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.

EEG Microstates in Social and Affective Neuroscience.

Social interactions require both the rapid processing of multifaceted socio-affective signals (e.g., eye gaze, facial expressions, gestures) and their integration with evaluations, social knowledge, and expectations. Researchers interested in understanding complex social cognition and behavior face a "black box" problem: What are the underlying mental processes rapidly occurring between perception and action and why are there such vast individual differences? In this review, we promote electroencephalography (EEG) microstates as a powerful tool for both examining socio-affective states (e.g., processing whether someone is in need in a given situation) and identifying the sources of heterogeneity in socio-affective traits (e.g., general willingness to help others). EEG microstates are identified by analyzing scalp field maps (i.e., the distribution of the electrical field on the scalp) over time. This data-driven, reference-independent approach allows for identifying, timing, sequencing, and quantifying the activation of large-scale brain networks relevant to our socio-affective mind. In light of these benefits, EEG microstates should become an indispensable part of the methodological toolkit of laboratories working in the field of social and affective neuroscience.

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.

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.

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.

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.

Zazen meditation and no-task resting EEG compared with LORETA intracortical source localization.

Meditation is a self-induced and willfully initiated practice that alters the state of consciousness. The meditation practice of Zazen, like many other meditation practices, aims at disregarding intrusive thoughts while controlling body posture. It is an open monitoring meditation characterized by detached moment-to-moment awareness and reduced conceptual thinking and self-reference. Which brain areas differ in electric activity during Zazen compared to task-free resting? Since scalp electroencephalography (EEG) waveforms are reference-dependent, conclusions about the localization of active brain areas are ambiguous. Computing intracerebral source models from the scalp EEG data solves this problem. In the present study, we applied source modeling using low resolution brain electromagnetic tomography (LORETA) to 58-channel scalp EEG data recorded from 15 experienced Zen meditators during Zazen and no-task resting. Zazen compared to no-task resting showed increased alpha-1 and alpha-2 frequency activity in an exclusively right-lateralized cluster extending from prefrontal areas including the insula to parts of the somatosensory and motor cortices and temporal areas. Zazen also showed decreased alpha and beta-2 activity in the left angular gyrus and decreased beta-1 and beta-2 activity in a large bilateral posterior cluster comprising the visual cortex, the posterior cingulate cortex and the parietal cortex. The results include parts of the default mode network and suggest enhanced automatic memory and emotion processing, reduced conceptual thinking and self-reference on a less judgmental, i.e., more detached moment-to-moment basis during Zazen compared to no-task resting.

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.

Functional brain network efficiency predicts intelligence.

The neuronal causes of individual differences in mental abilities such as intelligence are complex and profoundly important. Understanding these abilities has the potential to facilitate their enhancement. The purpose of this study was to identify the functional brain network characteristics and their relation to psychometric intelligence. In particular, we examined whether the functional network exhibits efficient small-world network attributes (high clustering and short path length) and whether these small-world network parameters are associated with intellectual performance. High-density resting state electroencephalography (EEG) was recorded in 74 healthy subjects to analyze graph-theoretical functional network characteristics at an intracortical level. Ravens advanced progressive matrices were used to assess intelligence. We found that the clustering coefficient and path length of the functional network are strongly related to intelligence. Thus, the more intelligent the subjects are the more the functional brain network resembles a small-world network. We further identified the parietal cortex as a main hub of this resting state network as indicated by increased degree centrality that is associated with higher intelligence. Taken together, this is the first study that substantiates the neural efficiency hypothesis as well as the Parieto-Frontal Integration Theory (P-FIT) of intelligence in the context of functional brain network characteristics. These theories are currently the most established intelligence theories in neuroscience. Our findings revealed robust evidence of an efficiently organized resting state functional brain network for highly productive cognitions.

EEG microstates during resting represent personality differences.

We investigated the spontaneous brain electric activity of 13 skeptics and 16 believers in paranormal phenomena; they were university students assessed with a self-report scale about paranormal beliefs. 33-channel EEG recordings during no-task resting were processed as sequences of momentary potential distribution maps. Based on the maps at peak times of Global Field Power, the sequences were parsed into segments of quasi-stable potential distribution, the 'microstates'. The microstates were clustered into four classes of map topographies (A-D). Analysis of the microstate parameters time coverage, occurrence frequency and duration as well as the temporal sequence (syntax) of the microstate classes revealed significant differences: Believers had a higher coverage and occurrence of class B, tended to decreased coverage and occurrence of class C, and showed a predominant sequence of microstate concatenations from A to C to B to A that was reversed in skeptics (A to B to C to A). Microstates of different topographies, putative "atoms of thought", are hypothesized to represent different types of information processing.The study demonstrates that personality differences can be detected in resting EEG microstate parameters and microstate syntax. Microstate analysis yielded no conclusive evidence for the hypothesized relation between paranormal belief and schizophrenia.

A neural marker of costly punishment behavior.

Human readiness to incur personal costs to punish norm violators is a key force in the maintenance of social norms. The willingness to punish is, however, characterized by vast individual heterogeneity that is poorly understood. In fact, this heterogeneity has so far defied explanations in terms of individual-level demographic or psychological variables. Here, we use resting electroencephalography, a stable measure of individual differences in cortical activity, to show that a highly specific neural marker--baseline cortical activity in the right prefrontal cortex--predicts individuals' punishment behavior. The analysis of task-independent individual variation in cortical baseline activity provides a new window into the neurobiology of decision making by bringing dispositional neural markers to the forefront of the analysis.

Tonic activity level in the right prefrontal cortex predicts individuals' risk taking.

Human risk taking is characterized by a large amount of individual heterogeneity. In this study, we applied resting-state electroencephalography, which captures stable individual differences in neural activity, before subjects performed a risk-taking task. Using a source-localization technique, we found that the baseline cortical activity in the right prefrontal cortex predicts individual risk-taking behavior. Individuals with higher baseline cortical activity in this brain area display more risk aversion than do other individuals. This finding demonstrates that neural characteristics that are stable over time can predict a highly complex behavior such as risk-taking behavior and furthermore suggests that hypoactivity in the right prefrontal cortex might serve as a dispositional indicator of lower regulatory abilities, which is expressed in greater risk-taking behavior.

Meditators and non-meditators: EEG source imaging during resting.

Many meditation exercises aim at increased awareness of ongoing experiences through sustained attention and at detachment, i.e., non-engaging observation of these ongoing experiences by the intent not to analyze, judge or expect anything. Long-term meditation practice is believed to generalize the ability of increased awareness and greater detachment into everyday life. We hypothesized that neuroplasticity effects of meditation (correlates of increased awareness and detachment) would be detectable in a no-task resting state. EEG recorded during resting was compared between Qigong meditators and controls. Using LORETA (low resolution electromagnetic tomography) to compute the intracerebral source locations, differences in brain activations between groups were found in the inhibitory delta EEG frequency band. In the meditators, appraisal systems were inhibited, while brain areas involved in the detection and integration of internal and external sensory information showed increased activation. This suggests that neuroplasticity effects of long-term meditation practice, subjectively described as increased awareness and greater detachment, are carried over into non-meditating states.

Theta resting EEG in the right TPJ is associated with individual differences in implicit intergroup bias.

Why are some people more biased than others in their implicit evaluations during social interaction? The dispositional determinants of individual differences in implicit intergroup bias are poorly understood. Here, we explored whether such variability might be explained by stable neural traits. For that purpose, we used the source-localized resting electroencephalograms of 83 members of naturalistic social groups to explain their bias in an in-/outgroup implicit association test. Lower levels of resting theta current density in the right temporo-parietal junction (TPJ) were associated with stronger implicit intergroup bias and explained unique variability in bias beyond relevant personality questionnaires. These findings demonstrate the added value of the neural trait approach in predicting inter-individual differences in implicit social cognition. Given that low levels of resting theta current density during wakefulness likely reflect increased cortical activation, our results suggest that individuals with an efficiently working right TPJ possess capacities to mediate specific cognitive processes that predispose them towards stronger implicit intergroup bias. As the human species has evolved living in distinct social groups, the capacity to quickly differentiate friend from foe became highly adaptive and might thus constitute an essential part of human nature.

Frequency of everyday pro-environmental behaviour is explained by baseline activation in lateral prefrontal cortex.

Humankind faces a plethora of environmental problems, many of which are directly influenced by individual human behaviour. To better understand pro-environmental behaviour, we here try to identify interindividual markers that explain variance in the frequency of every-day pro-environmental behaviour. So far, research on this topic has mainly relied on subjective self-report measures and has yielded mixed results. In this study, we applied a neural trait approach to assess stable, objective individual differences. Using source-localised electroencephalography, we measured cortical activation at rest and combined our neural task-independent data with an ecologically valid assessment of everyday pro-environmental behaviour. We find whole-brain-corrected evidence that task-independent baseline activation in the right lateral prefrontal cortex, a brain area known to be involved in cognitive control and self-control processes, explains individual differences in pro-environmental behaviour. The higher the cortical baseline activation in this area, the higher the frequency of everyday pro-environmental behaviour. Implications for the promotion of pro-environmental behaviour are discussed.