MEDICINE

REVIEW ARTICLE

Brain Development During Adolescence

Neuroscientific Insights Into This Developmental Period

Kerstin Konrad, Christine Firk, Peter J. Uhlhaas

dolescence is the phase of life between late

SUMMARY
Background: Adolescence is the phase of life between late childhood and
A childhood and adulthood. It is a time not only of
physical maturation, but also of mental and emotional
adulthood. Typically, adolescents seek diversion, new experiences, and strong development into an independent, responsible adult.
emotions, sometimes putting their health at serious risk. In Germany, for
The major developmental tasks of adolescence
example, 62% of all deaths among persons aged 15 to 20 are due to traumatic
include the establishment and nurturing of intimate
injuries. Neuroscientific explanations have been proposed for typical adoles-
relationships and the development of identity, future
cent behavior; with these explanations in mind, one can derive appropriate
perspectives, independence, self-confidence, self-
ways of dealing with adolescents.
control, and social skills (1).
Method: We selectively review pertinent articles retrieved from the PubMed
database about the structural and functional development of the brain in Heightened risk-taking behavior
adolescence. Many adolescents and young adults are prone to take
Results: New findings in developmental psychology and neuroscience reveal risks and enjoy having extreme emotions (2, 3). This
that a fundamental reorganization of the brain takes place in adolescence. In is reflected in statistics showing that risky behavior
postnatal brain development, the maximum density of gray matter is reached in adolescence is linked with an elevated risk to
first in the primary sensorimotor cortex, and the prefrontal cortex matures last. health (4). In Germany, for example, 62% of all deaths
Subcortical brain areas, especially the limbic system and the reward system, among persons aged 15 to 20 are due to traumatic in-
develop earlier, so that there is an imbalance during adolescence between the juries. The most common causes of death are motor
more mature subcortical areas and less mature prefrontal areas. This may vehicle accidents, other accidents, violence, and self-
account for typical adolescent behavior patterns, including risk-taking. injury (5). The high mortality is attributable to drunk
Conclusion: The high plasticity of the adolescent brain permits environmental driving, driving without a seatbelt, carrying
influences to exert particularly strong effects on cortical circuitry. While this weapons, substance abuse, and unprotected sexual
makes intellectual and emotional development possible, it also opens the door intercourse (4).
to potentially harmful influences.
Boys and girls in comparison
►Cite this as: As can be seen in the Table, boys and girls engage in
Konrad K, Firk C, Uhlhaas PJ: Brain development during adolescence: risky behavior at similar frequencies. In recent years,
neuroscientific insights into this developmental period. for example, the prevalence of smoking among boys
Dtsch Arztebl Int 2013; 110(25): 425–31. DOI: 10.3238/arztebl.2013.0425
and girls has become nearly equal, although some
qualitative differences remain: Boys smoke more
cigarettes, and they also more commonly smoke
“harder” tobacco products such as cigars, black to-
bacco, and unfiltered cigarettes. Boys and girls also
drink different alcoholic beverages: Boys tend to
drink beer and hard liquor, while girls tend to drink
wine, sparkling wine, etc. Boys drink alcohol more
frequently and in larger amounts. They also consume
illegal drugs more commonly than girls. Boys are
more prone to accidents, and they take more risks
when driving. Girls, on the other hand, are more
likely to engage in health-endangering behavior in
the area of nutrition (e.g., dieting, eating disorders).

Method
Division of Child and Adolescent Clinical Neuropsychology at the Department of Child and Adolescent This review concerns new neurobiological insights into
Psychiatry and Psychotherapy, Aachen University Hospital, and Institute of Cognitive Neuroscience,
Institute of Neuroscience and Medicine (INM-3), Forschungszentrum Jülich: Prof. Dr. rer. nat. Konrad typical adolescent behavior and their implications for the
Department of Child and Adolescent Psychiatry and Psychotherapy, Aachen University Hospital: best ways to deal with adolescents. We studied these
Dr. Firk PhD issues with a selective search for relevant publications in
Institute of Neuroscience and Psychology, University of Glasgow: Dr. Uhlhaas PhD German library catalogues, in the PubMed database using

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TABLE

Risky behavior among German adolescents, in percent

Behavioral domains Boys Girls Age range

Substance abuse KiGGS*1:
Current smoking 20.5 20.3 11−17 years
Regular alcohol use 38.6 22.2
Current cannabis use 9.2 6.2
Curent use of other illegal drugs <0.5 <0.5
Delinquency BELLA*2:
Aggressive and dissocial behavior 7.9 7.2 7−17 years
Violence KiGGS*3:
Commission of violent acts 19.6 9.9 11−17 years
Being the victim of violent acts 5.2 3.9
Both of the above 7.6 3.6
Sexual behavior BZgA*4:
Sexual intercourse at or before age 14 14 12 14−17 years
Too early sexual intercourse in own estimation 38 22
No protection the first time 15 9
Sexual intercourse or other sexual conduct against one’s will 3 13
Mental abnormalities BELLA*3:
Depression 5.4 5.3 7−17 years
Anxiety 10.1 10.0
Attention deficit hyperactivity disorder (ADHD) 2.9 1.4
Scholastic problems Shell Youth Study*5:
Ever at risk for repeating a grade 28 24 12−21 years
Repeated a grade 20 14
Physical inactivity
Physically active less than once per week 10.1 21.5 KiGGS*6:
3 or more hours of television watching per day 22.1 23.6 11−17 years
KiGGS*7:
11−17 years
Nutrition KIGGS*8:
Overweight 9.0 8.1 14−17 years
Obese 8.2 8.9
Markedly underweight 2.4 1.4

*1
Lampert and Thamm 2007 (e9); *2Ravens-Sieberer et al. 2007 (e11); *3Schlack and Hölling 2007 (e10); *4BZgA 2006 (e12); *5Shell Deutschland Holding (e13);
*6Lampert et al. 2007 (e14); *7Lampert et al. 2007 (e15); *8Kurth and Schaffrath Rosario 2007 (e16). KiGGS, German Health Interview and Examination Survey for
Children and Adolescents; BELLA, BELLA Study (mental health module within KiGGS), BZgA, German Federal Centre for Health Education.
From (39) Bühler A: Risikoverhalten in der Jugend. In: Uhlhaas PJ. Konrad K (eds): Strukturelle Hirnentwicklung in der Adoleszenz. Stuttgart: Kohlhammer 2011;
189–205. Reprinted with the kind permission of Kohlhammer, Stuttgart

the search terms “adolescence/puberty,” “brain/neural,” as well (11, e1, e2). Thus, the anatomical and
and “development.” Cited publications were also physiological maturation processes that take place in
considered. Special attention was paid to human neuro- adolescence are much more dynamic than originally
imaging studies. thought. It can be concluded that a reorganization of
cortical circuits takes place in adolescence and is re-
Background flected in the changes in cognitive functioning and
Until just a few years ago, there was a general assump- affect regulation that are typical of this period of life
tion in developmental psychology and neuroscience (12).
that major changes in the architecture and functioning Interestingly, this pattern of human brain development
of the brain were limited to the prenatal period and the differs from that of nonhuman primates. Although, for
first five or six years of life. (For a historical overview, example, rhesus monkeys and chimpanzees (like human
see [6].) In the meantime, however, new scientific dis- beings) are born with immature brains, all cortical brain
coveries have compelled a revision of this assumption. areas in macaques mature at the same rate (13). In man,
Large-scale longitudinal studies have shown that a autopsy studies have shown that synaptogenesis reaches a
basic reorganization of the brain occurs during adoles- maximum in the visual and auditory cortices a few months
cence (7). Many synapses are eliminated (8) while, at after birth, while synapses are formed much more slowly
the same time, there is an increase in white matter (9, in the prefrontal cortex. Thus, over the course of human
10), and there are changes in neurotransmitter systems evolution, there was a switch from a synchronous to a

426 Deutsches Ärzteblatt International | Dtsch Arztebl Int 2013; 110(25): 425−31
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heterochronous pattern of cortical development (8). This FIGURE 1

protracted developmental process presumably facilitates
the development of specifically human skills, especially
those acquired through embedding in a highly stimulating
sociocultural environment, e.g., by schooling, music, ver- 30
bal communication, and social interaction (14) (Figure 1).


 2
The current understanding of brain
development in adolescence 15
Brain structure
The brain is fully grown relatively soon after birth, in the
sense that the cerebral cortex soon reaches its maximal
volume. Nonetheless, important structural maturation
0
processes continue to occur in adolescence, as structural 0 1 2.5 5 10 20
imaging studies have shown (15, e3–e5). In the brain, the Years
gray matter matures from back to front, so to speak: The
maximum density of gray matter is reached first in the The development of the prefrontal cortex is protracted in man compared to other primates.
primary sensorimotor cortex and last in higher associ- The Figure shows the synaptic density per 100 µm2 in the prefrontal cortex as a function of
ation areas such as the dorsolateral prefrontal cortex, the age in man (red), chimpanzees (blue), and rhesus macaques (olive green) (error bar = 95%
inferior parietal gyrus, and the superior temporal gyrus. confidence interval). From (40) Lui et al.: Extension of cortical synaptic development distin-
guishes humans from chimpanzees and macaques. Genome Research 2012; 22: 611–22.
This means that, in particular, brain areas such as the
Reprinted with the kind permission of Cold Spring Harbor Laboratory Press, New York
prefrontal cortex—which subserves higher cognitive
functions such as behavioral control, planning, and
assessing the risk of decisions—mature later than the
cortical areas associated with sensory and motor tasks
(16) (Figure 2). At the neural level, functional imaging studies of
Autopsy findings suggest that these gray matter brain development have shown that children and
changes are due to synaptic pruning (17). Many adolescents often have a broader, less focal activation
synapses are formed in childhood that are later pattern than adults, and that the effective recruitment of
removed in adolescence. This occurs in an experience- neural resources increases with age, so that neural ac-
dependent way, i.e., the synapses that survive are the tivity decreases in brain regions other than those that
ones that are more often “in use.” There are also other are relevant to the task at hand (22). It is not yet clear to
cellular mechanisms that might account for gray matter what extent this pattern of neural development is due to
changes in this phase of life, e.g., a reduction in the experience-dependent or biologically determined in-
number of glial cells and an increase of myelination fluences. Imaging studies have also shown that adoles-
(18). cents have heightened activity in limbic areas in
As the gray matter decreases in volume, the white emotional situations: For example, Galvan et al. (23)
matter increases in volume. The white matter is found that the anticipation of a reward is associated
composed of myelinated axons that conduct neural with a more marked activation in the nucleus
information rapidly. The volume of white matter accumbens in adolescents than in children and adults.
increases continually from childhood into early adult- Interestingly, these researchers also found a positive
hood (19). This expansion is presumed to be due, in correlation between activation in the nucleus
large part, to the progressive myelination of axons by accumbens and the adolescents’ individual risk-taking
oligodendrocytes (10). Myelination tends to proceed tendency (24).
from inferior to superior brain areas, and from posterior Moreover, both structural and functional imaging
to anterior. studies have shown that the prefrontal cortex becomes
more strongly linked to sensory and subcortical struc-
Brain function tures during adolescence (25, 26, e6). This implies a
The anatomical reorganization processes of the adoles- greater influence of frontal brain regions on cognitive
cent brain that are described above are associated with and affective processes. The development of cognitive
profound emotional and cognitive changes. In particu- and affective neural circuits should not be regarded as
lar, there is progressive development of executive the sole determinant of structural neurobiological
functions, i.e., cognitive processes that control thought maturation; rather, there appears to be a strong interac-
and behavior and thereby allow the individual to adapt tion of genetic factors with environmental demands.
flexibly to new, complex situational tasks (20). In For example, affect regulation and the brain structures
adolescence, at the same time that these basic cognitive subserving it are influenced by the parent-child interac-
skills are developing, there are also changes in social- tion (27).
affective abilities such as face recognition, the so-called Further findings showing that a profound reorgani-
theory of mind (i.e., the ability to put oneself mentally zation of neural circuitry takes place in adolescence are
in another’s place), and empathy (21). derived from electrophysiological studies, including

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FIGURE 2 electroencephalographic (EEG) studies of changes in

high-frequency and synchronous brain waves (28).
White matter a Brain development in adolescence is associated with a
450 decline of oscillatory activity at rest in the delta (0–3
Hz) and theta (4–7 Hz) bands, and an increase in the
alpha (8–12 Hz) and beta bands (13–30 Hz). With
task-dependent oscillations, the precision of synchro-
400 nization of oscillatory activity in the theta, alpha, and
beta bands increases. The late development of syn-
Volume (cm3)

chronized oscillations in adolescence is closely linked

350 to structural (anatomical) maturation processes as well
as to fundamental changes in neurotransmitter systems,
which have been intensively researched in the past few
years.
300

A neurobiological explanatory model for typical adolescent

behavior
250 One of the more influential neurobiological models to
4 6 8 10 12 14 16 18 20 22
explain typical adolescent behavior was developed by
Age (years) the group of Casey in New York (29, e7) (Figure 3).
The main premise of this model, based on neuroana-
tomical findings and data from functional imaging
Gray matter in frontal cortex b studies (23, 24, 30, 31), is that adolescence is a period
of neural imbalance caused by the relatively early
250
maturation of subcortical brain areas and the relatively
240 delayed maturation of prefrontal control areas (Figure
230 3), with the result that, in emotional situations, the
more mature limbic and reward systems gain the upper
220
hand, so to speak, over the still relatively immature pre-
210 frontal control system. This should not be taken to
Volume (cm3)

200 imply that adolescents are by nature unable to make

rational decisions. Rather, in situations that are particu-
190
larly emotionally laden (e.g., in the presence of other
180 adolescents or when there is the prospect of a reward),
170
the probability rises that rewards and emotions will
affect behavior more strongly than rational decision-
160 making processes (23, 24, 32). This model has been
150 tested in a series of experimental studies (Box).
4 6 8 10 12 14 16 18 20 22 It has been found, for example, that adolescents can
Age (years) assess the risk of certain behaviors just as well as adults
can when asked about them in a questionnaire. On the
other hand, ecologically valid behavioral tests clearly
Male Maximum show that adolescents make more risky decisions in
Female
groups than they do when alone (33). The reason is
95% confidence interval
presumably that, at this age, the benefit of risky behav-
ior—the social approbation of peers—is rated much
Development of the white matter and gray matter of the frontal cortex over a human lifetime; more highly than the risk itself. This may be associated
separate curves for each sex. From (7) Giedd JN, et al.: Brain development during childhood with the nonlinear maturation pattern of the prefrontal
and adolescence: a longitudinal MRI study. Nature Neuroscience 1999; 2: 861–3. Reprinted and limbic brain areas. In accordance with this model,
with the kind permission of Nature Publishing Group, London research on preventive programs has shown that pro-
grams based on imparting knowledge about risks are
less effective than those focusing on individual benefits
and on the training of social competence and resistance
(34).
It is intriguing to ask what functional benefit, if any,
might accrue to the individual from this temporary im-
balance between cortical and subcortical brain struc-
tures. From the evolutionary perspective, adolescence
is the developmental period in which a young person
acquires independence. This process is not unique to

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the human species; increased novelty-seeking and in- FIGURE 3

creased social interactions with other individuals of the
same age can be observed in many other species as well Maturation
(35). Risky behavior among adolescents can be seen as
the product of a biological dysequilibrium between the Limbic reward areas
search for diversion and new experiences (“sensation
seeking”) on the one hand, and as yet immature self-
regulatory capabilities on the other (2); its purpose may Prefrontal cortex
be to enable adolescents to break away from the
familial security zone, so that they can, for example,
find a partner outside their primary family. Immaturity Risk phase
of the prefrontal cortex seems to favor certain types of
learning and flexibility (1).
In fact, over the lifetime of an individual, there are Age
probably multiple developmental windows in which
the brain is particularly well prepared for certain types Adolescence
of learning experience. From the evolutionary perspec-
tive, the cognitive style typical of adolescence, which is Nonlinear maturation processes of subcortical and prefrontal brain
especially sensitive to social-affective stimuli and areas lead to an imbalance of neural networks in adolescence.
Modified from (12) Casey BJ, Jones RM, Hare TA: The adolescent
flexible in the assignment of goal priorities, may be op-
brain. Annals of the New York Academy of Sciences 2008; 1124:
timally suited to the social developmental tasks facing 111–26. Reprinted with the kind permission of John Wiley and Sons
the adolescent. This also implies that the adult brain
cannot be considered the optimal functional system in
an absolute sense, and that adolescence should not be
considered a state of deficient brain performance.

BOX
The influence of pubertal hormones
on adolescent brain development
The maturation of the reproductive system during The influence of peers on risky
puberty is associated with rising concentrations of the behavior
gonadal steroid hormones. The brain has a high density
Researchers recruited persons in three age groups (13 to
of steroid receptors, and it is thus plausible that the sex
16 years, 18 to 22 years, and over 24 years) to study
hormones exert an effect on neural networks in adoles-
whether the influence of contemporaries (peers) on risky
cence. Sisk and Foster (36, e8) have proposed that a
decisions depended on the age of the probands. Partici-
second wave of cerebral restructuring occurs in adoles-
pants were put in a type of driving simulator in which they
cence, building on an earlier, perinatal phase of sexual
had to drive as far as possible until a traffic light turned red
differentiation. According to this model, the hormones
of puberty affect the further structuring of the adoles-
Risky behavior
cent brain, so that a permanent reorganization of the
3.5
brain results, with the effect that neural networks are Age (years)
sensitized to activating hormonal effects. The rising 3 13–16
concentrations of pubertal hormones have different ef- 2.5 18–22
fects on the developing hypothalamic-pituitary-adrenal 24+
(HPA) axis in boys and girls: The rise in androgens in 2
boys apparently inhibits the hypothalamic secretion of 1.5
corticotropin-releasing hormone (CRH), while
1
estrogens in girls regulate the HPA axis upward. Es-
trogens may make girls more susceptible to stress, 0.5
while androgens make boys more resilient to it (37). 0
Alone With peers
Overview
Until now, research on early childhood has received the and a wall appeared. If the car was not stopped soon
most attention from the scientific community and the enough, it crashed into the wall, and the driver lost points.
media. Recent findings show, however, that the The participants were either alone or in groups of three
continuing psychological and biological changes of persons in the simulator. The 13- to 16-year-olds were
adolescence exert a powerful influence on cerebral found to be more likely to make risky decisions than the
structure and function. The brain of the adolescent goes participants in the other age groups, but only in the pres-
through a new phase of plasticity in which environ- ence of their peers. Adult driving behavior was indepen-
mental factors can have major, lasting effects on dent of the presence or absence of peers (33).
cortical circuitry. This opens up new opportunities for

Deutsches Ärzteblatt International | Dtsch Arztebl Int 2013; 110(25): 425−31 429
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education. For example, for the very reason that adoles- account. Major current issues of this type include the
cents are so readily influenced by emotions, they stand question of legalizing cannabis consumption and the
to profit from learning experiences taking place in a applicability of juvenile delinquency law in adoles-
positive emotional context that are intentionally de- cence.
signed to train emotional regulation. Given that risky
behavior in adolescence has a neurobiological basis, Conflict of interest statement
attempts to suppress such behavior completely seem Prof. Konrad has received lecture honoraria from the Medice, Lilly, and Novartis
companies and research support (outside funding) from Vifor Pharma Ltd.
bound to fail. It would be more reasonable to enable
The other authors state that no conflict of interest exists.
adolescents to have emotional experiences in a safe
environment, and to increase the social rewards associ-
ated with non-risky behaviors through regulatory legis- Manuscript submitted on 7 August 2012, revised version accepted on
27 March 2013.
lation (e.g., prohibition of certain kinds of advertising)
and the provision of emotionally positive models. For
Translated from the original German by Ethan Taub, M.D.
instance, the teenage lead character in a television soap
opera might decide to opt out of a hard-drinking contest
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E: The development of neural synchrony reflects late maturation Lehr- und Forschungsgebiet Klinische Neuropsychologie
des Kindes- und Jugendalters
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[email protected]
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MEDICINE

REVIEW ARTICLE

Brain Development During Adolescence

Neuroscientific Insights Into This Developmental Period

Kerstin Konrad, Christine Firk, Peter J. Uhlhaas

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