MINI REVIEW

published: 22 February 2022

doi: 10.3389/fnint.2022.827097

Understanding the Dynamics of the

Developing Adolescent Brain
Through Team Science
Kiki Zanolie 1,2 *† , Ili Ma 1,2† , Marieke G. N. Bos 1,2 , Elisabeth Schreuders 1,2 ,
Annelinde R. E. Vandenbroucke 1,2 , Jorien van Hoorn 1,2 , Anna C. K. van Duijvenvoorde 1,2 ,
Lara Wierenga 1,2 , Eveline A. Crone 1,3 and Berna Güroğlu 1,2
1
Department of Developmental and Educational Psychology, Institute of Psychology, Leiden University, Leiden, Netherlands,
2
Leiden Institute for Brain and Cognition (LIBC), Leiden University, Leiden, Netherlands, 3 Erasmus School of Social and
Behavioral Sciences, Erasmus University Rotterdam, Rotterdam, Netherlands

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
Edited by: perspectives article, we highlight the promising role of team science in achieving this
Marie-Ève Tremblay,
University of Victoria, Canada
goal. To illustrate our point, we describe meso (peer relations) and micro (social learning)
Reviewed by:
approaches to understand social development in adolescence as crucial aspects of
Juneman Abraham, adolescent mental health. Finally, we provide an overview of how our team has extended
Binus University, Indonesia
our collaborations beyond scientific partners to multiple societal partners for the purpose
*Correspondence: of informing and including policymakers, education and health professionals, as well as
Kiki Zanolie
[email protected] adolescents themselves when conducting and communicating research.
† Keywords: adolescence, brain development, social development, mental wellbeing, team science
These authors have contributed
equally to this work and share first
authorship

Received: 01 December 2021 INTRODUCTION

Accepted: 31 January 2022
Published: 22 February 2022 Adolescence is a developmental phase between the ages of 10 and 24 years (Sawyer et al., 2018).
Citation: Adolescence starts with puberty, setting off a cascade of hormonal changes signaling the start of
Zanolie K, Ma I, Bos MGN, biological maturation (Dahl et al., 2018), and is characterized by major physical, psychological,
Schreuders E, Vandenbroucke ARE, and social changes (Blakemore and Mills, 2014). Adolescents navigate an increasingly complex
van Hoorn J, van Duijvenvoorde ACK, social network in which peer relations become more salient and are an important source for social
Wierenga L, Crone EA and Güroğlu B learning (e.g., learning about, with, and from peers to adjust to changing social environments;
(2022) Understanding the Dynamics
Westhoff et al., 2020a). Both peer relations and social learning have a great impact on mental
of the Developing Adolescent Brain
Through Team Science.
well-being (Nelson et al., 2005, 2016; Vitaro et al., 2009). Moreover, adolescence is considered
Front. Integr. Neurosci. 16:827097. a period of heightened sensitivity to mental health problems, with approximately 75% of adult
doi: 10.3389/fnint.2022.827097 mental health problems first appearing during adolescence (Kessler et al., 2007; Solmi et al., 2021).

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Zanolie et al. Understanding the Adolescent Brain

As biological, psychological, and social changes occur understanding the complex dynamics of adolescent development
concurrently in adolescence, it is crucial to understand how requires rich measurements and we highlight the promising role
these changes are intertwined and contribute to successful of team science in achieving this goal.
developmental outcomes, such as resilience and mental
health, as well as to maladaptive outcomes, such as risky Peer Relations
behaviors and psychopathology (Davey et al., 2008; Crone Adolescence is characterized by a significant shift in focus from
and Dahl, 2012; Güroğlu, 2021). We further argue that parents toward peers, also referred to as social reorientation
understanding adolescence as a developmental phase with (Nelson et al., 2005). Compared to children, adolescents spend
risks and opportunities requires incorporating a transactional increasingly more time without adult supervision and in the
perspective with measurements at multiple levels (genetic, company of their peers, where fitting in the peer group and peers’
hormonal, neural, behavioral) and across different social opinions become vital for adolescents’ self-identity development
settings (e.g., school, parent relationships, peer relationships; see (Laursen and Veenstra, 2021). Social goals, such as acceptance
Figure 1). Considering the multitude of factors influencing by the peer group and forming and maintaining friendships,
development and the interlinked complexity of their are particularly important in the school context, as they are
corresponding measurement levels, we propose that team consistently linked with markers of positive social adjustment
science is a fruitful approach to understanding the dynamics of and academic achievement (Dawes, 2017). Recently, evidence
adolescent development. from neuroimaging studies corroborate the significance of the
To understand individual differences in optimal conditions peer context for adolescents by finding that adolescents show
for growing up in an increasingly complex social world, heightened neural responses during social decision-making in
we use a variety of neurobiological and behavioral methods. brain regions related to reward and motivation, such as the
Current influential models of adolescent brain development ventral striatum, and in social cognition, such as dorsomedial
describe an asynchronous development of the limbic ‘‘socio- prefrontal cortex (for reviews see Van Hoorn et al., 2019;
affective system’’ and cortical ‘‘cognitive control system’’ Andrews et al., 2021). For example, in early- and mid-
during adolescence (Steinberg, 2008; Somerville et al., 2010). adolescence, mere peer presence, when being observed by an
These models emphasize that faster maturation of the limbic unfamiliar peer, results in heightened neural activation in the
system compared to the slower maturation of the cortical medial prefrontal cortex (mPFC; Somerville et al., 2013), and
system underlies heightened reward sensitivity and risk-taking when being observed by a friend, adolescents show increased
tendencies, leading to risky and impulsive behaviors such as risk-taking behavior, with heightened activation of the ventral
alcohol use (Peters et al., 2017). Recent accounts of adolescent striatum (Chein et al., 2011). In an Event-Related Potential (ERP)
development also include the impact of individual differences study, we have shown that manipulation of participants’ social
in hormonal, genetic, behavioral, and neural influences which rank (high vs. low rank) modulated neural responses during
are intertwined in a social context (Crone and Dahl, 2012; social exchanges in mid-adolescents but not in children or adults,
Pfeifer and Allen, 2021). Specifically, adolescence is seen as signifying that even transient social interactions are particularly
a time for heightened goal flexibility, where social goals can salient for mid-adolescents (Zanolie and Crone, 2021). In
influence pathways for development. Hence, the asynchronous another study, compared to being alone, the presence of a
development between the limbic and cortical system, together group of spectators (consisting of adolescent confederate actors)
with increasingly complex and influential social experiences, led to increased prosocial behavior which was accompanied by
such as peer relations and social learning, makes adolescence a enhanced activation in social brain areas such as mPFC, temporal
sensitive window for socio-affective development which can lead parietal junction (TPJ), precuneus and superior temporal sulcus
to multiple pathways, such as risky behaviors and mental illness, (STS; Van Hoorn et al., 2016). Taken together, these studies of
or prosocial behavior and mental resilience (Crone and Dahl, peer presence illustrate the importance of capturing the peer
2012; Güroğlu, 2021; see Figure 1). context when studying adolescents’ social behavior (Figure 1,
In this article, we highlight the unique position of our highly micro level).
collaborative multidisciplinary research program and focus on Neuroimaging studies aiming to capture real-life peer context
its contributions to the field. We provide an overview of our (Figure 1, meso level), however, face the challenge of bringing
research focusing on: (1) meso level peer relations; and (2) micro peer relationships into the highly controlled experimental
level social learning and next describe their combined influence laboratory setting, such as in the MRI scanner (see Güroğlu and
on mental health. We propose that these two themes are crucial in Veenstra, 2021 for a more extensive review of this research line).
charting the complexity of the dynamically interlinked biological, In tackling this challenge, sociometric assessments of the peer
psychological, and social changes in adolescence. We provide network provide a useful tool to classify an individual’s peer
examples of our research designs with controlled experimental status within a real-life peer group (see Box 1). Combinations
settings at the micro-level and assessments of real-life social of sociometric assessments with neuroimaging and/or economic
relationships at the meso level (see Figure 1). Finally, we exchange paradigms assessing social decision-making (see Box 2)
demonstrate how collaborations can be extended to multiple led to insights into how social interactions and their neural
societal partners to inform and include policy makers, education underpinnings may depend on peer context (Güroğlu et al.,
and health professionals, and adolescents themselves when 2014). For example, we found in adults that interactions
conducting and communicating research. We conclude that with familiar peers relate to heightened activation of brain

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Zanolie et al. Understanding the Adolescent Brain

FIGURE 1 | Overview of the measures required to chart the complexity of developmental changes during adolescence and their impact. Note: This figure illustrates
the richness of measures needed in studies that aim to capture the complexity of developmental changes during adolescence (purple). Findings from such studies
will subsequently have a scientific and societal impact (green). Impact on the scientific field and on society are interlinked, as collaborating and communicating with
societal stakeholders (e.g., policymakers, teachers, parents, and adolescents) also informs new research questions.

regions of affect and reward (including the ventral striatum in a social context, future studies need to incorporate measures
and amygdala) and social cognition (including the mPFC, of social networks with assessments of brain function and
TPJ, STS, and precuneus; Güroğlu et al., 2008). Recently, we structure (Lamblin et al., 2017; Baek et al., 2021). Additionally,
showed that the developmental trajectories of ventral striatum the complexity of social dynamics has in recent years only
responses to rewards are modulated by friendship stability been amplified through the addition of the online social
across a 5-year period (Schreuders et al., 2018) and that ventral layer where young people can have meaningful connections.
striatum responses to winning money are also (negatively) Future studies aiming to understand the dynamics of adolescent
related to acceptance by the peer group (Meuwese et al., development need to include assessments of both offline and
2018). We also showed that in young adults (Schreuders et al., online connections.
2018) and mid-adolescents (Schreuders et al., 2019), prosocial
decisions toward friends compared to disliked or unfamiliar Social Learning
peers, are related to increased activation of the putamen, part Peer relations interact with individual and social learning.
of the reward circuitry, and the posterior temporoparietal Social learning encompasses learning about, with, and from
regions that are involved in social-cognitive processes. Moreover, others. In the peer context, it involves learning about the
neural responses to social rejection depend on the excluder’s characteristics and preferences of a peer or a peer group, such
peer status relative to the adolescent’s own status (De Water as their trustworthiness or cooperativeness (Nelson et al., 2005;
et al., 2017). Our longitudinal studies further showed that the Blakemore and Mills, 2014; Sawyer et al., 2018). Adaptive social
history of peer experiences across childhood modulates neural behavior requires adolescents to learn about these characteristics
responses to social exclusion and during social decision-making and adjust their own behavior accordingly, such as learning when
in adolescence (Will and Güroğlu, 2016; Will et al., 2016, 2018; to be prosocial and towards whom (Steinberg and Morris, 2001;
Asscheman et al., 2019). Van den Bos et al., 2011; Lockwood et al., 2016; Crone and
Taken together, increasing evidence shows both current and Fuligni, 2020). These social learning processes are crucial for
long-term patterns of social experiences with peers modulate fostering healthy relationships with peers, which are predictive
adolescent social behavior and their underlying neural processes of adolescents’ long-term well-being (Paus et al., 2008; Crone and
(Güroğlu, 2022). In order to understand the developing brain Dahl, 2012; Dahl et al., 2018; Sawyer et al., 2018).

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Zanolie et al. Understanding the Adolescent Brain

BOX 1 | Using sociometric assessments to study social experiences.

Sociometric assessments based on nominations of classmates on various criteria (e.g., “who do you like?”, “who do you dislike?”, “who are your friends?”) are most
valuable for assessing social experiences, and more specifically for assessing peer relationships, in an efficient manner. Crucially, these social assessments require
access to a closed network (e.g., classmates, sports team, an orchestra) where the nominations can be made. These nominations can be used to assess social
experiences at two levels. At the dyadic level, they reveal reciprocal relationships, such as mutual friendships between two people who nominate each other as
a friend (see, e.g., Güroğlu et al., 2008; Schreuders et al., 2019). At the group level, they reveal information on the status of the individual within the peer group,
such as accepted or rejected status based on the total number of received nominations from like and dislike nominations (Will et al., 2016; Will and Güroğlu,
2016). Finally, by applying graph theory, these nominations can be used to calculate social network characteristics that can be used to characterize individuals’
positions in the network, such as centrality, as well as identifying group level characteristics, such as social cohesion (Van den Bos et al., 2018). An increasing
number of studies combine assessments of peer relations with neuroscientific designs to investigate how peer relations modulate brain activity (see for review
Güroğlu and Veenstra, 2021).

BOX 2 | Using economic games to study social interactions.

Economic games create social contexts that reveal fundamental aspects about the participant’s social preferences in ways that are quantifiable. They have proven
highly efficient in assessing various forms of (pro) social behavior both in adults and in development (Camerer, 2011; Will and Güroğlu, 2016). These paradigms are
based on an economic exchange between at least two players where the participant’s decisions have actual consequences for their own and their interaction partner’s
payoff. The simplest example is the Dictator Game in which the first player is given valued goods (e.g., money, toys, candies) and can share a portion of those goods
with a second player. While game theoretical models assume that humans are rational players who aim to maximize their own profit, findings consistently show that
people typically share some portion of the goods, thereby revealing other-regarding preferences. In the Ultimatum game, the first player (proposer) is a variant where
the second player can either accept or reject the share given by the first player. If accepted, the goods are divided as proposed by the first player. If rejected, both
players receive nothing. The reward maximizing strategy is to accept any offer greater than zero, but again consistent findings show that offers viewed as unfair are
rejected (Fehr and Schmidt, 1999). Offers made by the first player are typically higher in the Ultimatum Game than Dictator Game, revealing strategic considerations
to reduce the probability of rejection. Another variant is the Trust Game, in which the first player (investor) can again share a portion of goods with the second player
(trustee). The portion received by the trustee is multiplied by the experimenters. The trustee then chooses to either share the profit with the investor (reciprocation;
both profit from the exchange) or keep all the profit (betrayal; only the trustee profits, the investor loses the entrusted amount; Berg et al., 1995). These paradigms can
be presented in a repeated fashion, such that participants play multiple rounds of these games with the same partner(s). Feedback received during these repeated
interactions facilitates learning about the social preferences of other individuals or groups. Economic exchange paradigms are simple enough to administer to a wide
age range (from 3 years old to adults) and enable studying developmental patterns in social behavior (Güroğlu et al., 2009; Meuwese et al., 2015; Zanolie et al., 2015;
Ma et al., 2017). Moreover, their structured nature makes them further suitable for neuroimaging research.

Social learning is often studied with repeated behavioral mid-adolescence. These findings are consistent with the idea that
economic paradigms, where participants play multiple rounds of learning about the consequences of actions in an interpersonal
an economic game with the same partner or multiple partners context is especially salient for adolescents and furthermore
from one experimentally selected group (see Box 2). Peer highlight early to mid-adolescence as a sensitive period for
characteristics or peer evaluations are typically experimentally learning about others (Blakemore and Mills, 2014; Nelson et al.,
manipulated, allowing participants to learn through positive 2016; Sawyer et al., 2018; Andrews et al., 2021).
and negative feedback (Ma et al., 2020; Westhoff et al., 2020b; Considering that learning at school takes place in the peer
Zanolie and Crone, 2021). Reinforcement learning models can context (i.e., in classrooms and group assignments), learning
then be used to characterize individual differences in learning about, with, and from peers are also crucial research lines
strategies, learning speed, and the underlying cognitive processes to identify the optimal conditions of learning at school. In
that cause age differences in learning about others (Sutton and ongoing studies, we focus on determining which individuals
Barto, 2018; Nussenbaum and Hartley, 2019; Wilson and Collins, work well together by combining reinforcement learning or
2019). For example, we used an information sampling paradigm feedback processing paradigms with sociometric assessments.
in which participants were able to sample information about Such studies form the first steps of identifying optimal conditions
a peer’s history of trustworthiness before deciding to trust or for learning in the context of peers by informing how peer
not trust them (Ma et al., 2020). We found that behavioral relationships in classrooms and differences in learning strategies
adaptation to the gathered evidence improved with age, especially between collaborating students may influence (social) learning.
from early to mid-adolescence. In other studies, we manipulated
the cooperativeness of groups (Westhoff et al., 2020b) and Social Experiences and Mental Health
used a probabilistic learning task in which participants could It is well-established that social experiences influence mental
sometimes earn rewards for themselves and sometimes for others health and well-being. For example, close friendships during
(Westhoff et al., 2021). We found that probabilistic learning adolescence are a protective factor against mental health
to benefit others showed age-related improvement across problems across adolescence and later in life (Van Harmelen
adolescence and was associated with ventromedial prefrontal et al., 2017, 2021). However, being rejected by peers is associated
cortex responses to unexpected outcomes. Learning for the self with self-harm (Esposito et al., 2019) and depressive symptoms
was stable across adolescence and associated with ventral striatal (Platt et al., 2013). Also, epidemiological studies have shown
responses to unexpected outcomes. Together, these findings a peak in the emergence of mental health problems across
suggest that adolescents show rapid improvements in behavioral adolescence (Dalsgaard et al., 2020). Showing symptoms of
adjustments to the social environment, especially from early to psychopathology in childhood or adolescence are found to

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Zanolie et al. Understanding the Adolescent Brain

be a key predictor of mental health problems and other adolescents via targeted websites designed for youth1 and
adverse outcomes later in life (Zisook et al., 2007; Caspi et al., scientific articles for children (Westhoff et al., 2020a). We
2020). These findings indicate that there are developmental also inform youth professionals by designing educational
processes enhancing or bringing about vulnerabilities to develop material for elementary and high schools about the developing
mental health problems. Theoretical models propose a complex brain (e.g., http://www.breinkennisleiden.nl/onderwijs),
interplay between brain development, hormonal changes, and and through our contributions to science-translation
social development in interaction with environmental factors reports such as those on differences between boys and
that may explain the emergence and maintenance of mental girls in learning (report Dutch Educational Council),
health problems across adolescence (Pfeifer and Allen, 2021). and UNESCO’s International Science and Evidence-based
So far, only a few studies have directly linked the relationship Education Assessment on the social-emotional learning
between social context, brain development, and mental health (Gotlieb et al., 2022).
outcomes. One study showed that greater subgenual anterior It is particularly important to include adolescents themselves
cingulate activity (sgACC) during a social exclusion game when forming policies and designing interventions in order
was associated with an increase in parent-reported depressive to make their participation efforts optimal and to contribute
symptoms 1 year later (Masten et al., 2013). Social interactions to their sense of autonomy which benefits their mental health
with friends have also been related to activation of the sgACC (Fuligni, 2019). Especially during mid-adolescence interventions
and the ventral striatum. These brain regions are associated with typically tend to fail when they do not align with adolescents’
the reward circuitry, speculatively providing indirect evidence desired feeling to be respected and accorded status (Yeager
linking positive peer interactions with mental health (Güroğlu et al., 2018). Peer-led interventions to generate positive
et al., 2008; Schreuders et al., 2021). Research is needed to behavioral changes can be powerful when the complexity of
elucidate the complex interplay between brain development, peer relations and social networks are taken into account as
social context, and mental health (Davey et al., 2008; Pfeifer well as social learning (e.g., imitation, norms, and positive
and Allen, 2021). To better understand mental health and reinforcement; Veenstra and Laninga-Wijnen et al., 2022).
the transition from mental health to mental illness, our The next step toward improving these efforts is setting up
ongoing studies aim to contextualize individual differences in projects in which adolescents are involved in co-designing and
relation to social development and genetic factors (e.g., by co-creating research (Whitmore and Mills, 2021). Not only
using twin designs; Crone et al., 2020), and their association does this enrich the context in which scientific findings
with mental health outcomes in a developmental context can be launched and interpreted, but crucially informs
(Ferschmann et al., 2021). researchers in important ways, helping them to improve
Crucial in contextualizing individual differences in etiology their research designs and paradigms. In our current projects,
and maintenance of mental health problems is investigating adolescent volunteers are also involved in disseminating
neurobiological mechanisms of psychopathology from a knowledge to their peers, thereby increasing the likelihood
longitudinal perspective. Specifically, non-linear developmental that the information is relevant and interesting for the target
changes in cortical and subcortical structures may explain audience (e.g., http://www.instagram.com/breinboost). Finally,
why cross-sectional developmental neuroimaging studies in several innovative projects we involve societal stakeholders
may find mixed results depending on the age range of the (e.g., teachers, adolescents, policymakers) in the research
participants (Wierenga et al., 2014a,b; Mills et al., 2021). With consortium from the start of the project and create research
longitudinal designs, we, for example, found that heightened and knowledge dissemination projects together throughout
scores on externalizing symptoms were associated with smaller the project (see e.g., http://www.neurolab.nl/startimpuls,
developmental changes in brain structure (Vijayakumar Vandenbroucke et al., 2021).
et al., 2014; Oostermeijer et al., 2016; Ambrosino et al.,
2017; Bos et al., 2018a; but see Ducharme et al., 2011). CONCLUSIONS
Likewise, for internalizing symptoms, such as depression,
longitudinal studies revealed associations with aberrant brain In this perspectives article, we provided an overview of ways
development (Whittle et al., 2014; Luby et al., 2016; Bos et al., of characterizing developmental changes during adolescence
2018b). Together, these studies highlight the importance of and their relation to the developing brain. We highlighted the
longitudinal studies for understanding mental health problems importance of capturing social contextual factors, as social
and their development. experiences play a crucial role in shaping many developmental
trajectories. We further emphasized the importance of
INTEGRATING SCIENCE AND SOCIETY longitudinal approaches in developmental studies in identifying
predictors of mental health. The social context is increasingly
Integrating scientific knowledge about adolescent development important given that young people today grow up in a highly
into society can be achieved in multiple ways. Popular science socially complex environment. The recent COVID-19 pandemic
books such as ‘‘The Adolescent Brain’’ (Het Puberende Brein, showed how strong the effects of the changing social context
Crone, 2018) and ‘‘Inventing ourselves’’ (Blakemore, 2018) are on adolescents’ mental health has been (Orben et al., 2020;
help to reach a wider audience, including policymakers
(‘‘science for policy’’). In our team, we aim to reach 1 http://www.kijkinjebrein.nl/

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Zanolie et al. Understanding the Adolescent Brain

Van de Groep et al., 2020; Asscheman et al., 2021; Breaux et al., FUNDING
2021; Green et al., 2021; Klootwijk et al., 2021). An increased
understanding of the effect of social contextual factors on KZ was supported by the AXA Research Fund. IM was
the development and neurobiological mechanisms underlying supported by the National Science Foundation. ES and MB were
mental health will inform high stake policy questions, and supported by the Research Council of Norway (RCN; 288083).
find their way to daily practice. Throughout this overview, we ES and AV were supported by the The Netherlands Organization
illustrated the value of collaborative team science to understand for Scientific Research (NWO; NWA 400.17.602 Startimpulse
adolescent development and the value of integration of science grant). JH was supported by NWO [406-11-019 ResearchTalent
and society to be able to inform policy and practice. Grant]. AD was supported by NWO [464-15-176 Open Research
Area (ORA) grant] and the Social Resilience and Security
AUTHOR CONTRIBUTIONS Program (Leiden University). LW was supported by NWO
(024.001.003 Gravitation grant). EC was supported by the
All authors co-designed the aims and wrote the article. KZ and European Research Council (ERC; StG-263234 Starting grant
IM designed the article aims, outline, figure, and integrated and CoG-681632 Consolidator grant) and NWO (453-14-
individual author contributions. All authors contributed to the 001 Vici grant). BG was supported by NWO (VENI 451-10-
article and approved the submitted version. 021 Veni grant).

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Westhoff, B., Molleman, L., Viding, E., van den Bos, W., and van Copyright © 2022 Zanolie, Ma, Bos, Schreuders, Vandenbroucke, van Hoorn,
Duijvenvoorde, A. C. (2020b). Developmental asymmetries in learning to van Duijvenvoorde, Wierenga, Crone and Güroğlu. This is an open-access article
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