Neural dynamics between anterior insular cortex and right supramarginal gyrus dissociate genuine affect sharing from perceptual saliency of pretended pain
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Evaluation Summary:
Lamm and colleagues explore brain dynamics of empathy in response to facial expressions of simulated versus genuine pain in others. Using a novel experimental fMRI task and dynamic causal modelling, the findings suggest that activation of the anterior insula for genuine pain reflects affect sharing rather than automatic responses triggered by the perceptual salience of events. The paper is of broad interest to an audience of social and affective neuroscientists interested in how humans track the emotional responses of others.
(This preprint has been reviewed by eLife. We include the public reviews from the reviewers here; the authors also receive private feedback with suggested changes to the manuscript. The reviewers remained anonymous to the authors.)
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Abstract
Empathy for pain engages both shared affective responses and self-other distinction. In this study, we addressed the highly debated question of whether neural responses previously linked to affect sharing could result from the perception of salient affective displays. Moreover, we investigated how the brain network involved in affect sharing and self-other distinction underpinned our response to a pain that is either perceived as genuine or pretended (while in fact both were acted for reasons of experimental control). We found stronger activations in regions associated with affect sharing (anterior insula [aIns] and anterior mid-cingulate cortex) as well as with affective self-other distinction (right supramarginal gyrus [rSMG]), in participants watching video clips of genuine vs. pretended facial expressions of pain. Using dynamic causal modeling, we then assessed the neural dynamics between the right aIns and rSMG in these two conditions. This revealed a reduced inhibitory effect on the aIns to rSMG connection for genuine pain compared to pretended pain. For genuine pain only, brain-to-behavior regression analyses highlighted a linkage between this inhibitory effect on the one hand, and pain ratings as well as empathic traits on the other. These findings imply that if the pain of others is genuine and thus calls for an appropriate empathic response, neural responses in the aIns indeed seem related to affect sharing and self-other distinction is engaged to avoid empathic over-arousal. In contrast, if others merely pretend to be in pain, the perceptual salience of their painful expression results in neural responses that are down-regulated to avoid inappropriate affect sharing and social support.
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Author Response:
Reviewer #2:
Weaknesses of the Methods and Results:
- In my view, the experiment does not allow to unambiguously disentangle self vs. other distinction (as mentioned in the abstract "..we investigated how affect sharing and self-other distinction interact.."). For example, genuine vs. pretended pain could be distinguished from the participants own experience in a comparable way. The higher rating of unpleasantness for genuine pain in others does not necessary mean that the participants cannot separate own from others experiences.
We thank the reviewer for raising this issue and for prompting us to further clarify and better state our research purpose. In terms of its original theoretical foundation and motivation, the current study aimed to investigate whether and how neural signatures underlying two essential …
Author Response:
Reviewer #2:
Weaknesses of the Methods and Results:
- In my view, the experiment does not allow to unambiguously disentangle self vs. other distinction (as mentioned in the abstract "..we investigated how affect sharing and self-other distinction interact.."). For example, genuine vs. pretended pain could be distinguished from the participants own experience in a comparable way. The higher rating of unpleasantness for genuine pain in others does not necessary mean that the participants cannot separate own from others experiences.
We thank the reviewer for raising this issue and for prompting us to further clarify and better state our research purpose. In terms of its original theoretical foundation and motivation, the current study aimed to investigate whether and how neural signatures underlying two essential components of empathy, namely affect sharing and self-other distinction, track individual responses to genuine vs. pretended pain. We agree though that our experimental design does not allow to disentangle unequivocally the precise aspects of self- and other-related processing in the two main conditions of interest (genuine pain or pretended pain). We thus modified any wording suggesting otherwise, so as to avoid further misunderstanding by readers.
Accordingly, we have provided a more elaborate theoretical clarification in the Abstract and Introduction about our particular interest in studying self-other distinction and its neural correlates in the right supramarginal gyrus (rSMG) during empathy. We also mention as a potential limitation that our design did not aim to explicitly quantify self-other distinction.
Action taken: In the manuscript, we have made the following changes:
- We modified the sentence "[...] we investigated how affect sharing and self-other distinction interact [...]" to
“[...] we investigated how the brain network involved in affect sharing and self-other distinction underpinned [...] ” in the Abstract (P. 1).
Besides, we modified another sentence “[...] to investigate the hypothesized distinct interactions between affective response and self-other distinction [...]” to
“[...] to investigate the hypothesized brain patterns of affective responses and self-other distinction [...]” in the Introduction (P. 4).
- We added sentences in the Discussion (P. 13): “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.”
- The experimental design does not unambiguously allows to disentangle genuine vs pretended pain from other factors, such as the differences in pain expression, painful feeling in others and higher unpleasantness in these two conditions. I understand that the intensity pain expression, painful feeling in others and unpleasantness for others is inherently tied to genuine vs. pretended pain. But the author already saw that the instruction of "genuine vs. pretented" influenced the ratings of pain expression. Hence, this allows two interpretation of the results: either the influence from the anterior Insula on the rSMG is driven by higher perceived pain expression, painful feeling in others and unpleasantness or by the conditions of genuine vs. pretended pain. Or (more likely) by an interaction between these factors. It would, for example help to explore the association between the aIns-rSMG interaction pain expression ratings (or painful feeling in others or higher unpleasantness) in videos with genuine pain und pretended pain separately. The author should further discuss this point that different factors (pain expression, etc) contribute to the differences between genuine vs. pretended pain.
We thank the reviewer for the thoughtful consideration of different factors that might contribute to disentangle genuine pain vs. pretend pain. One thing we would like to address beforehand: to disentangle the specific contributors underlying the manipulation is not the main focus for the current study, as 1) our primary aim was to study the effects of the experimental manipulation as a whole; we thus used the three behavioral ratings mainly to collect additional information on and to interpret the expected effects of the manipulation, and 2) these factors (and their behavioral measures) are inherently (cor)related and hard to be disentangled precisely anyways, as mentioned by the reviewer and as shown by extensive previous research both by our and other groups.
Action taken: Nonetheless, in the revised manuscript, we have now:
- discussed how different factors possibly interact and in this way contribute to the differences (in the modulatory effect) between genuine vs. pretended pain, in the Discussion (P. 11):
“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.”
- performed additional linear regression models and model comparison (see details in the response to comment #3) to investigate whether an interaction between behavioral measures could be a potential contributor to the modulatory effect of genuine pain and pretended pain; in short, the model without interactions is the winning model both for genuine pain and pretended pain.
We have now discussed this result (P. 11):
“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 [...]”
We thank the reviewer for this suggestion for further analysis.
We performed additional linear regression models (with and without interaction) and model comparison to explore whether any interaction between behavioral ratings heavily contributed to the modulatory effect. Results showed that the model without interaction was the most efficient model for both conditions.
We report the additional analyses as follows:
In the Methods section (P. 24-25): “Considering that interactions between behavioral ratings might contribute to the regression model, we tested five regression models (with and without interaction; see Supplementary Table 1) for both genuine pain and pretended pain. Results showed that for both genuine pain and pretend pain, the model without any interaction outperformed other models.”
Supplementary Table 1. Model comparison of linear regression models with three behavioral ratings (independent variables) and the inhibitory effect (dependent variable) for genuine pain and pretended pain. Smaller AIC/BIC indicates better model fit. Results showed that M1 (without interaction; highlighted with underlining) was the best fitting model for both genuine pain and pretended pain.
Accordingly, we now report the results of the winning model of the multiple regression analyses, instead of the original stepwise regression. These analyses found that only the rating of painful feelings in others was significant for genuine pain, while no significant effects whatsoever were found for pretended pain.
Action taken:
In the manuscript, we have made the following changes:
We modified “stepwise linear regression” to “multiple linear regression” in the Methods, Results, and Figure 3 legend (P. 24, P. 7, and P. 37)
We added the sentence “The results of the winning multiple regression model are reported in the Results section.” in the Methods (P. 25).
We added the results of the multiple regression analyses for genuine pain and pretended pain, in the Results section (P. 7-8): “For the genuine pain condition, we find that the modulatory effect was significantly related to the rating of painful feelings in others (t = 2.317, p = 0.026) but not related to the rating of either painful expressions in others (t = -1.492, p = 0.144) or unpleasantness in self (t = 0.058, p = 0.954). For the pretended pain condition, none of the ratings was significantly related to the modulatory effect (Figure 3D).”
We moved the results of the original stepwise regression analyses with behavioral ratings into the supplementary data (see Supplementary File 2):
“Results of the stepwise regression analyses on modulatory effects and behavioral ratings are shown below. Note that this analysis reflects our original analysis approach; prompted by a reviewer comment, we however changed the analysis plan and performed and reported the findings of multiple regression analyses in the main text. Importantly, the conclusions of the two analysis approaches are consistent.
To examine how the modulatory effects from the DCM were related to the behavioral ratings, we computed two stepwise linear regression models for each condition. The regression model was significant for the genuine pain condition (F model (1, 41) = 4.639, p = 0.037, R2 = 0.104), when painful feelings in others were added to the model and the other two ratings were excluded (B = 0.079, beta = 0.322, p = 0.037). However, the model was not significant for the pretended pain condition. The variance inflation factors (VIFs) for three ratings in both models were calculated to diagnose collinearity, showing no severe collinearity problem (all VIFs < 5; the smallest VIF =1.132 and the largest VIF = 4.387).”
- The multiple regression analyses revealed an association between the unpleasantness for the participants and the aIns, when accounting for the painful expression and the pain experienced by the other. This, however, does not reveal the specificity of the aIns for encoding the unpleasantness for the participants. It might well be that variance is shared in the association between the aIns and pain expression and pain by the other and unpleasantness for the participants, but simply strongest for unpleasantness. Such ambiguity could be resolved by additional multiple regressions of 1) pain expression (controlling for pain by the other and unpleasantness for the participants) and 2) pain for the other (controlling for pain expression and unpleasantness for the participants).
We thank the reviewer for this comment. As an overall premise, please note that we would not want to claim that the aIns is specifically engaged in encoding affective activities without any engagement of other processes; instead, we are entirely aware that the aIns activation participates in a variety of affective and cognitive processes. Nonetheless, our original multiple regression models were performed as a second-level group analysis with all three ratings as independent variables. Results showed that only the rating of “unpleasantness in self” was significant, rather than all ratings that were universally influenced by domain-general factors.
As the reviewer suggested, we additionally performed five multiple regression analyses with all possible orders of three behavioral measures to test whether the order matters. In the end, we found consistent results across all six regression analyses, suggesting that the selective correlation of aIns and the rating of unpleasantness in self was robust.
Action taken: In the manuscript, we have:
Modified “specifically” to “selectively” in the Results (P. 6).
Added the content in the Methods (P. 22) “To test whether the order of entering ratings into the regression model influence the results, we performed five additional regression analyses with all possible orders of three ratings. The results were consistent across all six regression models, and we only showed the result for one regression (i.e., expression + feeling + unpleasantness) in the Results section.
Modified the sentence in the Results (P. 6) “We found significant clusters in bilateral aIns, visual cortex, and cerebellum (Figure 2B); notably, when statistically accounting for ratings of painful expressions in others and painful feelings in others, all three clusters were exclusively explained by the ratings of self-unpleasantness.” to
“We found significant clusters in bilateral aIns, visual cortex, and cerebellum that could be selectively explained by the ratings of self-unpleasantness and could not be explained by either the ratings of painful expressions in others or painful feelings in others (Figure 2B).”
- Modified the sentence in the Discussion (P. 9) “[...] but the increased activation in aIns was also selectively correlated with ratings of self-oriented unpleasantness (i.e., after statistically accounting for painful expressions and painful feelings in others) [...]” to
“[...] but the increased activation in aIns was also selectively correlated with ratings of self- oriented unpleasantness and was not correlated with neither other-related painful expressions nor painful feelings in terms of the regression analysis [...]”
and added the sentence “[...] (otherwise the increased aIns activation should also be explained by other behavioral ratings in the sense of shared influence by domain-general effects).”
- Modified the legend for Figure 3 (P. 37) “[...] revealed a positive correlation between the inhibitory effect and painful feelings in others (after accounting for the other two ratings) for genuine pain [...]” to
“[...] revealed a positive correlation between the inhibitory effect and painful feelings in others and not with other two ratings for genuine pain [...]”
- Is the regression biased by the differences between conditions in the aIns in both fMRI signals and the ratings?
We thank the reviewer for this comment. The reason that we compared the differences between conditions was mainly aimed to control for potential effects of perceptual salience. This aim was consistent for both fMRI signals and behavioral ratings. Note that, as the aIns activation and all behavioral ratings were higher for genuine pain as opposed to pretended pain, the current result could not be explained by an inverse effect (i.e., higher aIns activation and higher ratings of unpleasantness in self for pretended pain). Therefore, we do not consider it is problematic to use differences between conditions when performing the multiple regression analysis.
Action taken:
We have more explicitly specified “differences between conditions for” three behavioral ratings as independent variables for the multiple regression model in the Methods (P. 22).
We added the sentence “The reason that we used the comparison between conditions for both brain signals and behavioral ratings was to control for potential effects of perceptual salience.” In the Methods (P. 22).
- The inclusion of the rSMG into the DCM model is not straight forward for me. It could have been based on previous literature, but then the aMCC should have been added as well. Furthermore, while the implication of the rSMG in distinction of self vs. others is established, the actual process in this experiment cannot be revealed. The authors state that the rSMG is involved in action observation or imitating emotions (page 9, line 200).
We appreciate the reviewer’s comment that shows we seemed not to convey clearly why we have postulated a role of rSMG. We have now made our rationale more explicit and clear.
Action taken:
We have now:
Modified the clarification of rSMG in the Discussion (P. 10): “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 et al., 2017; Hawco et al., 2017). Importantly, a specific role in affective rather than cognitive self-other distinction has been consistently identified for rSMG (Silani et al., 2013; Steinbeis et al., 2015; Bukowski et al., 2020). [...]”
Added further clarification in the Discussion (P. 12) after the sentence “ [...] the correlation findings provide further evidence that the modulation of aIns to rSMG is implicated in encoding others’ emotional states,”
with “which serves as a functional foundation for self-other processing [...] This regulation cannot be totally attributed to domain-general processes, otherwise other ratings should have also explained this variation.”
Additionally, we agree re: aMCC, which we also predicted to play a role; but it was not the case at least in our data. In fact, we have already addressed this in the original version of the ms. (maintained on P. 7 of revised ms.): “Our original analysis plan was to include aMCC in the DCM analyses, but based on the fact that aMCC did not show as strong evidence (in terms of the multiple regression analysis) as the aIns of being involved in our task, we decided to use a more parsimonious DCM model without the aMCC.”
Whether Results support their conclusions:
The results support the distinction between the experimental conditions of genuine vs. pretended pain in the aIns and as a modulatory influence on the connectivity between the aIns and the rSMG. However, the authors aimed to test if genuine vs. pretended pain modulate regulatory influences from the aIns on the rSMG that are connected to self-other distinction (as proposed in the discussion page 8, line 170). Yet, any insights about self-other distinction are only inferred reversely, since there is no outcome that indicates how well participants distinguished between themselves and the other person. For example in the discussion the authors state that: " we thus propose that the higher rSMG engagement in genuine pain conditions reflects an increasing demand for self-other distinction imposed by the stronger shared negative affect experiences in this condition". This is not supported by the results. Furthermore, the title mentioned automated responses to pretended pain, which I could not understand, given the current results.
We thank the reviewer for this comment, which somewhat follows up on similar arguments made and replied to in comment #1 above. Indeed, we fully agree that our design did not allow us to quantify self-other distinction, but that we inferred its engagement based on a strong theoretical motivation and the replication of previous findings on rSMG involvement during self-other distinction. As outlined above (cf. #1), this limitation was added to the revised manuscript.
We also adjusted the way of reasoning for which we put the theoretical explanation ahead of the inference so that readers can better realize this statement is supported by stronger theoretical motivation in the Discussion (P. 10):
First “Theoretical models of empathy [...]” and then “Concerning the current finding, we thus propose that [...]”
We thank the reviewer for pointing out the potential ambiguity in the title. We agree it may be somewhat “imprecise”, and have revised the title accordingly (P. 1):
“Neural dynamics between anterior insular cortex and right supramarginal gyrus dissociate genuine affect sharing from perceptual saliency of pretended pain”.
Likely impact of the work on the field:
These results are expected to advance the field, since they allow to disentangle visual expressions of pain from genuine pain in others. Thereby, this work could resolves the question about neural processes that are specific to pain in others beyond other salient cues.
We thank the reviewer for this positive acclaim of our study.
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Evaluation Summary:
Lamm and colleagues explore brain dynamics of empathy in response to facial expressions of simulated versus genuine pain in others. Using a novel experimental fMRI task and dynamic causal modelling, the findings suggest that activation of the anterior insula for genuine pain reflects affect sharing rather than automatic responses triggered by the perceptual salience of events. The paper is of broad interest to an audience of social and affective neuroscientists interested in how humans track the emotional responses of others.
(This preprint has been reviewed by eLife. We include the public reviews from the reviewers here; the authors also receive private feedback with suggested changes to the manuscript. The reviewers remained anonymous to the authors.)
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Reviewer #1 (Public Review):
Lamm and colleagues explore how the brain regulates our empathic response to others in genuine versus simulated pain. Using a clever experimental design, the authors sought to adjudicate between two conflicting perspectives in the literature - whether the empathic response reflects the alignment of neural activity typically engaged for the affective component of self-experienced pain or whether such activity is better accounted by an automatic response to salient perceptual cues (pain vividly expressed on the face). They used carefully designed videos of actors responding to painful stimuli or pretending to respond in pain. The experimental design provided an elegant way of teasing apart underlying affect, controlling for the perceptual saliency of the facial pain expression.Empathy-related activity in …
Reviewer #1 (Public Review):
Lamm and colleagues explore how the brain regulates our empathic response to others in genuine versus simulated pain. Using a clever experimental design, the authors sought to adjudicate between two conflicting perspectives in the literature - whether the empathic response reflects the alignment of neural activity typically engaged for the affective component of self-experienced pain or whether such activity is better accounted by an automatic response to salient perceptual cues (pain vividly expressed on the face). They used carefully designed videos of actors responding to painful stimuli or pretending to respond in pain. The experimental design provided an elegant way of teasing apart underlying affect, controlling for the perceptual saliency of the facial pain expression.Empathy-related activity in the anterior insula was found to reflect affective sharing; a fundamental component of the empathic response. Using dynamic causal modelling, the authors explored the interactions between affective responses and self-other differentiation, focusing on the right supramarginal gyrus as a major hub previously been implicated in the self-other distinction. They found a dynamic partnership between the anterior insula and the right supramarginal gyrus which enables humans to track the genuine emotional response of another person and to regulate one's own response accordingly.
This study is very well-conducted with clear hypotheses that are grounded in theory and novel methodology that enables the authors to explore their hypotheses in a carefully controlled manner. The interpretation of findings is measured, limitations are appropriately discussed and directions for future study are presented.
Overall, these findings provide a novel lens through which we can understand the processes by which humans respond to the genuine suffering of others and how we regulate our responses accordingly.
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Reviewer #2 (Public Review):
The authors provide an experiment in which videos with painful and non-painful stimulation of unknown persons are shown. In the genuine condition one person is shown who undergoes a painful experience, whereas another person in the pretentious condition is shown who undergoes a non-painful experience, but has a facial expression of pain (both persons are only pretending). The authors found higher rating of pain expression, painful feeling in others and higher unpleasantness related to the participant for videos with painful stimulation in the genuine condition. Mass-univariate fMRI activity in a regression analysis revealed a relation between higher unpleasantness related to the participant that was associated with activation in the anterior insular cortex (aIns). The authors further build a dynamic causal …
Reviewer #2 (Public Review):
The authors provide an experiment in which videos with painful and non-painful stimulation of unknown persons are shown. In the genuine condition one person is shown who undergoes a painful experience, whereas another person in the pretentious condition is shown who undergoes a non-painful experience, but has a facial expression of pain (both persons are only pretending). The authors found higher rating of pain expression, painful feeling in others and higher unpleasantness related to the participant for videos with painful stimulation in the genuine condition. Mass-univariate fMRI activity in a regression analysis revealed a relation between higher unpleasantness related to the participant that was associated with activation in the anterior insular cortex (aIns). The authors further build a dynamic causal model (DCM) that included the aIns and the rSMG and showed that neural activity during videos with painful stimulation that was modulated in genuine vs. pretended condition. The authors interpret these findings by a model in which genuine (vs. pretented) pain in others modulates the influence of aIns activity on the rSMG to regulate self-other distinction.
An account of the major strengths:
The experimental design is well controlled for pain vs. no pain condition. The genuine vs. the pretended condition allows to disentangle visual expression of pain from actual pain in others (at least in the pilot study).
The DCM is an elegant method to test the modulatory influence of genuine vs. pretended pain on two brain regions.Weaknesses of the Methods and Results:
In my view, the experiment does not allow to unambiguously disentangle self vs. other distinction (as mentioned in the abstract "..we investigated how affect sharing and self-other distinction interact.."). For example, genuine vs. pretended pain could be distinguished from the participants own experience in a comparable way. The higher rating of unpleasantness for genuine pain in others does not necessary mean that the participants cannot separate own from others experiences.
The experimental design does not unambiguously allows to disentangle genuine vs pretended pain from other factors, such as the differences in pain expression, painful feeling in others and higher unpleasantness in these two conditions. I understand that the intensity pain expression, painful feeling in others and unpleasantness for others is inherently tied to genuine vs. pretended pain. But the author already saw that the instruction of "genuine vs. pretented" influenced the ratings of pain expression. Hence, this allows two interpretation of the results: either the influence from the anterior Insula on the rSMG is driven by higher perceived pain expression, painful feeling in others and unpleasantness or by the conditions of genuine vs. pretended pain. Or (more likely) by an interaction between these factors.
It would, for example help to explore the association between the aIns-rSMG interaction pain expression ratings (or painful feeling in others or higher unpleasantness) in videos with genuine pain und pretended pain separately.
The author should further discuss this point that different factors (pain expression, etc) contribute to the differences between genuine vs. pretended pain.The multiple regression analyses revealed an association between the unpleasantness for the participants and the aIns, when accounting for the painful expression and the pain experienced by the other. This, however, does not reveal the specificity of the aIns for encoding the unpleasantness for the participants.
It might well be that variance is shared in the association between the aIns and pain expression and pain by the other and unpleasantness for the participants, but simply strongest for unpleasantness.
Such ambiguity could be resolved by additional multiple regressions of 1) pain expression (controlling for pain by the other and unpleasantness for the participants) and 2) pain for the other (controlling for pain expression and unpleasantness for the participants).Is the regression biased by the differences between conditions in the aIns in both fMRI signals and the ratings?
The inclusion of the rSMG into the DCM model is not straight forward for me. It could have been based on previous literature, but then the aMCC should have been added as well. Furthermore, while the implication of the rSMG in distinction of self vs. others is established, the actual process in this experiment cannot be revealed. The authors state that the rSMG is involved in action observation or imitating emotions (page 9, line 200).
Whether Results support their conclusions:
The results support the distinction between the experimental conditions of genuine vs. pretended pain in the aIns and as a modulatory influence on the connectivity between the aIns and the rSMG. However, the authors aimed to test if genuine vs. pretended pain modulate regulatory influences from the aIns on the rSMG that are connected to self-other distinction (as proposed in the discussion page 8, line 170). Yet, any insights about self-other distinction are only inferred reversely, since there is no outcome that indicates how well participants distinguished between themselves and the other person. For example in the discussion the authors state that: " we thus propose that the higher rSMG engagement in genuine pain conditions reflects an increasing demand for self-other distinction imposed by the stronger shared negative affect experiences in this condition". This is not supported by the results.
Furthermore, the title mentioned automated responses to pretended pain, which I could not understand, given the current results.Likely impact of the work on the field:
These results are expected to advance the field, since they allow to disentangle visual expressions of pain from genuine pain in others. Thereby, this work could resolves the question about neural processes that are specific to pain in others beyond other salient cues.
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