The Neural Correlates and Evolutionary Benefits of Embarrassment
By Amanda Yang
A common fear amongst humans is the failure to meet social expectations and uphold one’s external, or social, image (Leary & Kowalski, 1995). Such fear is a primary cause of feelings of embarrassment (Miller, 1996). Embarrassment itself can be thought of as a self-conscious emotion that arises when the self is brought to the forefront of awareness after violation of a social expectation (Keltner, 1995). Notably, many physiological changes are associated with the feeling of embarrassment such as increased pulse, blood pressure, and perspiration (Keltner, 1995). In addition to physiological effects, several behavioral changes are also associated with embarrassment such as gaze aversion and increased face touching (e.g., Shearn et al., 1990). At a more granular level, research in the past two decades has made progress in identifying neural pathways of embarrassment, although this area is not well studied. Previous studies indicate that the anterior cingulate cortex and angular insular play critical roles in embarrassment, but further implications of these findings remain unknown. I argue that in the context of behavioral and social studies of embarrassment, the neural pathways identified may represent not just brain regions associated with embarrassment but also biological sites performing evolutionarily beneficial functions and thus molded by evolutionary selection.
In 2008, an fMRI study by Morita and colleagues was one of the first to investigate and identify the brain regions associated with embarrassment. 19 healthy participants were recruited for this two-day study. On the first day of experimentation, the participants’ faces were video recorded as they gave speeches in front of a video camera. 21 images were then selected from the video recording for each participant. The experimenters then gave each image a photogenic score with high scores indicating most attractive and low scores indicating least attractive. On the second day of experimentation, participants underwent fMRI scanning and were presented with the 21 self-images and 21 images of another participant. They then rated how embarrassed they felt when they saw each image. For each image, as the experimenters’ photogenic score decreased, the participant’s embarrassment rating increased significantly for self-images in comparison to images of others. Additionally, the researchers saw a significant increase in the activation of the anterior cingulate cortex (ACC) and anterior insular cortex (AI) when participants viewed self-images versus images of others. Since the self-images had a significantly higher mean embarrassment rating in comparison to that of others’ images, the findings suggest that the AI and ACC likely play an important role during embarrassing experiences.
Later, in 2013, Morita and colleagues conducted a follow-up study on 32 healthy participants to understand the specific roles of the ACC and AI during embarrassment. Their experimental paradigm was a modified version of their previous investigation (2008). This time, to heighten feelings of embarrassment and mimic an authentic social situation, Morita et al. (2013) added observers to the experimental design so that half of the participants would be watched by an assigned partner as they viewed the images. Each participant-partner pair underwent simultaneous scanning in different fMRI scanners, such that each person could each see the other’s face through a live video as their brains were scanned. The other half of participants did not have a partner and underwent testing without such observation.
The researchers found that in comparison to those in the non-observation condition, the participants in the observation condition self-reported higher embarrassment ratings and exhibited increased activity in the right AI, suggesting that the right AI makes an important contribution to the subjective feeling of embarrassment. Moreover, in comparison to those in the non-observation condition, those in the observation condition displayed increased functional connectivity between the caudal ACC and the bilateral medial prefrontal cortex (mPFC). Since both their findings and previous research implicate the ACC and mPFC in the assessment of the self (Ochsner et al. as cited in Morita et al., 2013), Morita and colleagues suggest that in social situations, the ACC might have “increased access to information” needed in self-evaluating processes. Though one can question whether viewing unattractive images of oneself constitutes a strong enough embarrassment stimulus, the findings of Morita et al. (2008) and Morita et al. (2013) identify the ACC and the AI as possible neurological bases of embarrassment.
Morita et al.’s research, however, is also important for reasons beyond the neuroscience they describe in their paper. In particular, their study potentially implicates the ACC and the AI as important sites conferring evolutionary benefits. Some psychologists have conceptualized embarrassment as an anticipation of negative evaluation by others (Miller, 1995) — an uncomfortable feeling but one necessary to train ourselves to abide by social norms. Thought of in this way, embarrassment represents an internalized cost of socially undesirable behavior, inhibiting people from committing actions that would gain the disapproval of others. This inhibition could conceivably have provided a selective advantage, enabling humans to cooperate and help each other more. Because of this hypothesized evolutionary benefit of embarrassment, it follows that the ACC and the AI—as neural mediators of embarrassment—are locations in the brain whose functions have been molded by selective pressure. Morita et al. (2013) do not mention evolutionary psychology in their study, but when their data is interpreted in the context of the social role of embarrassment, it suggests that the neurobiology of ACC and AI are products of evolutionary selection, opening up a potential area of investigation into the evolutionary genetics and physiology of those brain regions as they relate to behavior.
Beyond the cognitive science of healthy participants, the role of the ACC and AI in the neural pathways of embarrassment has been further elucidated by the study of dementia patients with frontotemporal lobar degeneration (FTLD). FTLD predominantly atrophies the frontal and temporal lobes, amygdala, and other regions including the ACC and AI (Craik et al., 1999). Patients with this condition repeatedly commit social gaffes, making them interesting subjects for the study of embarrassment (Neary et al., 1998).
In one study, for instance, FTLD patients performed significantly worse in detecting social faux paus in short stories compared to controls (Gregory et al., 2002), suggesting that these patients have deficits in self-conscious emotions, chiefly embarrassment. In another study conducted by Sturm et al. (2008), emotional responding was examined in 24 FTLD patients and 16 controls during a karaoke task known to be reliably embarrassing for healthy individuals. The task involved participants belting out a song and then having them listen to their own singing without the background music playing. When watching themselves sing, the FTLD patients reported significantly lower embarrassment ratings than the controls, and the FTLD patients also exhibited diminished physiological responses characteristic of embarrassment (e.g., increased heart rate, blood pressure, skin conductance). These findings suggest that FTLD patients have deficits in self-conscious emotional responding.
In a follow up to their 2008 study, Sturm et al. (2013) investigated the specific neural correlates of embarrassment using the karaoke task described above. Both FTLD patients and healthy controls were recruited, and all participants underwent MRI scanning. Levels of embarrassment were assessed based on both physiological measures known to be heightened during embarrassment (e.g., heart rate, blood pressure, etc.) and behavioral measures (e.g., gaze aversion, blushing), which were quantified using the Emotional Expressive Behavior coding system (Gross & Levenson, 1993 as cited in Sturm et al., 2013). The researchers found that the ACC was the only brain region that could significantly predict one’s level of embarrassment. In particular, the smaller the ACC gray matter volume, the more diminished the physiological and behavioral emotional responses were. This finding is significant because it presents a predictive relationship between the ACC and embarrassment, adding support to aforementioned studies like Morita et al. (2008) and Sturm et al. (2008) relating ACC to embarrassment.
Despite its neurological focus, this study of FTLD patients and their compromised ability to feel embarrassed, also helps us to think about the evolutionary benefits of embarrassment and why humans feel this emotion in the first place. Although Sturm et al. do not mention this explicitly, their study of FTLD patients sheds light on the social function of embarrassment by depicting how those with a deficit in its neural pathway behave. Indeed, FTLD patients frequently act in socially inappropriate ways (e.g., inappropriateness of affect) often leaving caregivers disturbed (Neary et al., 1998). Taken together with other studies, therefore, their work helps flesh out an evolutionary hypothesis on the role of ACC and AI neural functioning in the context of embarrassment. Previous cognitive scientists, for instance, have noted that embarrassment is vital to the assimilation of social norms, a remarkably complex yet mostly subconscious task all humans must perform in order to live with others (Feinberg, Willer, & Keltner, 2012). In particular, embarrassment likely helps humans remember social gaffes and instances of social disapproval so that they do not repeat their mistakes and risk alienating others. It may also serve as a signal to others that the mistake was unintentional, committed out of naivete rather than ill-will—an important message for retaining others’ trust and repairing the social damage of the mistake. Embarrassment would therefore conceivably confer reproductive advantages by helping people understand how to interact with others and obtain aid from others to survive. In this context, the study of FTLD patients therefore implies that the function of the ACC and AI may have been the object of significant selective pressure.
More research, however, is needed. While we have made important progress in identifying the neural correlates of embarrassment, more work is required in order to understand this phenomenon, including its evolutionary underpinnings. The relatively small number of studies conducted to date and their varying methodologies prevent us from making strong conclusions. Few studies, for instance, have compared the neural correlates of embarrassment to that of other negative self-conscious emotions (e.g., guilt, shame) in a single experiment. Future studies should address this gap in the literature to help elucidate the extent of neurobiological overlap among these emotions and determine possible emotion-specific regions (e.g., embarrassment-specific region, guilt-specific region, etc.). Moreover, future research should also investigate how social anxiety may modulate the neural underpinnings of embarrassment since excessive concern for others’ negative judgment is a primary characteristic of social anxiety. This line of research would help us understand the potential altered neural activation in social anxiety disorders, which could provide insight into developing treatment for these patients.
Despite the limited research in this area, current studies have come to the consensus that the ACC and AI serve critical functions in the experience of embarrassment. Moreover, when neurological studies in this field are interpreted alongside behavioral models and physiological studies of embarrassment, they suggest a possible beneficial role for embarrassment in our evolutionary history. Although feelings of embarrassment may make one feel vulnerable or awkward, they signal to others that we care about their opinions and teach us to not commit the same social mistakes again—benefits which could have led to superior socialization and greater reproductive fitness.
About the Author
Amanda Yang is a junior at Harvard College concentrating in Psychology with a secondary field in Molecular and Cellular Biology.
References
Craik, F., Moroz, T., Moscovitch, M., Stuss, D., Winocur, G., Tulving, E., & Kapur, S. (1999).
In Search of the Self: A Positron Emission Tomography Study. Psychological Science, 10(1), 26-34. Retrieved December 7, 2020, from http://www.jstor.org/stable/40063373
Feinberg, M., Willer, R., & Keltner, D. (2012). Flustered and faithful: Embarrassment as a signal
of prosociality. Journal of Personality and Social Psychology, 102(1), 81–97.
Gregory, C., Lough, S., Stone, V., Erzinclioglu, S., Martin, L., Baron-Cohen, S., & Hodges, J. R.
(2002). Theory of mind in patients with frontal variant frontotemporal dementia and Alzheimer's disease: theoretical and practical implications. Brain: a journal of neurology, 125(Pt 4), 752–764. https://doi.org/10.1093/brain/awf079
Keltner, D., Buswell, B.N. (1997). Embarrassment: Its distinct form and appeasement functions.
Psychological Bulletin, 122(3), 250–270.
Leary, M. R., & Kowalski, R. M. (1995). The self-presentation model of social phobia. In R. G.
Heimberg, M. R. Liebowitz, D. A. Hope, & F. R. Schneier (Eds.), Social phobia: Diagnosis, assessment, and treatment (p. 94–112). The Guilford Press.
Miller R. S. (1995). On the nature of embarrassability: shyness, social evaluation, and social
skill. Journal of personality, 63(2), 315–339. https://doi.org/10.1111/j.1467-6494.1995.tb00812.x
Miller, R. S. (1996). Emotions and social behavior. Embarrassment: Poise and peril in everyday
life. The Guilford Press.
Morita, T., Itakura, S., Saito, D. N., Nakashita, S., Harada, T., Kochiyama, T., & Sadato, N.
(2008). The role of the right prefrontal cortex in self-evaluation of the face: a functional magnetic resonance imaging study. Journal of cognitive neuroscience, 20(2), 342–355.
Morita, T., Tanabe, H. C., Sasaki, A. T., Shimada, K., Kakigi, R., & Sadato, N. (2014). The
anterior insular and anterior cingulate cortices in emotional processing for self-face recognition. Social cognitive and affective neuroscience, 9(5), 570–579. https://doi.org/10.1093/scan/nst011
Müller-Pinzler, L., Rademacher, L., Paulus, F. M., & Krach, S. (2016). When your friends make
you cringe: Social closeness modulates vicarious embarrassment-related neural activity. Social cognitive and affective neuroscience, 11(3), 466–475. https://doi.org/10.1093/scan/nsv130
Neary, D., Snowden, J. S., Gustafson, L., Passant, U., Stuss, D., Black, S., Freedman, M.,
Kertesz, A., Robert, P. H., Albert, M., Boone, K., Miller, B. L., Cummings, J., & Benson, D. F. (1998). Frontotemporal lobar degeneration: a consensus on clinical diagnostic criteria. Neurology, 51(6), 1546–1554. https://doi.org/10.1212/wnl.51.6.1546
Shearn, D., Bergman, E., Hill, K., Abel, A., Hinds, L. (1990). Facial coloration and temperature
responses in blushing. Psychophysiology, 27, 687–93.
Sturm, V. E., Ascher, E. A., Miller, B. L., & Levenson, R. W. (2008). Diminished self-conscious
emotional responding in frontotemporal lobar degeneration patients. Emotion (Washington, D.C.), 8(6), 861–869. https://doi.org/10.1037/a0013765
Sturm, V. E., Sollberger, M., Seeley, W. W., Rankin, K. P., Ascher, E. A., Rosen, H. J., Miller,
B. L., & Levenson, R. W. (2013). Role of right pregenual anterior cingulate cortex in self-conscious emotional reactivity. Social cognitive and affective neuroscience, 8(4), 468–474. https://doi.org/10.1093/scan/nss023
Tangney, J. P., Stuewig, J., & Mashek, D. J. (2007). Moral emotions and moral behavior. Annual
review of psychology, 58, 345–372. https://doi.org/10.1146/annurev.psych.56.091103.070145
In 2008, an fMRI study by Morita and colleagues was one of the first to investigate and identify the brain regions associated with embarrassment. 19 healthy participants were recruited for this two-day study. On the first day of experimentation, the participants’ faces were video recorded as they gave speeches in front of a video camera. 21 images were then selected from the video recording for each participant. The experimenters then gave each image a photogenic score with high scores indicating most attractive and low scores indicating least attractive. On the second day of experimentation, participants underwent fMRI scanning and were presented with the 21 self-images and 21 images of another participant. They then rated how embarrassed they felt when they saw each image. For each image, as the experimenters’ photogenic score decreased, the participant’s embarrassment rating increased significantly for self-images in comparison to images of others. Additionally, the researchers saw a significant increase in the activation of the anterior cingulate cortex (ACC) and anterior insular cortex (AI) when participants viewed self-images versus images of others. Since the self-images had a significantly higher mean embarrassment rating in comparison to that of others’ images, the findings suggest that the AI and ACC likely play an important role during embarrassing experiences.
Later, in 2013, Morita and colleagues conducted a follow-up study on 32 healthy participants to understand the specific roles of the ACC and AI during embarrassment. Their experimental paradigm was a modified version of their previous investigation (2008). This time, to heighten feelings of embarrassment and mimic an authentic social situation, Morita et al. (2013) added observers to the experimental design so that half of the participants would be watched by an assigned partner as they viewed the images. Each participant-partner pair underwent simultaneous scanning in different fMRI scanners, such that each person could each see the other’s face through a live video as their brains were scanned. The other half of participants did not have a partner and underwent testing without such observation.
The researchers found that in comparison to those in the non-observation condition, the participants in the observation condition self-reported higher embarrassment ratings and exhibited increased activity in the right AI, suggesting that the right AI makes an important contribution to the subjective feeling of embarrassment. Moreover, in comparison to those in the non-observation condition, those in the observation condition displayed increased functional connectivity between the caudal ACC and the bilateral medial prefrontal cortex (mPFC). Since both their findings and previous research implicate the ACC and mPFC in the assessment of the self (Ochsner et al. as cited in Morita et al., 2013), Morita and colleagues suggest that in social situations, the ACC might have “increased access to information” needed in self-evaluating processes. Though one can question whether viewing unattractive images of oneself constitutes a strong enough embarrassment stimulus, the findings of Morita et al. (2008) and Morita et al. (2013) identify the ACC and the AI as possible neurological bases of embarrassment.
Morita et al.’s research, however, is also important for reasons beyond the neuroscience they describe in their paper. In particular, their study potentially implicates the ACC and the AI as important sites conferring evolutionary benefits. Some psychologists have conceptualized embarrassment as an anticipation of negative evaluation by others (Miller, 1995) — an uncomfortable feeling but one necessary to train ourselves to abide by social norms. Thought of in this way, embarrassment represents an internalized cost of socially undesirable behavior, inhibiting people from committing actions that would gain the disapproval of others. This inhibition could conceivably have provided a selective advantage, enabling humans to cooperate and help each other more. Because of this hypothesized evolutionary benefit of embarrassment, it follows that the ACC and the AI—as neural mediators of embarrassment—are locations in the brain whose functions have been molded by selective pressure. Morita et al. (2013) do not mention evolutionary psychology in their study, but when their data is interpreted in the context of the social role of embarrassment, it suggests that the neurobiology of ACC and AI are products of evolutionary selection, opening up a potential area of investigation into the evolutionary genetics and physiology of those brain regions as they relate to behavior.
Beyond the cognitive science of healthy participants, the role of the ACC and AI in the neural pathways of embarrassment has been further elucidated by the study of dementia patients with frontotemporal lobar degeneration (FTLD). FTLD predominantly atrophies the frontal and temporal lobes, amygdala, and other regions including the ACC and AI (Craik et al., 1999). Patients with this condition repeatedly commit social gaffes, making them interesting subjects for the study of embarrassment (Neary et al., 1998).
In one study, for instance, FTLD patients performed significantly worse in detecting social faux paus in short stories compared to controls (Gregory et al., 2002), suggesting that these patients have deficits in self-conscious emotions, chiefly embarrassment. In another study conducted by Sturm et al. (2008), emotional responding was examined in 24 FTLD patients and 16 controls during a karaoke task known to be reliably embarrassing for healthy individuals. The task involved participants belting out a song and then having them listen to their own singing without the background music playing. When watching themselves sing, the FTLD patients reported significantly lower embarrassment ratings than the controls, and the FTLD patients also exhibited diminished physiological responses characteristic of embarrassment (e.g., increased heart rate, blood pressure, skin conductance). These findings suggest that FTLD patients have deficits in self-conscious emotional responding.
In a follow up to their 2008 study, Sturm et al. (2013) investigated the specific neural correlates of embarrassment using the karaoke task described above. Both FTLD patients and healthy controls were recruited, and all participants underwent MRI scanning. Levels of embarrassment were assessed based on both physiological measures known to be heightened during embarrassment (e.g., heart rate, blood pressure, etc.) and behavioral measures (e.g., gaze aversion, blushing), which were quantified using the Emotional Expressive Behavior coding system (Gross & Levenson, 1993 as cited in Sturm et al., 2013). The researchers found that the ACC was the only brain region that could significantly predict one’s level of embarrassment. In particular, the smaller the ACC gray matter volume, the more diminished the physiological and behavioral emotional responses were. This finding is significant because it presents a predictive relationship between the ACC and embarrassment, adding support to aforementioned studies like Morita et al. (2008) and Sturm et al. (2008) relating ACC to embarrassment.
Despite its neurological focus, this study of FTLD patients and their compromised ability to feel embarrassed, also helps us to think about the evolutionary benefits of embarrassment and why humans feel this emotion in the first place. Although Sturm et al. do not mention this explicitly, their study of FTLD patients sheds light on the social function of embarrassment by depicting how those with a deficit in its neural pathway behave. Indeed, FTLD patients frequently act in socially inappropriate ways (e.g., inappropriateness of affect) often leaving caregivers disturbed (Neary et al., 1998). Taken together with other studies, therefore, their work helps flesh out an evolutionary hypothesis on the role of ACC and AI neural functioning in the context of embarrassment. Previous cognitive scientists, for instance, have noted that embarrassment is vital to the assimilation of social norms, a remarkably complex yet mostly subconscious task all humans must perform in order to live with others (Feinberg, Willer, & Keltner, 2012). In particular, embarrassment likely helps humans remember social gaffes and instances of social disapproval so that they do not repeat their mistakes and risk alienating others. It may also serve as a signal to others that the mistake was unintentional, committed out of naivete rather than ill-will—an important message for retaining others’ trust and repairing the social damage of the mistake. Embarrassment would therefore conceivably confer reproductive advantages by helping people understand how to interact with others and obtain aid from others to survive. In this context, the study of FTLD patients therefore implies that the function of the ACC and AI may have been the object of significant selective pressure.
More research, however, is needed. While we have made important progress in identifying the neural correlates of embarrassment, more work is required in order to understand this phenomenon, including its evolutionary underpinnings. The relatively small number of studies conducted to date and their varying methodologies prevent us from making strong conclusions. Few studies, for instance, have compared the neural correlates of embarrassment to that of other negative self-conscious emotions (e.g., guilt, shame) in a single experiment. Future studies should address this gap in the literature to help elucidate the extent of neurobiological overlap among these emotions and determine possible emotion-specific regions (e.g., embarrassment-specific region, guilt-specific region, etc.). Moreover, future research should also investigate how social anxiety may modulate the neural underpinnings of embarrassment since excessive concern for others’ negative judgment is a primary characteristic of social anxiety. This line of research would help us understand the potential altered neural activation in social anxiety disorders, which could provide insight into developing treatment for these patients.
Despite the limited research in this area, current studies have come to the consensus that the ACC and AI serve critical functions in the experience of embarrassment. Moreover, when neurological studies in this field are interpreted alongside behavioral models and physiological studies of embarrassment, they suggest a possible beneficial role for embarrassment in our evolutionary history. Although feelings of embarrassment may make one feel vulnerable or awkward, they signal to others that we care about their opinions and teach us to not commit the same social mistakes again—benefits which could have led to superior socialization and greater reproductive fitness.
About the Author
Amanda Yang is a junior at Harvard College concentrating in Psychology with a secondary field in Molecular and Cellular Biology.
References
Craik, F., Moroz, T., Moscovitch, M., Stuss, D., Winocur, G., Tulving, E., & Kapur, S. (1999).
In Search of the Self: A Positron Emission Tomography Study. Psychological Science, 10(1), 26-34. Retrieved December 7, 2020, from http://www.jstor.org/stable/40063373
Feinberg, M., Willer, R., & Keltner, D. (2012). Flustered and faithful: Embarrassment as a signal
of prosociality. Journal of Personality and Social Psychology, 102(1), 81–97.
Gregory, C., Lough, S., Stone, V., Erzinclioglu, S., Martin, L., Baron-Cohen, S., & Hodges, J. R.
(2002). Theory of mind in patients with frontal variant frontotemporal dementia and Alzheimer's disease: theoretical and practical implications. Brain: a journal of neurology, 125(Pt 4), 752–764. https://doi.org/10.1093/brain/awf079
Keltner, D., Buswell, B.N. (1997). Embarrassment: Its distinct form and appeasement functions.
Psychological Bulletin, 122(3), 250–270.
Leary, M. R., & Kowalski, R. M. (1995). The self-presentation model of social phobia. In R. G.
Heimberg, M. R. Liebowitz, D. A. Hope, & F. R. Schneier (Eds.), Social phobia: Diagnosis, assessment, and treatment (p. 94–112). The Guilford Press.
Miller R. S. (1995). On the nature of embarrassability: shyness, social evaluation, and social
skill. Journal of personality, 63(2), 315–339. https://doi.org/10.1111/j.1467-6494.1995.tb00812.x
Miller, R. S. (1996). Emotions and social behavior. Embarrassment: Poise and peril in everyday
life. The Guilford Press.
Morita, T., Itakura, S., Saito, D. N., Nakashita, S., Harada, T., Kochiyama, T., & Sadato, N.
(2008). The role of the right prefrontal cortex in self-evaluation of the face: a functional magnetic resonance imaging study. Journal of cognitive neuroscience, 20(2), 342–355.
Morita, T., Tanabe, H. C., Sasaki, A. T., Shimada, K., Kakigi, R., & Sadato, N. (2014). The
anterior insular and anterior cingulate cortices in emotional processing for self-face recognition. Social cognitive and affective neuroscience, 9(5), 570–579. https://doi.org/10.1093/scan/nst011
Müller-Pinzler, L., Rademacher, L., Paulus, F. M., & Krach, S. (2016). When your friends make
you cringe: Social closeness modulates vicarious embarrassment-related neural activity. Social cognitive and affective neuroscience, 11(3), 466–475. https://doi.org/10.1093/scan/nsv130
Neary, D., Snowden, J. S., Gustafson, L., Passant, U., Stuss, D., Black, S., Freedman, M.,
Kertesz, A., Robert, P. H., Albert, M., Boone, K., Miller, B. L., Cummings, J., & Benson, D. F. (1998). Frontotemporal lobar degeneration: a consensus on clinical diagnostic criteria. Neurology, 51(6), 1546–1554. https://doi.org/10.1212/wnl.51.6.1546
Shearn, D., Bergman, E., Hill, K., Abel, A., Hinds, L. (1990). Facial coloration and temperature
responses in blushing. Psychophysiology, 27, 687–93.
Sturm, V. E., Ascher, E. A., Miller, B. L., & Levenson, R. W. (2008). Diminished self-conscious
emotional responding in frontotemporal lobar degeneration patients. Emotion (Washington, D.C.), 8(6), 861–869. https://doi.org/10.1037/a0013765
Sturm, V. E., Sollberger, M., Seeley, W. W., Rankin, K. P., Ascher, E. A., Rosen, H. J., Miller,
B. L., & Levenson, R. W. (2013). Role of right pregenual anterior cingulate cortex in self-conscious emotional reactivity. Social cognitive and affective neuroscience, 8(4), 468–474. https://doi.org/10.1093/scan/nss023
Tangney, J. P., Stuewig, J., & Mashek, D. J. (2007). Moral emotions and moral behavior. Annual
review of psychology, 58, 345–372. https://doi.org/10.1146/annurev.psych.56.091103.070145