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].”

“5A), we focus on the 30-min time window beginning 5 min into the completion of each injection hereafter [first 30 min: BLA, F(1,24) = 4.”

“, 2006); and a spiteful, emotion-driven system including the mPFC, insula and striatum, which represent the immediately reinforcing aspects of sanctioning norm violators (de Quervain et al., 2004; Koenigs & Tranel, 2007; Sanfey et al., 2003; Singer et al., 2006; Tabibnia et al., 2008).”

“Keywords: Cooperation; Altruism; Sacrifice; Trust; Touch”

“For instance, this region is more engaged in prosocial individuals when they defect, whereas pro-self individuals engage the AI more when they reciprocate [King-Casas et al., 2008; Singer et al., 2006].”

“Alongside other characteristics of the observer-target relationship such as the affective link (Singer et al., 2006), familiarity was repeatedly shown to modulate the amount of empathy in a given situation (Cialdini et al., 1997; de Vignemont and Singer, 2006; Hein and Singer, 2008).”

“, 2009), by social relationship with the target (Singer et al., 2006), by familiarity for the target (Cialdini et al.”

“... interpretation of this finding as showing the existence of “a common mechanism for understanding the emotion in others and feeling the same emotions in ourselves.” Now, advocates of the direct-perception modelofempathymight argue thatthisfinding isevidenceinfavor of the second-personal interactionist view of socia…”

“However, prior work has shown a that males have a stronger tendency towards mentalizing, and females towards empathic concern [26, 106].”

“, 2004), as well as the extent of empathy experiencedwith another subject (Singer et al., 2006).”

“…between neurophysiological measures and various subjective measures encompassing empathy- and pain-related emotional and sensory indices (Singer et al. 2004, 2006; Avenanti et al. 2005, 2006; Jackson et al. 2005, 2006; Morrison et al. forthcoming), but so far, no study has linked neural…”

“On the other hand, of course, when relational quality is high, empathic concern will be more naturally felt (Bartal, 1976; Singer et al., 2006), and, thus, such interactions will be marked by what Ashforth and Humphrey describe as “instances whereby one spontaneously and genuinely experiences the…”

“The ventral striatum is known to be involved in the processing of reward and positive affect (Elliot et al., 2000; Roy et al., 2012), learning (Willems et al., 2010) and social cognition (Singer et al., 2006; Tricomi et al., 2010).”

“Research carried out by Singer et al. (2006) is one of the best known examples of this.”

“The physical pain of cheaters activates both pain matrix and reward- related areas in the observer's brain (Singer et al., 2006), while early traumatic social experience virtually blocks both emotion recognition and automatic facial mimicry of a perceived emotion (Ardizzi et al., 2013, 2015).”

“The consideration of fairness for others in pain also modulates the pain matrix (Singer et al., 2006).”

“Others have suggested that beyond the use of shared representations, additional higher-level inferential processes may be necessary for complete empathic understanding and social cognition more generally (Decety and Jackson, 2004; Beer and Ochsner, 2006; Mitchell, 2006; Singer et al., 2006).”

“Key brain regions thought to be involved in emotional empathy include the anterior cingulate and insula, ventromedial prefrontal cortex, and amygdala (13, 69, 70).”

“We also know something of the brain function underlying the emotions joy, ecstasy, rage or fear[24,42,43], all of which may be part of ecstatic experience.”

“In Study 31 (Singer et al., 2006), four nonindependent correlations that are described in the text were not included, though they were listed numerically in the supplementary materials (as indicated in the main text).”

“A very recent fMRI study (Singer et al., 2006) employing the Prisoner’s Dilemma paradigm has shown that the brain activity of male subjects during observing an opponent in pain was modulated by the perceived fairness of the opponent.”

“…or psychological traits such as alexithymia (Bird et al., 2010; Silani et al., 2008), expertise (Cheng et al., 2007), the perception of fairness (Singer et al., 2006), group membership of the other (Avenanti et al., 2010; Hein et al., 2010), contextual appraisal of the situation (Lamm et al.,…”

“For instance, fMRI research shows that there is activation in the reward areas of the brain when bystanders witness leaders being punished when they behave unfairly (Singer et al., 2006).”

“…shared neural representations for the perception and experience of disgust [Jabbi et al., 2007; Wicker et al., 2003], somatosensation [Bufalari et al., 2007; Masten et al., 2011; Singer et al., 2006], emotion [Carr et al., 2003; Pfeifer et al., 2008], and motor behavior [Keysers and Fadiga, 2008].”

“For instance, observing a prosocial subject receiving pain stimulation triggers empathy responses reflected in increased bilateral activity of the AI and the ACC, compared to observing an antisocial subject (Singer et al., 2006).”

“NAc encodes reward value and decision-making, and, as noted, both IC and NAc are implicated in rodent and human social behaviors (Lamm & Singer, 2010; Trezza et al., 2011; Singer et al., 2006; Lichtenberg et al., 2018).”

“This is reminiscent of findings in humans that men and women show similar levels of elevated activity in pain-related regions for fair demonstrators receiving shocks, but only women do for unfair demonstrators [68].”

“Recent studies investigating the neural correlates of empathy have implicated two primary regions; the anterior insula and dorsal anterior cingulate cortex (Botvinick et al., 2005; Singer et al., 2004, 2006; Jackson et al., 2005, 2006; Saarela et al., 2007; Lamm et al., 2007).”

“Recent research in neuroeconomics has also found that such perceived unfairness has neural correlates in pain-related brain areas (Sanfey et al. 2003; Singer et al. 2006) and that the opportunity to punish a norm-violator activates reward-related areas of the brain (De Quervain et al. 2004).”

“Social neuroscience has recently experienced a growing interest in the neural correlates of prosocial and moral behavior (e.g., de Greck et al., 2012; Decety et al., 2011; Güroğlu et al., 2011; Hsu et al., 2008; Mathur et al., 2010; Rameson et al., 2011; Shen and Han, 2012; Singer et al., 2006; Sommer et al., 2010; Yo…”

“Punishers often feel angry at being cheated (e.g., Pillutla and Murnighan, 1996; Ben-Shakhar et al., 2007) and people (men, at least) will show neurological signatures of pleasure when seeing someone who cheated them apparently receive a painful stimulus (Singer et al., 2006).”

“It has also been reported that women are more empathic in conflict-resolution situations, and that men manifest a lower activation of brain areas related to empathy when they observe a painful stimulus received by a person known to be unfair (de Wied, Branje, & Meeus, 2007; Singer et al., 2006).”

“In the case of conflict, network theory suggests the importance of building ‘‘bridging’’ social capital among members of different groups (Adler and Kwon 2002) in order to facilitate better understanding of diverse viewpoints and empathy (Singer et al. 2006).”

“…studies have shown activation of medial prefrontal areas in response to pain observation, including the anterior cingulate (ACC), supplementary motor area (SMA) and presupplementary motor area (pre-SMA) (Morrison et al., 2004, Singer et al., 2004, 2006; Jackson et al., 2005; Lamm et al., 2007).”

“…of complex social interactions that have been amplified in the human lineage, such as increased empathy, cooperation, understanding of other individual’s mental states, and language (Singer et al., 2004, 2006, 2009; Rilling et al., 2008; Ackermann and Riecker, 2010; Rilling and Sanfey, 2011).”

“In our study, the only gender difference we found was more malevolent reciprocity of male participants towards an unfair interaction partner, which could reflect a greater inclination of males to punish unfair behaviour (Singer et al., 2006).”

“The gender difference in empathy toward friends and enemies found in the present study appears to be at odds with results reported by Singer et al. (2006). Using fMRI, they found that women and men showed an equally strong empathic response in pain-related areas of the brain when they saw a “fair” confederate receive…”

“…schadenfreude is more likely to arise when people deserve their misfortunes (Feather, 2008; Feather & Naim, 2005; Feather & Sherman, 2002; Portmann, 2000; Singer et al., 2006; van Dijk et al., 2005) and when misfortunes happen to people who are disliked (Hareli & Weiner, 2002; Heider, 1958).”

“Research has demonstrated that the insula is more active when participants witness a close other experience a negative event, specifically a mild shock (Singer et al., 2006).”

“Consistent with our first hypothesis, we found that when participants watched a friend be rejected their brain activation showed striking overlap with regions activated previously for the actual experience of social rejection, as well as for actual and empathic experiences of physical pain (Eisenberger et al, 2003 ; S…”

“Furthermore, recent imaging studies have found that social and contextual factors can also regulate empathic neural responses to others’ pain [10,11,24–28], including race of the target person [29–34].”

“Relevant to this issue is an fMRI study demonstrating that empathic responses to the pain of others’ are dramatically lower for unfair than fair individuals (Singer et al., 2006).”

“The right anterior insula has indeed been typically associated with empathy (Singer et al. 2004, 2006 ; Singer & Lamm, 2009; Lamm & Singer, 2010).”

“The nucleus accumbens was activated instead of empathy-related region like the anterior cingulate cortex (ACC) or anterior insula (AI) when seeing the betrayers suffering from pain (Singer et al., 2006).”

“, [52,53]).”

“…" (e.g., Avenanti et al., 2010; Botvinick et al., 2005; Cheng et al., 2007; Gu and Han, 2007; Jackson et al., 2005; Lamm et al., 2010), overlapping cortical structures that include affective areas (e.g., Lamm et al., 2007; Singer et al., 2004, 2006) and sensory areas (e.g., Avenanti et al., 2006).”

“…(BOLD) signal recorded using functional magnetic resonance imaging (fMRI) [Jackson et al., 2005, 2006; Morrison et al., 2004; Saarela et al., 2007; Singer et al., 2004, 2006], the amplitudes of event-related potentials (ERPs) [Fan and Han, 2008; Han et al., 2008], the nonphase-locked…”

“Moreover, empathic responses can be modulated by the subjective attitude held toward suffering individuals [7], as well as by personal experience [8].”

“…studies have shown that brain reward regions, including the caudate nucleus and related structures in the ventral striatum, are activated when subjects successfully punish others who have previously treated them unfairly in a trust game or a PD game (de Quervain et al., 2004; Singer et al., 2006).”

“…a loved one leads to more activity in insula and cingulate regions (Cheng, Chen, Lin, Chou, & Decety, 2010; Singer et al., 2004), while pain experienced by a disliked individual or group leads to less activity in these regions (Hein, Silani, Preuschoff, Batson, & Singer, 2010; Singer et al., 2006).”

“Moreover, empathy may be modulated by the interpersonal context in which individuals are engaged (Lanzetta and Englis, 1989; Singer et al., 2006) and so the lack of interaction with the demonstrator in the video replay group may have also influenced the relationship between empathy and placebo…”

“As reviewed (see Figure 1), the empathic event may influence the likelihood of empathizing [69] and the power of empathic responses [20].”

“Both male and female participants recruited empathy-related AI/FO and ACC activity when they observed fair players in pain, which was positively correlated with self-reported empathy.(129) However, male participants recruited reduced empathy-related AI/FO activity and increased rewardrelated VS activity when they obse…”

“cover story with an unfair ultimatum game offer allowed us to create an antagonist as the third player (Braams et al., 2013; Sanfey et al., 2003; Singer et al., 2006).”

“It has also been demonstrated that the anterior insula plays a crucial role for empathic experience related to pain (Singer et al., 2004, 2006).”

“Outgroup chimpanzees possibly elicited a hostile response, which interfered with empathybased engagement [49], whereas the gelada baboons were viewed as a socially meaningless stimulus.”

“…others in pain and may be linked to activation in insula and sensorimotor areas (Avenanti et al., 2009; Betti et al., 2009; Cheng et al., 2008; Singer et al., 2004, 2006), whereas cognitive empathy (PT) might be involved when watching others being nonpainfully touched, associated with…”

“, 1987; Davis, 1996; Preston and de Waal, 2002; Decety and Jackson, 2004), even to the extent that not only the direct observation of another person’s pain (Jackson et al., 2006; Singer et al., 2006) but also the perception of a signal that pain is being experienced is suffi cient to induce empathy-related brain activ…”

“role in the response to othersʼ outcomes (Cooper, Kreps, Wiebe, Pirkl, & Knutson, 2010; Mobbs et al., 2009; Singer et al., 2006; Delgado, Frank, & Phelps, 2005), and varia-”