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Testosterone is a steroid hormone which the male testes and, to a lesser extent, the female ovaries secrete in mammals. Testosterone is present in the central nervous system throughout life [...], Both biosociological and psychological models, as well as animal research, suggest that testosterone has a key role in social interactions (1-7). Evidence from animal studies in rodents shows that testosterone causes aggressive behaviour towards conspecifics (7). Folk wisdom generalizes and adapts these findings to humans, suggesting that testosterone induces antisocial, egoistic, or even aggressive human behaviours. However, many researchers have questioned this folk hypothesis (1-6), arguing that testosterone is primarily involved in status-related behaviours in challenging social interactions, but causal evidence that discriminates between these views is sparse. Here we show that the sublingual administration of a single dose of testosterone in women causes a substantial increase in fair bargaining behaviour, thereby reducing bargaining conflicts and increasing the efficiency of social interactions. However, subjects who believed that they received testosterone--regardless of whether they actually received it or not--behaved much more unfairly than those who believed that they were treated with placebo. Thus, the folk hypothesis seems to generate a strong negative association between subjects' beliefs and the fairness of their offers, even though testosterone administration actually causes a substantial increase in the frequency of fair bargaining offers in our experiment.

Hair steroid analysis is increasingly recognized for its ability to capture cumulative hormone secretion, thought to reflect an individual's response to long-term environmental conditions. Despite its growing use, the influence of single, isolated events on hair steroid concentrations remains underexplored. Our study therefore examined the effects of a single-dose transdermal testosterone administration (150 mg) and acute laboratory stress induction on hair testosterone and cortisol levels in a randomized, between-subject, placebo-controlled, and double-blind design. Liquid chromatography-tandem mass spectrometry analysis revealed a significant increase in hair testosterone levels seven weeks post-testosterone administration, underscoring the efficacy of hair analysis in detecting single-use hormone intake. Moreover, we observed a significant elevation in hair cortisol levels seven weeks post-exposure to the laboratory somatic stressor (Cold pressor test), highlighting the efficacy of hair analysis in capturing experimentally induced hormonal responses. The exploration of contextual factors revealed that individuals in committed relationships exhibited lower levels of both hair cortisol and testosterone compared to those who were single or in uncommitted relationships. Our study provides new insights into the sensitivity of hair analysis for detecting hormonal changes following single-dose hormone administration and experimentally induced short-term stress events. The exploratory findings emphasize the importance of individual contextual factors in influencing hair hormone concentrations and lay the groundwork for further investigation into the dynamics of cumulative hair hormone measurements., Competing Interests: Declaration of Competing Interest The authors report no conflicts of interest., (Copyright © 2024. Published by Elsevier Ltd.)

Rewards are a broad category of stimuli inducing approach behavior to aid survival. Extensive evidence from animal research has shown that wanting (the motivation to pursue a reward) and liking (the pleasure associated with its consumption) are mostly regulated by dopaminergic and opioidergic activity in dedicated brain areas. However, less is known about the neuroanatomy of dopaminergic and opioidergic regulation of reward processing in humans, especially when considering different types of rewards (i.e., social and nonsocial). To fill this gap of knowledge, we combined dopaminergic and opioidergic antagonism (via amisulpride and naltrexone administration) with functional neuroimaging to investigate the neurochemical and neuroanatomical bases of wanting and liking of matched nonsocial (food) and social (interpersonal touch) rewards, using a randomized, between-subject, placebo-controlled, double-blind design. While no drug effect was observed at the behavioral level, brain activity was modulated by the administered compounds. In particular, opioid antagonism, compared to placebo, reduced activity in the medial orbitofrontal cortex during consumption of the most valued social and nonsocial rewards. Dopamine antagonism, however, had no clear effects on brain activity in response to reward anticipation. These findings provide insights into the neurobiology of human reward processing and suggest a similar opioidergic regulation of the neural responses to social and nonsocial reward consumption., (© 2024 The Authors. Human Brain Mapping published by Wiley Periodicals LLC.)

Humans are strategically more prosocial when their actions are being watched by others than when they act alone. Using a psychopharmacogenetic approach, we investigated the endocrinological and computational mechanisms of such audience-driven prosociality. One hundred and ninety-two male participants received either a single dose of testosterone (150 mg) or a placebo and performed a prosocial and self-benefitting reinforcement learning task. Crucially, the task was performed either in private or when being watched. Rival theories suggest that the hormone might either diminish or strengthen audience-dependent prosociality. We show that exogenous testosterone fully eliminated strategic, i.e., feigned, prosociality and thus decreased submission to audience expectations. We next performed reinforcement-learning drift-diffusion computational modeling to elucidate which latent aspects of decision-making testosterone acted on. The modeling revealed that testosterone compared to placebo did not deteriorate reinforcement learning per se. Rather, when being watched, the hormone altered the degree to which the learned information on choice value translated to action selection. Taken together, our study provides novel evidence of testosterone's effects on implicit reward processing, through which it counteracts conformity and deceptive reputation strategies., (© 2023. The Author(s).)

The ability to learn about other people is crucial for human social functioning. Dopamine has been proposed to regulate the precision of beliefs, but direct behavioural evidence of this is lacking. In this study, we investigate how a high dose of the D2/D3 dopamine receptor antagonist sulpiride impacts learning about other people's prosocial attitudes in a repeated Trust game. Using a Bayesian model of belief updating, we show that in a sample of 76 male participants sulpiride increases the volatility of beliefs, which leads to higher precision weights on prediction errors. This effect is driven by participants with genetically conferred higher dopamine availability (Taq1a polymorphism) and remains even after controlling for working memory performance. Higher precision weights are reflected in higher reciprocal behaviour in the repeated Trust game but not in single-round Trust games. Our data provide evidence that the D2 receptors are pivotal in regulating prediction error-driven belief updating in a social context., (© 2023. The Author(s).)

Human behaviour requires flexible arbitration between actions we do out of habit and actions that are directed towards a specific goal. Drugs that target opioid and dopamine receptors are notorious for inducing maladaptive habitual drug consumption; yet, how the opioidergic and dopaminergic neurotransmitter systems contribute to the arbitration between habitual and goal-directed behaviour is poorly understood. By combining pharmacological challenges with a well-established decision-making task and a novel computational model, we show that the administration of the dopamine D2/3 receptor antagonist amisulpride led to an increase in goal-directed or 'model-based' relative to habitual or 'model-free' behaviour, whereas the non-selective opioid receptor antagonist naltrexone had no appreciable effect. The effect of amisulpride on model-based/model-free behaviour did not scale with drug serum levels in the blood. Furthermore, participants with higher amisulpride serum levels showed higher explorative behaviour. These findings highlight the distinct functional contributions of dopamine and opioid receptors to goal-directed and habitual behaviour and support the notion that even small doses of amisulpride promote flexible application of cognitive control., Competing Interests: NM, SK, CM, CG, IG, MW, CE, CL, GS, CM No competing interests declared, (© 2022, Mikus et al.)

Art, as a prestigious cultural commodity, concerns aesthetic and monetary values, personal tastes, and social reputation in various social contexts-all of which are reflected in choices concerning our liking, or in other contexts, our actual willingness-to-pay for artworks. But, how do these different aspects interact in regard to the concept of social reputation and our private versus social selves, which appear to be essentially intervening, and potentially conflicting, factors driving choice? In our study, we investigated liking and willingness-to-pay choices using-in art research-a novel, forced-choice paradigm. Participants (N = 123) made choices from artwork-triplets presented with opposing artistic quality and monetary value-labeling, thereby creating ambiguous choice situations. Choices were made in either private or in social/public contexts, in which participants were made to believe that either art-pricing or art-making experts were watching their selections. A multi-method design with eye-tracking, neuroendocrinology (testosterone, cortisol), and motivational factors complemented the behavioral choice analysis. Results showed that artworks, of which participants were told were of high artistic value were more often liked and those of high monetary-value received more willingness-to-pay choices. However, while willingness-to-pay was significantly affected by the presumed observation of art-pricing experts, liking selections did not differ between private/public contexts. Liking choices, compared to willingness-to-pay, were also better predicted by eye movement patterns. Whereas, hormone levels had a stronger relation with monetary aspects (willingness-to-pay/ art-pricing expert). This was further confirmed by motivational factors representative for reputation seeking behavior. Our study points to an unexplored terrain highlighting the linkage of social reputation mechanisms and its impact on choice behavior with a ubiquitous commodity, art., Competing Interests: The authors have declared that no competing interests exist.

The sex hormone estradiol is hypothesized to play a key role in human cognition, and reward processing specifically, via increased dopamine D1-receptor signalling. However, the effect of estradiol on reward processing in men has never been established. To fill this gap, we performed a double-blind placebo-controlled study in which men (N = 100) received either a single dose of estradiol (2 mg) or a placebo. Subjects performed a probabilistic reinforcement learning task where they had to choose between two options with varying reward probabilities to maximize monetary reward. Results showed that estradiol administration increased reward sensitivity compared to placebo. This effect was observed in subjects' choices, how much weight they assigned to their previous choices, and subjective reports about the reward probabilities. Furthermore, effects of estradiol were moderated by reward sensitivity, as measured through the BIS/BAS questionnaire. Using reinforcement learning models, we found that behavioral effects of estradiol were reflected in increased learning rates. These results demonstrate a causal role of estradiol within the framework of reinforcement learning, by enhancing reward sensitivity and learning. Furthermore, they provide preliminary evidence for dopamine-related genetic variants moderating the effect of estradiol on reward processing., (Copyright © 2021 Elsevier Inc. All rights reserved.)

Recent research suggests that when we lack a sense of control, we are prone to motivational failures and early quitting in competitions. Testosterone, on the other hand, is thought to boost competitiveness. Here we investigate the interaction between these factors, testing the testosterone's potential to enhance persistence in a competition against a stronger opponent, depending on experimentally manipulated perceived control. Healthy participants were administered a single dose of testosterone or placebo. They first underwent a task designed to either induce low or high perceived control and then entered a costly competition against a progressively stronger opponent that they could quit at any time. In the placebo group, men with low perceived control quitted twice as early as those with high perceived control. Testosterone countered this effect, making individuals with low control persist in the competition for as long as those with high perceived control, and did so also despite raising participants' explicit awareness of the opponents' advantage. This psychoendocrinological effect was not modulated by basal cortisol levels, CAG repeat polymorphism of the androgen receptor gene, or trait dominance. Our results provide the first causal evidence that testosterone promotes competitive persistence in humans and demonstrate that this effect depends on the psychological state elicited prior to the competition, broadening our understanding of the complex relationships between testosterone and social behaviors., (Copyright © 2021 The Authors. Published by Elsevier Ltd.. All rights reserved.)

The observation of animal orofacial and behavioral reactions has played a fundamental role in research on reward but is seldom assessed in humans. Healthy volunteers (N = 131) received 400 mg of the dopaminergic antagonist amisulpride, 50 mg of the opioidergic antagonist naltrexone, or placebo. Subjective ratings, physical effort, and facial reactions to matched primary social (affective touch) and nonsocial (food) rewards were assessed. Both drugs resulted in lower physical effort and greater negative facial reactions during reward anticipation, especially of food rewards. Only opioidergic manipulation through naltrexone led to a reduction in positive facial reactions to liked rewards during reward consumption. Subjective ratings of wanting and liking were not modulated by either drug. Results suggest that facial reactions during anticipated and experienced pleasure rely on partly different neurochemical systems, and also that the neurochemical bases for food and touch rewards are not identical., Competing Interests: SK, SG, CM, PS, IG, MW, CE, GS No competing interests declared, (© 2020, Korb et al.)

Persisting even when the rewards of continued effort are fading is essential for achieving long-term goals, skills, and good health, alike. Yet, we often quit when things get hard. Here, we tested whether augmenting the feeling of control through external measures increases persistence under such discouraging circumstances. In two laboratory experiments, we first induced illusory control by manipulating the base-rate of positive outcomes and then tested the effect of this elevation of participants' perceived control upon their persistence under diminishing returns and in a competition against a stronger opponent. Induced illusory control significantly enhanced people's persistence in both of these motivationally challenging situations. Our findings demonstrate that motivation is dependent upon perceived, rather than objective, control, and reveal that this can be leveraged to counteract quitting behavior when things get hard, for instance in rehabilitation, physical activity interventions, or other training settings.

Higher testosterone levels in males have previously been linked to decreased stress reactivity, but in other cases, testosterone has been reported to increase the stress response. We addressed these inconsistencies in a placebo-controlled single-dose testosterone administration study, in which 120 male participants were randomly assigned to undergo a cold-pressor test (CPT, a non-social somatic stressor), a socially evaluated cold-pressor test (SECPT, a social-somatic stressor), or a lukewarm water test (LWT, a non-stressful control condition). Throughout the experiment, blood pressure and interbeat intervals were measured continuously, and saliva samples for hormonal analyses were taken repeatedly at predefined time points. When comparing the groups treated with placebo, the SECPT elicited a larger increase in the systolic blood pressure than CPT, in agreement with previous studies. However, testosterone administration altered this pattern. Compared to placebo, testosterone increased systolic blood pressure during the CPT, whereas the opposite effect was found during the SECPT. Cortisol reactivity was not affected by testosterone administration. The CAG repeat polymorphism of the androgen receptor gene was unrelated to the effects of testosterone on the stress response, but it was correlated with blood pressure across the whole sample. Our findings demonstrate that testosterone's effects on the stress response are dependent on the social context. Testosterone's ability to flexibly influence the response to stressors may be an important mechanism through which the hormone promotes adaptive behavior. Our results are also in line with research showing that testosterone decreases social anxiety and suggest it may help to modulate the effects of stress in socially challenging situations., Competing Interests: Declaration of Competing Interest None., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Whether cognitive, motivational and hedonic aspects of reward anticipation and consumption can be reliably assessed with explicit and implicit measures, and if different motivational (decision utility) and hedonic (experienced utility) processes get recruited by distinct reward types, remain partly unsolved questions that are relevant for theories of social and non-social decision-making. We investigated these topics using a novel experimental paradigm, including carefully matched social and nonsocial rewards, and by focusing on facial responses. Facial expressions are indeed an often-cited implicit measure of rewards' hedonic impact. For example, food rewards elicit powerful facial responses - characterized by lip smacking, tongue protrusion, and relaxation of the middle face - in human newborns, juvenile monkeys, and adult rats. The same stimuli elicit more nuanced facial reactions in adult humans, which can be best captured with facial electromyography (fEMG). However, little is known about facial expressions preceding reward consumption, reflecting the motivation to obtain and possibly the expected pleasantness of a reward, and whether similar facial expressions are elicited by different types of rewards. To investigate these questions, a novel within-subject experimental paradigm was developed. During the anticipation and consumption of social (affective touch) and nonsocial (food) rewards, explicit (ratings of wanting and liking, physical effort) and implicit (fEMG) measures of wanting and liking were taken in 43 healthy adult participants. Reduced activation of the Corrugator Supercilii (CS) muscle (reflecting less frowning and indicating greater positive response) was found in trials with higher wanting and effort during the anticipation of food rewards, as well as in trials with higher liking and effort during the consumption of food rewards. The CS muscle is thus a sensitive measure of wanting and liking of food rewards both during their anticipation and consumption. Crucially, thanks to careful reward matching, these results cannot be explained by differences in subjective wanting, liking, or effort produced to obtain the two types of rewards. No significant modulation of the Zygomaticus Major (ZM) muscle was found for social or food rewards. Explorative analyses however indicated that the ZM may activate during the delivery of the most wanted touch, but not for the most wanted food. The absence of significant effects of social rewards on the activation of CS and ZM muscles are discussed in relation to the specifics of this innovative task comparing two types of matched rewards in the same participants. The present findings contribute to the understanding of the processes underlying motivational and hedonic aspects of rewards, and may therefore inform models of social and non-social decision-making., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Psychopathic offenders have a high propensity to violate social norms, as indicated for instance by their widespread lying and cheating behaviour. The reasons for their norm violations are not well understood, though, as they are able to recognise norms in a given situation and also punish norm violators. In this study, we investigated whether psychopathic offenders would violate fairness norms during a repeated trust game because of increased profit-maximising concerns. We measured back-transfer decisions in the repeated trust game, and affective arousal by means of skin conductance responses (SCR) in violent offenders with varying degrees of psychopathy, and non-offenders with low-trait psychopathy. Psychopathy in offenders was measured with the Psychopathy Checklist-Revised (PCL-R). In the task, a participant and an interaction partner entrusted each other money for multiple rounds with the goal to earn as much money as possible. Fairness norm violations were positively associated with Factor 2 scores (the lifestyle/anti-social psychopathy subscale) of the PCL-R, but this was not accompanied by clear profit-maximising behaviour. In addition, anticipatory arousal to self-advantageous decisions was higher in all offenders, independent of their degree of psychopathy, compared with non-offenders. The results of our study widen our understanding of social decision-making in psychopathy. They also suggest treatment possibilities in offenders scoring high on Factor 2, targeting empathic concern and related prosocial intentions to overcome norm-violating behaviour.