Negative Reciprocity
A widely used behavioral task in the research of negative reciprocity is the ultimatum game (UG). In a typical UG, participants act as a responder and decide whether to accept a fair or unfair division of money suggested by a proposer (Sanfey et al., 2003). If the division is accepted, the money would be split as proposed; but if the division is rejected, neither one would receive anything. Participants commonly accepted offers when the divisions comply with the fairness norm (fair offers). Although participants could have obtained a certain amount of money by accepting the unfair offers, they rejected more offers (i.e., receiving nothing) as the extent of the proposer's norm violation increase (i.e., the offers become less fair), indicating the negative reciprocity and negative cost enforcement. In one line of research, neuroimaging studies using this task have consistently demonstrated the involvements of brain areas related to the initial evaluation of norm compliance/violation (Aoki et al., 2015; Feng et al., 2015; Gabay et al., 2014). Specifically, responders gave higher happiness ratings to more equal offers (Tabibnia et al., 2008); this observation was consistent with the greater responses in the vmPFC to fair (vs. unfair) offers, suggesting that the vmPFC contributed to the processing of the social rewards of fairness norm compliance (Baumgartner et al., 2011; Dawes et al., 2012; Tabibnia et al., 2008; Xiang et al., 2013). In contrast, compared with fair offers, unfair offers would activate the anterior insula, an area implicated in detecting norm violation (Cheng et al., 2017; Civai, 2013; Civai et al., 2012; Guo et al., 2013; Strobel et al., 2011; Xiang et al., 2013) or signaling emotional processing via representations of aversive internal states (Chang and Sanfey, 2011; Corradi-Dell'Acqua et al., 2012; Guo et al., 2013; Sanfey et al., 2003), and the amygdala, which was linked to signal negative emotional response to norm violation (Gospic et al., 2013; Haruno and Frith, 2010; Yu et al., 2014).
Another line of research revealed greater activations in brain regions related to the integration of social norms and economic self-interest in favor of flexible decision-making in the unfair condition as compared to the fair condition (Aoki et al., 2014; Feng et al., 2015; Gabay et al., 2014). Specifically, the unfairness-evoked aversive responses and the self-interest that would be obtained by acceptance contradict each other, resulting in a motivational conflict that was suggested to be monitored by the dACC (Fehr and Camerer, 2007; Sanfey et al., 2003). Neural evidence suggested two ways to resolve this conflict: first, the unfairness-evoked aversive responses may be suppressed, probably implemented by brain regions associated with emotion regulation such as the vlPFC and dmPFC, resulting in an increase in acceptance rates (Civai et al., 2012; Grecucci et al., 2012; Tabibnia et al., 2008). Second, the conflict may be resolved by inhibiting selfish motives to promote norm compliance; this would rely on the cognitive control functions in the right dlPFC (Knoch et al., 2006; Ruff et al., 2013; Zhu et al., 2014). In addition, it was shown that, as compared to the gain frame used in the traditional UG, participants were more likely to reject unfair offers in the loss frame, where the proposers proposed unfair offers to share the loss (Zhou and Wu, 2011). Neuroimaging data indicated that loss reduced the responsiveness of the dopamine system (ventral striatum) to fairness while enhancing the motivation to reject the offer. This process was complemented by increased responses of dlPFC to insultingly unfair offers (Guo et al., 2013; Wu et al., 2014).
Notably, the reciprocal behaviors in UG are based not only on the preference for fair outcomes (i.e., egalitarianism) but also on reciprocal considerations regarding the others' intentions (i.e., intention-based reciprocity) (Charness and Rabin, 2002; Dufwenberg and Kirchsteiger, 2004; Falk et al., 2003; Rabin, 1993; Zheng et al., 2014). For example, the same unfair offers are more likely to be accepted if the proposer demonstrates good intentions by choosing the inequitable division over an even more unfair division (Falk et al., 2003). This increase in acceptance rates is associated with activity in the anterior medial prefrontal cortex and the TPJ, implying that higher demands in moral mentalizing are required in social decision-making when the decision to reject could not be readily justified (Güroğlu et al., 2010). Moreover, a gradual shift in other-regarding preferences was observed from simple rule-based egalitarianism to complex intention-based reciprocity from early childhood to young adulthood (Sul et al., 2017). The preference shift was associated with cortical thinning of the dmPFC and posterior temporal cortex, which were involved in social inference as indicated by the meta-analytic reverse-inference analysis.
Moreover, Yu et al. (2015) investigated the neural substrates underlying the processing of both intention and consequence of the other's harm using an interactive game. In the task, participants interacted with anonymous co-players, who decided to deliver pain stimulation either to him/herself or to the paired participant to earn a monetary reward. In some cases, the decision was reversed by the computer. Unbeknownst to the co-player, the participant was then allowed to punish the co-player by reducing his/her monetary reward after seeing the co-player's intention. Behaviorally, the punishment was lower in the accidental condition (unintended harm relative to intended harm) but higher in the failed-attempt condition (intended no-harm relative to unintended no-harm). Neurally, the left amygdala was activated in conditions with blameworthy intention (i.e., intentional harm and failed attempt). The accidental (relative to intentional) harm activated the right TPJ and IFG, while the failed attempt (relative to genuine no-harm) activated the anterior insula and posterior IFG. Effective connectivity analysis revealed that in the unintentional conditions (i.e., accidental and failed attempt) the IFG received input from the TPJ and AI and sent regulatory signals to the amygdala. These findings demonstrate that the processing of intention may gate the emotional responses to transgression and regulate subsequent reactive punishment.