Cited by 653Psychology of Moral and Emotional JudgmentNeural and Behavioral Psychology StudiesAction Observation and Synchronization

Parsing the neural correlates of moral cognition: ALE meta-analysis on morality, theory of mind, and empathy

Open access
Bzdok, Danilo, Schilbach, Leonhard, Vogeley, Kai, Schneider, Karla, Laird, Angela R., Langner, Robert, Eickhoff, Simon B.·2012·Brain Structure and Function

Abstract

Morally judicious behavior forms the fabric of human sociality. Here, we sought to investigate neural activity associated with different facets of moral thought. Previous research suggests that the cognitive and emotional sources of moral decisions might be closely related to theory of mind, an abstract-cognitive skill, and empathy, a rapid-emotional skill. That is, moral decisions are thought to crucially refer to other persons' representation of intentions and behavioral outcomes as well as (vicariously experienced) emotional states. We thus hypothesized that moral decisions might be implemented in brain areas engaged in 'theory of mind' and empathy. This assumption was tested by conducting a large-scale activation likelihood estimation (ALE) meta-analysis of neuroimaging studies, which assessed 2,607 peak coordinates from 247 experiments in 1,790 participants. The brain areas that were consistently involved in moral decisions showed more convergence with the ALE analysis targeting theory of mind versus empathy. More specifically, the neurotopographical overlap between morality and empathy disfavors a role of affective sharing during moral decisions. Ultimately, our results provide evidence that the neural network underlying moral decisions is probably domain-global and might be dissociable into cognitive and affective sub-systems.

AI snapshot

Written from the paper’s full text.

TL;DR

This large-scale ALE meta-analysis found that moral cognition recruits brain areas overlapping more with theory of mind than with empathy, supporting a domain-global view with dissociable cognitive and affective subsystems. It identified dmPFC, right MTG, and bilateral TPJ as common across moral cognition, theory of mind, and empathy.

Key findings

  • Across 2,607 peak coordinates from 247 experiments in 1,790 participants, moral cognition consistently engaged bilateral vmPFC/FP/dmPFC, precuneus, TPJ, PCC, right temporal pole, right MTG, and left amygdala.
  • Conjunction analyses showed morality overlapped with ToM-minus-empathy in bilateral vmPFC/FP/dmPFC and TPJ, right MTG, and right temporal pole; morality overlapped with empathy-minus-ToM only in dmPFC; all three domains converged in dmPFC, right MTG, and bilateral TPJ.
  • Empathy’s most prominent convergence was in aMCC/SMA and AI/IFG, which did not overlap with morality, leading the authors to conclude that affective sharing is unlikely to be involved in moral judgments and that moral cognition is closer to ToM than to empathy.
  • BrainMap functional characterization linked the common network to ToM, semantic processing, imagination, social cognition, and episodic/explicit memory.

Methods

  • PubMed search for fMRI/PET studies on moral cognition, ToM, and empathy up to 2010; inclusion required full-brain coverage, no pharmacological manipulations, lesions, or mental/neurological disorders, and whole-brain group coordinates in Talairach/MNI space. Exclusions included ROI analyses, empathy-for-pain studies, and static facial emotion-recognition tasks. The search yielded 67 moral experiments/507 foci, 68 ToM experiments/724 foci, and 112 empathy experiments/1,376 foci.
  • Revised ALE coordinate-based meta-analysis modeled foci as 3D Gaussian probability distributions, computed modeled activation maps, and thresholded at cluster-level-corrected p < 0.05. Conjunction analyses used the minimum statistic; difference analyses used 10,000 randomizations with p < 0.001; BrainMap metadata were tested with binomial tests Bonferroni-corrected for multiple comparisons.

Limitations

  • The paper states that meta-analyses are affected by publication bias against null results, and that BrainMap contains only about 21% of published neuroimaging studies, so its functional characterization is not strictly representative.
  • The paper states that some moral cognition tasks have limited ecological validity; artificial scenarios such as the trolley dilemma may have systematically overestimated cognitive versus emotional processes and contributed to the similarity between moral cognition and ToM.
  • The paper states that no statistical method to correct for multiple comparisons in ALE difference maps has yet been established, and it notes that meta-analytic spatial resolution limits interpretation of adjacent but non-overlapping PCC clusters.

Model-written summary. Check it against the paper before relying on it.

Integrity signals

Registry

Crossref · 10.1007/s00429-012-0380-y

Cites

140 of 140 references verified0 citation errors · 0 not checkable

Statistics

Nothing to recompute.

Claims the paper makes

0 of 103 cited sources checked · 0 supported · 0 flagged

  • Findingown data

    Convergence in the posterior cingulate cortex was located slightly more rostro-dorsally in the ALE on moral cognition compared to the ALE on empathy.

    “convergence in the PCC was located slightly more rostro-dorsally in the ALE on moral cognition compared to the ALE on empathy”
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  • Findingown data

    Meta-analysis of studies related to theory of mind (ToM) revealed convergence in the bilateral vmPFC/FP/dmPFC, precuneus, TPJ, temporal pole, middle temporal gyrus, posterior superior temporal sulcus (pSTS), and inferior frontal gyrus (IFG), as well as the right MT/V5.

    “Meta-analysis of studies related to ToM revealed convergence in the bilateral vmPFC/FP/dmPFC, Prec, TPJ, TP, MTG, posterior superior temporal sulcus (pSTS), and inferior frontal gyrus (IFG), as well as the right MT/V5.”
    Quote in the paperSubstantial claim
  • Findingown data

    The conjunction analysis across the results of all three individual meta-analyses on moral cognition, ToM, and empathy yielded convergence in the dmPFC, right MTG, and bilateral TPJ.

    “the conjunction analysis across the results of all three individual meta-analyses on moral cognition, ToM, and empathy yielded convergence in the dmPFC, right MTG, and bilateral TPJ”
    Quote in the paperSubstantial claim
  • Findingown data

    The conjunction across the neural network linked to moral cognition and the activation more robustly linked to empathy versus ToM revealed an overlap in the dmPFC.

    “the conjunction across the neural network linked to moral cognition and the activation more robustly linked to empathy versus ToM revealed an overlap in the dmPFC”
    Quote in the paperSubstantial claim
  • Findingown data

    The common network observed across all three task domains was selectively associated with tasks tapping ToM, semantic processing, imagination, and social cognition, as well as episodic and explicit memory.

    “the common network observed across all three task domains was selectively associated with tasks tapping ToM, semantic processing, imagination, and social cognition, as well as episodic and explicit memory”
    Quote in the paperSubstantial claim
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  • Classified as probably not a substantial claim. 27
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    Findingown data

    The left amygdala was found in the analysis of moral cognition-related brain activity, while the right amygdala was found in the analysis of empathy-related brain activity.

    “the left AM was found in the analysis of moral cognition-related brain activity, while the right AM was found in the analysis of empathy-related brain activity”
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    Findingown data

    The AI/IFG and aMCC/SMA were revealed as the most significant points of convergence in the ALE on non-pain empathy but did not show any overlap with the ALE on morality.

    “Furthermore, it is important to note that the AI/IFG and aMCC/SMA were revealed as the most significant points of convergence in the ALE on non-pain empathy but did not show any overlap with the ALE on morality.”
    Quote in the paperSubstantial claim
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    Findingown data

    Brain activity related to both moral cognition and non-pain empathy converged significantly in an area of the dmPFC, which was not revealed by the ALE on ToM.

    “Brain activity related to both moral cognition and non-pain empathy converged significantly in an area of the dmPFC, which was not revealed by the ALE on ToM.”
    Quote in the paperSubstantial claim
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    Findingown data

    The neural correlates of moral judgments are much closer related to the neural correlates of ToM than to those of empathy, as evidenced by the quantity of overlap in the conjunction analyses.

    “Third, the neural correlates of moral judgments are much closer related to the neural correlates of ToM than to those of empathy, as evidenced by the quantity of overlap in the conjunction analyses.”
    Quote in the paperSubstantial claim
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    Findingown data

    The AM was significantly involved in the individual ALEs on morality and non-pain empathy, although in contralateral hemispheres.

    “It is interesting to note that the AM was significantly involved in the individual ALEs on morality and non-pain empathy, although in contralateral hemispheres.”
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    Interpretationreasoning

    Empathy for pain- and non-pain-related affect appears to be implemented by an overlapping network that might recruit supplementary areas/networks depending on the specific affective modality with which participants are to empathize.

    “empathy for pain- and non-pain-related affect appears to be implemented by an overlapping network that might recruit supplementary areas/networks depending on the specific affective modality with which participants are to empathize”
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    Study design and conductnone given

    Study selection was grounded on whether the original authors claimed to have isolated brain activity related to moral cognition, ToM, or empathy.

    “study selection was grounded on the objective measure whether or not the authors claimed to have isolated brain activity that relates to moral cognition, ToM, or empathy”
    Quote in the paperSubstantial claim
    • Rests on an assumption: The task categories used by the original neuroimaging studies validly isolate moral cognition, theory of mind, or empathy.
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    Study design and conductnone given

    The meta-analysis included only studies with full brain coverage and absence of pharmacological manipulations, brain lesions, or mental/neurological disorders.

    “The inclusion criteria comprised full brain coverage as well as absence of pharmacological manipulations, brain lesions or mental/neurological disorders.”
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    Only neuroimaging studies that aimed at eliciting the conscious and isomorphic experience of somebody else's affective state were included in the empathy category.

    “we only included those neuroimaging studies into the “empathy” category that aimed at eliciting the conscious and isomorphic experience of somebody else’s affective state”
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    Study design and conductnone given

    Only neuroimaging studies that required participants to adopt an intentional stance towards others were included in the ToM category.

    “we only included those neuroimaging studies into the “ToM” category that required participants to adopt an intentional stance towards others”
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    • Rests on an assumption: The task categories used by the original neuroimaging studies validly isolate moral cognition, theory of mind, or empathy.
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    Premisecitation

    ToM and empathic skills precede mature moral reflection in primate evolution and in ontogeny.

    “ToM and empathic skills precede mature moral reflection in primate evolution (Greene and Haidt 2002) and in ontogeny (Tomasello 2001; Kohlberg et al. 1983; Frith and Frith 2003; Piaget 1932)”
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    Only neuroimaging studies that required participants to make appropriateness judgments on actions of one individual towards others were included in the morality category.

    “we only included those neuroimaging studies into the “morality” category that required participants to make appropriateness judgments on actions of one individual towards others”
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    Premisecitation

    More automatic, bottom-up driven mapping and awareness of others' emotional states in empathy might be modulated by more controlled, top-down processes involved in attributing mind states (ToM).

    “more automatic, bottom-up driven mapping and awareness of others emotional states in the context of self–other distinction (i.e., empathy) might be modulated by more controlled, top-down processes involved in attributing mind states (i.e., ToM)”
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    Premisecitation

    ToM and empathy differ in representational content (mental states versus affect), yet both might be similarly brought about by interaction between top-down and bottom-up processes.

    “ToM and empathy differ in the representational content (mental states versus affect), yet both might be similarly brought about by interaction between top-down and bottom-up processes”
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    Premisecitation

    Deficient empathy skills in psychopathic populations are believed to contribute to morally inappropriate behavior.

    “the deficient empathy skills in psychopathic populations are believed to contribute to morally inappropriate behavior”
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    Experiencing empathy was shown to alleviate harmful actions towards others.

    “experiencing empathy was shown to alleviate harmful actions towards others”
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    Study design and conductnone given

    Studies were only considered if they reported results of whole-brain group analyses as coordinates corresponding to a standard reference space (Talairach/Tournoux, MNI).

    “studies were only considered, if they reported results of whole-brain group analyses as coordinates corresponding to a standard reference space (Talairach/Tournoux, MNI)”
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    Premisecitation

    Embodied representation and meta-representation might not constitute two mutually exclusive processes.

    “embodied representation and meta-representation might not constitute two mutually exclusive processes”
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    Experiments assessing neural effects in a priori defined regions of interest were excluded.

    “experiments assessing neural effects in a priori defined regions of interest were excluded”
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    Study design and conductcitation

    A random-effects inference was invoked, focusing on above-chance convergence across different experiments.

    “a random-effects inference was invoked, focusing on the above-chance convergence across different experiments”
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    Premisecitation

    The PCC is thought important for the modality-independent retrieval of autobiographical memories and their integration with current emotional states.

    “The PCC is thought important for the modality-independent retrieval of autobiographical memories and their integration with current emotional states (Fink et al. 1996; Maddock et al. 2001; Maddock 1999; Schilbach et al. 2008a).”
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    Interoception-related meta-representation of emotion has been suggested to underlie the concomitant involvement of the aMCC/SMA and AI/IFG in neuroimaging studies on empathy as reflecting affective sharing.

    “Interoception-related meta-representation of emotion has thus been suggested to underlie the concomitant involvement of the aMCC/SMA and AI/IFG in neuroimaging studies on empathy as reflecting affective sharing (Singer and Lamm 2009; Fan et al. 2011).”
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    Premisecitation

    AM activity typically increases in the left hemisphere in elaborate social-cognitive processes and in the right hemisphere in automatic, basic emotional processes.

    “In particular, AM activity typically increases in the left hemisphere in elaborate social-cognitive processes and in the right hemisphere in automatic, basic emotional processes (Phelps et al. 2001; Bzdok et al. 2012; Markowitsch 1998; Gläscher and Adolphs 2003).”
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    The AM is thought to automatically extract biological significance from the environment.

    “This brain region is thought to automatically extract biological significance from the environment (Ball et al. 2007; Sander et al. 2003; Bzdok et al. 2011; Müller et al. 2011).”
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    Study design and conductcitation

    The resulting non-parametric p values for difference analyses were thresholded at p < 0.001.

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    Functional characterization used a binomial test, Bonferroni-corrected for multiple comparisons, to identify functional roles of the convergent network.

    “Using a binomial test, Bonferroni-corrected for multiple comparisons, the functional roles of the convergent network as a whole were identified”
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    Interpretationown data

    General affective processes might play a role in both moral judgments and empathy, given that the respective meta-analyses individually revealed the dmPFC (direct overlap) and the amygdala (oppositely lateralized).

    “Second, general affective processes might play a role in both moral judgments and empathy, given that the respective meta-analyses individually revealed the dmPFC (direct overlap) and the amygdala (oppositely lateralized).”
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    Premisecitation

    This network, especially the anterior insula, is not only implicated in meta-representation of emotional states but also in interoceptive awareness.

    “Moreover, this network, especially the anterior insula, is not only implicated in meta-representation of emotional states but also in interoceptive awareness (Craig 2002, 2009; Kurth et al. 2010).”
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    Premisecitation

    The AI/IFG and aMCC/SMA form a network that is widely believed to represent one's own and others' emotional states regardless of the actual affective or sensory modality.

    “In particular, the AI/IFG and aMCC/SMA form a network that is widely believed to represent one’s own and others’ emotional states regardless of the actual affective or sensory modality (Lamm et al. 2011; Wicker et al. 2003; Singer et al. 2004; Lamm and Singer 2010; Fan et al. 2011).”
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    The relevant fMRI studies crucially hinge on the repeatedly criticized functional localizer technique, which rests on strong a priori hypotheses.

    “Second, the relevant fMRI studies crucially hinge on the repeatedly criticized functional localizer technique, which rests on strong a priori hypotheses (Friston et al. 2006; Mitchell 2008).”
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    Study design and conductcitation

    The resulting ALE scores were tested against the earlier calculated true ALE scores and cut off at a cluster-level-corrected threshold of p < 0.05.

    “The resulting ALE scores were tested against the earlier calculated ‘true’ ALE scores and cut off at a cluster-level-corrected threshold of p < 0.05.”
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    Study design and conductnone given

    The randomization procedure employed to compute the contrast between ALE analyses is highly conservative as it estimates the probability for a true difference between the two datasets.

    “the randomization procedure employed to compute the contrast between ALE-analyses is in itself highly conservative as it estimates the probability for a true difference between the two datasets”
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    Premisecitation

    Studies on empathy for pain were disregarded because pain is not considered a classic emotion and watching painful scenes does not induce isomorphic vicarious experiences.

    “we disregarded studies on empathy for pain because pain, although possessing an affective dimension, is not considered a classic emotion (Izard 1971; Ekman 1982). Rather, it is a bodily sensation mediated by distinct sensory receptors (Craig 2002; Saper 2000), and watching painful scenes does not induce isomorphic vicarious experiences”
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    A statistical method to correct for multiple comparisons when assessing differences between ALE maps has not yet been established.

    “Unfortunately, a statistical method to correct for multiple comparisons when assessing the differences between ALE maps has not yet been established.”
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    Premisecitation

    Meta-analyses are necessarily based on the available literature and hence may be affected by the potential publication-bias disfavoring null results.

    “Meta-analyses are necessarily based on the available literature and hence may be affected by the potential publication-bias disfavoring null results (Rosenthal 1979).”
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    Similar brain mechanisms in remembering past episodes and envisioning future circumstances is supported by their engagement of identical brain areas, as evidenced by a quantitative meta-analysis.

    “Similar brain mechanisms in remembering past episodes and envisioning future circumstances is supported by their engagement of identical brain areas, as evidenced by a quantitative meta-analysis (Spreng et al. 2009).”
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    Interpretationcitation

    These four psychological categories (ToM, explicit memory retrieval, language, and imagination of objects/scenes) summarize what sets humans probably apart from non-human primates.

    “Intriguingly, these four seemingly disparate psychological categories summarize what sets humans probably apart from non-human primates (Tomasello 2001; Frith and Frith 2010).”
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    These isolated conceptual scripts may be reassembled to enable the forecasting of future events.

    “These isolated conceptual scripts may be reassembled to enable the forecasting of future events (Tulving 1983, 1985; Schacter et al. 2007).”
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    Construction of detached probabilistic scenes has been argued to influence ongoing decision making by estimating outcomes of behavioral choices.

    “Indeed, construction of detached probabilistic scenes has been argued to influence ongoing decision making by estimating outcomes of behavioral choices (Boyer 2008; Suddendorf and Corballis 2007; Schilbach et al. 2008b).”
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    Retrograde amnesic patients were reported to be impaired not only in prospection but also in imagining novel experience.

    “Moreover, retrograde amnesic patients were reported to be impaired not only in prospection but also in imagining novel experience (Hassabis et al. 2007).”
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    Neither Greene's nor Moll's concept can exhaustively explain why vmPFC patients demonstrated a rationally biased approach to solving moral dilemmas, yet an emotionally biased approach to moral cognition in an economic game.

    “Apart from that, neither Greene’s nor Moll’s concept can exhaustively explain why vmPFC patients demonstrated a rationally biased approach to solving moral dilemmas (Koenigs et al. 2007; Moretto et al. 2010), yet, an emotionally biased approach to moral cognition in an economic game (Koenigs and Tranel 2007).”
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    Interpretationcitation

    This caveat might have contributed to the similar results in the meta-analyses on moral cognition and on ToM (rather than on empathy), given that emotion processing is thought to play a paramount role in real-world moral cognition.

    “This caveat might have contributed to the similar results in the meta-analyses on moral cognition and on ToM (rather than on empathy), given that emotion processing is thought to play a paramount role in real-world moral cognition (Krebs 2008; Tangney et al. 2007; Haidt 2001).”
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    Interpretationown data

    The present large-scale meta-analysis provides evidence for a domain-global view of moral cognition, rather than for a distinct moral module, by showing its functional integration of distributed brain networks.

    “The present large-scale meta-analysis provides evidence for a domain-global view of moral cognition, rather than for a distinct moral module (Hauser 2006), by showing its functional integration of distributed brain networks.”
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    Autobiographical memory supplies numerous building blocks of social semantic knowledge.

    “In particular, autobiographical memory supplies numerous building blocks of social semantic knowledge (Bar 2007; Binder et al. 2009).”
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    Interpretationown data

    Our results support the notion that moral reasoning is related to both seeing things from other persons' points of view and to grasping others' feelings.

    “Ultimately, our results support the notion that moral reasoning is related to both seeing things from other persons’ points of view and to grasping others’ feelings (Piaget 1932; Tomasello 2001; Decety et al. 2012).”
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    • cites Decety et al. 2012

      Not checked in the free run.

45 of 45
  • The paper relies on this without support.

    The task categories used by the original neuroimaging studies validly isolate moral cognition, theory of mind, or empathy.

    “study selection was grounded on the objective measure whether or not the authors claimed to have isolated brain activity that relates to moral cognition, ToM, or empathy”
  • The paper relies on this without support.

    Convergence of activation across independent neuroimaging studies indicates shared neural processes or common network involvement.

    “Moral cognition-, ToM-, and empathy-related brain activity converged in the bilateral TPJ, dmPFC, and right MTG, which therefore form a common network potentially involved in social-cognitive processes.”
  • The paper relies on this without support.

    Lack of consistent involvement of a brain region in a meta-analysis of moral paradigms indicates that the corresponding psychological process is unlikely to be involved in moral judgments.

    “First, affective sharing, one core aspect of empathy, is unlikely to be involved in moral judgments, given the lack of consistent involvement of the AI/IFG or aMCC/SMA in paradigms probing the latter.”
  • The paper relies on this without support.

    The BrainMap database's metadata categories validly characterize the functional associations of the converging network.

    “In line with this, metadata profiling demonstrates solid associations of this network with neuroimaging studies related to ToM, explicit memory retrieval, language, and imagination of objects/scenes.”
4 of 4

Claims other papers make about this one

Each is checked against this paper’s own text: does it say what the citing paper says it does?

Cites

Flagged first, then by citations

Research works printed but not linked: 1

  1. Izard CE (1971) The face of emotion. Appleton-Century-Crofts, New York

Cited by

Sorted by citations

Authors
Bzdok, D., Schilbach, L., Vogeley, K., Schneider, K., Laird, A. R., Langner, R., & Eickhoff, S. B.
Affiliations
  • RWTH Aachen University (DE)
  • University of Cologne (DE)
  • Forschungszentrum Jülich (DE)
  • Jülich Aachen Research Alliance (DE)
  • The University of Texas at San Antonio Health Science Center (US)
Corresponding author
Eickhoff, Simon B.
Venue
Brain Structure and Function
Published
2012-01-24
Volume
217
Issue
4
Pages
783-796
ISSN
  • 1863-2653 (ISSN-L)
  • 1863-2661
OpenAlex
W2055132787
Cited by
653
References
140
Citations by year
2026: 32 · 2025: 22 · 2024: 48 · 2023: 38 · 2022: 53 · 2021: 85 · 2020: 74 · 2019: 65 · 2018: 45 · 2017: 52 · 2016: 43 · 2015: 37 · 2014: 33 · 2013: 21 · 2012: 5
FWCI
15.52
Type
journal article
Language
en
Open access
Locations
  • Brain Structure and Function (publishedVersion, cc-by-nc)
  • Europe PMC (PubMed Central) (publishedVersion)
  • PubMed Central (publishedVersion)
  • PubMed (publishedVersion)
  • Kölner Universitäts PublikationsServer (Universität zu Köln) (publishedVersion)
  • Max Planck Institute for Plasma Physics (submittedVersion)
  • JuSER (Forschungszentrum Jülich) (submittedVersion)
  • RWTH Publications (RWTH Aachen) (submittedVersion)
  • MPG.PuRe (Max Planck Society) (submittedVersion)
Funders
  • Deutsche Forschungsgemeinschaft (IRTG1328, DFG. IRTG 1328, R01-MH074457)
  • National Institute of Mental Health (R01MH074457)
Topics
  • Life Sciences › Neuroscience › Cognitive Neuroscience › Psychology of Moral and Emotional Judgment
  • Life Sciences › Neuroscience › Cognitive Neuroscience › Neural and Behavioral Psychology Studies
  • Social Sciences › Psychology › Social Psychology › Action Observation and Synchronization
Keywords
moral cognition, moral judgment, theory of mind, empathy, neural correlates of morality, affective sharing
MeSH terms
Brain Mapping, Cognition, Empathy, Humans, Morals, Nerve Net, Likelihood Functions, Theory of Mind, Functional Neuroimaging

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Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.

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