Daytime and season do not affect reinforcement learning capacity in a response time adjustment task.

Seasonal and circadian rhythms have a broad impact on physiological aspects, such as dopamine neurotransmission, and may be involved in the etiology of mood disorders. Considering this, studies on the influence of season and daytime on cognitive function are rare. The present study aimed to assess the impact of seasonal and diurnal effects on the ability to maximize reward outcomes by optimizing response times adaptively. For this purpose, a reward-based learning task that required an adaptation of response time to either a fast or a slow response was used. Eighty German participants (mean age ± SD = 21.86 ± 1.89 years, 41 women) were examined twice, in the morning and in the evening. Half of the participants were tested during the summer, while the other half performed the test in the winter. No impact of daytime, season or of the external factors photoperiodicity and temperature on reinforcement learning could be found. However, a generally slower response speed in the morning compared to the evening appeared. Previously conducted tasks could not display behavioral differences in both times of season and daytime, although neurophysiological findings suggest it.

Endogenous testosterone correlates with parochial altruism in relation to costly punishment in different social settings.

Testosterone plays a key role in shaping human social behavior. Recent findings have linked testosterone to altruistic behavior in economic decision tasks depending on group membership and intergroup competition. The preferential treatment of ingroup members, while aggression and discrimination is directed towards outgroup members, has been referred to as parochial altruism. Here we investigated in two consecutive studies, whether testosterone is associated with parochial altruism depending on individual tendency for costly punishment. In the first study, 61 men performed a single-shot ultimatum game (UG) in a minimal group context, in which they interacted with members of an ingroup and outgroup. In the second study, 34 men performed a single-shot UG in a more realistic group context, in which they responded to the proposals of supporters of six political parties during the German election year 2017. Political parties varied in their social distance to the participants' favorite party as indicated by an individual ranking. Participants of study 2 also performed a cued recall task, in which they had to decide whether they had already encountered a face during the previous UG (old-new decision). In order to make the UG data of study 2 most comparable to the data of study 1, the rejection rates of several parties were combined according to the social distance ranking they achieved. Parties ranked 1 to 3 formed the relatively close and favored 'ingroup' that shared similar political values with the participant (e.g., left wing parties), while the 'outgroup' consisted of parties ranked from 4 to 6 with more distant or even antagonistic political views (e.g., conservative to right wing parties). In both studies, results showed a parochial pattern with higher rejection rates made in response to outgroup compared to ingroup offers. Interestingly, across studies higher salivary testosterone was associated with higher rejection rates related to unfair outgroup offers in comparison to the unfair offers made by ingroup members. The present findings suggest that latent intergroup biases during decision-making may be positively related to endogenous testosterone. Similar to previous evidence that already indicated a role of testosterone in shaping male parochial altruism in male soccer fans, these data underscore the general, yet rather subtle role of male testosterone also in other social settings.

DAT1-Genotype and Menstrual Cycle, but Not Hormonal Contraception, Modulate Reinforcement Learning: Preliminary Evidence.

Hormone by genotype interactions have been widely ignored by cognitive neuroscience. Yet, the dependence of cognitive performance on both baseline dopamine (DA) and current 17ß-estradiol (E2) level argues for their combined effect also in the context of reinforcement learning. Here, we assessed how the interaction between the natural rise of E2 in the late follicular phase (FP) and the 40 base-pair variable number tandem repeat polymorphism of the dopamine transporter (DAT1) affects reinforcement learning capacity. 30 women with a regular menstrual cycle performed a probabilistic feedback learning task twice during the early and late FP. In addition, 39 women, who took hormonal contraceptives (HC) to suppress natural ovulation, were tested during the "pill break" and the intake phase of HC. The present data show that DAT1-genotype may interact with transient hormonal state, but only in women with a natural menstrual cycle. We found that carriers of the 9-repeat allele (9RP) experienced a significant decrease in the ability to avoid punishment from early to late FP. Neither homozygote subjects of the 10RP allele, nor subjects from the HC group showed a change in behavior between phases. These data are consistent with neurobiological studies that found that rising E2 may reverse DA transporter function and could enhance DA efflux, which would in turn reduce punishment sensitivity particularly in subjects with a higher transporter density to begin with. Taken together, the present results, although based on a small sample, add to the growing understanding of the complex interplay between different physiological modulators of dopaminergic transmission. They may not only point out the necessity to control for hormonal state in behavioral genetic research, but may offer new starting points for studies in clinical settings.

Neural substrates of male parochial altruism are modulated by testosterone and behavioral strategy.

Parochial altruism refers to ingroup favoritism and outgroup hostility and has recently been linked to testosterone. Here, we investigated the neurobiological mechanism of parochial altruism in male soccer fans playing the ultimatum game (UG) against ingroup and outgroup members (i.e., fans of the favorite or of a rivalling team) using functional magnetic resonance imaging. Our results suggest that individual differences in altruistic tendency influence the tendency for parochialism. While altruistic subjects rejected unfair offers independent of team membership, the more self-oriented 'pro-selfs' displayed a stronger ingroup bias and rejected outgroup offers more often. However, during a second session that introduced a team competition the altruists adapted to this parochial pattern. Behavioral strategy was further characterized by dissociable and context-dependent correlations between endogenous testosterone and neural responses in the anterior insula and the ventromedial prefrontal cortex. In sum, the present findings indicate that parochial altruism is shaped by individual differences in testosterone and behavioral strategy. In that way the results are in line with evolutionary theories of both individual and group selection.

Endogenous Testosterone and Exogenous Oxytocin Modulate Attentional Processing of Infant Faces.

Evidence indicates that hormones modulate the intensity of maternal care. Oxytocin is known for its positive influence on maternal behavior and its important role for childbirth. In contrast, testosterone promotes egocentric choices and reduces empathy. Further, testosterone decreases during parenthood which could be an adaptation to increased parental investment. The present study investigated the interaction between testosterone and oxytocin in attentional control and their influence on attention to baby schema in women. Higher endogenous testosterone was expected to decrease selective attention to child portraits in a face-in-the-crowd-paradigm, while oxytocin was expected to counteract this effect. As predicted, women with higher salivary testosterone were slower in orienting attention to infant targets in the context of adult distractors. Interestingly, reaction times to infant and adult stimuli decreased after oxytocin administration, but only in women with high endogenous testosterone. These results suggest that oxytocin may counteract the adverse effects of testosterone on a central aspect of social behavior and maternal caretaking.

Testosterone is associated with cooperation during intergroup competition by enhancing parochial altruism.

The steroid hormone testosterone is widely associated with negative behavioral effects, such as aggression or dominance. However, recent studies applying economic exchange tasks revealed conflicting results. While some point to a prosocial effect of testosterone by increasing altruistic behavior, others report that testosterone promotes antisocial tendencies. Taking into account additional factors such as parochial altruism (i.e., ingroup favoritism and outgroup hostility) might help to explain this contradiction. First evidence for a link between testosterone and parochial altruism comes from recently reported data of male soccer fans playing the ultimatum game. In this study high levels of endogenous testosterone predicted increased altruistic punishment during outgroup interactions and at the same time heightened ingroup generosity. Here, we report findings of another experimental task, the prisoner's dilemma, applied in the same context to examine the role of testosterone on parochial tendencies in terms of cooperation. In this task, 50 male soccer fans were asked to decide whether or not they wanted to cooperate with partners marked as either fans of the subject's own favorite team (ingroup) or fans of other teams (outgroups). Our results show that high testosterone levels were associated with increased ingroup cooperation during intergroup competition. In addition, subjects displaying a high degree of parochialism during intergroup competition had significantly higher levels of testosterone than subjects who did not differentiate much between the different groups. In sum, the present data demonstrate that the behavioral effects of testosterone are not limited to aggressive and selfish tendencies but may imply prosocial aspects depending on the context. By this means, our results support the previously reported findings on testosterone-dependent intergroup bias and indicate that this social hormone might be an important factor driving parochial altruism.

How to be patient. The ability to wait for a reward depends on menstrual cycle phase and feedback-related activity.

Dopamine (DA) plays a major role in reinforcement learning with increases promoting reward sensitivity (Go learning) while decreases facilitate the avoidance of negative outcomes (NoGo learning). This is also reflected in adaptations of response time: higher levels of DA enhance speeding up to get a reward, whereas lower levels favor slowing down. The steroid hormones estradiol and progesterone have been shown to modulate dopaminergic tone. Here, we tested 14 women twice during their menstrual cycle, during the follicular (FP) and the luteal phase (LP), applying functional magnetic resonance imaging while they performed a feedback learning task. Subsequent behavioral testing assessed response time preferences with a clock task, in which subjects had to explore the optimal response time (RT) to maximize reward. In the FP subjects displayed a greater learning-related change of their RT than during the LP, when they were required to slow down. Final RTs in the slow condition were also predicted by feedback-related brain activation, but only in the FP. Increased activation of the inferior frontal junction and rostral cingulate zone was thereby predictive of slower and thus better adapted final RTs. Conversely, final RT was faster and less optimal for reward maximization if activation in the ventromedial prefrontal cortex was enhanced. These findings show that hormonal shifts across the menstrual cycle affect adaptation of response speed during reward acquisition with higher RT adjustment in the FP in the condition that requires slowing down. Since high estradiol levels during the FP increase synaptic DA levels, this conforms well to our hypothesis that estradiol supports Go learning at the expense of NoGo learning. Brain-behavior correlations further indicated that the compensatory capacity to counteract the follicular Go bias may be linked to the ability to more effectively monitor action outcomes and suppress bottom-up reward desiring during feedback processing.

Does competition really bring out the worst? Testosterone, social distance and inter-male competition shape parochial altruism in human males.

Parochial altruism, defined as increased ingroup favoritism and heightened outgroup hostility, is a widespread feature of human societies that affects altruistic cooperation and punishment behavior, particularly in intergroup conflicts. Humans tend to protect fellow group members and fight against outsiders, even at substantial costs for themselves. Testosterone modulates responses to competition and social threat, but its exact role in the context of parochial altruism remains controversial. Here, we investigated how testosterone influences altruistic punishment tendencies in the presence of an intergroup competition. Fifty male soccer fans played an ultimatum game (UG), in which they faced anonymous proposers that could either be a fan of the same soccer team (ingroup) or were fans of other teams (outgroups) that differed in the degree of social distance and enmity to the ingroup. The UG was played in two contexts with varying degrees of intergroup rivalry. Our data show that unfair offers were rejected more frequently than fair proposals and the frequency of altruistic punishment increased with increasing social distance to the outgroups. Adding an intergroup competition led to a further escalation of outgroup hostility and reduced punishment of unfair ingroup members. High testosterone levels were associated with a relatively increased ingroup favoritism and also a change towards enhanced outgroup hostility in the intergroup competition. High testosterone concentrations further predicted increased proposer generosity in interactions with the ingroup. Altogether, a significant relation between testosterone and parochial altruism could be demonstrated, but only in the presence of an intergroup competition. In human males, testosterone may promote group coherence in the face of external threat, even against the urge to selfishly maximize personal reward. In that way, our observation refutes the view that testosterone generally promotes antisocial behaviors and aggressive responses, but underlines its rather specific role in the fine-tuning of male social cognition.