 6332 (https://neurovault.org/collections/6332/). Statistical and pattern weight images are available on Neurovault.

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Author details

1. Feng Zhou

   1. Clinical Hospital of Chengdu Brain Science Institute, MOE Key Laboratory for Neuroinformation, University of Electronic Science and Technology of China, Chengdu, China
   2. Department of Psychological and Brain Sciences, Dartmouth College, Hanover, United States

   Contribution

   Formal analysis, Visualization, Methodology, Writing - original draft, Writing - review and editing

   For correspondence

   zhou.feng@live.com

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-4509-4026

2. Jialin Li

   Clinical Hospital of Chengdu Brain Science Institute, MOE Key Laboratory for Neuroinformation, University of Electronic Science and Technology of China, Chengdu, China

   Contribution

   Data curation, Formal analysis, Writing - original draft

   Competing interests

   No competing interests declared

3. Weihua Zhao

   Clinical Hospital of Chengdu Brain Science Institute, MOE Key Laboratory for Neuroinformation, University of Electronic Science and Technology of China, Chengdu, China

   Contribution

   Data curation, Project administration

   Competing interests

   No competing interests declared

4. Lei Xu

   Clinical Hospital of Chengdu Brain Science Institute, MOE Key Laboratory for Neuroinformation, University of Electronic Science and Technology of China, Chengdu, China

   Contribution

   Data curation, Project administration

   Competing interests

   No competing interests declared

5. Xiaoxiao Zheng

   Clinical Hospital of Chengdu Brain Science Institute, MOE Key Laboratory for Neuroinformation, University of Electronic Science and Technology of China, Chengdu, China

   Contribution

   Data curation, Project administration

   Competing interests

   No competing interests declared

6. Meina Fu

   Clinical Hospital of Chengdu Brain Science Institute, MOE Key Laboratory for Neuroinformation, University of Electronic Science and Technology of China, Chengdu, China

   Contribution

   Data curation, Project administration

   Competing interests

   No competing interests declared

7. Shuxia Yao

   Clinical Hospital of Chengdu Brain Science Institute, MOE Key Laboratory for Neuroinformation, University of Electronic Science and Technology of China, Chengdu, China

   Contribution

   Data curation, Project administration, Writing - review and editing

   Competing interests

   No competing interests declared

8. Keith M Kendrick

   Clinical Hospital of Chengdu Brain Science Institute, MOE Key Laboratory for Neuroinformation, University of Electronic Science and Technology of China, Chengdu, China

   Contribution

   Conceptualization, Supervision, Project administration, Writing - review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-0371-5904

9. Tor D Wager

   Department of Psychological and Brain Sciences, Dartmouth College, Hanover, United States

   Contribution

   Resources, Software, Supervision, Methodology, Writing - review and editing

   Competing interests

   No competing interests declared

10. Benjamin Becker

    Clinical Hospital of Chengdu Brain Science Institute, MOE Key Laboratory for Neuroinformation, University of Electronic Science and Technology of China, Chengdu, China

    Contribution

    Conceptualization, Funding acquisition, Writing - original draft, Project administration, Writing - review and editing

    For correspondence

    ben_becker@gmx.de

    Competing interests

    No competing interests declared

    "This ORCID iD identifies the author of this article:" 0000-0002-9014-9671

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