Human Prosocial Preferences Are Related to Slow-Wave Activity in Sleep.

Prosocial behavior is crucial for the smooth functioning of the society. Yet, individuals differ vastly in the propensity to behave prosocially. Here, we try to explain these individual differences under normal sleep conditions without any experimental modulation of sleep. Using a portable high-density EEG, we measured the sleep data in 54 healthy adults (28 females) during a normal night's sleep at the participants' homes. To capture prosocial preferences, participants played an incentivized public goods game in which they faced real monetary consequences. The whole-brain analyses showed that a higher relative slow-wave activity (SWA, an indicator of sleep depth) in a cluster of electrodes over the right temporoparietal junction (TPJ) was associated with increased prosocial preferences. Source localization and current source density analyses further support these findings. Recent sleep deprivation studies imply that sleeping enough makes us more prosocial; the present findings suggest that it is not only sleep duration, but particularly sufficient sleep depth in the TPJ that is positively related to prosociality. Because the TPJ plays a central role in social cognitive functions, we speculate that sleep depth in the TPJ, as reflected by relative SWA, might serve as a dispositional indicator of social cognition ability, which is reflected in prosocial preferences. These findings contribute to the emerging framework explaining the link between sleep and prosocial behavior by shedding light on the underlying mechanisms.

The path of dishonesty: identification of mental processes with electrical neuroimaging.

Much research finds that lying takes longer than truth-telling. Yet, the source of this response time difference remains elusive. Here, we assessed the spatiotemporal evolution of electrical brain activity during honesty and dishonesty in 150 participants using a sophisticated electrical neuroimaging approach-the microstate approach. This uniquely positioned us to identify and contrast the entire chain of mental processes involved during honesty and dishonesty. Specifically, we find that the response time difference is the result of an additional late-occurring mental process, unique to dishonest decisions, interrupting the antecedent mental processing. We suggest that this process inhibits the activation of the truth, thus permitting the execution of the lie. These results advance our understanding of dishonesty and clarify existing theories about the role of increased cognitive load. More broadly, we demonstrate the vast potential of our approach to illuminate the temporal organization of mental processes involved in decision-making.

Interindividual differences in intergenerational sustainable behavior are associated with cortical thickness of the dorsomedial and dorsolateral prefrontal cortex.

Intergenerational sustainability requires people of the present generation to make sacrifices today to benefit others of future generations (e.g. mitigating climate change, reducing public debt). Individuals vary greatly in their intergenerational sustainability, and the cognitive and neural sources of these interindividual differences are not yet well understood. We here combined neuroscientific and behavioral methods by assessing interindividual differences in cortical thickness and by using a common-pool resource paradigm with intergenerational contingencies. This enabled us to look for objective, stable, and trait-like neural markers of interindividual differences in consequential intergenerational behavior. We found that individuals behaving sustainably (vs. unsustainably) were marked by greater cortical thickness of the dorsomedial and dorsolateral prefrontal cortex. Given that these brain areas are involved in perspective-taking and self-control and supported by mediation analyses, we speculate that greater cortical thickness of these brain areas better enable individuals to take the perspective of future generations and to resist temptations to maximize personal benefits that incur costs for future generations. By meeting recent calls for the contribution of neuroscience to sustainability research, it is our hope that the present study advances the transdisciplinary understanding of interindividual differences in intergenerational sustainability.

Local slow-wave activity over the right prefrontal cortex reveals individual risk preferences.

In everyday life, we have to make decisions under varying degrees of risk. Even though previous research has shown that the manipulation of sleep affects risky decision-making, it remains unknown whether individual, temporally stable neural sleep characteristics relate to individual differences in risk preferences. Here, we collected sleep data under normal conditions in fifty-four healthy adults using a portable high-density EEG at participants' home. Whole-brain corrected for multiple testing, we found that lower slow-wave activity (SWA, an indicator of sleep depth) in a cluster of electrodes over the right prefrontal cortex (PFC) is associated with higher individual risk propensity. Importantly, the association between local sleep depth and risk preferences remained significant when controlling for total sleep time and for time spent in deep sleep, i.e., sleep stages N2 and N3. Moreover, the association between risk preferences and SWA over the right PFC was very similar in all sleep cycles. Because the right PFC plays a central role in cognitive control functions, we speculate that local sleep depth in this area, as reflected by SWA, might serve as a dispositional indicator of self-regulatory ability, which in turn reflects risk preferences.

Different Behavioral Types of Distributional Preferences Are Characterized by Distinct Neural Signatures.

There are many situations where resources are distributed between two parties and where the deciding party has information about the initial distribution and can change its outcome, for example, the allocation of budget for funds or bonuses, where the deciding party might have self-interested motives. Although the neural underpinnings of distributional preferences of resources have been extensively studied, it remains unclear if there are different types of distributional preferences and if these types underlie different disposing neural signatures. We used source-localized resting EEG in combination with a data-driven clustering approach to participants' behavior in a distribution game in order to disentangle the neural sources of the different types of distributional preferences. Our findings revealed four behavioral types: Maximizing types always changed initial distributions to maximize their personal outcomes, and compliant types always left initial distributions unchanged. Disadvantage-averse types only changed initial distributions if they received less than the other party did, and equalizing types primarily changed initial distributions to fair distributions. These behavioral types differed regarding neural baseline activation in the right inferior frontal gyrus. Maximizing and compliant types showed the highest baseline activation, followed by disadvantage-averse types and equalizing types. Furthermore, maximizing types showed significantly higher baseline activation in the left orbitofrontal cortex compared to compliant types. Taken together, our findings show that different types of distributional preferences are characterized by distinct neural signatures, which further imply differences in underlying psychological processes in decision-making.

Who initiates punishment, who joins punishment? Disentangling types of third-party punishers by neural traits.

The act of punishing unfair behavior by unaffected observers (i.e., third-party punishment) is a crucial factor in the functioning of human societies. In everyday life, we see different types of individuals who punish. While some individuals initiate costly punishment against an unfair person independently of what other observers do (independent punishers), others condition their punishment engagement on the presence of another person who punishes (conditional punishers). Still others do not want to partake in any sort of punishment (nonpunishers). Although these distinct behavioral types have a divergent impact on human society, the sources of heterogeneity are poorly understood. We present novel laboratory evidence on the existence of these three types. We use anatomical brain characteristics in combination with stated motives to characterize these types. Findings revealed that independent punishers have larger gray matter volume in the right temporo-parietal junction compared to conditional punishers and nonpunishers, an area involved in social cognition. Conditional punishers are characterized by larger gray matter volume in the right dorsolateral prefrontal cortex, a brain area known to be involved in behavioral control and strategic reasoning, compared to independent punishers and nonpunishers. Finally, both independent punishers and nonpunishers are characterized by larger gray matter volume in an area involved in the processing of social and monetary rewards, that is, the bilateral caudate. By using a neural trait approach, we were able to differentiate these three types clearly based on their neural signatures, allowing us to shed light on the underlying psychological mechanisms.

Neural Mechanisms Underlying Individual Differences in Control-Averse Behavior.

When another person tries to control one's decisions, some people might comply, but many will feel the urge to act against that control. This control aversion can lead to suboptimal decisions and it affects social interactions in many societal domains. To date, however, it has been unclear what drives individual differences in control-averse behavior. Here, we address this issue by measuring brain activity with fMRI while healthy female and male human participants made choices that were either free or controlled by another person, with real consequences to both interaction partners. In addition, we assessed the participants' affects, social cognitions, and motivations via self-reports. Our results indicate that the social cognitions perceived distrust and lack of understanding for the other person play a key role in explaining control aversion at the behavioral level. At the neural level, we find that control-averse behavior can be explained by functional connectivity between the inferior parietal lobule and the dorsolateral prefrontal cortex, brain regions commonly associated with attention reorientation and cognitive control. Further analyses reveal that the individual strength of functional connectivity complements and partially mediates the self-reported social cognitions in explaining individual differences in control-averse behavior. These findings therefore provide valuable contributions to a more comprehensive model of control aversion.SIGNIFICANCE STATEMENT Control aversion is a prevalent phenomenon in our society. When someone tries to control their decisions, many people tend to act against the control. This can lead to suboptimal decisions such as noncompliance to medical treatments or disobeying the law. The degree to which individuals engage in control-averse behavior, however, varies significantly. Understanding the proximal mechanisms that underlie individual differences in control-averse behavior has potential policy implications, for example, when designing policies aimed at increasing compliance with vaccination recommendations, and is therefore a highly relevant research goal. Here, we identify a neural mechanism between parietal and prefrontal brain regions that can explain individual differences in control-averse behavior. This mechanism provides novel insights into control aversion beyond what is accessible through self-reports.

Neural traits characterize unconditional cooperators, conditional cooperators, and noncooperators in group-based cooperation.

Contributing to and maintaining public goods are important for a functioning society. In reality, however, we see large variations in contribution behavior. While some individuals are not cooperative, others are highly so. Still others cooperate only to the extent they believe others will. Although these distinct behavioral types clearly have a divergent social impact, the sources of heterogeneity are poorly understood. We used source-localized resting electroencephalography in combination with a model-free clustering approach to participants' behavior in the Public Goods Game to explain heterogeneity. Findings revealed that compared to noncooperators, both conditional cooperators and unconditional cooperators are characterized by higher baseline activation in the right temporo-parietal junction, an area involved in social cognition. Interestingly, conditional cooperators were further characterized by higher baseline activation in the left lateral prefrontal cortex, an area involved in behavioral control. Our findings suggest that conditional cooperators' better capacities for behavioral control enable them to control their propensity to cooperate and thus to minimize the risk of exploitation by noncooperators.

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.

Understanding Individual Differences in Domain-General Prosociality: A Resting EEG Study.

Prosocial behavior is of vital importance for the smooth functioning of society. However, the propensity to behave in a prosocial manner is characterized by vast individual differences. In order to reveal the sources of these differences, some studies have used objective, task-independent neural traits, for instance resting electroencephalography (EEG). Despite providing valuable insights into the neural signatures of several domains of prosociality, each of these studies has only focused on one single domain. Here, we exposed 137 participants to different social dilemma situations in order to obtain a measure of the individuals' domain-general prosociality and recorded multi-channel task-independent, resting EEG. Using a source-localization technique, we found that resting current density within the temporo-parietal junction in two beta bands (beta2 and beta3) was positively associated with domain-general prosociality. This is the first demonstration of neural signatures underlying individual differences in the propensity to behave in a prosocial manner across different social situations.

Clocking the social mind by identifying mental processes in the IAT with electrical neuroimaging.

Why do people take longer to associate the word "love" with outgroup words (incongruent condition) than with ingroup words (congruent condition)? Despite the widespread use of the implicit association test (IAT), it has remained unclear whether this IAT effect is due to additional mental processes in the incongruent condition, or due to longer duration of the same processes. Here, we addressed this previously insoluble issue by assessing the spatiotemporal evolution of brain electrical activity in 83 participants. From stimulus presentation until response production, we identified seven processes. Crucially, all seven processes occurred in the same temporal sequence in both conditions, but participants needed more time to perform one early occurring process (perceptual processing) and one late occurring process (implementing cognitive control to select the motor response) in the incongruent compared with the congruent condition. We also found that the latter process contributed to individual differences in implicit bias. These results advance understanding of the neural mechanics of response time differences in the IAT: They speak against theories that explain the IAT effect as due to additional processes in the incongruent condition and speak in favor of theories that assume a longer duration of specific processes in the incongruent condition. More broadly, our data analysis approach illustrates the potential of electrical neuroimaging to illuminate the temporal organization of mental processes involved in social cognition.

Intrinsic connectivity networks underlying individual differences in control-averse behavior.

When people sense that another person tries to control their decisions, some people will act against the control, whereas others will not. This individual tendency to control-averse behavior can have far-reaching consequences, such as engagement in illegal activities or noncompliance with medical treatments. Although individual differences in control-averse behavior have been well documented in behavioral studies, their neurological basis is less well understood. Here, we use a neural trait approach to examine whether individual differences in control-averse behavior might be linked to stable brain-based characteristics. To do so, we analyze the association between intrinsic connectivity networks as measured by resting state functional magnetic resonance imaging and control-averse behavior in an economic exchange game. In this game, subjects make choices that are either free or controlled by another person, with real consequences to both interaction partners. We find that the individual level of control-averse behavior can be positively predicted by intrinsic connectivity within the salience network, but not the central executive network or the default mode network. Specifically, subjects with a more prominent connectivity hub in the dorsal anterior cingulate cortex show greater levels of control-averse behavior. This finding provides the first evidence that the heterogeneity in control-averse behavior might originate in systematic differences of the stable functional brain organization.

The scent of attractiveness: levels of reproductive hormones explain individual differences in women's body odour.

Individuals are thought to have their own distinctive body odour which reportedly plays an important role in mate choice. In the present study we investigated individual differences in body odours of women and examined whether some women generally smell more attractive than others or whether odour preferences are a matter of individual taste. We then explored whether levels of reproductive hormones explain women's body odour attractiveness, to test the idea that body odour attractiveness may act as a chemosensory marker of reproductive fitness. Fifty-seven men rated body odours of 28 healthy, naturally cycling women of reproductive age. We collected all odours at peak fertility to control for menstrual cycle effects on body odour attractiveness. Women's salivary oestradiol, progesterone, testosterone and cortisol levels were assessed at the time of odour collection to test whether hormone levels explain body odour attractiveness. We found that the men highly agreed on how attractive they found women's body odours. Interestingly, women's body odour attractiveness was predicted by their oestradiol and progesterone levels: the higher a woman's levels of oestradiol and the lower her levels of progesterone, the more attractive her body odour was rated. In showing that women's body odour attractiveness is explained by levels of female reproductive hormones, but not by levels of cortisol or testosterone, we provide evidence that body odour acts as a valid cue to potential fertility.

Men's preferences for women's body odours are not associated with human leucocyte antigen.

Body odours reportedly portray information about an individual's genotype at the major histocompatibility complex (MHC, called human leucocyte antigen, HLA, in humans). While there is strong experimental support for MHC-associated mating behaviour in animals, the situation in humans is more complex. A lot of effort has been spent on testing HLA-associated odour preferences of women. To date, only very few studies have looked at HLA-linked olfactory preferences in men and these studies have revealed inconsistent results. Here, we investigate men's HLA-associated preferences for women's body odours. Importantly, and in contrast to previous studies, these odours were gathered at peak fertility (i.e. just before ovulation) when any HLA-associated odour preferences should be strongest. We scrutinized whether men's preference for women's body odours is influenced by (i) the number of shared HLA alleles between men and women, (ii) HLA heterozygosity, and (iii) the frequency of rare HLA alleles. We found that men could readily differentiate between odours they found attractive and odours they found less attractive, but that these preferences were not associated with HLA. Specifically, men did not prefer odours from women who are HLA dissimilar, HLA heterozygous, or who have rare HLA alleles. Together, these findings suggest that HLA has no effect on men's odour preferences.

Temporal Characteristics of EEG Microstates Mediate Trial-by-Trial Risk Taking.

People seem to have difficulties when perceiving events whose outcome has no influence on the outcome of future events. This illusion that patterns exist where there are none may lead to adverse consequences, such as escalating losses in financial trading or gambling debt. Despite the enormous social consequences of these cognitive biases, however, their neural underpinnings are poorly understood. Attempts to investigate them have so far relied on evoked neural activity, whereas spontaneous brain activity has been treated as noise to be averaged out. Here, we focus on the spontaneous electroencephalographic (EEG) activity during inter-trial-intervals (ITI) in a sequential risky decision-making task. Using multilevel mediation analyses, our results show that the percentage of time covered by two EEG microstates (i.e., functional brain-states of coherent activity) mediate the influence of outcomes of prior decisions on subsequent risk taking on a trial-by-trial basis. The devised multilevel mediation analysis of the temporal characteristics of EEG microstates during ITI provides a new window into the neurobiology of decision making by bringing the spontaneous brain activity to the forefront of the analysis.

Neuroanatomy of intergroup bias: A white matter microstructure study of individual differences.

Intergroup bias-the tendency to behave more positively toward an ingroup member than an outgroup member-is a powerful social force, for good and ill. Although it is widely demonstrated, intergroup bias is not universal, as it is characterized by significant individual differences. Recently, attention has begun to turn to whether neuroanatomy might explain these individual differences in intergroup bias. However, no research to date has examined whether white matter microstructure could help determine differences in behavior toward ingroup and outgroup members. In the current research, we examine intergroup bias with the third-party punishment paradigm and white matter integrity and connectivity strength as determined by diffusion tensor imaging (DTI). We found that both increased white matter integrity at the right temporal-parietal junction (TPJ) and connectivity strength between the right TPJ and the dorsomedial prefrontal cortex (DMPFC) were associated with increased impartiality in the third-party punishment paradigm, i.e., reduced intergroup bias. Further, consistent with the role that these brain regions play in the mentalizing network, we found that these effects were mediated by mentalizing processes. Participants with greater white matter integrity at the right TPJ and connectivity strength between the right TPJ and the DMPFC employed mentalizing processes more equally for ingroup and outgroup members, and this non-biased use of mentalizing was associated with increased impartiality. The current results help shed light on the mechanisms of bias and, potentially, on interventions that promote impartiality over intergroup bias.

Enhanced emotional empathy after psychosocial stress in young healthy men.

Empathy is a core prerequisite for human social behavior. Relatively, little is known about how empathy is influenced by social stress and its associated neuroendocrine alterations. The current study was designed to test the impact of acute stress on emotional and cognitive empathy. Healthy male participants were exposed to a psychosocial laboratory stressor (trier social stress test, (TSST)) or a well-matched control condition (Placebo-TSST). Afterwards they participated in an empathy test measuring emotional and cognitive empathy (multifaceted empathy test, (MET)). Stress exposure caused an increase in negative affect, a rise in salivary alpha amylase and a rise in cortisol. Participants exposed to stress reported more emotional empathy in response to pictures displaying both positive and negative emotional social scenes. Cognitive empathy (emotion recognition) in contrast did not differ between the stress and the control group. The current findings provide initial evidence for enhanced emotional empathy after acute psychosocial stress.

Individual differences in inhibitory control--relationship between baseline activation in lateral PFC and an electrophysiological index of response inhibition.

The capacity to inhibit inappropriate responses is crucial for goal-directed behavior. Inhibiting such responses seems to come more easily to some of us than others, however. From where do these individual differences originate? Here, we measured 263 participants' neural baseline activation using resting electroencephalogram. Then, we used this stable neural marker to predict a reliable electrophysiological index of response inhibition capacity in the cued Continuous Performance Test, the NoGo-Anteriorization (NGA). Using a source-localization technique, we found that resting delta, theta, and alpha1 activity in the left middle frontal gyrus and resting alpha1 activity in the right inferior frontal gyrus were negatively correlated with the NGA. As a larger NGA is thought to represent better response inhibition capacity, our findings demonstrate that lower levels of resting slow-wave oscillations in the lateral prefrontal cortex, bilaterally, are associated with a better response inhibition capacity.

Cathodal HD-tDCS above the left dorsolateral prefrontal cortex increases environmentally sustainable decision-making.

Environmental sustainability is characterized by a conflict between short-term self-interest and longer-term collective interests. Self-control capacity has been proposed to be a crucial determinant of people's ability to overcome this conflict. Yet, causal evidence is lacking, and previous research is dominated by the use of self-report measures. Here, we modulated self-control capacity by applying inhibitory high-definition transcranial current stimulation (HD-tDCS) above the left dorsolateral prefrontal cortex (dlPFC) while participants engaged in an environmentally consequential decision-making task. The task includes conflicting and low conflicting trade-offs between short-term personal interests and long-term environmental benefits. Contrary to our preregistered expectation, inhibitory HD-tDCS above the left dlPFC, presumably by reducing self-control capacity, led to more, and not less, pro-environmental behavior in conflicting decisions. We speculate that in our exceptionally environmentally friendly sample, deviating from an environmentally sustainable default required self-control capacity, and that inhibiting the left dlPFC might have reduced participants' ability to do so.

Interindividual differences in mindfulness are linked to sleep-electroencephalographic characteristics.

STUDY OBJECTIVES: Mindfulness describes the ability to focus on the presence, including one's thoughts and feelings. Trait mindfulness-a person's inherent tendency to be mindful-has been connected to increased subjective sleep quality, but evidence from objective EEG-based sleep measures is lacking. Here, we investigate whether objective EEG-based sleep parameters explain interindividual differences in trait mindfulness. METHODS: Whole-night polysomnographic data were gathered from 52 healthy adults (27 females; agemean = 21.5 [SE = 0.28]) in their homes using a portable high-density EEG device. Trait mindfulness was assessed using the Five Facet Mindfulness Questionnaire short form (FFMQ-SF). RESULTS: Trait mindfulness was positively correlated at trend level with the percentage of rapid eye movement (REM), but not N1, N2, or slow wave sleep. Additionally, those exhibiting less REM beta/gamma power and NREM beta power displayed higher trait mindfulness and vice versa. Lastly, we replicated findings connecting higher trait mindfulness to better subjective sleep quality. CONCLUSIONS: REM sleep is pivotal for emotional processing. Decreased REM high-frequency activity was suggested to reflect adrenergic reduction that defuses affective experiences. Increased NREM high-frequency activity is a marker for cognitive hyperarousal in insomnia. We speculate that differences in trait mindfulness might be explained by differences in REM- and NREM-sleep functions that promote ideal emotional regulation and prevent hyperarousal.