Familiarity and Audience Effects on Giving: A Functional Magnetic Resonance Imaging Study.

Giving is often characterized by the conflicting decision to give up something of value to benefit others. Recent evidence indicated that giving is highly context-dependent. To unravel the neural correlates of social context, in this study, young adults (n = 32) performed a novel giving fMRI paradigm, in which they divided coins between self and known (friends) or unknown (unfamiliar) others. A second manipulation included presence of others; giving decisions were made with an audience or anonymously. Results showed that participants gave more coins to a friend than to an unfamiliar other and generally gave more in the presence of an audience. On a neural level, medial prefrontal cortex and the right insula were most active for relatively generous decisions. These findings possibly reflect that aversion of norm deviation or fairness concerns drive differences in the frequency of giving. Next, activation in separate subregions of the TPJ-IPL (i.e., a region that comprises the TPJ and inferior parietal lobule) was found for target and audience contexts. Overall, our findings suggest that donation size and social contextual information are processed in separable brain regions and that TPJ-IPL plays an important role in balancing self- and other-oriented motives related to the social context.

Giving to Friends, Classmates, and Strangers in Adolescence.

This study examined how perspective taking and sensitivity to social rewards predict giving to friends, classmates, and strangers in adolescence. Five hundred and twenty adolescents aged 12-17 years completed questionnaires on perspective taking and social rewards and played three Dictator Games in which they divided coins between themselves and a friend, classmate, and stranger. We found that, irrespective of age, adolescents donated most to a friend, less to their classmate, and least to a stranger, and females donated more than males. Individual differences in perspective taking and social reward sensitivity moderated how much adolescents donated, especially to strangers. These findings suggest that perspective taking and sensitivity to social rewards influence giving behavior in adolescence, especially to unknown others.

Developmental Changes and Individual Differences in Trust and Reciprocity in Adolescence.

A Trust Game was used to examine trust and reciprocity development in 12-18-year-old-adolescents (N = 496), as findings have been conflicting and transitions in adolescence remain elusive. Furthermore, this study tested the roles of gender, risk, and individual differences in empathy, impulsivity, and antisocial tendencies in trust and reciprocity. Results indicate stability in trust and a decrease in reciprocity across adolescence, but also show that trust and reciprocity choices were influenced by risk, and that empathy mediated the age-related decrease in reciprocity. Males trusted more than females, but there were no gender differences in reciprocity. These findings highlight the importance of considering individual differences and adolescents' sensitivities to varying contexts in explaining trust and reciprocity development in adolescence.

Mighty metaphors: behavioral and ERP evidence that power shifts attention on a vertical dimension.

Thinking about the abstract concept power may automatically activate the spatial up-down image schema (powerful up; powerless down) and consequently direct spatial attention to the image schema-congruent location. Participants indicated whether a word represented a powerful or powerless person (e.g. 'king' or 'servant'). Following each decision, they identified a target at the top or bottom of the visual field. In Experiment 1 participants identified the target faster when their spatial position was congruent with the perceived power of the preceding word than when it was incongruent. In Experiment 2 ERPs showed a higher N1 amplitude for congruent spatial positions. These results support the view that attention is driven to the image schema congruent location of a power word. Thus, power is partially understood in terms of vertical space, which demonstrates that abstract concepts are grounded in sensory-motor processing.

Growing in generosity? The effects of giving magnitude, target, and audience on the neural signature of giving in adolescence.

Giving is essential for forming and maintaining social relationships, which is an important developmental task for adolescents. This pre-registered fMRI study investigated behavioral and neural correlates of adolescents' (N = 128, ages 9 - 19 years) small versus large size giving in different social contexts related to target (i.e., giving to a friend or unfamiliar peer) and peer presence (i.e., anonymous versus audience giving). Participants gave more in the small size than large size condition, more to friends than to unfamiliar peers, and more in the audience compared to anonymous condition. Giving very small or large amounts was associated with increased activity in the medial prefrontal cortex (mPFC) and anterior insula (AI), and older adolescents showed increased lateral and anterior PFC activation for small size giving. We observed activity in the intraparietal cortex (IPL), dorsolateral prefrontal cortex, and AI for giving to friends, but no age-related differences in this activity. Behaviorally, in contrast, we observed that older adolescents differentiated more in giving between friends and unfamiliar peers. Finally, we observed interactions between peer presence and target in the AI, and between giving magnitude and target in the precuneus. Together, findings reveal higher context-dependency of giving and more lateral PFC activity for small versus large giving in older adolescents.

Understanding the Dynamics of the Developing Adolescent Brain Through Team Science.

One of the major goals for research on adolescent development is to identify the optimal conditions for adolescents to grow up in a complex social world and to understand individual differences in these trajectories. Based on influential theoretical and empirical work in this field, achieving this goal requires a detailed understanding of the social context in which neural and behavioral development takes place, along with longitudinal measurements at multiple levels (e.g., genetic, hormonal, neural, behavioral). In this perspectives article, we highlight the promising role of team science in achieving this goal. To illustrate our point, we describe meso (peer relations) and micro (social learning) approaches to understand social development in adolescence as crucial aspects of adolescent mental health. Finally, we provide an overview of how our team has extended our collaborations beyond scientific partners to multiple societal partners for the purpose of informing and including policymakers, education and health professionals, as well as adolescents themselves when conducting and communicating research.

What motivates the adolescent? Brain regions mediating reward sensitivity across adolescence.

The relation between brain development across adolescence and adolescent risky behavior has attracted increasing interest in recent years. It has been proposed that adolescents are hypersensitive to reward because of an imbalance in the developmental pattern followed by the striatum and prefrontal cortex. To date, it is unclear if adolescents engage in risky behavior because they overestimate potential rewards or respond more to received rewards and whether these effects occur in the absence of decisions. In this study, we used a functional magnetic resonance imaging paradigm that allowed us to dissociate effects of the anticipation, receipt, and omission of reward in 10- to 12-year-old, 14- to 15-year-old, and 18- to 23-year-old participants. We show that in anticipation of uncertain outcomes, the anterior insula is more active in adolescents compared with young adults and that the ventral striatum shows a reward-related peak in middle adolescence, whereas young adults show orbitofrontal cortex activation to omitted reward. These regions show distinct developmental trajectories. This study supports the hypothesis that adolescents are hypersensitive to reward and adds to the current literature in demonstrating that neural activation differs in adolescents even for small rewards in the absence of choice. These findings may have important implications for understanding adolescent risk-taking behavior.

Ranking status differentially affects rejection sensitivity in adolescence: An event-related potential study.

Adolescents are sensitive to peer rejection but this may be dependent on their status. This study examined the role of ranking status on rejection sensitivity in adolescence using an experimental bargaining design. To manipulate ranking status, participants between ages 9-22-years (final sample n = 102) performed a reaction time task with two peers to induce high and low status. Next, participants played an iterative Ultimatum Game as high or low status proposer with an opposite status responder. Rejection of fair offers was associated with larger Medial Frontal Negativity (MFN) compared to acceptance of fair offers. An interaction between age and status group revealed that after rejection of fair offers, mid-adolescents showed a larger MFN when having a low status and smaller MFN when having a high status, relative to children and adults. These findings suggest that the MFN reacts as a neural alarm system to social prediction errors, signaling a need for vigilance to deviations from the norm, which is influenced by ranking status especially during mid-adolescence.

Mood Variability Among Early Adolescents in Times of Social Constraints: A Daily Diary Study During the COVID-19 Pandemic.

The COVID-19 pandemic and associated governmental regulations have drastically changed the daily social lives of children, adolescents, and adults. Changes in the social context may particularly affect children who are in the transition to adolescents (henceforth referred to as early adolescents) as adolescence is a crucial period for peer interactions and development of independence and autonomy. Yet, the impact of the pandemic and associated governmental regulations on early adolescents' emotional well-being has yet to be clarified. In the current study, we explored daily fluctuations in mood in 54 early adolescents (M age = 11.07) during the first few months (April 2020-June 2020) of the COVID-19 pandemic. Moreover, the role of parents and peers on adolescents' mood variability was investigated. Adolescents rated their mood (i.e., happiness, anger, sadness, anxiety) and peer interactions once a day during four separate weeks across different weeks of containment measures in the Netherlands. Moreover, adolescents reported on their experienced attachment to parents and peers and internalizing problems during baseline and the final measurement, respectively. Results showed relatively stable levels of mood during the first few months of the COVID-19 pandemic. However, individual differences in mood variability during the first assessment week were negatively associated with the experienced level of attachment to both parents and peers. Moreover, heightened levels of mood variability did not mediate the link between attachment and internalizing problems. Lastly, the quality of offline contact, but not online contact, was negatively related to adolescents' mood variability. Overall, this study suggests that mood of early adolescents did not heavily fluctuated across the first few months of the COVID-19 pandemic. Our findings add to the growing body of literature aiming to understand how adolescent's life are affected by the COVID-19 crisis and illustrates that social connectedness to parents or peers may facilitate resilience to distress and daily mood fluctuation in early adolescents.

A daily diary study on adolescents' mood, empathy, and prosocial behavior during the COVID-19 pandemic.

Adolescence is a formative phase for social development. The COVID-19 pandemic and associated regulations have led to many changes in adolescents' lives, including limited opportunities for social interactions. The current exploratory study investigated the effect of the first weeks of COVID-19 pandemic lockdown on Dutch adolescents' (N = 53 with attrition, N = 36 without attrition) mood, empathy, and prosocial behavior. Longitudinal analyses comparing pre-pandemic measures to a three-week peri-pandemic daily diary study showed (i) decreases in empathic concern, opportunities for prosocial actions, and tension, (ii) stable levels of social value orientation, altruism, and dire prosociality, and (iii) increased levels of perspective-taking and vigor during the first weeks of lockdown. Second, this study investigated peri-pandemic effects of familiarity, need, and deservedness on giving behavior. To this end, we utilized novel hypothetical Dictator Games with ecologically valid targets associated with the COVID-19 pandemic. Adolescents showed higher levels of giving to a friend (a familiar other, about 51% of the total share), a doctor in a hospital (deserving target, 78%), and individuals with COVID-19 or a poor immune system (targets in need, 69 and 63%, respectively) compared to an unfamiliar peer (39%) This suggests that during the pandemic need and deservedness had a greater influence on adolescent giving than familiarity. Overall, this study demonstrates detrimental effects of the first weeks of lockdown on adolescents' empathic concern and opportunities for prosocial actions, which are important predictors of healthy socio-emotional development. However, adolescents also showed marked resilience and a willingness to benefit others as a result of the lockdown, as evidenced by improved perspective-taking and mood, and high sensitivity to need and deservedness in giving to others.

Evaluating the negative or valuing the positive? Neural mechanisms supporting feedback-based learning across development.

How children learn from positive and negative performance feedback lies at the foundation of successful learning and is therefore of great importance for educational practice. In this study, we used functional magnetic resonance imaging (fMRI) to examine the neural developmental changes related to feedback-based learning when performing a rule search and application task. Behavioral results from three age groups (8-9, 11-13, and 18-25 years of age) demonstrated that, compared with adults, 8- to 9-year-old children performed disproportionally more inaccurately after receiving negative feedback relative to positive feedback. Additionally, imaging data pointed toward a qualitative difference in how children and adults use performance feedback. That is, dorsolateral prefrontal cortex and superior parietal cortex were more active after negative feedback for adults, but after positive feedback for children (8-9 years of age). For 11- to 13-year-olds, these regions did not show differential feedback sensitivity, suggesting that the transition occurs around this age. Pre-supplementary motor area/anterior cingulate cortex, in contrast, was more active after negative feedback in both 11- to 13-year-olds and adults, but not 8- to 9-year-olds. Together, the current data show that cognitive control areas are differentially engaged during feedback-based learning across development. Adults engage these regions after signals of response adjustment (i.e., negative feedback). Young children engage these regions after signals of response continuation (i.e., positive feedback). The neural activation patterns found in 11- to 13-year-olds indicate a transition around this age toward an increased influence of negative feedback on performance adjustment. This is the first developmental fMRI study to compare qualitative changes in brain activation during feedback learning across distinct stages of development.

Switching between colors and shapes on the basis of positive and negative feedback: an fMRI and EEG study on feedback-based learning.

A crucial element of testing hypotheses about rules for behavior is the use of performance feedback. In this study, we used fMRI and EEG to test the role of medial prefrontal cortex (PFC) and dorsolateral (DL) PFC in hypothesis testing using a modified intradimensional/extradimensional rule shift task. Eighteen adults were asked to infer rules about color or shape on the basis of positive and negative feedback in sets of two trials. Half of the trials involved color-to-color or shape-to-shape trials (intradimensional switches; ID) and the other half involved color-to-shape or shape-to-color trials (extradimensional switches; ED). Participants performed the task in separate fMRI and EEG sessions. ED trials were associated with reduced accuracy relative to ID trials. In addition, accuracy was reduced and response latencies increased following negative relative to positive feedback. Negative feedback resulted in increased activation in medial PFC and DLPFC, but more so for ED than ID shifts. Reduced accuracy following negative feedback correlated with increased activation in DLPFC, and increased response latencies following negative feedback correlated with increased activation in medial PFC. Additionally, around 250msec following negative performance feedback participants showed a feedback-related negative scalp potential, but this potential did not differ between ID and ED shifts. These results indicate that both medial PFC and DLPFC signal the need for performance adjustment, and both regions are sensitive to the increased demands of set shifting in hypothesis testing.

Verifying visual properties in sentence verification facilitates picture recognition memory.

According to the perceptual symbols theory (Barsalou, 1999), sensorimotor simulations underlie the representation of concepts. We investigated whether recognition memory for pictures of concepts was facilitated by earlier representation of visual properties of those concepts. During study, concept names (e.g., apple) were presented in a property verification task with a visual property (e.g., shiny) or with a nonvisual property (e.g., tart). Delayed picture recognition memory was better if the concept name had been presented with a visual property than if it had been presented with a nonvisual property. These results indicate that modality-specific simulations are used for concept representation.

Predicting reading and mathematics from neural activity for feedback learning.

Although many studies use feedback learning paradigms to study the process of learning in laboratory settings, little is known about their relevance for real-world learning settings such as school. In a large developmental sample (N = 228, 8-25 years), we investigated whether performance and neural activity during a feedback learning task predicted reading and mathematics performance 2 years later. The results indicated that feedback learning performance predicted both reading and mathematics performance. Activity during feedback learning in left superior dorsolateral prefrontal cortex (DLPFC) predicted reading performance, whereas activity in presupplementary motor area/anterior cingulate cortex (pre-SMA/ACC) predicted mathematical performance. Moreover, left superior DLPFC and pre-SMA/ACC activity predicted unique variance in reading and mathematics ability over behavioral testing of feedback learning performance alone. These results provide valuable insights into the relationship between laboratory-based learning tasks and learning in school settings, and the value of neural assessments for prediction of school performance over behavioral testing alone. (PsycINFO Database Record

Rejection in Bargaining Situations: An Event-Related Potential Study in Adolescents and Adults.

The neural correlates of rejection in bargaining situations when proposing a fair or unfair offer are not yet well understood. We measured neural responses to rejection and acceptance of monetary offers with event-related potentials (ERPs) in mid-adolescents (14-17 years) and early adults (19-24 years). Participants played multiple rounds of the Ultimatum Game as proposers, dividing coins between themselves and a second player (responder) by making a choice between an unfair distribution (7 coins for proposer and 3 for responder; 7/3) and one of two alternatives: a fair distribution (5/5) or a hyperfair distribution (3/7). Participants mostly made fair offers (5/5) when the alternative was unfair (7/3), but made mostly unfair offers (7/3) when the alternative was hyperfair (3/7). When participants' fair offers (5/5; alternative was 7/3) were rejected this was associated with a larger Medial Frontal Negativity (MFN) compared to acceptance of fair offers and rejection of unfair offers (7/3; alternative was 3/7). Also, the MFN was smaller after acceptance of unfair offers (7/3) compared to rejection. These neural responses did not differ between adults and mid-adolescents, suggesting that the MFN reacts as a neural alarm system to social prediction errors which is already prevalent during adolescence.

Corrigendum: Number-induced shifts in spatial attention: a replication study.

[This corrects the article on p. 987 in vol. 5, PMID: 25309472.].

Number-induced shifts in spatial attention: a replication study.

In a spatial attention paradigm, Fischer et al. (2003) showed that merely perceiving a number shifted attention according to the magnitude of the number. Low numbers shifted attention to the left and high numbers shifted attention to the right. This suggests that numbers are represented by the mental number line - a spatial image schema that is ordered from left to right with increasing magnitude. In six experiments, we used the spatial attention paradigm of Fischer et al. (2003) to investigate if and when such mental representations are activated. Participants detected visual targets that were preceded by low and high numbers. Between experiments we manipulated how participants processed the number. Participants either merely perceived the number, as in the experiments by Fischer et al. (2003) processed the number's parity, or processed the number's magnitude. Our results provide little support for the idea that numbers shift spatial attention. Only in one of the two experiments in which participants processed number magnitude did participants respond faster to targets in congruent locations (left for low magnitudes and right for high magnitudes) than in incongruent locations. In the other five experiments number magnitude did not affect spatial attention. This shows, in contrast to Fischer et al.'s (2003) results, that the mental number line is not activated automatically but at best only when it is contextually relevant. Furthermore, these results suggest that image schemas in general may be context-dependent rather than fundamental to mental concepts.

The relation between resting state connectivity and creativity in adolescents before and after training.

An important component of creativity is divergent thinking, which involves the ability to generate novel and useful problem solutions. In this study, we tested the relation between resting-state functional connectivity of brain areas activated during a divergent thinking task (i.e., supramarginal gyrus, middle temporal gyrus, medial frontal gyrus) and the effect of practice in 32 adolescents aged 15-16. Over a period of two weeks, an experimental group (n = 16) conducted an 8-session Alternative Uses Task (AUT) training and an active control group (n = 16) conducted an 8-session rule switching training. Resting-state functional connectivity was measured before (pre-test) and after (post-test) training. Across groups at pre-test, stronger connectivity between the middle temporal gyrus and bilateral postcentral gyrus was associated with better divergent thinking performance. The AUT-training, however, did not significantly change functional connectivity. Post hoc analyses showed that change in divergent thinking performance over time was predicted by connectivity between left supramarginal gyrus and right occipital cortex. These results provide evidence for a relation between divergent thinking and resting-state functional connectivity in a task-positive network, taking an important step towards understanding creative cognition and functional brain connectivity.

The relation between gray matter morphology and divergent thinking in adolescents and young adults.

Adolescence and early adulthood are developmental time periods during which creative cognition is highly important for adapting to environmental changes. Divergent thinking, which refers to generating novel and useful solutions to open-ended problems, has often been used as a measure of creative cognition. The first goal of this structural neuroimaging study was to elucidate the relationship between gray matter morphology and performance in the verbal (AUT; alternative uses task) and visuo-spatial (CAT; creative ability test) domain of divergent thinking in adolescents and young adults. The second goal was to test if gray matter morphology is related to brain activity during AUT performance. Neural and behavioral data were combined from a cross-sectional study including 25 adolescents aged 15-17 and 20 young adults aged 25-30. Brain-behavior relationships were assessed without a priori location assumptions and within areas that were activated during an AUT-scanner task. Gray matter volume and cortical thickness were not significantly associated with verbal divergent thinking. However, visuo-spatial divergent thinking (CAT originality and fluency) was positively associated with cortical thickness of the right middle temporal gyrus and left brain areas including the superior frontal gyrus and various occipital, parietal, and temporal areas, independently of age. AUT brain activity was not associated with cortical thickness. The results support an important role of a widespread brain network involved in flexible visuo-spatial divergent thinking, providing evidence for a relation between cortical thickness and visuo-spatial divergent thinking in adolescents and young adults. However, studies including visuo-spatial divergent thinking tasks in the scanner are warranted.