Empathy enables us to share and understand the emotional states of other people, often based on their facial expressions. This empathic response involves being able to distinguish our own emotional state from someone else’s, and it is influenced by how we recognize that person’s emotion. In real life, knowing and identifying whether the facial expression we are witnessing reflects genuine or pretended pain is particularly important so that we can appropriately react to someone’s emotions and avoid unnecessary personal distress.

How our brains manage to do this is still heavily debated. Two areas, the anterior insular (aIns for short) and the mid-cingulate cortex, appear to be activated when someone ‘feels’ someone else’s pain. However, these regions might just automatically be triggered by vivid emotional facial expressions, regardless of whether we really respond to that pain.

To examine this question, Zhao et al. measured brain activity as healthy adults watched video clips of people either feeling or pretending to feel pain. The activation of aIns was particularly related to the emotional component that someone shared with another person’s genuine pain, but not to pretended pain. This suggests that neurons in the aIns track a truly empathic response when seeing someone who is actually experiencing pain.

Effective connectivity analyses which reflect how brain areas ‘crosstalk’ also revealed distinct patterns when people viewed expressions of genuine, as opposed to pretended pain. Zhao et al. focused on the interactions between the alns and the right supramarginal gyrus, a brain region which helps to distinguish another person’s emotions from our own. This crosstalk tracked others’ feelings when participants viewed expressions of genuine but not of pretended pain.

Put together, these findings provide a more refined model of empathy and its neural underpinnings. This will help further our understanding of conditions such as autism or depression, in which a person’s social skills and emotional processing are impaired.

As social beings, our own affective states are influenced by other people’s feelings and affective states. The facial expression of pain by others acts as a distinctive cue to signal their pain to others, and thus results in sizeable affective responses in the observer. Certifying such responses as evidence for empathy, however, requires successful self-other distinction, the ability to distinguish the affective response experienced by ourselves from the affect experienced by the other person.

Studies using a wide variety of methods convergently have shown that observing others in pain engages neural responses aligning with those coding for the affective component of self-experienced pain, with the anterior insula (aIns) and the anterior mid-cingulate cortex (aMCC) being two key areas in which such an alignment has been detected (Lamm et al., 2011; Rütgen et al., 2015; Jauniaux et al., 2019; Xiong et al., 2019; Zhou et al., 2020; Fallon et al., 2020, for meta-analyses). However, there is consistent debate on whether activity observed in these areas should indeed be related to the sharing of pain affect or whether it may not rather result from automatic responses to salient perceptual cues – with pain vividly expressed on the face being one particularly prominent example (Zaki et al., 2016, for review). It was thus one major aim of our study to address this question. In this respect, contextual factors, individuals’ appraisals, and attentional processes would all impact their exact response to the affective states of others (Gu and Han, 2007; Hein and Singer, 2008, for review; Lamm et al., 2010; Forbes and Hamilton, 2020; Zhao et al., 2021). Recently, Coll et al., 2017 have thus proposed a framework that attempts to capture these influences on affect sharing and empathic responses. This model posits that individuals who see identical negative facial expressions of others may have different empathic responses due to distinct contextual information and that this may depend on identification of the underlying affective state displayed by the other. In the current functional magnetic resonance imaging (fMRI) study, we therefore created a situation where we varied the genuineness of the pain affect felt by participants while keeping the perceptual saliency (i.e., the quality and strength of pain expressions) identical. To this end, participants were shown video clips of other persons who supposedly displayed genuine pain on their face vs. merely pretended to be in pain. Note that for reasons of experimental control, all painful expressions on the videos had been acted. This enabled us to interpret possible differences between conditions to the observers’ appraisal of the situation rather than to putative visual and expressive differences. This way, we sought to identify the extent to which responses in affective nodes (such as the aIns and the aMCC) genuinely track the pain of others, rather than resulting predominantly from the salient facial expressions associated with the pain.

Another major aim of our study was to assess how self-other distinction allowed individuals to distinguish between the sharing of actual pain vs. regulating an inappropriate and potentially misleading ‘sharing’ of what in reality is only a pretended affective state. We focused on the right supramarginal gyrus (rSMG), which has been suggested to act as a major hub selectively engaged in affective self-other distinction (Silani et al., 2013; Steinbeis et al., 2015; Hoffmann et al., 2016; Bukowski et al., 2020). Though previous studies have indicated that rSMG is functionally connected with areas associated with affect processing (Mars et al., 2012; Bukowski et al., 2020), we lack more nuanced insights into how exactly rSMG interacts with these areas, and thus how it supports accurate empathic responses. Hence, we used dynamic causal modeling (DCM) to investigate the hypothesized brain patterns of affective responses and self-other distinction for the genuine and pretended pain situations, focusing on the aIns, aMCC, and their interaction with rSMG. Furthermore, we investigated the relationship between neural activity and behavioral responses as well as empathic traits. In line with the literature reviewed above, we expected that, on the behavioral level, genuine pain would result in – alongside the obvious other-oriented higher pain ratings – higher self-oriented unpleasantness ratings. On the neural level, we predicted aIns and aMCC to show a stronger response to the genuine expressions of pain, but that these areas would also respond to the pretended pain, but to a lower extent. Differences in rSMG engagement and distinct patterns of this area’s effective connectivity with aIns and aMCC were expected to relate to self-other distinction, and thus to explain the different empathic responses to genuine pain vs. pretended pain.

In this study, we developed and used a novel experimental paradigm in which participants watched video clips of persons who supposedly either genuinely experienced pain or merely pretended to be in strong pain. Combining mass-univariate analysis with effective connectivity (DCM) analyses, our study provides evidence on the distinct neural dynamics between regions suggestive of affect processing (i.e., aIns and aMCC) and self-other distinction (i.e., rSMG) for genuinely sharing vs. responding to pretended, non-genuine pain. With this, we aimed to clarify two main questions: First, whether neural responses in areas such as the aIns and aMCC to the pain of others are indeed related to a veridical sharing of affect, as opposed to simply tracking automatic responses to salient affective displays. And second, how processes related to self-other distinction, implemented in the rSMG, enable appropriate empathic responses to genuine vs. merely pretended affective states.

The mass-univariate analyses suggest that the increased activity in aIns for genuine pain as opposed to pretended pain properly reflects affect sharing. As aforementioned, the network of affective sharing and certain domain-general processes (e.g., salience detection and automatic emotion processing) overlap in aIns and aMCC (Zaki et al., 2016, for review). This indicates that indeed, part of the activation in these areas could be related to perceptual salience, which is why it has been widely debated as a potential confound of empathy and affect sharing models (Zaki et al., 2016, for review; Lamm et al., 2019, for review). However, when comparing genuine pain versus pretended pain, activity in these areas was not only found to be stronger in response to genuine pain, but the increased activation in aIns was also selectively correlated with ratings of self-oriented unpleasantness and was not correlated with either other-related painful expressions or painful feelings in terms of the regression analysis. That only aIns and not also aMCC shows such correlation may be explained by previous studies, according to which aIns is more specifically associated with affective representations, while the role of aMCC rather seems to evaluate and regulate emotions that arise due to empathy (Fan et al., 2011; Lamm et al., 2011; Jauniaux et al., 2019). Taken together, the activation and brain-behavior findings provide evidence that responses in aIns (and to a lesser extent also the aMCC) are not simply automatic responses triggered by perceptually salient events (otherwise the increased aIns activation should also be explained by other behavioral ratings in the sense of shared influence by domain-general effects). Rather, they seem to track the actual affective states of the other person, and thus the shared neural representation of that response (see Zhou et al., 2020, for similar recent conclusions based on multi-voxel pattern analyses). Our findings are also in line with the proposed model of Coll et al., 2017, which suggests that affect sharing is the consequence of emotion identification. More specifically, while part of the activation in the aIns and aMCC is indeed related to an (presumably earlier) automatic response, the added engagement of these areas once they have identified the pain as genuine shows that only in this condition, they then also engage in proper affect sharing. Ideally, one should be able to discern these processes in time, but neither the temporal resolution of our fMRI measurements nor the paradigm in which we always announced the conditions beforehand would have been sensitive enough to do so. Thus, future studies including complementary methods, such as EEG and MEG, and tailored experimental designs are needed to pinpoint the exact sequence of processes engaged in automatic affective responses vs. proper affect sharing.

Beyond higher activation in affective nodes supporting (pain) empathy, increased activation was also found in rSMG. The inferior parietal lobule was shown to be generally engaged in selective attention, action observation, and imitating emotions (Bach et al., 2010; Pokorny et al., 2015; Gola, 2017; Hawco et al., 2017). Importantly, a specific role in affective rather than cognitive self-other distinction has been identified for rSMG (Silani et al., 2013; Steinbeis et al., 2015; Bukowski et al., 2020). Based on such findings, it has been proposed that the rSMG allows for a rapid switching between or the integration of self- and other-related representations, as two processes that may underpin the functional basis of successful self-other distinction (Lamm et al., 2016, for review). Theoretical models of empathy and related socio-affective responses suggest that such regulation is especially important to avoid so-called empathic over-arousal, which would shift the focus away from empathy and the other’s needs, toward taking care of one’s own personal distress (Batson et al., 1987, for review; Decety and Lamm, 2011, for review). Concerning the current findings, we thus propose that the higher rSMG engagement in the genuine pain condition reflects an increasing demand for self-other distinction imposed by the stronger shared negative affect experienced in this condition.

Beyond these differences in the magnitude of rSMG activation, the DCM analysis demonstrated less inhibition on the aIns-to-rSMG connection for genuine pain compared to pretended pain. Note that our focus on the right aIns rather than bilateral aIns was because it is located in the same hemisphere as the right SMG. Various theoretical accounts suggest that areas such as the aIns and rSMG may play a key role in comparing self-related information with the sensory evidence (Decety and Lamm, 2007, for review; Seth, 2013, for review). According to recent theories on predictive processing (Clark, 2013, for review) and active inference (Friston, 2010, for review), the brain can be regarded as a “prediction machine”, in which the top-down signals pass over predictions and the bottom-up signals convey prediction errors across different levels of cortical hierarchies (Chen et al., 2009; Friston, 2010, for review; Bastos et al., 2015). It is suggested that these top-down predictions are mediated by inhibitory neural connections (Zhang et al., 2008; Bastos et al., 2015; Miska et al., 2018). Our findings align with such views, by suggesting that the inhibitory connection from aIns to rSMG can be explained as the predictive mismatch between the top-down predictions of self-related information (e.g., personal affect) and sensory inputs (e.g., pain facial expressions). This suppression of neural activity leads to an explaining away of incoming bottom-up prediction error. This is reflected by the absence of any condition-dependent modulatory effects on the rSMG to aIns connection, suggesting that the influence of the task conditions is sufficiently modeled by the predictions from aIns to rSMG. Therefore, the stronger inhibition for pretended pain, compared to genuine pain, could indicate a higher demand to overcome the mismatch between the visual inputs and the agent’s prior beliefs and contextual information about the situation (i.e., “this person looks like in pain, but I know he/she does not actually feel it”). We speculate that a dynamic interaction between sensory-driven and control processes is underlying the modulatory effect: when individuals realized after an initial sensory-driven response to the facial expression that it was not genuinely expressing pain, control, and appraisal processes led to a reappraisal of the triggered emotional response, and thus a dampening of the unpleasantness. The reduced inhibition in the genuine pain condition could moreover be a mechanism that explains the higher rSMG activation in this condition.

Model comparison showed that the best model to explain the inhibitory effect with the behavioral ratings for both the genuine and pretended pain is the model without interactions between ratings. That is, if any behavioral rating contributed to the modulation of aIns to rSMG, the effect would be more likely coming from single ratings rather than their interactions. Specifically, we found the strength of the inhibitory effect in the genuine pain condition to correlate with ratings of painful feelings in others, but not with the ratings of pain expression in others or unpleasantness in self. For the pretended pain condition, none of the ratings showed a correlation. The latter could in principle be due to a lack of variation in the ratings (which by way of the design were mostly close to zero or one). We deem it more plausible, though, that the correlation findings provide further evidence that the modulation of aIns to rSMG is implicated in encoding others’ emotional states, which serves as a functional foundation for self-other processing when participants engaged in genuine affect sharing. This regulation cannot be totally attributed to domain-general processes, otherwise other ratings should have also explained this variation. It is also interesting to note that the found correlation relates to cognitive evaluations of the other’s pain rather than to own affect, as tracked by the unpleasantness in self-ratings. This would to some extent be in line with DCM findings by Kanske et al., 2016. These authors found that the inhibition of the temporoparietal junction (TPJ) by the aIns was linked to interactions between Theory of Mind (ToM) and empathic distress, that is the interaction of ‘cognitive’ vs. ‘affective’ processes engaged in understanding others’ cognitive and affective states. Note that the right TPJ is an overarching area involved in self-other distinction of which rSMG is considered a part or at least closely connected to Decety and Lamm, 2007, for review.

The correlations between the DCM inhibitory effect and empathic traits assessed via questionnaires provide further refinements for the relevance of rSMG in implementing self-other distinction to allow for an appropriate empathic response. When participants shared genuine affect, the inhibitory effect on the aIns to rSMG connection was positively correlated with affective ability and negatively correlated with affective reactivity. Affective ability reflects the capacity to subjectively share emotions with others, while affective reactivity plays a role in the susceptibility to vicarious distress and thus to more automatic responses to another’s emotion (Batchelder et al., 2017). Again, as for the correlations with the three rating scales, we did not find correlations of empathic traits for the pretended pain condition. Taken together, the DCM results and their qualification by the correlation findings suggest that in the genuine pain condition, which requires an accurate sharing of pain, rSMG interacts with aIns to achieve ‘affective-to-affective’ self-other distinction – that is disambiguating affective signals originating in the self from those attributable to the other person. The aIns to rSMG connection in the pretended pain condition may reflect a related, yet slightly distinct mechanism. Here, it seems that ‘cognitive-to-affective’ self-other distinction is at play, which helps resolve conflicting information between the top-down contextual information (i.e., that the demonstrator is not actually in pain) from what seems an unavoidable affective response to the highly salient perceptual cue of the facial expression of pain. Given our behavioral and trait data did not allow us to distinguish more precisely between these different types of self-other distinction, this however remains an interpretation and a hypothesis that will require further investigation. This inhibitory effect might be related to socioemotional disturbances of individuals with autism spectrum disorders (ASD), who show impairments in social cognition, including self- and other-related processing (Hoffmann et al., 2016; Lamm et al., 2016, for review). It is thus likely that ASD individuals exhibit distinct inhibition of the aIns-to-rSMG connectivity pattern compared to healthy controls. Further research with ASD individuals is required.

One potential limitation of the study could be the slightly higher ratings of other-oriented pain expressions for genuine pain, which were hypothesized to have no difference, as compared to pretended pain. As we found the enhanced aIns activation in the genuine pain condition mainly tracked personal unpleasantness rather than perceptually domain-general processes, and because the effect size of the pain expression difference was much smaller than for the affect ratings, we consider this difference did not fundamentally influence the interpretation of our findings. An additional limitation was that our study design did not aim to explicitly quantify self-other distinction. Rather, in line with previous research and based on our theoretical framework and rationale, we inferred the engagement of this process from the experimental conditions and the associated behavioral and neural responses. We expect our findings to prompt and inform future research designed to quantify and experimentally disentangle self- and other-related processes more explicitly. Note though that our participants and the targets shown in the videos were balanced with respect to their sex/gender. Yet sex/gender effects (for a review, though, see Christov-Moore, 2014) were outside the scope of the current study, and we thus did not perform any sex/gender-related analyses.

In conclusion, the current study advances our understanding of two main aspects of empathy. First, we provide evidence that empathy-related responses in the aIns can indeed be linked to affective sharing, rather than attributing them to responses triggered only by perceptual saliency. Second, we show how aIns and rSMG are orchestrated to track what another person really feels, thus enabling us to appropriately respond to their actual needs. Beyond these basic research insights, our study provides novel avenues for clinical application, and the investigation of contextual and interpersonal factors in the accurate diagnosis of pain and its expression.

All data needed to evaluate the conclusions in the paper are present in the paper. Raw functional imaging and behavioral data are deposited at https://doi.org/10.5281/zenodo.4783235. Processed behavioral data and individual DCM parameters are accessible at https://github.com/Yili-Zhao/Genuine_pretended-pain-task.git. Unthresholded statistical maps are available at https://identifiers.org/neurovault.collection:9949.

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Author details

1. Yili Zhao

   Social, Cognitive and Affective Neuroscience Unit, Department of Cognition, Emotion, and Methods in Psychology, Faculty of Psychology, University of Vienna, Vienna, Austria

   Contribution

   Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Software, Validation, Visualization, Writing - original draft, Writing - review and editing

   Competing interests

   none

   "This ORCID iD identifies the author of this article:" 0000-0003-4535-6636

2. Lei Zhang

   Social, Cognitive and Affective Neuroscience Unit, Department of Cognition, Emotion, and Methods in Psychology, Faculty of Psychology, University of Vienna, Vienna, Austria

   Contribution

   Conceptualization, Formal analysis, Methodology, Resources, Software, Supervision, Validation, Writing - original draft, Writing - review and editing

   Competing interests

   none

   "This ORCID iD identifies the author of this article:" 0000-0002-9586-595X

3. Markus Rütgen

   1. Social, Cognitive and Affective Neuroscience Unit, Department of Cognition, Emotion, and Methods in Psychology, Faculty of Psychology, University of Vienna, Vienna, Austria
   2. Vienna Cognitive Science Hub, University of Vienna, Vienna, Austria

   Contribution

   Conceptualization, Formal analysis, Funding acquisition, Methodology, Resources, Software, Supervision, Validation, Writing - original draft, Writing - review and editing

   Competing interests

   none

   "This ORCID iD identifies the author of this article:" 0000-0003-4947-7734

4. Ronald Sladky

   Social, Cognitive and Affective Neuroscience Unit, Department of Cognition, Emotion, and Methods in Psychology, Faculty of Psychology, University of Vienna, Vienna, Austria

   Contribution

   Conceptualization, Formal analysis, Methodology, Resources, Software, Supervision, Validation, Writing - original draft, Writing - review and editing

   Competing interests

   none

   "This ORCID iD identifies the author of this article:" 0000-0002-5986-1572

5. Claus Lamm

   1. Social, Cognitive and Affective Neuroscience Unit, Department of Cognition, Emotion, and Methods in Psychology, Faculty of Psychology, University of Vienna, Vienna, Austria
   2. Vienna Cognitive Science Hub, University of Vienna, Vienna, Austria

   Contribution

   Conceptualization, Formal analysis, Funding acquisition, Methodology, Resources, Supervision, Validation, Writing - original draft, Writing - review and editing

   For correspondence

   claus.lamm@univie.ac.at

   Competing interests

   none

   "This ORCID iD identifies the author of this article:" 0000-0002-5422-0653

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