Empathy incites a stable prosocial decision bias.

Empathy toward suffering individuals serves as potent driver for prosocial behavior. However, it remains unclear whether prosociality induced by empathy for another person's pain persists once that person's suffering diminishes. To test this, participants underwent functional magnetic resonance imaging while performing a binary social decision task that involved allocation of points to themselves and another person. In block one, participants completed the task after witnessing frequent painful stimulation of the other person, and in block two, after observing low frequency of painful stimulation. Drift-diffusion modeling revealed an increased initial bias toward making prosocial decisions in the first block compared with baseline that persisted in the second block. These results were replicated in an independent behavioral study. An additional control study showed that this effect may be specific to empathy as stability was not evident when prosocial decisions were driven by a social norm such as reciprocity. Increased neural activation in dorsomedial prefrontal cortex was linked to empathic concern after witnessing frequent pain and to a general prosocial decision bias after witnessing rare pain. Altogether, our findings show that empathy for pain elicits a stable inclination toward making prosocial decisions even as their suffering diminishes.

Connected in Bad Times and in Good Times: Empathy Induces Stable Social Closeness.

Humans need social closeness to prosper. There is evidence that empathy can induce social closeness. However, it remains unclear how empathy-related social closeness is formed and how stable it is as time passes. We applied an acquisition-extinction paradigm combined with computational modeling and fMRI, to investigate the formation and stability of empathy-related social closeness. Female participants observed painful stimulation of another person with high probability (acquisition) and low probability (extinction) and rated their closeness to that person. The results of two independent studies showed increased social closeness in the acquisition block that resisted extinction in the extinction block. Providing insights into underlying mechanisms, reinforcement learning modeling revealed that the formation of social closeness is based on a learning signal (prediction error) generated from observing another's pain, whereas maintaining social closeness is based on a learning signal generated from observing another's pain relief. The results of a reciprocity control study indicate that this feedback recalibration is specific to learning of empathy-related social closeness. On the neural level, the recalibration of the feedback signal was associated with neural responses in anterior insula and adjacent inferior frontal gyrus and the bilateral superior temporal sulcus/temporoparietal junction. Together, these findings show that empathy-related social closeness generated in bad times, that is, empathy with the misfortune of another person, transfers to good times and thus may form one important basis for stable social relationships.

Can monetary incentives overturn fairness-based decisions?

Fairness norms and resulting behaviours are an important prerequisite for cooperation in human societies. At the same time, financial incentives are commonly used to motivate social behaviours, yet it remains unclear how financial incentives affect fairness-based behaviours. Combining a decision paradigm from behavioural economics with hierarchical drift-diffusion modelling, we investigated the effect of different financial incentives on two types of fairness-based decisions in four experimental groups. In two groups, participants divided points between themselves and a disadvantaged person, inciting fairness-based compensation behaviour, in two other groups they divided points between themselves and a fairness violator, inciting fairness-based punishment behaviour. In addition, each group received financial incentives that were either aligned or in conflict with the respective fairness-based behaviour. This design allowed us to directly investigate how different incentives shape the cognitive mechanism of fairness-based decisions and whether these effects are comparable across different fairness domains (fairness-based punishment versus fairness-based compensation). Results showed that offering conflicting incentives diminished fairness-congruent decision behaviour and rendered the fairness-congruent decision process less efficient. These findings demonstrate that financial incentives can undermine fairness-based behaviour, and thus are relevant for the development of incentive schemes aimed at fostering cooperative behaviour.

Financial incentives facilitate stronger neural computation of prosocial decisions in lower empathic adult females.

Financial incentives are commonly used to motivate behaviors. However, there is also evidence that incentives can impede the behavior they are supposed to foster, for example, documented by a decrease in blood donations if a financial incentive is offered. Based on these findings, previous studies assumed that prosocial motivation is shaped by incentives. However, so far, there is no direct evidence showing an interaction between financial incentives and a specific prosocial motive. Combining drift-diffusion modeling and fMRI, we investigated the effect of financial incentives on empathy, i.e., one of the key motives driving prosocial decisions. In the empathy-alone condition, participants made prosocial decisions based on empathy. In the empathy-bonus condition, they were offered a financial bonus for prosocial decisions, in addition to empathy induction. On average, the bonus enhanced the information accumulation in empathy-based decisions. On the neural level, this enhancement was related to the anterior insula, the same region that also correlated with empathy ratings. Moreover, the effect of the financial incentive on anterior insula activation was stronger the lower a person scored on empathy. These findings show that financial incentives enhance prosocial motivation in the absence of empathy.

The neural computation of human prosocial choices in complex motivational states.

Motives motivate human behavior. Most behaviors are driven by more than one motive, yet it is unclear how different motives interact and how such motive combinations affect the neural computation of the behaviors they drive. To answer this question, we induced two prosocial motives simultaneously (multi-motive condition) and separately (single motive conditions). After the different motive inductions, participants performed the same choice task in which they allocated points in favor of the other person (prosocial choice) or in favor of themselves (egoistic choice). We used fMRI to assess prosocial choice-related brain responses and drift diffusion modeling to specify how motive combinations affect individual components of the choice process. Our results showed that the combination of the two motives in the multi-motive condition increased participants' choice biases prior to the behavior itself. On the neural level, these changes in initial prosocial bias were associated with neural responses in the bilateral dorsal striatum. In contrast, the efficiency of the prosocial decision process was comparable between the multi-motive and the single-motive conditions. These findings provide insights into the computation of prosocial choices in complex motivational states, the motivational setting that drives most human behaviors.

Frequency of helping friends and helping strangers is explained by different neural signatures.

Acts of helping friends and strangers are part of everyday life. However, people vary significantly with respect to how often they help others and with respect to whom they actually help on a day-to-day basis. Despite everyday helping being so pervasive, these individual differences are poorly understood. Here, we used source-localized resting electroencephalography to measure objective and stable individual differences in neural baseline activation in combination with an ecologically valid method that allows assessment of helping behavior in the field. Results revealed that neural baseline activation in the right dorsolateral prefrontal cortex (DLPFC) - a brain region associated with self-control and strategic social behavior - predicts the daily frequency of helping friends, whereas the daily frequency of helping strangers was predicted by neural baseline activation in the dorsomedial prefrontal cortex (DMPFC) - a brain region associated with social cognition processes. These findings offer evidence that distinct neural signatures and associated psychological and cognitive processes may underlie the propensity to help friends and strangers in daily life.

Distinct Patterns of Cognitive Conflict Dynamics in Promise Keepers and Promise Breakers.

On a daily basis, we see how different people can be in keeping or breaking a given promise. However, we know very little about the cognitive conflict dynamics that underlie the decision to keep or break a promise and whether this is shaped by inter-individual variability. In order to fill this gap, we applied an ecologically valid promise decision task with real monetary consequences for all involved interaction partners and used mouse tracking to identify the dynamic, on-line cognitive processes that underlie the decision to keep or break a promise. Our findings revealed that on average, the process of breaking a promise is associated with largely curved mouse trajectories, while the process of keeping a promise was not, indicating that breaking a promise is associated with a larger conflict. Interestingly, however, this conflict pattern was strongly shaped by individual differences. Individuals who always kept their promises did not show any signs of conflict (i.e., straight mouse trajectories), indicating that they were not tempted by the monetary benefits associated with breaking the promise. In contrast, individuals who did not always keep their promise exhibited a large conflict (i.e., curved mouse trajectories), irrespective of whether they broke or kept their promise. A possible interpretation of these findings is that these individuals were always tempted by the unchosen decision option - the desire to act in a fair manner when breaking the promise and the monetary benefits when keeping the promise. This study provides the first piece of evidence that there are substantial inter-individual differences in cognitive conflict dynamics that underlie the decision to keep or break promises and that mouse tracking is able to illuminate important insights into individual differences in complex human's decision processes.

Structural Differences in Insular Cortex Reflect Vicarious Injustice Sensitivity.

Sensitivity to injustice inflicted on others is a strong motivator of human social behavior. There are, however, enormous individual differences in vicarious injustice sensitivity. Some people are strongly affected when witnessing injustice, while others barely notice it, but the factors behind this heterogeneity are poorly understood. Here we examine the neuroanatomical basis of these differences using voxel-based morphometry and Freesurfer image analysis suite. Whole brain corrected analyses show that a person's propensity to be vicariously affected by injustice to others is reflected by the gray matter volume and thickness of the bilateral mid insular cortex. The larger a person's gray matter volume and thickness of the mid insula, the higher that person's sensitivity to injustice experienced by others. These findings show that the individual neuroanatomy of the mid insular cortex captures a person's predisposition to be vicariously affected by injustice, and thus adds a novel aspect to previous functional work that has linked this region to the processing of transient vicarious states.

Serotonin and molecular neuroimaging in humans using PET.

The serotonergic system is one of the most important modulatory neurotransmitter systems in the human brain. It plays a central role in major physiological processes and is implicated in a number of psychiatric disorders. Along with the dopaminergic system, it is also one of the phylogenetically oldest human neurotransmitter systems and one of the most diverse, with 14 different receptors identified up to this day, many of whose function remains to be understood. The system's functioning is even more diverse than the number of its receptors, since each is implicated in a number of different processes. This review aims at illustrating the distribution and summarizing the main functions of the serotonin (5-hydroxytryptamin, 5-HT) receptors as well as the serotonin transporter (SERT, 5-HTT), the vesicular monoamine transporter 2, monoamine oxidase type A and 5-HT synthesis in the human brain. Recent advances in in vivo quantification of these different receptors and enzymes that are part of the serotonergic system using positron emission tomography are described.