Empathic neural responses are modulated by the perceived fairness of others

“Increased pain-related activation (asterisk indicates whole-brain corrected) for women in ACC* [9, 18, 27], left FI* [242, 15,23], right FI* [30, 18,218], left SII* [260,230, 18], right SII* [63,230, 24] and brainstem* [3,218,218]; for men in left FI* [233, 33, 3] right FI [42, 33, 3] and brainstem [3,233,230].”

“Furthermore, a few studies have suggested sex differences in these activations (Leibenluft et al. 2004; Azim et al. 2005; Platek et al. 2005; Singer et al. 2006).”

“For example, although both men and women exhibited empathy-related cortical activation after seeing study confederates in pain, only in men were those activations reduced if the confederates had played unfairly in an economic game played prior to pain testing (Singer et al. 2006).”

“Based on previous findings indicating that the impact of an unfair partner’s behaviour is stronger in men compared to women [46], only male participants were selected.”

“Singer et al. (2006) found that for male participants, seeing the pain of a cooperative confederate activated neural networks associated with first-hand experience of pain.”

“…of empathy for pain—in particular, when our attention is directed to the somatosensory aspects of the pain experience (Avenanti, Bueti, Galati, & Aglioti, 2005; Avenanti, Minio-Paluello, Bufalari, & Aglioti, 2006; Gu & Han, 2007; Lamm, Nusbaum, Meltzoff, & Decety, 2007b; Singer et al., 2006).”

“This idea is of ancient origin, but is most clearly articulated in Adam Smith’s Theory of Moral Sentiments.”

“Substantial evidence shows that when reasoning about others, women are more inclined to empathize than men (Toussaint and Webb, 2005), with a corresponding gender difference in neural activation (Singer et al., 2006).”

“…(Yang et al. 2009). fMRI studies have not consistently revealed a greater activity in pain-related regions in females when watching others in pain (Lamm et al. 2011), but there is evidence that females modulate their vicarious pain less than men according to social context (Singer et al. 2006).”

“…behavior-based and neural activation differences, showing for instance that empathy in women is less susceptible to previous unfairness of the observed other (Singer et al. 2006) and that emotion recognition activates mirror regions to a stronger degree in women (Schulte-Ruther et al. 2008).”

“, 2004) are acknowledged to be related to the intrinsically hedonic value of mutual cooperation, while the orbitofrontal cortex has been linked with the desire for revenge toward unfair partners (Singer et al., 2006; for an overview see Fehr and Camerer, 2007).”

“…and social decision making (Lee, 2008), romantic love (Bartels & Zeki, 2006; Fisher, Aron, & Brown, 2005), empathy (Lamm, Batson, & Decety, 2007; Singer et al., 2006), envy (Shamay-Tsoory, Tibi-Elhanany, & Aharon-Peretz, 2007), moral decision making (Casebeer, 2003), aesthetic judgment (Kirk,…”

“Studies using brain-scanning technology suggest that people experience pleasure when non-cooperators receive punishment (de Quervain et al., 2004; Singer et al., 2006).”

“…(Lahnakoski et al. 2012), revealing bilateral activation clusters in regions including anterior insulae and S2 that are related to both fi rst-hand and vicarious experiences of pain (Singer et al. 2004, 2006; Jackson et al. 2005, 2006; Hein et al. 2010; Lamm et al. 2011; Morelli et al. 2014).”

“In a renowned experiment conducted by the German neuroscientist Tania Singer and her research team (Singer et al. 2006), subjects were privy to the alleged fictional corporal punishment of individuals who had either played fairly in a game or had cheated.”

“Neurophysiological studies indicate that when people see or even imagine the pain of others, the brain activity is the same as if they were experiencing the observed pain themselves (Singer et al., 2006).”

“…et al. 2002, 2004); during third-party punishment with money or symbolic points (De Quervain et al. 2004), especially for male punishment observers (Singer et al. 2006); in charitable donations (Moll et al. 2006, Harbaugh et al. 2007, Zaki et al. 2013); in the rejection of ultimatum offers based…”

“This ‘‘pain matrix’’ includes the insula, the anterior and middle cingulate gyrus, the somatosensory cortices (SI and SII), and perhaps premotor areas as well (Singer et al. 2004, 2006; Jackson et al. 2006; Bufalari et al. 2007; Valeriani et al. 2008).”

“For example, knowledge of the cause of the emotion and moral evaluation of another’s behaviour can increase or decrease the empathy (Lamm, Batson, & Decety, 2007; Singer et al., 2006).”

“The structure of social interaction meets indeed particular, often irrepressible, expectations, such as those provided by reputational knowledge [9,10,37].”

“about the perceived fairness of an individual (Singer et al., 2006) or the successful outcome of a painful treatment (Lamm et al.”

“, 2004) and when evaluating the perceived ‘fairness’ of others’ actions (Singer et al., 2006).”

“Previous studies on ultimatum bargaining suggest that these activations can be interpreted in terms of registering both norm and status concern/self-interest violations (Sanfey et al., 2003; Knoch et al., 2006; Singer et al., 2006).”

“insula) when seeing an “unfair” player in pain in comparison to a “fair” player (Singer et al., 2006), suggesting that perception of fairness may affect the degree of empathy experienced towards others.”

“Neuroimaging studies of economic exchange games showa relationship between costly punishment of unfair behavior and neural structures associated with reward processing such as the dorsal and ventral striatum (De Quervain et al., 2004; Singer et al., 2006; Crockett et al., 2013).”

“…has been more closely linked with tracking the rewards 8 and emotional experiences of others 46 to support psychological processes including empathy 46 and social learning, in both monkeys and humans 8,47 Nevertheless, our observation that some neurons in ACCg encode strategic social information…”

“…medial prefrontal cortex and temporo-parietal junction, also play a vital role in prosocial behaviors—although traditionally they are not considered to be a part of the RLDM neural circuitry (Singer et al., 2006; Hare et al., 2010; Morishima et al., 2012; Waytz et al., 2012; Smith et al., 2014).”

“sity of the desire for revenge, admitted in a questionnaire filled after the experiment [25].”

“For example, even though “the meeting of minds” (Amodio and Frith, 2006) certainly has hedonic qualities and may “feel good” (see Schilbach et al., 2010a), the ventral striatum or other reward-related brain areas cannot be seen as specific neural correlates of the secondperson perspective or of mutual social interacti…”

“In line with these studies [5,7] (see also [16,17] and [6] for a”

“in someone else - is less pronounced when observers dislike the observed person in pain [33].”

“This dissociation is supported by previous studies on vicarious physical pain, which demonstrated that the affective responses within the AI and ACC are modulated by perceivers’ evaluations of the social setting (Singer et al. 2006; Cheng et al. 2007; Lamm et al. 2007).”

“There is limited evidence that failure to exhibit empathic responses to others’ pain is related to lower self-reported empathic concern or aggressive or antisocial behavior (Singer et al., 2004, 2006).”

“On the other hand, we did not address the modulation of empathy by contextual factors, such as perceived fairness (Singer et al. 2006), appraisal (Lamm, Batson et al. 2007; Lamm, Nusbaum et al. 2007; Hein and Singer 2008; Engen and Singer 2012), or perceived group membership (Hein et al. 2010).”

“Neuroscientific evidence also points in this direction: Empathy-related brain activity is observed in participants when another person who had previously acted in a fair manner receive a painful stimulus [26].”

“Previous research has generated results suggesting that empathic responses are modulated by situational characteristics like the affective link between individuals [50], social relationship [56], or being simultaneously in pain [9].”

“2004b ; Singer et al. 2006 ), consumer preferences ( McClure et al.”

“Previous neurological studies have shown a link between self-oriented and other-oriented feelings (e.g., Decety & Gr ezes, 2006; Jackson, Meltzoff, & Decety, 2005; Singer et al., 2004; Singer et al., 2006; for reviews, see Gallese, 2003 and Preston & deWaal, 2002).”

“…2008) on the ultimatum and dictator games, we did not use a “one-shot” paradigm but a repeatedgame paradigm where the same proposer repeatedly interacted with the same responder (for examples, see Civai, Corradi-Dell’Acqua, Gamer, & Rumiati, 2010; Polezzi et al., 2008; Smith & Silberberg, 2010).”

“The notion that schadenfreude is evoked when someone gets his or her comeuppance is clearly demonstrated by the theoretical and empirical work of Feather (eg, Feather, 1994 , 1999 , 2006 ; Feather & Sherman, 2002 ) and received further support from other studies (eg, Singer et al, 2006 ; Van Dijk, Goslinga, & Ouwerker…”

“Accompanied by AI activity, ACC also showed increased activations during unfair offers with the increment of self-contribution, indicating the involvement of AI/ACC network in norm violations (Singer et al., 2006; Güroğlu et al., 2010, 2011).”

“…feelings when they experience different emotions or sensations such as happiness, sadness, pain, touch, or tickling (Gallese, 2003). de Vignemont and Singer (2006) defined the four essential elements of empathy: (1) an affective state (2) that mirrors (3) and is induced by observing or imagining…”

“Interestingly, there is evidence that fairness-directed conducts, such as mutual cooperation with a human partner [24], donating to a charity [25,26], altruistic punishment and revenge [27,28] are related to neural activation of the mesolimbic dopaminergic system.”

“, unfair behaviors in a monetary game) have been shown 151 to reduce empathic responses from other players [42], supporting the main role of 152 approach/avoidance behaviors to promote social relationships.”

“In one functional magnetic resonance imaging (fMRI) study (Singer et al., 2006), male volunteers who observed an unfair confederate receiving pain showed lower empathy-related activation, accompanied by reward-related activation that might represent the desire for revenge.”

“Singer et al. (2006), using functionalmagnetic resonance imaging (fMRI), found that the activity in theACC decreaseswhen people believe that the punishment another person receives is considered fair; this was particularly true of the male participants.”

“While it is well-established that Schadenfreude does in fact occur in such cases (e.g., Feather and Nairn, 2005; Singer et al., 2006; Leach and Spears, 2009 ) and that this can also influence facial muscle activity recorded using EMG (Cikara and Fiske, 2012), to our knowledge no research has been…”

“Follow-up research by Singer et al. (2006) also indicates a clear gender difference in this pattern.”

“As mentioned, these secondary mechanisms may include, but are not limited to, narrative thinking (Breithaupt, 2011; Keen, 2007; Voss, 2004), fairness perception (Singer et al., 2006), causal fault attribution, past experiences, and temporal development (Saarela et al., 2007).”

“A recent follow-up study showed that the AI activity in response to pain in another person was modulated by the perceived fairness of that person (Singer et al, 2006 ).”

“When a defector in an economic game was punished, empathy-related brain responses were reduced, and reward-related brain responses were increased, in men but not women (Singer et al., 2006).”

“On the other hand, other studies revealed the modulation of contextual factors on empathy for other’s pain, such as the affective link between individuals (Singer et al., 2006), the intentionality of the perceived agency who induced the pain (Akitsuki & Decety, 2009; Decety et al.”

“The anterior insular region has also been found to activate during awareness (Singer et al., 2004, Singer et al., 2006) or viewing (Jackson et al., 2006; Lamm et al., 2007; Morrison et al., 2004) of another individual’s pain.”

“Activation in this network was also observed when participants saw an unknown but likeable person suffering pain (Singer et al., 2006) or watched videos showing body”