Next Article in Journal
Strategic Processing of Gender Stereotypes in Sentence Comprehension: An ERP Study
Previous Article in Journal
Improving Effects of Peripheral Nerve Decompression Microsurgery of Lower Limbs in Patients with Diabetic Peripheral Neuropathy
Font Type:
Arial Georgia Verdana
Font Size:
Aa Aa Aa
Line Spacing:
Column Width:
Systematic Review

The Neural Signatures of Shame, Embarrassment, and Guilt: A Voxel-Based Meta-Analysis on Functional Neuroimaging Studies

Clinical and Affective Neuroscience Laboratory, Department of Psychology and Cognitive Sciences, University of Trento, 38068 Rovereto, Italy
Marica de Vincenzi Onlus Foundation, 38122 Trento, Italy
Mental Health Services in the Capital Region of Denmark, Psychiatric Center Ballerup, 2750 Ballerup, Denmark
Neuroscience and Society Laboratory, Neuroscience Area, SISSA, 34136 Trieste, Italy
Center for Medical Sciences, University of Trento, 38122 Trento, Italy
Author to whom correspondence should be addressed.
Brain Sci. 2023, 13(4), 559;
Submission received: 30 December 2022 / Revised: 10 March 2023 / Accepted: 16 March 2023 / Published: 26 March 2023
(This article belongs to the Section Social Cognitive and Affective Neuroscience)


Self-conscious emotions, such as shame and guilt, play a fundamental role in regulating moral behaviour and in promoting the welfare of society. Despite their relevance, the neural bases of these emotions are uncertain. In the present meta-analysis, we performed a systematic literature review in order to single out functional neuroimaging studies on healthy individuals specifically investigating the neural substrates of shame, embarrassment, and guilt. Seventeen studies investigating the neural correlates of shame/embarrassment and seventeen studies investigating guilt brain representation met our inclusion criteria. The analyses revealed that both guilt and shame/embarrassment were associated with the activation of the left anterior insula, involved in emotional awareness processing and arousal. Guilt-specific areas were located within the left temporo-parietal junction, which is thought to be involved in social cognitive processes. Moreover, specific activations for shame/embarrassment involved areas related to social pain (dorsal anterior cingulate and thalamus) and behavioural inhibition (premotor cortex) networks. This pattern of results might reflect the distinct action tendencies associated with the two emotions.

1. Introduction

Moral emotions are crucial in regulating social interactions, as they promote the welfare of society or of other people [1] Indeed, they provide the emotional drive to properly behave in social interactions [2,3,4], forcing individuals to implement strategies that are optimal over a long period, even though they do not appear functional in the short period [5,6].
It has been proposed that moral cognition depends on the prefrontal, temporal, and limbic circuits, and is associated with the integration of context-independent and -dependent information and with the accompanying emotional reactions (event-feature-emotion complexes model, EFEC) [7,8]. Specifically, the prefrontal cortex seems to be responsible for representing context-dependent knowledge of event sequences [9,10], the temporal lobes for perceiving social cues and for representing context-independent social semantic knowledge [11,12,13], and the limbic system for the generation of emotional and motivational states [14]. Hence, according to the EFEC model, the generation of moral emotions, including self-conscious emotions, relies on the integrity of a network including prefrontal, temporal, and limbic areas [7,8].
Several studies investigating the neural substrates of moral cognition [15,16] confirmed the anatomical predictions of this model and better defined the topography of the brain areas associated with moral processing. Indeed, they showed that the ventromedial and dorsomedial prefrontal cortices (vmPFC and dmPFC, respectively), the temporo-parietal junction (TPJ), the precuneus, the posterior cingulate cortex, the left amygdala, the anterior temporal lobes (ATL), and the lateral orbitofrontal cortex were consistently found activated in neuroimaging studies investigating moral processing [3,4,15,16].
Among moral emotions, a sub-group of emotions (e.g., shame, embarrassment, guilt, and pride) defined as self-conscious emotions helps individuals to navigate the complexities of fitting into groups [1], satisfying the human need of belonging to social groups [17]. Self-conscious emotions are evoked by self-reflection and self-evaluation [18], and occur when social norms or agreed-upon social rules are violated [19], providing an immediate feedback that promotes inhibition or reinforcement of behaviour [3,4,18] (see Table 1). One case in point is shame, which has been proposed as an algorithm the brain uses to inhibit socially and morally unwanted behaviours [3,4].
While the EFEC model might explain the cognitive processes underlying all self-conscious emotions, which are all induced by moral and social norm violation [19], it does not make any prediction about the different processes that might occur in different types of emotions such as the negative self-conscious emotions. Indeed, even though shame, embarrassment, and guilt are often (but culpably) used interchangeably, they appear to be substantially different [20]. Shame is typically elicited by the belief that the individual’s violation of standards of morality, aesthetics, or competence defines who the individual is [21]. Hence, it involves the way the individual perceives themselves and how they believe other people see them and their inadequacy to fulfil social standards [22]. The distinction between shame and embarrassment is still a matter of debate (for review see [23]). If, on the one hand, embarrassment might be considered a dimension of shame [24], on the other, it might represent a distinct emotional entity [25,26]. Embarrassment seems related to trivial social transgressions, occurring suddenly and in public contexts, especially in the presence of individuals with equal or higher hierarchical social status [1,26,27,28,29]. Conversely, shame emerges when one personally perceives the serious violation of a moral norm, which might be also experienced in private situations [26,29]. Furthermore, shame and embarrassment also differ in intensity (i.e., shame is more intense than embarrassment) [30], in duration (i.e., shame is more persistent than embarrassment) [31], and in the focus of attention (i.e., shame affects the self, embarrassment affects the persona, the apparent self). However, these two emotions also have some features in common. They are associated with the same specific physiological reactions (e.g., blushing) [32] and the same action tendency, leading people to hide and reduce their social presence, making movement and speech more difficult and less likely [25,26,33,34]. However, it has also been reported that, differently from shame, embarrassment leads to reparative behaviours to re-gain social approval [25,35,36]. At the neural level, shame has been selectively associated with the dorso-lateral prefrontal cortex (dlPFC), the posterior cingulate cortex, and the sensory-motor cortex, whereas embarrassment has been associated with the ventro-lateral prefrontal cortex (vlPFC), the amygdala, and occipital areas, and both emotions with the hippocampus and midbrain [19]. However, it must be acknowledged that, since the distinction between shame and embarrassment is not sharp, being classified according to the private-public, moral-conventional, or low-high intensity dimensions, it is not easy to establish which brain areas are involved in processing these emotions, and which areas might selectively process one of the two emotions.
If the difference between shame and embarrassment is not as clear-cut, the distinction between guilt and the other two emotions is more evident. Guilt occurs when the violation of social norms induces harm or suffering to other individuals [3,4,37], typically in a relationship or among members of the same group [38]. Differently from shame and embarrassment, in which the self and the persona, respectively, are perceived as defective, in guilt, a specific action is typically perceived as wrong [34,37,39]. The occurrence of guilt induces remorse and behavioral responses that aim to repair the wrong action [18]. This difference in the focus of shame and guilt, self-oriented and other-oriented, respectively, has important consequences on empathy for other people: while guilt tends to increase the empathic concern towards other people, empathic responses seem to be disrupted by the self-oriented distress associated with shame [18].
In a review, Bastin and collaborators [19] suggested that guilt processing was selectively associated with the ventral anterior cingulate cortex (ACC), the precuneus, and premotor and posterior temporal areas. In addition, both guilt and shame processing were associated with the anterior insula and the dorsal ACC, and both guilt and embarrassment processing were associated with the dorsomedial prefrontal cortex (dmPFC), the vlPFC, and the anterior temporal lobe (ATL) [19]. Moreover, a recent meta-analysis [40] partially confirmed the guilt neural substrates proposed by Bastin and collaborators (2014), reporting the activation of the precuneus, the dorsal ACC, the dmPFC, and posterior temporal areas in association with guilt processing [40].
However, it is worth noting that studies investigating self-conscious emotions used heterogeneous methods that prevent any firm conclusions from being drawn. For this reason, we have run a meta-analysis study including neuroimaging research on the neural substrates of negative self-conscious emotions, i.e., to pinpoint brain areas consistently associated with shame/embarrassment and guilt processing. Since the distinction between shame and embarrassment is still a matter of debate, and since some of the studies in the literature did not distinguish clearly between the two emotions, we decided to include shame and embarrassment in one category. We predicted that shame/embarrassment and guilt may show different brain activations mirroring behavioural differences related to the emotions, together with some shared activations in light of their moral-self-conscious nature.

2. Materials and Methods

In order to find studies investigating the neural underpinnings of shame, embarrassment, and guilt, we conducted a research on PubMed (URL = accessed on 1 January 2022) using the terms ((“fMRI” OR “functional magnetic resonance imaging” OR “PET”) AND (“shame” OR “embarrassment” OR “guilt” OR “moral emotions” OR “self-conscious emotions” OR “moral violations” OR “social standard violation”)) and setting a range of dates between 1 January 1995 and 13 February 2023. This research identified 169 studies.
Subsequently, we refined our research by applying the following criteria:
Papers originally published in English;
fMRI or PET studies including task-related whole brain analyses. Studies reporting a region of interest (ROI analyses, resting-state fMRI analyses, diffusion tensor imaging (DTI), or voxel-based morphometry (VBM)) were excluded;
Participants were healthy adults: in the case of studies involving neurological or psychiatric patients, children, or adolescents, we considered only contrasts involving healthy controls, if reported;
Studies investigating the neural underpinnings of shame and guilt were included into two different sets, for two distinct meta-analyses. Specifically, we included studies contrasting shame/embarrassment vs. neutral or other emotional conditions, and guilt vs. neutral or other emotional conditions. Studies failing to distinguish embarrassment/shame and guilt were excluded.
Since the difference between shame and embarrassment is not clear-cut, as they can be classified according to different criteria, since they elicit the same physiological reactions and the same action tendencies, and since their distinction is still a matter of debate, we decided to include both shame and embarrassment in the same set.
This method allowed us to identify 18 studies for the shame/embarrassment set (188 foci, 439 total subjects) and 22 studies (one study was excluded in both sets because the contrast of interest did not show any significant result) for the guilt set (134 foci, 571 total subjects) (see Table 2). The most used paradigm in the studies analysed was emotion induction through verbal scripts (shame/embarrassment = 6, guilt = 8), pictures (shame/embarrassment = 5), both scripts and pictures (guilt = 3), vignettes (shame/embarrassment = 5), or movies (guilt = 1), while a few studies used the recollection of autobiographical memories through verbal scripts (shame/embarrassment = 1; guilt = 3), interpersonal games (shame/embarrassment = 1, guilt = 5), obedience paradigm (guilt = 1), or implicit association task (guilt = 1).

Statistical Analysis

Analyses were conducted using the software GingerALE v3.0.2 (URL = accessed on 13 February 2023). The activation likelihood estimation method, implemented in the software [75,76,77], uses probability theory to define the spatial convergence of foci reported in the selected studies. Specifically, a Gaussian blur with an empirically-derived full width half maximum (dependent on the number of participants included in the study) is applied to each focus from a single study. Then, all the foci from a single study are represented in a modelled activation map and voxel-wise ALE scores are computed combining all the individual maps. To distinguish between true convergence of foci from random noise, a permutation test is applied. We adopted the method described by Turkeltaub et al. [77] that minimises within-study effects, preventing the summation of foci from the same experiment that are placed close to each other. For studies reporting between-subjects contrasts, we used the number of participants included in the smallest group as the total number of study participants.
The analyses were performed on the studies’ coordinates in MNI space. So, in the case of studies reporting coordinates in Talairach space, we converted them to MNI space using the coordinate converter of the GingerALE software, while we kept the same coordinates in studies reporting results in MNI space. For each set of studies, we performed the meta-analysis applying a cluster-level family-wise error correction using an uncorrected p-value < 0.001 for individual voxels, 1000 permutations, and a cluster-level threshold of p < 0.05, as suggested by Eickhoff and collaborators [78]. Finally, we performed further analyses to show possible overlaps or differences among the two emotions. We ran (1) a conjunction analysis aiming to elucidate common neural activations of shame/embarrassment and guilt; and (2) contrast analyses in order to highlight specific neural activations of either shame/embarrassment or guilt. Contrast analyses were performed subtracting one of the outputs of the previous analyses (ALE images) to the other (i.e., Shame/Embarrassment vs. Guilt, Guilt vs. Shame/Embarrassment). Then, simulations on data created by pooling the original data of both study groups into two new groups (same sample size as the original groups) were performed. Subsequently, a new subtraction map was computed with the two new datasets (subtracting one to the other) and it was compared to the true data. After 1000 permutations, a voxel-wise p-value image revealed, for each voxel, where the real data is located in the distribution of all the possible values (for that specific voxel). Values were converted into z-scores. We adopted an FDR correction (FDR pN in GingerALE) with p < 0.05 [76,79,80]. The specific contribution of each study to cluster formation can be found in the Supplementary Materials. Results are visualised using SurfICE (URL = accessed on 1 March 2023).

3. Results

3.1. Shame/Embarrassment

The meta-analysis on shame/embarrassment revealed five significant clusters (see Figure 1 and Table 3). One cluster included the left anterior insula and the pars orbitalis of the left inferior frontal gyrus (cluster 1), while three clusters were located within the frontal lobes and included the left anterior cingulate cortex (cluster 2), the right inferior frontal gyrus (IFG, cluster 3), and the right precentral gyrus (cluster 4). It is worth noting that clusters 3 and 4 received an important contribution from studies contrasting self- vs. other-face stimuli (see Supplementary Materials). The other clusters were located within the medial portion of the left thalamus (cluster 5).

3.2. Guilt

The meta-analysis on guilt revealed two significant clusters (see Figure 2 and Table 4). One cluster was located at the level of the insula, involving also the orbital part of IFG (cluster 1). The other cluster was located on the posterior part of the left middle temporal gyrus, extending also to the angular gyrus at the junction between the temporal and parietal lobe (cluster 2).

3.3. Contrast Analyses

Conjunction analyses (see Figure 3 and Table 5) showed that both shame/embarrassment and guilt shared the activation of one cluster located within the left dorsal anterior insula and the pars orbitalis of the left inferior frontal gyrus. Contrast analyses (see Figure 3 and Table 5) revealed two significant clusters for the contrast ‘shame/embarrassment vs. guilt’, both at the level of the anterior cingulate cortex. The contrast ‘guilt vs. shame/embarrassment’ yielded no significant result.

4. Discussion

In the current meta-analysis, we analysed the functional neuroimaging literature on shame/embarrassment and guilt with the aim to identify the brain areas consistently associated with the processing of the two emotions. The results show that both emotions are associated with the activation of the left anterior insula, but they also show specific sets of areas involved in the processing of shame/embarrassment.

4.1. The Shame/Embarrassment Network

The occurrence of self-emotional distress in association with shame/embarrassment [4,18,81] might explain the association of the processing of these emotions with the dorsal ACC (cluster 2), the left anterior insula (cluster 1), and the medial nuclei of the thalami (cluster 5). Neuropsychological studies have highlighted that patients with dorsal ACC lesions, typically made in order to treat drug-resistant pain [82], are still able to perceive and correctly localise painful sensations, but such sensations are not distressing anymore [83]. Moreover, it is worth noting that the surgical lesion of the dorsal ACC also leads to a reduced concern about the opinions or the social judgement of other people [84], and can be used in the treatment of drug-resistant obsessive-compulsive disorder, a psychiatric syndrome which is often associated with extremely intense shame experiences [85]. Medial thalamic nuclei are thought to be involved in affective aspects of physical pain perception and attachment-related processes [86]. This set of areas is highly overlapping with those involved in the processing of both physical and social pain [87]. Social pain is the unpleasant experience associated with damage to social bonds or to social values (e.g., rejection, negative social evaluations, bereavement), and is thought to be processed by part of the neural circuit involved in processing physical pain [88]. Shame and embarrassment are thought to be important aspects of social pain, since they might signal that the social standards of others are not met [88].
The meta-analysis on shame/embarrassment also revealed clusters within the right premotor area (cluster 4) and right IFG (cluster 3), which have been associated with motor and speech inhibition [89,90], and which is consistent with the action tendencies associated with shame/embarrassment. Indeed, in contrast with guilt, which is often associated with pro-social behaviour aiming to repair the transgression that has occurred [18], shame and embarrassment lead to a reduction of social presence, speech, and movements [25,33], which could explain the activation of areas involved in motor and speech inhibition in shame/embarrassment processing. Hence, the presentation of shameful or embarrassing stimuli might automatically activate behavioral motor scripts aiming to reduce social presence. However, it is worth noting that studies contrasting self- vs. other-faces were important contributors to the generation of these clusters, as previous meta-analyses on neuroimaging studies on self-face recognition have reported [91,92]. Their relevance in the self-face processing network, especially for the right IFG, might be associated with the ability to differentiate self from other information [92].

4.2. The Guilt Network

Differently from shame and embarrassment, guilt is thought to be associated with social abilities, such as empathy and theory of mind, which were proposed to be specifically related to guilt generation [19,93]. In our meta-analysis, we found an association between guilt processing and TPJ, which was reliably found as a crucial area for distinguishing self- and other-actions, and representing other individuals’ mental and affective states, see [94] for a meta-analysis. Although the association between guilt and TPJ did not reach the significance level in the contrast analyses (i.e., guilt vs. shame/embarrassment), considering the convergence with a previous meta-analysis of guilt processing [40] and the associations between guilt processing and theory of mind, and theory of mind and TPJ, we believe that TPJ should be taken into consideration as a crucial area in the processing of guilt. However, the association between guilt, empathy, and theory of mind is not univocal. On the one hand, guilt is thought to increase the understanding of others’ affective and mental states [18]; on the other, taking others’ perspective and empathising with others seem to be crucial in order to experience guilt [95]. Hence, our results might refer to functions that are cause or consequence of the emotional experience.

4.3. Common Areas

The anterior insula was found in association with a wide variety of tasks, see [96]. Among the cognitive functions associated with the anterior insula that also include interoception, pain perception, and body awareness, it is worth mentioning its role in emotional awareness [96], arousal, and self-reflection, e.g., [97,98]. In addition, the lesion of this area is associated with pain asymbolia [99], a condition in which patients are still able to localise a painful stimulation and to identify it as pain but they lose all the unpleasant aspects (e.g., bodily, emotional, and behavioural signs) of pain [100]. The same type of patients showed reduced arousal ratings and an attenuated valence rating to emotional stimuli compared to both pathological and healthy controls [101]. The interpretation of these findings is not univocal. If, on the one hand, they might reflect the impairment in arousal processing, on the other they might also be caused by a deficit in emotional awareness. In addition, functional neuroimaging studies on healthy individuals investigating self-referential processing found anterior insula activation, e.g., [97,98]. Given that in our meta-analysis a high number of the studies included contrasted shame/embarrassment or guilt vs. neutral baselines, their common association with the anterior insula might simply reflect their arousal. Hence, the association between negative self-conscious emotion processing and the activation of the left anterior insula in functional neuroimaging studies might reflect their intensity, the awareness of the subjective experience of shame/embarrassment and guilt, or self-directed evaluation processes that are necessary in order to generate both guilt and shame experiences.
Our conjunction analysis did not show the involvement of the mPFC in representing both shame/embarrassment and guilt. Although our meta-analysis on shame/embarrassment revealed the activation of the ACC and the left mPFC (cluster 2), the same analysis on guilt did not show this cluster of activation. It is worth noting that these clusters of activation overlap with the results of a previous meta-analysis on guilt processing [40]. The mPFC represents a high-level integration area and is thought to support different aspects of social and affective processing [102,103] ranging from self-reflection [104], person perception [105], affective appraisal [106,107], theory of mind [108], learning, and predicting actions outcomes [109]. Moreover, the same area was found to be active in functional neuroimaging studies investigating moral judgement when moral evaluations were contrasted with non-moral or neutral baselines [110]. If, on the one hand, it has been proposed that the mPFC associates external stimuli (e.g., context-based information) with their socio-emotional value through a connection with anterior temporal lobes [8], on the other, it might be involved in self-referential processing (e.g., representation of traits, abilities, attitudes, and behaviours regarding the self), which is necessary in order to generate self-conscious emotions. This latter hypothesis seems to be confirmed by neuropsychological studies showing that patients with damage in the mPFC were impaired in self-referential memory [111]), self-evaluation [112], and self-referential verbal production [113].

5. Conclusions

Our meta-analysis revealed common and distinct neural substrates for the processing of shame/embarrassment and guilt. While the activation of the left anterior insula was associated with both shame/embarrassment and guilt processing, the pain network, including the medial thalami, the dorsal ACC, and premotor areas, was specifically associated with shame/embarrassment processing, while the left TPJ was associated with specific guilt processing.

6. Limitations

The main limitation of our work is the small number of studies investigating shame and embarrassment separately, which did not allow us to perform distinct meta-analyses on embarrassment and shame, as well as the relatively small number of participants included in most of the studies. The wide variety of paradigms investigating self-conscious emotions, including reading scripts, viewing vignettes, and recalling autobiographical memories, might affect the reliability of the results. Further studies investigating self-conscious emotions are necessary to better characterise common and specific brain networks involved in their processing.

Supplementary Materials

The following supporting information can be downloaded at:, Table S1: Specific contribution of each study to cluster formation.

Author Contributions

Conceptualization, L.P., E.P. and A.G.; methodology, L.P. and A.G.; software, L.P. and E.P.; validation, all the authors; formal analysis, L.P., E.P., C.G. and A.G.; investigation, L.P., E.P. and A.G.; resources, L.P. and A.G.; data curation, L.P., E.P. and A.G.; writing—original draft preparation, L.P., E.P. and A.G.; writing—review and editing, C.G., R.I.R. and R.J.; visualization, L.P. and E.P.; supervision, A.G.; project administration, A.G.; funding acquisition, L.P. All authors have read and agreed to the published version of the manuscript.


This research was funded by the CARITRO Foundation (Rovereto, Trento, Italy).

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

Data are available upon request to the corresponding author.

Conflicts of Interest

The authors declare no conflict of interest.


  1. Haidt, J. The Moral Emotions. In Handbook of Affective Sciences; Oxford University Press: Oxford, UK, 2003; Volume 11, pp. 852–870. [Google Scholar]
  2. Kroll, J.; Egan, E. Psychiatry, moral worry, and the moral emotions. J. Psychiatr. Pract. 2004, 10, 352–360. [Google Scholar] [CrossRef] [PubMed]
  3. Piretti, L.; Pappaianni, E.; Lunardelli, A.; Zorzenon, I.; Ukmar, M.; Pesavento, V.; Rumiati, R.I.; Job, R.; Grecucci, A. The Role of Amygdala in Self-Conscious Emotions in a Patient With Acquired Bilateral Damage. Front. Neurosci. 2020, 14, 677. [Google Scholar] [CrossRef]
  4. Grecucci, A.; Neresini, A.; Job, R. The Moral Algorithm. Toward a Neuropsychological Model of Shame. In The Dark Side: Philosophical Reflections on the “Negative Emotions”; Giacomoni, P., Valentini, N., Dellantonio, S., Eds.; Springer: Cham, Switzerland, 2021. [Google Scholar]
  5. Ridley, M.; Waal FB, D. The origins of virtue. Nature 1996, 383, 785. [Google Scholar]
  6. Sober, E.; Wilson, D.S. Unto Others: Evolution and Psychology of Unselfish Behavior; Harvard University Press: Cambridge, MA, USA, 1998. [Google Scholar]
  7. Moll, J.; Zahn, R.; De Oliveira-Souza, R.; Krueger, F.; Grafman, J.H. The neural basis of human moral cognition. Nat. Rev. Neurosci. 2005, 6, 799–809. [Google Scholar] [CrossRef] [PubMed]
  8. Moll, J.; De Oliveira-Souza, R.; Zahn, R. The Neural Basis of Moral Cognition. Ann. N. Y. Acad. Sci. 2008, 1124, 161–180. [Google Scholar] [CrossRef] [PubMed]
  9. Grafman, J. Similarities and Distinctions among Current Models of Prefrontal Cortical Functions. In Structure and Functions of the Human Prefrontal Cortex; Grafman, J., Holyoak, K.J., Boller, F., Eds.; Academy of Sciences: New York, NY, USA, 1995; pp. 337–368. [Google Scholar]
  10. Wood, J.; Grafman, J. Human Prefrontal Cortex: Processing and Representational Perspectives. Nat. Rev. Neurosci. 2003, 4, 139–147. [Google Scholar] [CrossRef]
  11. Ralph, M.; Jefferies, E.; Patterson, K.; Rogers, T.T. The neural and computational bases of semantic cognition. Nat. Rev. Neurosci. 2017, 18, 42–55. [Google Scholar] [CrossRef]
  12. Olson, I.R.; McCoy, D.; Klobusicky, E.; Ross, L.A. Social cognition and the anterior temporal lobes: A review and theoretical framework. Soc. Cogn. Affect. Neurosci. 2013, 8, 123–133. [Google Scholar] [CrossRef] [Green Version]
  13. Haxby, J.V.; Hoffman, E.A.; Gobbini, M.I. The distributed human neural system for face perception. Trends Cogn. Sci. 2000, 4, 223–233. [Google Scholar] [CrossRef]
  14. Saper, C.B. Hypothalamic Connections with the Cerebral Cortex. In Progress in Brain Research; Elsevier: Amsterdam, The Netherlands, 2000; Volume 126, pp. 39–48. [Google Scholar] [CrossRef]
  15. Eres, R.; Louis, W.R.; Molenberghs, P. Common and distinct neural networks involved in fMRI studies investigating morality: An ALE meta-analysis. Soc. Neurosci. 2018, 13, 384–398. [Google Scholar] [CrossRef]
  16. Bzdok, D.; Schilbach, L.; Vogeley, K.; Schneider, K.; Laird, A.R.; Langner, R.; Eickhoff, S.B. Parsing the neural correlates of moral cognition: ALE meta-analysis on morality, theory of mind, and empathy. Brain Struct. Funct. 2012, 217, 783–796. [Google Scholar]
  17. Baumeister, R.F.; Leary, M.R. The need to belong: Desire for interpersonal attachments as a fundamental human motivation. Psychol. Bull. 1995, 117, 497. [Google Scholar] [CrossRef]
  18. Tangney, J.P.; Stuewig, J.; Mashek, D.J. Moral emotions and moral behavior. Annu. Rev. Psychol. 2007, 58, 345–372. [Google Scholar]
  19. Bastin, C.; Harrison, B.J.; Davey, C.G.; Moll, J.; Whittle, S. Feelings of shame, embarrassment and guilt and their neural correlates: A systematic review. Neurosci. Biobehav. Rev. 2016, 71, 455–471. [Google Scholar] [CrossRef] [PubMed]
  20. Gibson, M. Shame and guilt in child protection social work: New interpretations and opportunities for practice. Child Fam. Soc. Work 2015, 20, 333–343. [Google Scholar] [CrossRef] [Green Version]
  21. Wong, Y.; Tsai, J. Cultural Models of Shame and Guilt. In The Self-Conscious Emotions: Theory and Research; Guilford Press: New York, NY, USA, 2007; pp. 209–223. [Google Scholar]
  22. Tangney, J.P.; Wagner, P.; Gramzow, R. Proneness to shame, proneness to guilt, and psychopathology. J. Abnorm. Psychol. 1992, 101, 469–478. [Google Scholar] [CrossRef] [PubMed]
  23. Crozier, W.R. Differentiating shame from embarrassment. Emot. Rev. 2014, 6, 269–276. [Google Scholar] [CrossRef]
  24. Probyn, E. Blush: Faces of Shame; University of Minnesota Press: Minneapolis, MN, USA, 2005. [Google Scholar]
  25. Keltner, D.; Buswell, B.N. Embarrassment: Its distinct form and appeasement functions. Psychol. Bull. 1997, 122, 250. [Google Scholar] [CrossRef]
  26. Tangney, J.P.; Miller, R.S.; Flicker, L.; Barlow, D.H. Are shame, guilt, and embarrassment distinct emotions? J. Pers. Soc. Psychol. 1996, 70, 1256–1269. [Google Scholar] [CrossRef]
  27. Keltner, D. Evidence for the Distinctness of Embarrassment, Shame, and Guilt: A Study of Recalled Antecedents and Facial Expressions of Emotion. Cogn. Emot. 1996, 10, 155–172. [Google Scholar] [CrossRef]
  28. Buss, A. Psychological Dimensions of the Self; Sage Publications, Inc: Thousand Oaks, CA, USA, 2001. [Google Scholar]
  29. Tangney, J.P. Self-Relevant Emotions. In Handbook of Self and Identity; Leary, M.R., Tangney, J.P., Eds.; Guilford Press: New York, NY, USA, 2003; pp. 384–400. [Google Scholar]
  30. Rochat, P. Others in Mind: Social Origins of Self-Consciousness; Cambridge University Press: Cambridge, UK, 2009. [Google Scholar]
  31. Scheff, T.J. Microsociology: Discourse, Emotion, and Social Structure; University of Chicago Press: Chicago, IL, USA, 1994. [Google Scholar]
  32. Sabini, J.; Silver, M. TARGET ARTICLE: Why Emotion Names and Experiences Don’t Neatly Pair. Psychol. Inq. 2005, 16, 1–10. [Google Scholar] [CrossRef]
  33. Asendorpf, J.B. Development of inhibition during childhood: Evidence for situational specificity and a two-factor model. Dev. Psychol. 1990, 26, 721. [Google Scholar] [CrossRef]
  34. Lewis, M.; Haviland-Jones, J.M.; Feldman Barrett, L. Handbook of Emotions; Guilford Press: New York, NY, USA, 1993. [Google Scholar]
  35. Feinberg, M.; Willer, R.; Keltner, D. Flustered and faithful: Embarrassment as a signal of prosociality. J. Personal. Soc. Psychol. 2012, 102, 81. [Google Scholar] [CrossRef] [Green Version]
  36. Leary, M.R.; Landel, J.L.; Patton, K.M. The Motivated Expression of Embarrassment Following a Self-Presentational Predicament. J. Personal. 1996, 64, 619–636. [Google Scholar] [CrossRef]
  37. Hoffman, M.L. Development of Prosocial Motivation: Empathy and Guilt. In The Development of Prosocial Behavior; Academic Press: New York, NY, USA, 1982; pp. 281–313. [Google Scholar]
  38. Fiske, A.P. Structures of Social Life: The Four Elementary Forms of Human Relations: Communal Sharing, Authority Ranking, Equality Matching, Market Pricing; Free Press: New York, NY, USA, 1991. [Google Scholar]
  39. Lewis, H.B. Shame and guilt in neurosis. Psychoanal. Rev. 1971, 58, 419–438. [Google Scholar] [PubMed]
  40. Gifuni, A.J.; Kendal, A.; Jollant, F. Neural mapping of guilt: A quantitative meta-analysis of functional imaging studies. Brain Imaging Behav. 2017, 11, 1164–1178. [Google Scholar] [CrossRef]
  41. Bas-Hoogendam, J.M.; Van Steenbergen, H.; Kreuk, T.; Van Der Wee, N.J.A.; Westenberg, P.M. How embarrassing! The behavioral and neural correlates of processing social norm violations. PLoS ONE 2017, 12, e0176326. [Google Scholar] [CrossRef]
  42. Bastin, C.; Rakesh, D.; Harrison, B.J.; Davey, C.G.; Allen, N.B.; Muller, S.; Whittle, S. Feelings of shame and guilt are associated with distinct neural activation in youth. Biol. Psychol. 2021, 159, 108025. [Google Scholar] [CrossRef] [PubMed]
  43. Berthoz, S.; Armony, J.L.; Blair, R.J.R.; Dolan, R.J. An fMRI study of intentional and unintentional (embarrassing) violations of social norms. Brain 2002, 125, 1696–1708. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  44. Finger, E.C.; Marsh, A.A.; Kamel, N.; Mitchell, D.G.; Blair, J.R. Caught in the act: The impact of audience on the neural response to morally and socially inappropriate behavior. Neuroimage 2006, 33, 414–421. [Google Scholar] [CrossRef]
  45. Krach, S.; Cohrs, J.C.; Loebell, N.C.D.E.; Kircher, T.; Sommer, J.; Jansen, A.; Paulus, F.M. Your Flaws Are My Pain: Linking Empathy To Vicarious Embarrassment. PLoS ONE 2011, 6, e18675. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  46. Krach, S.; Kamp-Becker, I.; Einhäuser, W.; Sommer, J.; Frässle, S.; Jansen, A.; Rademacher, L.; Müller-Pinzler, L.; Gazzola, V.; Paulus, F.M. Evidence from pupillometry and fMRI indicates reduced neural response during vicarious social pain but not physical pain in autism. Hum. Brain Mapp. 2015, 36, 4730–4744. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  47. Laneri, D.; Krach, S.; Paulus, F.M.; Kanske, P.; Schuster, V.; Sommer, J.; Müller-Pinzler, L. Mindfulness meditation regulates anterior insula activity during empathy for social pain. Hum. Brain Mapp. 2017, 38, 4034–4046. [Google Scholar] [CrossRef] [Green Version]
  48. Melchers, M.; Markett, S.; Montag, C.; Trautner, P.; Weber, B.; Lachmann, B.; Buss, P.; Heinen, R.; Reuter, M. Reality TV and vicarious embarrassment: An fMRI study. Neuroimage 2015, 109, 109–117. [Google Scholar] [CrossRef]
  49. Mayer, A.V.; Müller-Pinzler, L.; Krach, S.; Paulus, F.M. Spinach in the teeth: How ego- and allocentric perspectives modulate neural correlates of embarrassment in the face of others’ public mishaps. Cortex 2020, 130, 275–289. [Google Scholar] [CrossRef] [PubMed]
  50. Michl, P.; Meindl, T.; Meister, F.; Born, C.; Engel, R.; Reiser, M.; Hennig-Fast, K. Neurobiological underpinnings of shame and guilt: A pilot fMRI study. Soc. Cogn. Affect. Neurosci. 2012, 9, 150–157. [Google Scholar] [CrossRef]
  51. Morita, T.; Itakura, S.; Saito, D.N.; Nakashita, S.; Harada, T.; Kochiyama, T.; Sadato, N. The Role of the Right Prefrontal Cortex in Self-evaluation of the Face: A Functional Magnetic Resonance Imaging Study. J. Cogn. Neurosci. 2008, 20, 342–355. [Google Scholar] [CrossRef] [PubMed]
  52. Morita, T.; Kosaka, H.; Saito, D.N.; Ishitobi, M.; Munesue, T.; Itakura, S.; Omori, M.; Okazawa, H.; Wada, Y.; Sadato, N. Emotional responses associated with self-face processing in individuals with autism spectrum disorders: An fMRI study. Soc. Neurosci. 2012, 7, 223–239. [Google Scholar] [CrossRef]
  53. Morita, T.; Tanabe, H.C.; Sasaki, A.T.; Shimada, K.; Kakigi, R.; Sadato, N. The anterior insular and anterior cingulate cortices in emotional processing for self-face recognition. Soc. Cogn. Affect. Neurosci. 2013, 9, 570–579. [Google Scholar] [CrossRef] [Green Version]
  54. Morita, T.; Kosaka, H.; Saito, D.N.; Fujii, T.; Ishitobi, M.; Munesue, T.; Inohara, K.; Okazawa, H.; Kakigi, R.; Sadato, N. Neural correlates of emotion processing during observed self-face recognition in individuals with autism spectrum disorders. Res. Autism Spectr. Disord. 2016, 26, 16–32. [Google Scholar] [CrossRef]
  55. Stroth, S.; Paye, L.; Kamp-Becker, I.; Wermter, A.-K.; Krach, S.; Paulus, F.M.; Müller-Pinzler, L. Empathy in Females With Autism Spectrum Disorder. Front. Psychiatry 2019, 10. Available online: (accessed on 1 March 2023).
  56. Takahashi, H.; Yahata, N.; Koeda, M.; Matsuda, T.; Asai, K.; Okubo, Y. Brain activation associated with evaluative processes of guilt and embarrassment: An fMRI study. Neuroimage 2004, 23, 967–974. [Google Scholar] [CrossRef]
  57. Wagner, U.; N’Diaye, K.; Ethofer, T.; Vuilleumier, P. Guilt-Specific Processing in the Prefrontal Cortex. Cereb. Cortex 2011, 21, 2461–2470. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  58. Zhu, R.; Feng, C.; Zhang, S.; Mai, X.; Liu, C. Differentiating guilt and shame in an interpersonal context with univariate activation and multivariate pattern analyses. Neuroimage 2018, 186, 476–486. [Google Scholar] [CrossRef]
  59. Basile, B.; Mancini, F.; Macaluso, E.; Caltagirone, C.; Frackowiak, R.S.; Bozzali, M. Deontological and altruistic guilt: Evidence for distinct neurobiological substrates. Hum. Brain Mapp. 2010, 32, 229–239. [Google Scholar] [CrossRef] [PubMed]
  60. Cheng, Y.; Chou, J.; Martínez, R.M.; Fan, Y.T.; Chen, C. Psychopathic traits mediate guilt-related anterior midcingulate activity under authority pressure. Sci. Rep. 2021, 11, 14856. [Google Scholar] [CrossRef] [PubMed]
  61. Domínguez D, J.F.; van Nunspeet, F.; Gupta, A.; Eres, R.; Louis, W.R.; Decety, J.; Molenberghs, P. Lateral orbitofrontal cortex activity is modulated by group membership in situations of justified and unjustified violence. Soc. Neurosci. 2018, 13, 739–755. [Google Scholar] [CrossRef]
  62. Fourie, M.M.; Thomas, K.G.F.; Amodio, D.M.; Warton, C.M.R.; Meintjes, E. Neural correlates of experienced moral emotion: An fMRI investigation of emotion in response to prejudice feedback. Soc. Neurosci. 2014, 9, 203–218. [Google Scholar] [CrossRef] [PubMed]
  63. Gilead, M.; Katzir, M.; Eyal, T.; Liberman, N. Neural correlates of processing “self-conscious” vs. “basic” emotions. Neuropsychologia 2016, 81, 207–218. [Google Scholar] [CrossRef]
  64. Gradin, V.B.; Pérez, A.; Macfarlane, J.A.; Cavin, I.; Waiter, G.; Tone, E.B.; Dritschel, B.; Maiche, A.; Steele, J.D. Neural correlates of social exchanges during the Prisoner’s Dilemma game in depression. Psychol. Med. 2016, 46, 1289–1300. [Google Scholar] [CrossRef] [Green Version]
  65. Green, S.; Ralph, M.A.L.; Moll, J.; Deakin, J.F.W.; Zahn, R. Guilt-Selective Functional Disconnection of Anterior Temporal and Subgenual Cortices in Major Depressive Disorder. Arch. Gen. Psychiatry 2012, 69, 1014–1021. [Google Scholar] [CrossRef] [Green Version]
  66. Kédia, G.; Berthoz, S.; Wessa, M.; Hilton, D.; Martinot, J.-L. An Agent Harms a Victim: A Functional Magnetic Resonance Imaging Study on Specific Moral Emotions. J. Cogn. Neurosci. 2008, 20, 1788–1798. [Google Scholar] [CrossRef]
  67. Li, Z.; Yu, H.; Zhou, Y.; Kalenscher, T.; Zhou, X. Guilty by association: How group-based (collective) guilt arises in the brain. NeuroImage 2020, 209, 116488. [Google Scholar] [CrossRef]
  68. Molenberghs, P.; Ogilvie, C.; Louis, W.R.; Decety, J.; Bagnall, J.; Bain, P.G. The neural correlates of justified and unjustified killing: An fMRI study. Soc. Cogn. Affect. Neurosci. 2015, 10, 1397–1404. [Google Scholar] [CrossRef]
  69. Morey, R.A.; McCarthy, G.; Selgrade, E.S.; Seth, S.; Nasser, J.D.; LaBar, K.S. Neural systems for guilt from actions affecting self versus others. Neuroimage 2012, 60, 683–692. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  70. Nihonsugi, T.; Numano, S.; Haruno, M. Functional connectivity basis and underlying cognitive mechanisms for gender differences in guilt aversion. Eneuro 2021, 8. [Google Scholar] [CrossRef] [PubMed]
  71. Peth, J.; Sommer, T.; Hebart, M.N.; Vossel, G.; Büchel, C.; Gamer, M. Memory detection using fMRI—Does the encoding context matter? Neuroimage 2015, 113, 164–174. [Google Scholar] [CrossRef] [PubMed]
  72. Shin, L.M.; Dougherty, D.D.; Orr, S.P.; Pitman, R.K.; Lasko, M.; Macklin, M.L.; Alpert, N.M.; Fischman, A.J.; Rauch, S.L. Activation of anterior paralimbic structures during guilt-related script-driven imagery. Biol. Psychiatry 2000, 48, 43–50. [Google Scholar] [CrossRef]
  73. Ty, A.; Mitchell, D.G.; Finger, E. Making amends: Neural systems supporting donation decisions prompting guilt and restitution. Pers. Individ. Differ. 2017, 107, 28–36. [Google Scholar] [CrossRef]
  74. Yu, H.; Hu, J.; Hu, L.; Zhou, X. The voice of conscience: Neural bases of interpersonal guilt and compensation. Soc. Cogn. Affect. Neurosci. 2013, 9, 1150–1158. [Google Scholar] [CrossRef] [Green Version]
  75. Eickhoff, S.B.; Laird, A.R.; Grefkes, C.; Wang, L.E.; Zilles, K.; Fox, P.T. Coordinate-based activation likelihood estimation meta-analysis of neuroimaging data: A random-effects approach based on empirical estimates of spatial uncer-tainty. Hum. Brain Mapp. 2009, 30, 2907–2926. [Google Scholar] [CrossRef] [Green Version]
  76. Eickhoff, S.B.; Bzdok, D.; Laird, A.R.; Kurth, F.; Fox, P.T. Activation likelihood estimation meta-analysis revisited. Neuroimage 2012, 59, 2349–2361. [Google Scholar] [CrossRef] [Green Version]
  77. Turkeltaub, P.E.; Eickhoff, S.B.; Laird, A.; Fox, M.; Wiener, M.; Fox, P. Minimizing within-experiment and within-group effects in activation likelihood estimation meta-analyses. Hum. Brain Mapp. 2012, 33, 1–13. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  78. Eickhoff, S.B.; Nichols, T.E.; Laird, A.R.; Hoffstaedter, F.; Amunts, K.; Fox, P.T.; Bzdok, D.; Eickhoff, C.R. Behavior, sensitivity, and power of activation likelihood estimation characterized by massive empirical simulation. Neuroimage 2016, 137, 70–85. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  79. Laird, A.R.; Fox, P.M.; Price, C.J.; Glahn, D.C.; Uecker, A.M.; Lancaster, J.L.; Turkeltaub, P.E.; Kochunov, P. ALE meta-analysis: Controlling the false discovery rate and performing statistical contrasts. Hum. Brain Mapp. 2005, 25, 155–164. [Google Scholar] [CrossRef] [PubMed]
  80. Zmigrod, L.; Garrison, J.R.; Carr, J.; Simons, J.S. The neural mechanisms of hallucinations: A quantitative meta-analysis of neuroimaging studies. Neurosci. Biobehav. Rev. 2016, 69, 113–123. [Google Scholar] [CrossRef] [Green Version]
  81. Piretti, L.; Pappaianni, E.; Rumiati, R.; Job, R.; Grecucci, A. Dissociating the role of dlPFC and dACC/dmPFC in emo-tional processing using tDCS. Cogn. Affect. Behav. Neurosci. 2021, 22, 304–315. [Google Scholar] [CrossRef] [PubMed]
  82. Yen, C.P.; Kung, S.S.; Su, Y.F.; Lin, W.C.; Howng, S.L.; Kwan, A.L. Stereotactic bilateral anterior cingulotomy for intractable pain. J. Clin. Neurosci. 2005, 12, 886–890. [Google Scholar] [CrossRef]
  83. Foltz, E.L.; White, L.E. Pain “Relief” by Frontal Cingulumotomy. J. Neurosurg. 1962, 19, 89–100. [Google Scholar] [CrossRef]
  84. Tow, P.M.; Whitty, C.W.M. Personality changes after operations on the cingulate gyrus in man. J. Neurol. Neurosurg. Psychiatry 1953, 16, 186–193. [Google Scholar] [CrossRef] [Green Version]
  85. Weingarden, H.; Renshaw, K.D. Shame in the obsessive compulsive related disorders: A conceptual review. J. Affect. Disord. 2014, 171, 74–84. [Google Scholar] [CrossRef] [Green Version]
  86. Price, D.D. Psychological and Neural Mechanisms of the Affective Dimension of Pain. Science 2000, 288, 1769–1772. [Google Scholar] [CrossRef] [Green Version]
  87. Eisenberger, N.I. The pain of social disconnection: Examining the shared neural underpinnings of physical and social pain. Nat. Rev. Neurosci. 2012, 13, 421–434. [Google Scholar] [CrossRef] [PubMed]
  88. MacDonald, G.; Leary, M.R. Why Does Social Exclusion Hurt? The Relationship Between Social and Physical Pain. Psychol. Bull. 2005, 131, 202–223. [Google Scholar] [CrossRef] [Green Version]
  89. Simmonds, D.J.; Pekar, J.J.; Mostofsky, S.H. Meta-analysis of Go/No-go tasks demonstrating that fMRI activation associated with response inhibition is task-dependent. Neuropsychologia 2008, 46, 224–232. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  90. Xue, G.; Aron, A.R.; Poldrack, R.A. Common Neural Substrates for Inhibition of Spoken and Manual Responses. Cereb. Cortex 2008, 18, 1923–1932. [Google Scholar] [CrossRef]
  91. Hu, C.; Di, X.; Eickhoff, S.B.; Zhang, M.; Peng, K.; Guo, H.; Sui, J. Distinct and common aspects of physical and psychological self-representation in the brain: A meta-analysis of self-bias in facial and self-referential judgements. Neurosci. Biobehav. Rev. 2016, 61, 197–207. [Google Scholar] [CrossRef] [PubMed]
  92. Platek, S.M.; Wathne, K.; Tierney, N.G.; Thomson, J.W. Neural correlates of self-face recognition: An effect-location meta-analysis. Brain Res. 2008, 1232, 173–184. [Google Scholar] [CrossRef] [PubMed]
  93. Leith, K.P.; Baumeister, R. Empathy, Shame, Guilt, and Narratives of Interpersonal Conflicts: Guilt-Prone People Are Better at Perspective Taking. J. Personal. 1998, 66, 1–37. [Google Scholar] [CrossRef]
  94. Decety, J.; Lamm, C. The Role of the Right Temporoparietal Junction in Social Interaction: How Low-Level Computational Processes Contribute to Meta-Cognition. Neurosci 2007, 13, 580–593. [Google Scholar] [CrossRef]
  95. Giammarco, E.A.; Vernon, P.A. Interpersonal Guilt and the Dark Triad. Pers. Individ. Differ. 2015, 81, 96–101. [Google Scholar] [CrossRef]
  96. Craig, A.D. How do you feel—Now? The anterior insula and human awareness. Nat. Rev. Neurosci. 2009, 10, 59–70. [Google Scholar] [CrossRef]
  97. Johnson, S.C.; Schmitz, T.W.; Kawahara-Baccus, T.N.; Rowley, H.A.; Alexander, A.L.; Lee, J.; Davidson, R.J. The Cerebral Response during Subjective Choice with and without Self-reference. J. Cogn. Neurosci. 2005, 17, 1897–1906. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  98. Modinos, G.; Ormel, J.; Aleman, A. Activation of Anterior Insula during Self-Reflection. PLoS ONE 2009, 4, e4618. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  99. Berthier, M.; Starkstein, S.; Leiguarda, R. Asymbolia for pain: A sensory-limbic disconnection syndrome. Ann. Neurol. 1988, 24, 41–49. [Google Scholar] [CrossRef]
  100. Aydede, M. (Ed.) Pain: New Essays on Its Nature and the Methodology of Its Study; MIT Press: Cambridge, MA, USA, 2005. [Google Scholar]
  101. Berntson, G.G.; Norman, G.J.; Bechara, A.; Bruss, J.; Tranel, D.; Cacioppo, J.T. The Insula and Evaluative Processes. Psychol. Sci. 2010, 22, 80–86. [Google Scholar] [CrossRef] [Green Version]
  102. Amodio, D.M.; Frith, C.D. Meeting of minds: The medial frontal cortex and social cognition. Nat. Rev. Neurosci. 2006, 7, 268–277. [Google Scholar] [CrossRef] [PubMed]
  103. Roy, M.; Shohamy, D.; Wager, T.D. Ventromedial prefrontal-subcortical systems and the generation of affective meaning. Trends Cogn. Sci. 2012, 16, 147–156. [Google Scholar] [CrossRef] [Green Version]
  104. Van der Meer, L.; Costafreda, S.; Aleman, A.; David, A.S. Self-reflection and the brain: A theoretical review and meta-analysis of neuroimaging studies with implications for schizophrenia. Neurosci. Biobehav. Rev. 2010, 34, 935–946. [Google Scholar] [CrossRef]
  105. Mitchell, J.P.; Heatherton, T.F.; Macrae, C.N. Distinct neural systems subserve person and object knowledge. Proc. Natl. Acad. Sci. USA 2002, 99, 15238–15243. [Google Scholar] [CrossRef] [Green Version]
  106. Scherer, K.R.; Schorr, A.; Johnstone, T. (Eds.) Appraisal Processes in Emotion: Theory, Methods, Research; Oxford University Press: Oxford, UK, 2001. [Google Scholar]
  107. Grecucci, A.; Frederickson, J.; Job, R. Advances in Emotion Regulation: From neuroscience to psychotherapy. Front. Psychol. 2017, 8, 985. [Google Scholar] [CrossRef] [Green Version]
  108. Frith, C.D.; Frith, U. The Neural Basis of Mentalizing. Neuron 2006, 50, 531–534. [Google Scholar] [CrossRef] [Green Version]
  109. Alexander, W.H.; Brown, J.W. Medial prefrontal cortex as an action-outcome predictor. Nat. Neurosci. 2011, 14, 1338–1344. [Google Scholar] [CrossRef] [PubMed]
  110. Garrigan, B.; Adlam, A.L.; Langdon, P.E. The neural correlates of moral decision-making: A systematic review and meta-analysis of moral evaluations and response decision judgements. Brain Cogn. 2016, 108, 88–97. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  111. Philippi, C.L.; Duff, M.C.; Denburg, N.L.; Tranel, D.; Rudrauf, D. Medial PFC Damage Abolishes the Self-reference Effect. J. Cogn. Neurosci. 2012, 24, 475–481. [Google Scholar] [CrossRef] [PubMed] [Green Version]
  112. Schmitz, T.W.; Rowley, H.A.; Kawahara, T.N.; Johnson, S.C. Neural correlates of self-evaluative accuracy after traumatic brain injury. Neuropsychologia 2006, 44, 762–773. [Google Scholar] [CrossRef]
  113. Kurczek, J.; Wechsler, E.; Ahuja, S.; Jensen, U.; Cohen, N.J.; Tranel, D.; Duff, M. Differential contributions of hippocampus and medial prefrontal cortex to self-projection and self-referential processing. Neuropsychologia 2015, 73, 116–126. [Google Scholar] [CrossRef] [PubMed] [Green Version]
Figure 1. Results of the meta-analysis on shame/embarrassment neural correlates. Bars represent ALE scores.
Figure 1. Results of the meta-analysis on shame/embarrassment neural correlates. Bars represent ALE scores.
Brainsci 13 00559 g001
Figure 2. Results of the meta-analysis on guilt neural correlates. Bars represent ALE scores.
Figure 2. Results of the meta-analysis on guilt neural correlates. Bars represent ALE scores.
Brainsci 13 00559 g002
Figure 3. Contrast analysis results. In purple, conjunction analysis (shame/embarrassment and guilt); in yellow, specific activations of shame/embarrassment vs. guilt.
Figure 3. Contrast analysis results. In purple, conjunction analysis (shame/embarrassment and guilt); in yellow, specific activations of shame/embarrassment vs. guilt.
Brainsci 13 00559 g003
Table 1. Differences between shame and guilt.
Table 1. Differences between shame and guilt.
TargetWhat we are: related to the entire self.
”I’m bad”
What we do: related to specific behaviours.
“What I did has been bad”
LevelInterpersonal—it occurs only with othersIntrapsychic—it occurs alone
Emotional activationPainfulLess painful
Emotional perceptionDifficult to recognizeEasy to recognize
Action tendencyMotivates hiding and inhibitionMotivates reparation of the situation
Relation with aggression, hostility, violence, externalizationIncreased for shame-proneness individualsDecreased for guilt-proneness individuals
ScapegoatBlame mainly othersBlame myself
ResponsibilityDeflected outwardAccepted
Adapted from Grecuccu et al., 2021 [4] and Tangney et al., 2007 [18].
Table 2. Studies investigating shame/embarrassment and guilt brain processing.
Table 2. Studies investigating shame/embarrassment and guilt brain processing.
SubsetAuthorsParadigmStimulus TypeContrastsFociSubjects (Females)
Bas-Hogendam et al., 2017 [41]InductionVerbal scriptsUnintentional violations > neutral521 (15)
Bastin et al., 2021 [42]InductionVerbal scriptsShame during reflectionNone42 (42)
Berthoz et al., 2002 [43]InductionVerbal scriptsUnintentional violations > normal1512 (0)
Finger et al., 2006 [44]InductionVerbal scriptsMoral and social with audience > social and neutral without audience216 (-)
Krach et al., 2011 [45]InductionVignettesVicarious embarrassment > neutral932 (17)
Krach et al., 2015 [46]InductionVignettesSocial pain > social neutral1716 (0)
Laneri et al., 2017 [47]InductionVignettesEmpathic embarrassment > neutral1451 (21)
Melchers et al., 2015 [48]InductionPicturesVicarious embarrassment > neutral660 (39)
Mayer et al., 2020 [49]InductionVignettes([allo: shared > neutral + non-shared > neutral] ∩ [ego: shared > neutral + non-shared > neutral]).1348 (26)
Michl et al., 2012 [50]InductionVerbal scriptsShame > neutral1014 (7)
Moll et al., 2008 [8]InductionVerbal scriptsEmbarrassment > neutral512 (6)
Morita et al., 2008 [51]InductionSelf- and other-facesSelf-face > other-face919 (10)
Morita et al., 2012 [52]InductionSelf- and other-facesSelf-face > other-face2915 (2)
Morita et al., 2013 [53]InductionSelf- and other-facesSelf-face > other-face1732 (16)
Morita et al., 2016 [54]InductionSelf- and other-facesSelf-face > other-face1318 (0)
Stroth et al., 2019 [55]InductionVignettesf-cg: (sks + nks) > ns29 (9)
Takahashi et al., 2004 [56]InductionVerbal scriptsEmbarrassment > neutral1019 (9)
Wagner et al., 2011 [57]RecollectionVerbal scriptsShame > neutral1015 (15)
Zhu et al., 2018 [58]Interpersonal gamePictorial stimuli (dots)Shame > happiness230 (17)
GuiltBasile et al., 2010 [59]InductionVerbal and facial stimuliGuilt > anger and sadness322 (13)
Bastin et al., 2021 [42]InductionVerbal scriptsGuilt during reflectionNone42 (42)
Cheng et al., 2021 [60]Obedience paradigmMoviePositive correlation of guilt ratings in the harm > neutral condition1061 (32)
Dominguez et al., 2018 [61]InductionPicturesIncorrect versus correct shooting decisions248 (35)
Finger et al., 2006 [44]InductionVerbal scriptsMoral > social and neutral516 (-)
Fourie et al., 2014 [62]Implicit association taskVerbal and facial stimuliPrejudice feedback > neutral feedback522 (22)
Gilead et al., 2016 [63]InductionVerbal scriptsGuilt > anger, joy, pride1019 (14)
Gradin et al., 2016 [64]Interpersonal gameVerbalDefection > cooperation625 (17)
Green et al., 2012 [65]InductionVerbal scriptsGuilt > indignation (within HC)722 (18)
Kedia et al., 2008 [66]InductionVerbal scriptsGuilt > self-anger429 (14)
Li et al., 2020 [67]Interpersonal gamePictorial stimuli (dots)Out-group_ Commit > Out-group_ Observe231 (19)
Michl et al., 2012 [50]InductionVerbal scriptsGuilt > neutral1914 (7)
Moll et al., 2008 [8]InductionVerbal scriptsGuilt > neutral612 (6)
Molenberghs et al., 2015 [68]InductionVideoCivilians > Soldiers348 (24)
Morey et al., 2012 [69]InductionVerbal scriptsPositive correlation of guilt616 (0)
Nihonsugi et al., 2021 [70]Interpersonal gamePictorial stimuli (cross) 652 (26)
Peth et al., 2015 [71]RecollectionVerbalGuilty action > neutral1020 (6)
Shin et al., 2000 [72]RecollectionVerbal scriptsGuilt > neutral88 (0)
Takahashi et al., 2004 [56]InductionVerbal scriptsGuilt > neutral519 (9)
Ty et al., 2017 [73]InductionVerbal and pictorial stimuliRestitution > harm118 (9)
Wagner et al., 2011 [57]RecollectionVerbal scriptsGuilt > neutral1015 (15)
Yu et al., 2013 [74]Interpersonal gamePictorial stimuli (dots)Self-incorrect > both incorrect124 (11)
Zhu et al., 2019 [58]Interpersonal gamePictorial stimuli (dots)Guilt > happiness530 (17)
Table 3. Results of the meta-analysis on shame/embarrassment processing.
Table 3. Results of the meta-analysis on shame/embarrassment processing.
ClusterVolume (mm3)CoordinatesALE Value (^103)LateralisationAnatomical LabelBA
−3820−1815.48 IFGorb47
21984−10282821.26LeftAnterior cingulate gyrus32
−6244020.42 Medial frontal gyrus8
48405082819.80RightPrecentral gyrus9/6
Note: the table shows the results of the meta-analysis on shame/embarrassment neural correlates. BA = Brodmann’s area, IFGorb = inferior frontal gyrus pars orbitalis, IFGtri = inferior frontal gyrus pars triangularis. Coordinates reported are in MNI space.
Table 4. Results of the meta-analysis on guilt processing.
Table 4. Results of the meta-analysis on guilt processing.
ClusterCluster Size (mm3)CoordinatesALE Value (^103)LateralisationAnatomical LabelBA
2904−46−582018.78LeftSTG/angular gyrus18
Note: the table shows the results of the meta-analysis on guilt neural correlates. BA = Brodmann’s area, IFGorb = inferior frontal gyrus pars orbitalis, STG = superior temporal gyrus. Coordinates are in MNI space.
Table 5. Contrast analyses results.
Table 5. Contrast analyses results.
ClusterCluster Size (mm3)CoordinatesLateralisationAnatomical LabelBA
Conjunction analysis
Contrast analysis—Shame/Emb vs. Guilt
180−62940LeftAnterior cingulate cortex
232−52033LeftAnterior cingulate cortex
Contrast analysis—Guilt vs. Shame/Emb
No significant results
Note: the table shows the results of contrast analyses. BA = Brodmann’s area, IFGorb = inferior frontal gyrus pars orbitalis. Coordinates are reported in MNI space.
Disclaimer/Publisher’s Note: The statements, opinions and data contained in all publications are solely those of the individual author(s) and contributor(s) and not of MDPI and/or the editor(s). MDPI and/or the editor(s) disclaim responsibility for any injury to people or property resulting from any ideas, methods, instructions or products referred to in the content.

Share and Cite

MDPI and ACS Style

Piretti, L.; Pappaianni, E.; Garbin, C.; Rumiati, R.I.; Job, R.; Grecucci, A. The Neural Signatures of Shame, Embarrassment, and Guilt: A Voxel-Based Meta-Analysis on Functional Neuroimaging Studies. Brain Sci. 2023, 13, 559.

AMA Style

Piretti L, Pappaianni E, Garbin C, Rumiati RI, Job R, Grecucci A. The Neural Signatures of Shame, Embarrassment, and Guilt: A Voxel-Based Meta-Analysis on Functional Neuroimaging Studies. Brain Sciences. 2023; 13(4):559.

Chicago/Turabian Style

Piretti, Luca, Edoardo Pappaianni, Claudia Garbin, Raffaella Ida Rumiati, Remo Job, and Alessandro Grecucci. 2023. "The Neural Signatures of Shame, Embarrassment, and Guilt: A Voxel-Based Meta-Analysis on Functional Neuroimaging Studies" Brain Sciences 13, no. 4: 559.

Note that from the first issue of 2016, this journal uses article numbers instead of page numbers. See further details here.

Article Metrics

Back to TopTop