Social brain development and the affective consequences of ostracism in adolescence.

Abstract

Recent structural and functional imaging studies have provided evidence for continued development of brain regions involved in social cognition during adolescence. In this paper, we review this rapidly expanding area of neuroscience and describe models of neurocognitive development that have emerged recently. One implication of these models is that neural development underlies commonly observed adolescent phenomena such as susceptibility to peer influence and sensitivity to peer rejection. Experimental behavioural evidence of rejection sensitivity in adolescence is currently sparse. Here, we describe a study that directly compared the affective consequences of an experimental ostracism manipulation (Cyberball) in female adolescents and adults. The ostracism condition led to significantly greater affective consequences in the adolescents compared with adults. This suggests that the ability to regulate distress resulting from ostracism continues to develop between adolescence and adulthood. The results are discussed in the context of models of neurocognitive development.

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Review Article
Social brain development and the affective consequences of ostracism
in adolescence
Catherine Sebastiana,*, Essi Vidingb, Kipling D. Williamsc, Sarah-Jayne Blakemorea
aInstitute of Cognitive Neuroscience, UCL, UK
bResearch Department of Clinical, Educational and Health Psychology, Division of Psychology and Language Sciences, UCL, UK
cDepartment of Psychological Sciences, Purdue University, UK
a r t i c l ei n f o
Article history:
Available online 22 July 2009
Keywords:
Adolescence
Ostracism
Social exclusion
Self
Social cognition
Social brain emotion regulation
Puberty
fMRI
Prefrontal cortex
a b s t r a c t
Recent structural and functional imaging studies have provided evidence for continued development of
brain regions involved in social cognition during adolescence. In this paper, we review this rapidly
expanding area of neuroscience and describe models of neurocognitive development that have emerged
recently. One implication of these models is that neural development underlies commonly observed ado-
lescent phenomena such as susceptibility to peer influence and sensitivity to peer rejection. Experimental
behavioural evidence of rejection sensitivity in adolescence is currently sparse. Here, we describe a study
that directly compared the affective consequences of an experimental ostracism manipulation (Cyber-
ball) in female adolescents and adults. The ostracism condition led to significantly greater affective con-
sequences in the adolescents compared with adults. This suggests that the ability to regulate distress
resulting from ostracism continues to develop between adolescence and adulthood. The results are dis-
cussed in the context of models of neurocognitive development.
? 2009 Elsevier Inc. All rights reserved.
1. Introduction
Human adolescence is a period of physical, psychological and
social transition between childhood and adulthood (Spear, 2000).
In recent years it has been established that substantial neural
development also occurs during this period of life (see Paus,
2005 for a review; Gogtay & Thompson, 2010; Paus, this issue;
Giedd & Lenroot, 2010; Schmithorst, 2010). There are significant
changes in grey matter and white matter volumes in brain regions
responsible for complex human behaviours, notably the prefrontal
cortex and temporo-parietal regions (Giedd et al., 1999; Gogtay
et al., 2004; Shaw et al., 2008; Sowell et al., 1999). These regions
are involved in a variety of cognitive functions, including social
cognition, mentalising (the attribution of mental states to oneself
and to other people) and self-related processing. In this paper,
we review developmental functional imaging studies of social cog-
nition, mentalising and self-processing, and discuss recent models
of adolescent neurocognitive development. We then describe a
behavioural study that investigated affective reactions to an in-
stance of experimentally induced ostracism in adolescents, com-
pared with adults. Finally, we evaluate how the results of our
study can inform models of adolescent development.
2. Developmental functional imaging studies of the social brain
2.1. The social brain
The social brain is defined as the network of brain regions sub-
serving social cognition, i.e. those enabling us to recognise others,
and to evaluate our own and others’ mental states (intentions, de-
sires and beliefs), feelings, enduring dispositions and actions
(Brothers, 1990; Frith & Frith, 2007). Many different brain regions
are involved in social cognition, including medial prefrontal cortex
(mPFC), anterior cingulate cortex (ACC), inferior frontal gyrus, pos-
terior superior temporal sulcus (pSTS), temporo-parietal junction
(TPJ), the amygdala and anterior insula (see Fig. 1). Some of these
brain regions are activated during the attribution of mental states
to oneself and to others. This ability, known as mentalising or the-
ory of mind, enables us to understand other people’s behaviour and
actions in terms of underlying mental states such as intentions, de-
sires and beliefs (Frith & Frith, 2007). Social cognitive processes
underlying mentalising range from basic perceptual processes such
as biological motion and face perception (Frith, 2007; Pelphrey &
Carter, 2008) to those enabling us to perceive and understand emo-
tional responses in ourselves and others (Olsson & Ochsner, 2008),
to more abstract meta-representational abilities enabling us to
hold an ‘intentional stance’, i.e. the idea that others’ act on the ba-
sis of their mental states (Dennett, 1987).
Using functional imaging and a wide range of stimuli, several
studies have shown remarkable consistency in identifying the
0278-2626/$ - see front matter ? 2009 Elsevier Inc. All rights reserved.
doi:10.1016/j.bandc.2009.06.008
* Corresponding author. Address: UCL Institute of Cognitive Neuroscience, 17
Queen Square, London, WC1N 3AR, UK. Fax: +44 20 7813 2835.
E-mail address: c.sebastian@ucl.ac.uk (C. Sebastian).
Brain and Cognition 72 (2010) 134–145
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brain regions that are involved in mentalising. These studies have
used stimuli such as stories (Fletcher et al., 1995; Gallagher et al.,
2000; Saxe & Kanwisher, 2003), sentences (den Ouden, Frith, Frith,
& Blakemore, 2005), cartoons (Brunet, Sarfati, Hardy-Bayle, & Dec-
ety, 2000; Gallagher et al., 2000) and animations (Castelli, Happe,
Frith, & Frith, 2000) designed to elicit the attribution of mental
states. In each case, the mentalising task resulted in the activation
of a network of regions including the amygdala, pSTS/TPJ, the tem-
poral poles and the mPFC. Each of these is hypothesised to contrib-
ute to different sub-processes involved in mentalising (Frith,
2007). For example, the amygdala is involved in processing emo-
tional facial expressions (Morris et al., 1998); the pSTS/TPJ is in-
volved in predicting complex movements (Pelphrey, Morris,
Michelich, Allison, & McCarthy, 2005; Saxe, Xiao, Kovacs, Perrett,
& Kanwisher, 2004); the temporal poles are thought to bind highly
processed perceptual inputs with an emotional response (Olson,
Plotzker, & Ezzyat, 2007); and the mPFC may have a special role
in understanding our own and others’ communicative intentions,
particularly the anterior rostral subregion (Frith, 2007).
There is considerable overlap between these social brain re-
gions and regions that are still developing structurally in adoles-
cence. Adolescence is an interesting time to investigate the
development of social cognition because this period of life is char-
acterised by changes in social behaviour (Brown, 2004) and in self-
awareness (Harter, 1990; Sebastian, Burnett, & Blakemore, 2008).
Adolescence is a time during which peers, rather than parents, be-
come influential in shaping social behaviour (Steinberg and Silver-
berg, 1986). In early adolescence, children become increasingly
self-conscious and more aware of, and concerned with, others’
opinions (Parker, Rubin, Erath, Wojslawowicz, & Buskirk, 2006;
Vartanian, 2000). Thus, social brain functions, including mentalis-
ing and self-awareness, might be expected to develop during ado-
lescence. In the next section, we review recent neuroimaging
studies that have investigated the functional development of the
social brain in adolescence.
2.2. Development of the mentalising network
A number of neuroimaging experiments have investigated the
development of mentalising during adolescence and have consis-
tently shown that mPFC activity decreases between adolescence
and adulthood. One fMRI study investigated the development of
communicative intent, using a task in which participants had to
decide whether a speaker was being sincere or ironic (Wang, Lee,
Sigman, & Dapretto, 2006). Understanding irony requires separat-
ing the literal from the intended meaning of a comment. In chil-
dren/young adolescents (aged 9–14) the mPFC and left inferior
frontal gyrus were more active during this task than in adults (aged
23–33). The authors interpreted the increased mPFC activity in
young adolescents as a reflection of the need to resolve the dis-
crepancy between the literal and intended meaning of an ironic re-
mark. The region of the mPFC that was more active in young
adolescents than in adults lies within the dorsal mPFC, an area that
is consistently activated by mentalising tasks in adults (Amodio &
Frith, 2006; Gilbert et al., 2006) (see Fig. 2; green dots).
A similar region of the dorsal mPFC in the right hemisphere was
found to be more active in adolescents than in adults in an fMRI
study that involved thinking about one’s own intentions (Blake-
more, den Ouden, Choudhury, & Frith, 2007). Adolescents (aged
12–18) and adults (aged 22–38) were presented with scenarios
about intentional causality (involving intentions and consequential
actions) or physical causality (involving natural events and their
consequences). The right dorsal mPFC was more active in adoles-
cents than in adults during intentional causality relative to physi-
cal causality (Fig. 2; blue dots). Conversely, a region in the right STS
was more active in adults than in adolescents when they were
thinking about intentional causality compared with physical cau-
sality. In this intentional causality study, the scenarios pertained
to the self insomuch as they asked about participants’ own hypo-
thetical intentions. In another developmental study that focused
on the processing of self-related sentences (Pfeifer, Lieberman, &
Dapretto, 2007), children (aged 9.5–10.8) and adults (aged 23–
31.7) read phrases about academic skills and social competence.
In the self condition, participants were asked to indicate whether
the phrases accurately described them. In the other condition they
were asked to indicate whether the phrases accurately described a
fictional, familiar other person (Harry Potter). The mPFC and ACC
were more active in the children than in adults during self-knowl-
edge retrieval compared with other-knowledge retrieval (see
Fig. 2; yellow dots). The authors suggested that, compared with
adults, early adolescents might rely more on ‘on-line’ self-reflec-
tive processing performed by the mPFC.
Fig. 1. Regions of the social brain. Studies using social cognitive tasks show consistent activation of a network of brain regions including the medial prefrontal cortex (mPFC),
temporo-parietal junction (TPJ), posterior superior temporal sulcus (pSTS), amygdala, anterior cingulate cortex (ACC), anterior insula (AI), inferior frontal gyrus (IFG) and
interparietal sulcus (IPS). See Frith and Frith (2007) for reviews of the function of these regions. Reproduced, with permission, from Blakemore (2008).
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To summarise, as yet there have only been a handful of develop-
mental neuroimaging studies of mentalising. However, there does
seem to be some consistency: the studies to date point to a devel-
opmental shift in the neural strategy for mentalising. In particular,
the studies have found that activity in mPFC decreases between
adolescence and adulthood. There are several different explana-
tions for changes in neural activity during social cognition tasks.
One possibility is that adolescents and adults use different cogni-
tive strategies to perform social cognition tasks. Another possibility
is that the functional differences are a consequence of the struc-
tural development that takes place during this period of life (see
Blakemore, 2008, for discussion).
2.3. Behavioural development of mentalising during adolescence
In the developmental neuroimaging studies of mentalising re-
viewed above, which showed that activity in mPFC decreased be-
tween adolescence and adulthood, task performance was equated
across age groups. Equating performance between groups is critical
for the interpretation of the functional neuroimaging data: if per-
formance between groups was significantly different, it would be
impossible to know whether a group difference in neural activity
was the cause, or simply a consequence of, the difference in perfor-
mance. However, matching performance in this way negates
important differences between adolescents and adults in terms of
social cognition. If the neural substrates for social cognition change
during adolescence, what are the consequences for social cognitive
behaviour? Most developmental studies of social cognition focus
on early childhood, possibly because children perform adequately
in even quite complex mentalising tasks by age five (Frith & Frith,
2007). It is a challenge therefore to design a task on which older
children and adolescents do not perform at ceiling level.
Recently, we used a computerised version of a mentalising task
on which even adults make significant errors (Keysar, Lin, & Barr,
2003). We gave our version of this task to 177 female participants
divided into five age groups: child I (7.3–9.7 years); child II (9.8–
11.4); adolescent I (11.5–13.9); adolescent II (14.0–17.7) and
adults (19.1–27.5) (Dumontheil, Apperly, & Blakemore, in press).
Participants viewed a set of shelves containing objects, which they
were instructed to move by a ‘‘director” who could see some but
not all of the objects. In the critical condition, participants need
to use the director’s perspective and move only objects that the
director can see, in order to make the correct response. The task in-
volves both mentalising (taking another’s perspective), and execu-
tive functions (the need for speeded response selection). The
results demonstrated improvement on this task even between
the adolescent II and adult groups. Thus, while theory of mind
tasks are passed by age four, these new data indicate that the inter-
action between theory of mind and executive functions continues
to develop in late adolescence (Dumontheil et al., in press).
2.4. Development of the affective processing network
The social brain also includes regions involved in processing
information about the affective states of self and other. However,
not all affective processing involves social stimuli, and a distinction
can be drawn between social emotions, which require the under-
standing of mental states (e.g. embarrassment and guilt), and basic
emotions, which do not (e.g. disgust, fear). A recent fMRI study
investigated changes during adolescence of the neural processing
of social emotion compared with basic emotion (Burnett, Bird,
Moll, Frith, & Blakemore, in press). Adult (age 22–32) and adoles-
cent (age 10–18) participants read scenarios that described either
social emotions (guilt or embarrassment) or basic emotions (fear
or disgust). Like the fMRI studies of mentalising reviewed above,
activity in the dorsal MPFC during social relative to basic emotion
was higher in the adolescent group than in the adult group (Fig. 2,
pink dot), while the opposite developmental pattern was found in
the left temporal pole.
One of the most important ways in which we have access to
others’ emotions is through interpreting facial expressions. Several
behavioural studies have shown developmental changes in this
ability during the course of adolescence (Herba, Landau, Russell,
Ecker, & Phillips, 2006; McGivern, Andersen, Byrd, Mutter, & Reilly,
2002; Thomas, De Bellis, Graham, & LaBar, 2007), and functional
neuroimaging studies have also found evidence of developmental
change. Thomas et al. (2001) compared young adolescents (mean
age 11 years) with adults using fMRI during the passive viewing
of fearful and neutral faces. They found that while adults activated
the amygdala only to fearful faces, adolescents activated the amyg-
dala more to neutral faces, possibly because the latter were more
ambiguous, or possibly because the amygdala is less selective ear-
lier in development. However, there are conflicting findings in this
area. For example Guyer et al. (2008) found greater amygdala acti-
vation to fearful faces in adolescents (aged 9–17 years) than in
adults. Development of prefrontal cortex response to faces has also
been found. Yurgelun-Todd and Killgore (2006) reported increased
activity in a number of lateral and superior prefrontal regions
(bilaterally for girls and right sided for boys) in response to fearful
faces between the ages of 8 and 15. Thus, frontal activity increased
between childhood and adolescence in this study. It would be
interesting to compare adult responses within a similar design,
as it might be predicted that PFC activity would decrease again,
in line with studies of mentalising.
Another important social cognitive skill is the ability to allocate
attention appropriately in a socio-emotional context. In one study,
adolescents (aged 9–17 years) showed activation of the ACC and
left OFC during passive viewing of fearful faces relative to neutral
Fig. 2. Activation of the medial prefrontal cortex (mPFC) during mentalising tasks
decreases during adolescence. The yellow shaded area indicates a section of the
dorsal mPFC that is activated in studies of mentalising. The coloured dots indicate
voxels in which decreased activity is observed between late childhood and
adulthood during participation in social cognition tasks. The red dot represents
the area of activation that was higher in adolescents than in adults during the
animations task used by Wang et al. (2006). The green dots represent areas that
were more active in adolescents than in adults during an irony-comprehension task
(Moriguchi, Ohnishi, Mori, Matsuda, & Komaki, 2007). The blue dots represent areas
of activation that were higher in adolescents than in adults during intention
understanding (Blakemore et al., 2007). The yellow dots represent areas that were
more active in children than in adults in a self–other evaluation task (Pfeifer et al.,
2007). The pink dot represents the area of activation that was higher in adolescents
than in adults in a social emotion task (Burnett et al., in press). The blue lines
indicate approximate borders between Brodmann areas, which are numbered on
the diagram. Adapted, with permission, from Blakemore (2008). (For interpretation
of the references to colour in this figure legend, the reader is referred to the web
version of this article.)
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faces, whereas adults (aged 25–36) did not (Monk et al., 2003).
When attention was directed to a non-emotional aspect of fearful
(relative to neutral) faces, activity in the ACC was higher in adoles-
cents compared with adults. Therefore, as in the mentalising stud-
ies reviewed above, frontal activity tended to decrease between
adolescence and adulthood. In addition, the findings suggest that,
whereas adults modulate brain activity based on attention de-
mands, adolescents modulate activity based on the emotional nat-
ure of a stimulus. This suggests that the neural basis of the ability
to pay attention to a non-salient stimulus (in this case, the nose of
a fearful face) in the presence of emotionally evocative, attention-
grabbing stimuli (the eyes of a fearful face) is still undergoing mat-
uration between adolescence and adulthood.
Related to this is the ability to regulate one’s own emotions,
which is a prerequisite for successful social interaction. Studies
in adults have shown that the regulation of emotions depends on
the PFC, particularly ventrolateral regions, and its connections with
limbic regions such as the amygdala (see Ochsner & Gross, 2005,
2007 for a review). Several studies have suggested functional
change in this network between childhood and adulthood. Lév-
esque et al. (2004) found that 8–10 year olds activated more pre-
frontal loci than adults when asked to suppress emotional
reactions to sad film clips (Lévesque et al., 2003) (though note that
the children and adults were not directly compared in the same
study).
More recently, Hare et al. (2008) scanned children, adolescents
and adults (age range 7–32) during a go-nogo task involving fear-
ful, happy, and calm facial expressions. Amygdala reactivity to
fearful faces was greater in adolescents than in younger or older
participants, and correlated with reaction time delays to fearful
compared with happy expressions. In contrast, ventral PFC activity
did not differ with age. Activity in ventral PFC was negatively cor-
related with reaction time difference, suggesting a regulatory role,
and stronger amygdala–ventral PFC connectivity was associated
with greater amygdala habituation over trials. The authors argue
that increased limbic activation relative to ventral PFC regulation
could contribute to the increased emotional reactivity and poor
decision making associated with adolescence.
Another approach is to look at how a measured behavioural dif-
ference in putative affective processing is reflected at the neural le-
vel. Grosbras et al. (2007) divided young adolescents (mean age
10 years), into groups with Low and High resistance to peer influ-
ence, as measured by the Resistance to Peer Influence (RPI) ques-
tionnaire (Steinberg & Monahan, 2007). These groups were then
compared in terms of their neural response to emotional and neu-
tral stimuli (passive viewing of angry and neutral hand gestures/fa-
cial expressions). For the angry stimuli only, individuals with High
RPI showed more co-ordinated activity across premotor and pre-
frontal regions than those with Low RPI. The authors suggested
that High RPI adolescents may spontaneously engage executive
processes in response to socially relevant stimuli, while Low RPI
adolescents do not. By taking an individual differences approach,
this study was able to suggest a neural mechanism that might ac-
count for differences in sensitivity to peer influence in early
adolescence.
While most of the above studies have used stimuli that tap into
a specific affective process (e.g. face or gesture processing), Guyer,
McClure-Tone, Shiffrin, Pine, and Nelson (in press) used an ecolog-
ically valid chat-room paradigm in order to investigate neural re-
sponses to anticipated peer evaluation in adolescents aged 9–17.
Participants were asked to think about how peers in the chat-room
would evaluate them, and these peers had previously been rated as
being of either high or low interest to the participant. For peers of
high interest, relative to low, significant interactions between age
and sex were seen. Specifically, in females only, there was increas-
ing activation with age in regions involved in affective processing
including the nucleus accumbens, hypothalamus, hippocampus
and insula. This differential response between males and females
mirrors behavioural data showing greater concern regarding peer
evaluation among adolescent females than males (La Greca & Lo-
pez, 1998), and neural data showing increasing sensitivity to social
stimuli with age in female adolescents (McClure et al., 2004).
To summarise, there is some indication of functional develop-
ment in brain regions involved in processing social stimuli be-
tween adolescence and adulthood. In particular, PFC activity
during mentalising and face-processing tasks decreases with age,
while PFC regions involved in emotion regulation exert increasing
top–down control between adolescence and adulthood. In contrast,
regions such as the amygdala, which are involved in the initial
emotional response to a stimulus may be more active and less
selective during adolescence than at other points during the life-
span. They may also show increased modulation by social salience,
for example in the chat-room paradigm in adolescent females. In
the following section, we describe recent models of adolescent
neurocognitive development that have been proposed to link
neurobiological changes initiated at the onset of puberty with so-
cio-emotional development. These models provide an important
theoretical framework in what has previously been a predomi-
nantly data-driven field.
3. Models of adolescent social neurocognitive development
Recently, several cognitive models have been proposed to ac-
count for behaviour associated with neurocognitive development
during adolescence. These focus on the links between the develop-
ment of executive functions (enabling flexible behaviour in pursuit
of a goal) and aspects of social cognition such as affect regulation,
with a view to explaining adolescent behavioural phenomena such
as risk-taking in the presence of peers. These models have yet to
incorporate explicitly the development of mentalising.
3.1. The Social Information Processing Network (SIPN) model
The SIPN model posits that social information processing occurs
by way of three interacting neural ‘‘nodes”, which afford the detec-
tionof socialstimulithatarethenintegratedintoa largeremotional
and cognitive framework (Nelson, Leibenluft, McClure, & Pine,
2005). The detection node, comprising the intraparietal sulcus, the
STS, the fusiform face area as well as temporal and occipital regions,
decipherssocialpropertiesofthestimulussuchasbiologicalmotion
(Haxby, Hoffman, & Gobbini, 2002; Pelphrey & Carter, 2008). The
affective node, comprising limbic areas including the amygdala,
ventral striatum, hypothalamus and OFC, is thought then to process
the emotional significance of the social stimulus (Whalen et al.,
1998; Winston, Strange, O’Doherty, & Dolan, 2002). Finally, the cog-
nitive-regulatory node, consisting of much of the PFC, is responsible
for response inhibition, goal-directed behaviour and complex social
behaviours (Frith, 2007). Changes in social cognitive behaviour dur-
ingadolescenceareproposedtoresultfromtheremodellingofthese
networks, particularly the affective and cognitive-regulatory nodes,
and connectivity between nodes. Remodelling may result in part
fromtheeffectofpubertalgonadalsteroidsonlimbicregions,which
are densely innervated by gonadal steroid receptors (Ernst, Romeo,
& Andersen, 2009; McEwen, 2001; Nelson et al., 2005); and partly
from the gradual maturation of the prefrontal cortex, which contin-
uesintothelateteensandearlytwenties(Gogtayetal.,2004;Sowell
et al., 1999).
The focus of the SIPN model is on how social stimuli are imbued
with emotional significance. The mentalising studies described
above (which often do not include an affective component) suggest
that the development of non-affective strategies for understanding
our own and others’ mental states may also represent an important
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contributor to changes in social cognition and behaviour during
adolescence.
3.2. Subcortical/cortical developmental mismatch models
Casey, Jones, and Hare (2008) have noted that children (and
adults) do not exhibit heightened emotional sensitivity or in-
creased risk-taking as do adolescents. This is suggestive of a non-
linear maturational trajectory for the brain networks subserving
these processes. In a recent review, these researchers suggested
that the limbic system (including the amygdala and nucleus
accumbens) matures earlier than the PFC, with the result that indi-
viduals are more greatly affected by the emotional context when
making decisions. The greatest mismatch in the development of
these systems occurs during adolescence.
A similar model proposed by Steinberg (2005, 2008), focuses on
the importance of social reward during adolescence. According to
this model, the changing reward value of positive social feedback
from peers and the development of executive functions interact
to explain well-documented phenomena such as adolescent risk-
taking and susceptibility to peer pressure. Remodelling of the
dopamine system (Andersen, Thompson, Rutstein, Hostetter, & Tei-
cher, 2000; Teicher, Andersen, & Hostetter, 1995) is hypothesised
to increase the salience of social rewards such as peer approval,
while gonadal steroid release is suggested to lead to an increasing
sensitivity to social stimuli, via effects on oxytocin receptors (Chib-
bar, Toma, Mitchell, & Miller, 1990; Insel, Young, Witt, & Crews,
1993). Concurrently, there is relatively gradual development of
prefrontal cognitive control mechanisms, and of connectivity be-
tween prefrontal and limbic regions (Hare et al., 2008). Therefore,
the PFC may not be as efficient at regulating social and emotional
responses as at other points in the lifespan.
An important strength of both of these models is that they sug-
gest neurobiological explanations for why adolescents may be par-
ticularly sensitive to social reward, and why this might result in
increased risk-taking. Self-report studies have shown that adoles-
cents find spending time with peers particularly rewarding and
are particularly influenced by their peers (Csikszentmihalyi, Lar-
son, & Prescott, 1977; Larson & Richards, 1991). Susceptibility to
peer influence is thought to contribute to adolescents’ greater pro-
pensity to engage in risky activities, compared with other age
groups (Steinberg, 2005, 2008). Empirical support for this theory
comes from a recent behavioural study measuring the incidence
of risky driving events in a car simulation video game, in which
adolescents and adults played either alone or with two friends
present. For adolescents, the presence of peers more than doubled
the number of risks taken, whereas for adults the presence of peers
had little effect on risky driving (Gardner & Steinberg, 2005).
This result has been interpreted as reflecting increased reward
salience of peers during adolescence, mediated by the remodelling
of dopaminergic circuitry (Steinberg, 2008). However, it is possible
that the continuing development of social cognitive abilities such
as mentalising (and its neural substrates) may also contribute.
For example, as individuals improve in their ability to represent
abstract social goals and the mental states of others during adoles-
cence, they become more aware of others’ reactions to them (Par-
ker et al., 2006). They also become more aware of the importance
of succeeding in social situations, and of the social costs of failure
(Davey, Yücel, & Allen, 2008). This might contribute to the in-
creased salience of peers, though this may not be restricted to re-
ward salience but would also stretch to include an increased
salience of potential negative consequences of social interaction.
This increased salience could be further compounded by the rela-
tive immaturity of prefrontal regions involved in emotion regula-
tion, resulting in heightened affective responses to negative
social interactions.
Indeed, another well-documented phenomenon during adoles-
cence is sensitivity to peer rejection (Kloep, 1999; Larson & Rich-
ards, 1994; O’Brien & Bierman, 1988). All the models discussed
are compatible with the hypothesis that brain development might
underlie this sensitivity. However, to date, studies have not ex-
plored rejection sensitivity empirically, comparing adolescents
with adults to ascertain whether affective consequences of a rejec-
tion episode really are greater during adolescence, or whether the
phenomenon could result from differences in environment (or
some other systematically varying external factor) with age. In
the next section, we describe a study that investigated the affective
consequences of ostracism in adolescence.
4. Ostracism in adolescence
As peer relationships become more important in adolescence,
the potential negative consequences of rejection or victimisation
by peers increase. For example, Crick et al. (1999) have studied
the effects of relational aggression (aggression based on damage,
or the threat of damage, to interpersonal relationships) on children
and adolescents. Often this takes the form of socially excluding the
victim using the ‘silent treatment’, or by spreading rumours about
the victim. Being a victim of relational aggression is associated
with social-psychological adjustment problems, including inter-
nalising problems (e.g. depression), and externalising problems
(e.g. lack of self-control).
Several studies using self-report methods have found hypersen-
sitivity to peer acceptance and rejection in adolescents compared
with younger children or adults (Kloep, 1999; Larson & Richards,
1994; O’Brien & Bierman, 1988). O’Brien and Bierman (1988) stud-
ied both preadolescent and adolescent attitudes to peer relation-
ships and found that, while both groups felt that peers provided
companionship and support, adolescents (aged 13–17) reported
that peer evaluations were more important in determining their
sense of personal self-worth than did younger children. Peer rejec-
tion was commonly viewed as an indication of their ‘unworthiness’
as an individual, and this peer effect on self-evaluation was most
apparent in girls aged 13–15. This sensitivity appears to decline be-
tween mid-adolescence and adulthood. For example, Kloep (1999)
found that the extent to which adolescent girls worried about peer
acceptance reached a peak at age 15–16, and declined sharply
thereafter.
Research in adults has led to the development of models of the
psychological consequences of ostracism (see Williams, 2007 for a
review). Most of these have necessarily focused on short term, iso-
lated instances of exclusion by others. Williams (1997, 2001) pro-
posed a need threat account, in which ostracism threatens four
fundamental psychological needs: self-esteem, belonging, control
and a sense of meaningful existence. Several studies have found
that these needs are threatened (a construct termed ‘need threat’)
following ostracism (Williams, Cheung, & Choi, 2000; Zadro, Bo-
land, & Richardson, 2006; Zadro, Williams, & Richardson, 2004),
and that people attempt to refortify these needs, for example, by
conforming more to group norms (Williams et al., 2000). This the-
ory has been generated on the basis of data from adult participants,
and it is unknown whether adolescents react to ostracism in the
same way.
The current study examined the affective consequences of an
experimental ostracism manipulation in adolescents compared
with adults. Ostracism was manipulated using Williams et al.’s
(2000) ‘Cyberball’ paradigm. In this task, participants play a ball
game over the internet with two other players, whose actions are
pre-programmed. However, participants are led to believe they
are playing with real individuals. After a few throws, the other
players stop throwing the ball to the participants. This paradigm
was chosen because it reliably induces feelings of rejection in
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adults (Eisenberger, Lieberman, & Williams, 2003; Williams et al.,
2000; Zadro et al., 2004, 2006). The affective consequences of
exclusion (low mood and anxiety) were measured via self-report.
Additionally, we compared adults and adolescents on Williams
et al.’s (2000) need threat scale.
In the current study we chose to test females for a number of
reasons. Social anxiety has been found to be greater in adolescent
girls than in boys, with higher social anxiety levels associated with
poorer social functioning (La Greca & Lopez, 1998). This is likely re-
lated to the greater extent to which self-worth is determined by
positive peer evaluations in adolescent girls than boys (O’Brien &
Bierman, 1988). Furthermore, it has been found that girls are both
more likely to use (Cairns, Cairns, Neckerman, & Ferguson, 1989)
and to be subjected to (Crick & Nelson, 2002), relational aggression,
and that adjustment problems are more strongly related to rela-
tional aggression in girls than in boys. Finally, adolescence is a
key time for the emergence of mood disorders such as depression
and anxiety (Costello et al., 2002; Paus, Keshavan, & Giedd, 2008),
and it is at this time that the pattern of higher incidence in girls
compared to boys is first seen (Angold et al., 1998). Therefore, it
is particularly important to characterise social information pro-
cessing and self-perception in adolescent girls.
To investigate whether the effects of ostracism change during
the course of adolescence, we included two adolescent groups,
which differed in terms of pubertal stage (verified by question-
naire): young adolescents (aged 11–13) and mid-adolescents (aged
14–15). Previous studies looking at the importance attached to
peer evaluations in every-day life have shown that it peaks be-
tween 13 and 16 years in girls (Kloep, 1999; O’Brien & Bierman,
1988). This study aimed to explore whether a similar trend would
be seen in an experimental paradigm. We predicted that the conse-
quences of social rejection on affective measures of mood and anx-
iety would be greater in adolescents than in adults on the basis of
previous reports of hypersensitivity to rejection in adolescence.
Similarly, the four needs might also be differentially threatened
by ostracism between the groups. However, an alternative possi-
bility is that the four needs would be threatened equally across
all age groups. One study (Zadro et al., 2006) has shown that the
extent to which the four needs are threatened is not moderated
by individual differences such as social anxiety immediately fol-
lowing ostracism, and that it is only after some time (45 min) that
differences in coping strategy based on individual differences
emerge. Therefore, the study further aims to explore whether there
are developmental differences in the initial experience of need
threat.
5. Methods
5.1. Participants
The study included 77 female participants divided into three
groups: young adolescents (YA) (N = 26, age range: 11.9–13.9,
mean = 12.8, SD = .59); mid-adolescents (MA) (N = 25, age range:
14.0–15.8, mean = 15.0, SD = .53), and adult (N = 26, age range:
22.2–47.1, mean = 27.4, SD = 6.2).
complete an adapted version of a developmental questionnaire
(Carskadon & Acebo, 1993). Only 61% of participants returned their
questionnaires; therefore Tanner stage data were used to confirm a
developmental difference between the groups, and was not used a
variable in subsequent analyses. Approximate Tanner stages were
calculated from the questionnaire data; the two adolescent groups
differed by one Tanner stage with a mean of 2.47 (SD = .67) in the
YA group and 3.36 (SD = .84) in the MA group.
All three groups were matched on standard age-appropriate
measures of verbal ability (one-way ANOVA: F(2, 74) = 1.31,
p = .28). Adolescents were tested using the British Picture Vocabu-
Participantswere askedto
lary Scale (BPVS-II: Dunn, Dunn, Whetton, & Burley, 1997). Adults
completed the National Adult Reading Test (NART-2; Nelson & Wil-
lison, 1991). Age Group means were YA: 116.3 (SD = 14.87), MA:
121.3 (SD = 17.43), and adult: 115.7 (SD = 5.58). Most of the adults
in our sample were university students, and the school from which
the adolescents were drawn is an academically selective school
from which over 90% go onto higher education. This suggests that
the groups were of similar socio-economic background. No partic-
ipant had a history of neurological or psychiatric disorder, deter-
mined by self and/or parent report.
5.2. Design
The design was a 3 ? 3 factorial with factors Age (YA, MA, A-
dult) and Condition (baseline, inclusion, ostracism).
5.3. Ostracism manipulation
Cyberball (Williams & Jarvis, 2006; Williams et al., 2000) was
used. In order to deflect attention from the true motivation of
the study, participants were told that the aim of the task was to
look at ‘mental visualisation ability’. Ethical permission for this
minor deception was obtained from the local research ethics
committee.
Players were represented on the computer screen by cartoon
drawings, with the participant’s character always located at the
bottom centre (see Fig. 3a). They could choose to throw the ball
to the players on either their left or their right by pressing corre-
sponding keyboard buttons. The game comprised 70 throws, last-
ing around 3 min. The probability that the other players would
throw the ball to the participant systematically varied according
to condition. Inclusion always preceded ostracism in order to avoid
negative spill-over effects, which would have been theoretically
more problematic than spill-over effects from inclusion in terms
of the hypotheses. This same fixed-order strategy has been used
whenever Cyberball has been employed in within-subjects designs
(Eisenberger, Way, Taylor, Welch, & Lieberman, 2007; Eisenberger
et al., 2003). In the inclusion condition, participants were in pos-
session of the ball 33% of the time (equal inclusion). In the ostra-
cism condition the confederates were initially programmed to
throw the ball to the participant with equal probability; however,
after the first eight throws, they stopped throwing it to the partic-
ipant altogether for the remainder of the game (around 50 throws).
5.4. Dependent measures
5.4.1. Mood
Participants rated how good/bad, happy/sad, friendly/un-
friendly and tense/relaxed they were currently feeling, on a scale
of 1–7. These anchors comprised the mood section of the need
threat questionnaire, devised by Williams et al. (2000). Mood rat-
ings were taken at baseline (before Cyberball) as well as after
inclusion and ostracism.
5.4.2. Anxiety
Anxiety levels were measured using the state/trait anxiety
inventory (Spielberger, 1983). This consists of 20 statements for
each subscale (State (STAI-S) and Trait (STAI-T)). Participants rated
how much each statement described them on a scale from 1 to 4.
Trait scores were measured only at baseline, while state anxiety
was measured both at baseline, and after inclusion and ostracism.
5.4.3. Need threat
The extent to which inclusion and ostracism affected the four
needs was measured using the need threat questionnaire (Wil-
liams et al., 2000). The need threat scale consists of 12 statements;
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three questions pertaining to each need (self-esteem, belonging,
control and meaningful existence). Participants rated how much
each statement described their reaction during Cyberball on a scale
from 1 to 5.
5.4.4. Manipulation check
At the end of the experiment, we wanted to be sure that partic-
ipants realised that they had been included/ostracised. Participants
rated the truth of two statements on a scale from 1 to 5 for each of
the two times they had played Cyberball (‘‘I was ignored” and ‘‘I
was excluded”). They were also asked to estimate what proportion
of the time they had received the ball, given that 33% was an equal
share.
5.5. Procedure
See Fig. 3b for a schematic of the experimental procedure. Par-
ticipants initially completed a measure of verbal ability; then base-
line self-report measures. These consisted of current mood ratings,
and the STAI. Participants then played Cyberball (inclusion condi-
tion). They were told that they would be playing with two other
girls over the internet, and that it was important for them to visu-
alise the experience as much as possible. Afterwards, they filled out
mood, state anxiety and need threat questionnaires. Participants
then played Cyberball in the ostracism condition, this time with
two different ‘girls’, and again completed the questionnaires. At
the end of the experiment, they completed the manipulation check
measures, and were then fully debriefed as to the aims of the
study. It was made clear that the computerised characters in
Cyberball were not real. No participant expressed regret at having
taken part.
6. Results
Data from one adult were excluded from state anxiety analyses,
becauseshedidnotcompletethemeasureafterall threeconditions.
Data were analysed using mixed model ANOVAs, with Age (YA, MA,
adult) as the between-subjects factor, and Condition (either (base-
line, inclusion and ostracism) or (inclusion and ostracism) as appro-
priate) as the within-subjects factor. Corrections for multiple
comparisons were made using Bonferroni corrections, and Green-
house Geisser corrections were used to correct for non-sphericity.
6.1. Manipulation check
All age groups realised that they were included and ostracised
in the appropriate conditions. Overall, participants estimated that
they received the ball 35.01% (SD = 11.72) of the time in the inclu-
sion condition, and 7.66% (SD = 5.39) in the ostracism condition.
These mean estimates were fairly accurate, with actual percent-
ages of 33% and 11%, respectively. In a mixed model ANOVA, there
was a main effect of Condition, with significantly higher estimates
of possession after inclusion than ostracism (F(1, 74) = 475.18,
p < .001, partial eta squared ðg2
of Age, and no Age ? Condition interaction.
Amalgamating responses to the items ‘I felt ignored’ and ‘I felt
excluded’, all participants reported feeling more excluded in the
ostracism condition (M = 4.64, SD = .49) than in the inclusion con-
dition (M = 1.68, SD = .78), (F(1, 74) = 779.95, p < .001, g2
There was no significant main effect of Age, and no Age ? Condi-
tion interaction. As the possible range of scores was 1–5, with five
meaning participants felt excluded, the mean scores suggest par-
ticipants reacted to the two conditions appropriately.
p¼ :87Þ). There was no main effect
p¼ :91).
3b)
IQ test
Baseline measures:
- Mood
- State/Trait anxiety
Cyberball
(Inclusion)
Inclusion measures:
- Mood
- State anxiety
- Need threat
Cyberball
(Ostracism)
Ostracism measures:
- Mood
- State anxiety
- Need threat
- Manipulation check
Debriefing
Timeline
3a)
Fig. 3. The Cyberball game and a schematic of the experimental procedure. (3a) The Cyberball game: participants are represented by a cartoon at the bottom of the screen,
and computerised characters stand on either side. In the ostracism condition, the other characters do not throw the ball to the participant. (3b) A schematic of the time course
of the experiment. On arrival, participants completed an IQ test and the baseline measures. They then played Cyberball (first inclusion, then ostracism). After each run of
Cyberball they completed a battery of self-report measures.
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6.2. Mood
For two of the mood anchors (good/bad and happy/sad), and
for overall mood, there was a main effect of Condition (good/
bad: F(1.60, 118.14) = 15.09, p < .001, g2
112.91) = 10.91, p < .001, g2
117.46) = 16.29, p < .001, g2
cantly lower mood after ostracism than after either baseline or
inclusion (all ps < .008). There were no main effects of Age. There
were significant interactions between Age and Condition for these
three anchors (good/bad: F(3.19, 118.14) = 3.01, p = .03, g2
happy/sad: F(3.05, 113) = 2.78, p = .044, g2
F(3.18, 117.5) = 3.00, p = .031, g2
showed that in all cases the interaction was due to significantly
lower mood after ostracism than baseline and inclusion in both
adolescent groups (all ps < .05). In contrast, there were no differ-
ences between conditions for adults (Fig. 4).
The remaining mood anchors (friendly/unfriendly and tense/re-
laxed) did not show an interaction between Condition and Age. For
friendly/unfriendly, there was a main effect of Condition (F(1.57,
116) = 21.45, p < .001, g2
cantly less friendly after ostracism compared with both baseline
and inclusion (all ps < .001), but no main effect of Age. For tense/
relaxed, there was a main effect of Condition (F(1.79, 132.72) =
5.11, p = .009, g2
nificantly less relaxed after ostracism in all age groups compared
with baseline (p = .022), but no significant difference between
ostracism and inclusion. There was also a main effect of Age
(F(2, 74) = 5.08, p = .009, g2
nificantly less relaxedthanadults
M = 3.84, SD = 1.49; Adults: M = 4.81, SD = 1.44; p = .011), with a
marginal effect in the same direction for YA (YA: M = 4.23,
SD = 1.61; p = .051).
p¼ :17); happy/sad: F(1.53,
andoverall mood:
p¼ :18). Post-hoc tests showed signifi-
p¼ :13)
F(1.59,
p¼ :08;
p¼ :07 and overall mood:
p¼ :08). Simple effects analyses
p¼ :23), due to participants feeling signifi-
p¼ :12), due to participants reporting feeling sig-
p¼ :12). This was due to MA being sig-
across conditions(MA:
6.3. State and trait anxiety
A one-way ANOVA comparing Age groups found no differences
in trait anxiety (measured at baseline).
For state anxiety, a mixed model ANOVA showed that there was
a main effect of Condition (F(1.7, 124.23) = 8.96, p < .001, g2
and a main effect of Age (F(2, 73) = 7.60, p = .001, g2
Fig. 5). Post-hoc tests showed that the main effect of Condition
was due to significantly greater anxiety after ostracism (M = 39.74,
p¼ :11),
p¼ :17) (see
SD = 10.12), compared with both baseline (M = 36.82, SD = 8.15;
p = .014) and inclusion (M = 36.45, SD = 8.70; p < .001). The main
effect of Age was due to significantly greater anxiety in the MA
group (M = 41.72, SD = 7.26) than in adults across conditions
(M = 33.63, SD = 7.89; p = .001). The Age ? Condition interaction
was not significant (F(3.4, 124.23) = 1.22, p = .31, g2
ever, because we had an a priori hypothesis that the effect of ostra-
cism on anxiety would be greater in adolescents than adults, we
conducted planned comparisons, but used Bonferroni correction
to be conservative. The results suggested that the main effect of
Condition was driven by the reactions of the YA group. Anxiety
was significantly higher after ostracism than either at baseline
(p = .034) or after inclusion (p < .001) in this group, while anxiety
in the MA and adult groups did not differ significantly between
conditions.
p¼ :03). How-
6.4. The four needs
For the need threat questionnaire, internal consistency between
the three items for each need (self-esteem, belonging, control, and
meaningful existence) was high (Cronbach’s a all >.80). Therefore a
single mean score was calculated for each need and this was used
in subsequent analyses.
The four needs were only measured after inclusion and ostra-
cism, as the questions pertained directly to the Cyberball experi-
ence. In separate mixed model ANOVAs for each need, there
were main effects of Condition for all four needs, with lower need
fulfilment after ostracism (self-esteem: F(1, 74) = 87.68, p < .001,
g2
F(1, 74) = 57.62, p < .001,
g2
F(1, 74) = 101.17, p < .001, g2
main effect of Age (F(1, 74) = 8.34, p = .001, g2
tests showed that this was due to significantly lower self-esteem
across conditions in the MA group (M = 2.83, SD = .54) compared
to adults (M = 3.39, SD = .69; p < .001). There were no other main
effects of Age, or interactions between Age and Condition.
p¼ :54; belonging: F(1, 74) = 136.81, p < .001, g2
p¼ :65; control:
meaningful
p¼ :44
p¼ :58. For self-esteem, there was a
and
existence:
p¼ :18). Post-hoc
7. Discussion
This study used an experimental ostracism manipulation
(Cyberball) to investigate the hypothesis that adolescents are
* Significant at p < .05, corrected
3.00
3.50
4.00
4.50
5.00
5.50
6.00
YAMA Adult
Group
Overall Mood
Baseline
Inclusion
Ostracism
**
Fig. 4. Overall mood ratings for each group under each condition. Mood was
significantly lowered by the ostracism condition compared with baseline and
inclusion in the two adolescent groups (YA and MA). Mood was lowered by
ostracism in the adult group, but this was not significant.
30
32
34
36
38
40
42
44
46
48
YAMA
Group
Adult
State Anxiety (STAI)
Baseline
Inclusion
Ostracism
*
* Significant at p < .05, corrected
Fig. 5. State anxiety scores for each group under each condition. There was a main
effect of Condition, with higher anxiety after ostracism in all groups. However,
planned comparisons showed that anxiety was significantly greater after ostracism
in the YA group only. There was also a main affect of Group, due to higher mean
state anxiety in the MA group compared to adults across conditions.
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hypersensitive to peer rejection. On some measures, affective reac-
tions to ostracism were greater in adolescents than in adults. We
found evidence for significantly lowered overall mood after ostra-
cism in both YA and MA groups, and significantly higher state anx-
iety in the YA group, but no differences between conditions on
either measure for adults. In contrast, each of Williams’ (1997,
2001) ‘four needs’ (self-esteem, belonging, control and meaningful
existence) was significantly threatened by ostracism in all groups.
Importantly, we found no group differences on any manipulation
check measure. Therefore, we found no evidence of a difference be-
tween the groups’ more general experience of being included and
ostracised.
7.1. Effects of ostracism on self-reported affective measures (mood and
anxiety)
Both YA and MA adolescents rated all moods (good, happy,
friendly and relaxed) as less positive after ostracism compared
with both baseline and inclusion conditions. In contrast, adults felt
less friendly and less relaxed after ostracism, but did not feel any
less good or happy. This stronger effect in the adolescents indicates
hypersensitivity to rejection, and also sheds light on previous stud-
ies looking at the effects of ostracism on mood in adults. Inconsis-
tent results have been seen with regard to mood effects following
ostracism in adults. While some studies have shown lowered mood
after ostracism (Stroud, Salovey, & Epel, 2002; Williams et al.,
2000), others have not (Twenge, Baumeister, Tice, & Stucke,
2001; Zadro et al., 2004). Our finding of lowered mood on only
some anchors (friendly/unfriendly and tense/relaxed) suggests
that not all mood states are equally affected by ostracism in adults.
Therefore, differences in how mood state has been measured in
previous studies may contribute to the inconsistencies seen. The
current data further suggest that adolescents may be more sensi-
tive than adults to the acute effects of short term ostracism on
mood, and that these effects may be more broad in scope, affecting
a wider range of measured mood states.
Following the Cyberball ostracism condition, there was a gen-
eral increase in state anxiety across age groups relative to baseline
and inclusion conditions. However, planned comparisons showed
that this difference was only significant in YA. This suggests a
heightened reactivity to ostracism in this group, particularly com-
pared to adults who reported comparable levels of anxiety to YA at
baseline and after inclusion. Although trait anxiety (measured at
baseline) did not differ between age groups, we did find age differ-
ences in state anxiety at baseline. Specifically, the MA group were
significantly more anxious than adults. This is in line with previous
work showing that social anxiety is at its peak at age 15, which also
corresponds to the peak onset of social phobia (Erath, Flanagan, &
Bierman, 2007; Mancini et al., 2005). This high baseline anxiety
makes anxiety levels reported after inclusion and ostracism diffi-
cult to interpret in MA since it may have masked the effects of
ostracism on anxiety.
7.2. Effects on the four needs
In line with Williams’ need threat model of ostracism, each of
the four needs was significantly threatened by the ostracism con-
dition, regardless of age group. This supports the idea that need
threat is an automatic, reflexive process that occurs independently
of individual differences in variables such as social anxiety (Wil-
liams, 2007; Zadro et al., 2006). The current results further suggest
that developmental stage (at least between adolescence and adult-
hood) does not significantly affect the experience of need threat,
and that increased need threat does not explain increased sensitiv-
ity to social ostracism in adolescence. Although the finding of low-
er self-esteem across conditions in the MA group was not explicitly
predicted, there are studies showing that girls are particularly
likely to suffer decreasing self-esteem over the course of adoles-
cence (Zimmerman, Copeland, Shope, & Dielman, 1997).
7.3. Methodological considerations
While results are consistent with our hypothesis that adoles-
cents would be hypersensitive to the affective consequences of
ostracism, there are alternative interpretations. The use of a
fixed-order within-subjects design meant that participants were
aware of the questionnaire measures that were being employed
following the inclusion condition, i.e. before they were ostracised.
It could be argued that adolescents were more sensitive than
adults to demand characteristics that this situation might create.
However, we think this is unlikely; most notably because adoles-
cents did not respond in the same way on all measures. They
showed hypersensitivity specifically on the affective measures,
but not on the need threat questionnaire. However, future studies
could eliminate this possibility by including physiological mea-
sures of affective arousal such as skin conductance responses or
cortisol levels (Blackhart, Eckel, & Tice, 2007).
In the current study, we did not have an accurate measure of
pubertal stage across all participants, and adolescent groups dif-
fered on both age and pubertal stage. Future studies could explore
sensitivity to ostracism in individuals matched for age but differing
in pubertal status to explore whether the effect is tied to pubertal
development.
7.4. Implications for adolescent social cognitive development
The current data replicate and extend previous findings in a
number of ways. Previous studies have used interviews and expe-
rience-sampling methods to explore how adolescents respond to
peer rejection in the context of their every-day lives (Coleman,
1974; Kloep, 1999; Larson & Richards, 1994; O’Brien & Bierman,
1988). The current study complements these studies, demonstrat-
ing hypersensitivity to general rejection (not necessarily by peers)
in an experimental context in adolescent females. The ostracism
manipulation was brief and without lasting consequences, yet
hypersensitivity, particularly on mood measures, was still seen.
Furthermore, as Cyberball was played over the internet, external
factors such as school environment, social status among peers or
participants’ physical appearance cannot fully account for the ef-
fect. It would be interesting to study whether the same effect
would be seen in adolescent males. A recent study in adults
showed lowered mood in both males and females following a so-
cial rejection task, but only females showed increased cortisol lev-
els, associated with a stress response (Stroud et al., 2002). This
complements developmental self-report studies suggesting that
adolescent females are more upset by negative peer evaluations
and relational aggression than males (O’Brien & Bierman; Crick &
Nelson, 2002). However, adolescent males and females have not
been directly compared on sensitivity to ostracism in an experi-
mental context.
7.5. Implications for models of adolescent neurocognitive development
The results of this study are in line with predictions derived
from the neurobiological models discussed above. Although the
models differ in terms of the precise developmental trajectories
of different neural systems, all have in common the idea that rela-
tively protracted development of prefrontal regulatory systems,
possibly in conjunction with a more rapidly developing limbic sys-
tem, may underlie commonly observed phenomenon such as risk-
taking in the presence of peers. These models would also predict
adolescent hypersensitivity to peer rejection in an experimental
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context. The continued development of mentalising might also
contribute to this effect by increasing the extent to which adoles-
cents are aware of others’ appraisals and of the importance of suc-
ceeding in social situations, and by increasing the extent to which
adolescents use social comparison to build a self-concept (Damon
& Hart, 1988; Davey et al., 2008; Harter, 1990; Parker et al., 2006;
Ruble, Boggiano, Feldman, & Loebl, 1980).
Functional imaging studies are needed in order to elucidate the
neural bases of sensitivity to ostracism in adolescence. In adults,
there is evidence suggesting that the neural substrates for social
pain overlap with those underlying physical pain (Eisenberger et
al., 2003). Using a range of paradigms, evidence has been found
for the involvement of the dorsal anterior cingulate cortex (dACC)
(Eisenberger et al., 2003, 2007), subgenual/ventral ACC (Somer-
ville, Heatherton, & Kelley, 2006), right ventrolateral PFC (Eisen-
berger et al., 2003), medial PFC, posterior cingulate, and insula
(Kross, Egner, Ochsner, Hirsch, & Downey, 2007). These regions
overlap considerably with those continuing to develop structurally
and functionally during adolescence (Giedd et al., 1999; Gogtay et
al., 2004; Shaw et al., 2008). One recent study has shown a similar
pattern of activation in adolescents aged 12.4–13.6 years (Masten
et al., 2009) to that seen in adults on the Cyberball task (Eisenber-
ger et al., 2003, 2007), and additionally showed the involvement of
the subgenual ACC, which was not activated in previous studies of
adults using the same task. However, adults and adolescents were
not compared within the same study, and this will be an important
next step in elucidating the neural correlates of increased sensitiv-
ity to social pain such as ostracism in adolescence.
8. Conclusions
We began this article by summarising recent work that has
explored the link between social cognition and functional brain
development during adolescence. This work suggests that many
aspects of social cognition and their neural substrates, including
face processing, social emotion processing and mentalising, are
still developing during the second decade of life. These findings
are compatible with neurocognitive models of adolescence, and
also make specific predictions about adolescent sensitivity to
negative social experiences. We conducted an experimental
manipulation to explore the affective consequences of ostracism
in adolescence. While the effects of ostracism on need threat
were broadly similar between age groups, as predicted by Wil-
liams’ need threat model, mood was more negatively affected
after an episode of ostracism in both groups of adolescents than
in adults, while anxiety was greater in the YA group. These
experimental findings complement self-report studies showing
hypersensitivity to social rejection during adolescence. The re-
search discussed in this review shows that an iterative approach
between data and theoretical models can be helpful in formulat-
ing and testing experimental hypotheses, particularly within a
relatively new field such as social developmental cognitive
neuroscience.
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