Mutual Liking and Social Anhedonia
Running head: MUTUAL LIKING AND SOCIAL ANHEDONIA
In Press, Brain and Cognition
Social Anhedonia and Medial Prefrontal Response to Mutual Liking in Late Adolescents
Kati L. Healey, Judith Morgan, Samuel C. Musselman, Thomas M. Olino, and Erika E. Forbes*
University of Pittsburgh, Department of Psychology
*Corresponding author. Email: firstname.lastname@example.org. Phone: 412-383-5438
3811 O’Hara St., WPIC—Loeffler 319, Pittsburgh, PA 15213
Mutual Liking and Social Anhedonia 1
Anhedonia, a cardinal symptom of depression defined as difficulty experiencing pleasure, is also
a possible endophenotype and prognostic factor for the development of depression. The onset of
depression typically occurs during adolescence, a period in which social status and affiliation are
especially salient. The medial prefrontal cortex (mPFC), a region implicated in reward, self-
relevant processing, and social cognition, exhibits altered function in adults with anhedonia, but
its association with adolescent anhedonia has yet to be investigated. We examined neural
response to social reward in 27 late adolescents, 18-21 years old, who varied in social anhedonia.
Participants reported their social anhedonia, completed ratings of photos of unfamiliar peers,
and underwent a functional magnetic resonance imaging task involving feedback about being
liked. Adolescents with higher social anhedonia exhibited greater mPFC activation in response
to mutual liking (i.e., being liked by someone they also liked) relative to received liking (i.e.,
being liked by someone whom they did not like). This association held after controlling for
severity of current depressive symptoms, although depressive severity was also associated with
greater mPFC response. Adolescents with higher levels of social anhedonia also had stronger
positive connectivity between the nucleus accumbens and the mPFC during mutual versus
received liking. These results, the first on the pathophysiology of adolescent anhedonia, support
altered neural reward-circuit response to social reward in young people with social anhedonia.
Medial Prefrontal Cortex
Mutual Liking and Social Anhedonia 2
Social Anhedonia and Medial Prefrontal Response to Mutual Liking in Late Adolescents
Anhedonia, or difficulty experiencing pleasure in anticipation or response to rewarding
stimuli, is a cardinal feature of depression. Across the lifespan, depression is a leading cause of
disability and suffering (World Health Organization, 2011). In addition to characterizing
depression, anhedonia is associated with the onset, course, and outcome of the disease. Low
levels of anhedonia in people with depression are protective, presaging better outcomes (Joiner
et al., 2002) and a lower chance of recurrence (Kasch et al., 2002). The clinical relevance of
anhedonia is particularly important in young people, as anhedonia predicts the onset of clinical-
level depression in children and adolescents (Pine et al., 1999; Wilcox & Anthony, 2004) and
resistance to standard depression treatment in adolescents (McMakin et al., 2012). Despite
anhedonia’s associations with the development of depression, a disorder that most commonly
begins during adolescence (Lewinsohn et al., 1994), relatively little work has been done to
examine the neural correlates of anhedonia in adolescence.
Anhedonia is relevant to appetitive and consummatory aspects of reward function, which
reflect the motivation to pursue rewarding experiences and the enjoyment of rewarding
experiences once obtained, respectively. Frontostriatal circuits, including dopamine-targeted
regions such as the ventral striatum (VS) and medial prefrontal cortex (mPFC), play critical
roles in reward function. Specifically, the striatum contributes to the motivation to approach
rewarding stimuli and the enjoyment of those stimuli, and a subregion of the mPFC (i.e.,
Brodmann Areas 24, 32, and medial 10) contributes to the regulation of reward function, partly
through its input to the striatum (Haber & Knutson, 2010). Broadly, meta-analytic evidence
indicates that depression involves low striatal response to monetary reward for youth and adults
(Zhang et al., 2013). More specifically, Wacker et al. (2009) found that self-reported
anhedonia—but not other symptoms of depression—were associated with nucleus accumbens
activation to reward receipt during a monetary incentive delay task. In that study, anhedonia
Mutual Liking and Social Anhedonia 3
was negatively correlated with self-reported positive affect, indicating that affective aspects of
anhedonia are related to neural response to reward.
Consistent with anhedonia’s subjective, behavioral, and neural aspects, adolescents with
depression experience lower subjective positive affect (Lonigan et al., 2003; Silk et al., 2011),
express shorter-duration, lower-intensity positive affect (Sheeber et al., 2009), and exhibit low
striatal response to reward (see Forbes & Dahl, 2012). Specific associations between anhedonia
and neural response to reward have yet to be reported in adolescents, but the literature on
clinical-developmental neuroscience suggest that adolescence is an important developmental
period for investigating anhedonia. Reward-related behavior and affect change markedly
during adolescence, with increased risky, reward-seeking activities (see Somerville et al.,
2010), more sensation-seeking (Steinberg et al., 2008), and stronger experience of rewards
(Ernst et al., 2005; Steinberg et al., 2008). Strikingly, these changes occur in tandem with
apparent decreases in reward responding, including low levels of subjective positive affect
(e.g., Larson et al., 2002), and increasing levels of depressive symptoms (Sawyer et al.,
2009). Developmental neuroimaging studies indicate that adolescents show altered
response to reward in the striatum (Bjork et al., 2004; Ernst et al., 2005; Forbes et al.,
2010; Galván et al., 2006) and mPFC (Bjork et al., 2004; Forbes et al., 2010).
Response in the mPFC may be particularly relevant to adolescent anhedonia. In addition
to its role in reward processing as the destination of the mesocortical dopamine pathway, the
mPFC is a key region in the brain’s default-mode network and is implicated in affect regulation
and in processing social and self-relevant stimuli (Amodio & Frith, 2006; Denny et al., 2012). In
responding to rewarding events, mPFC response could therefore indicate how relevant the
reward is to one’s preferences and whether the reward enhances one’s status among others.
Depression is associated with greater mPFC response to reward, which could reflect processes
such as difficulty shifting out of a negative, self-focused pattern of thought or over-regulation of
more basic, striatal response to reward (Forbes & Dahl, 2012). Notably, anhedonia itself is
Mutual Liking and Social Anhedonia 4
associated with greater mPFC response to positive autobiographical memories but less mPFC
response to negative stimuli (Keedwell et al., 2005b). This pattern is similar to that observed in
adults with depression but the opposite of that exhibited by healthy adults (Keedwell et al.,
2005a). This suggests that neural function underlying anhedonia, especially in the context of
depression, is disrupted in response to what are generally highly valued, self-defining
experiences. Furthermore, this disruption in the mPFC could reflect a pattern of responding to
such typically pleasant, poignant stimuli as somehow aversive.
In addition to specific regional responding, anhedonia is postulated to involve altered
connectivity between the mPFC and VS, such that the mPFC might serve to dampen VS
responses to reward (Forbes & Dahl, 2012). The disruption of functional connectivity between
the mPFC and VS during response to monetary reward in adolescents with a history of
depression (Morgan et al., under review) suggests that such functional connectivity—likely via
altered input to the VS from the ventral tegmental area—could be a mechanism for the
development of anhedonia. Neural response to reward and functional connectivity between
regions in rewarding contexts answer different questions: the first, about response in specific
regions in isolation, and the second, about coordination between regions during specific
contexts. Both of these can address function in reward circuitry, but they provide different
information. Given that previous conceptual models have postulated that both neural response
and frontostriatal connectivity are disrupted in anhedonia and in those who develop depression,
investigations with anhedonia in adolescents are crucial to critically test developmental
Although functional connectivity techniques such as psychophysiological interaction
(PPI) have been traditionally used to examine how regions coordinate in response to task
context, there is a burgeoning mental health literature in which investigations of brain circuitry
in psychopathology include examining whether psychopathology is related to functional
connectivity within the circuitry of interest. For example, studies of affective disorders have
Mutual Liking and Social Anhedonia 5
reported differences between subgroups of patients or between patients and healthy comparison
participants in functional connectivity between the amygdala and orbitofrontal or prefrontal
cortex during the processing of affective faces (e.g., Almeida et al., 2009; Almeida et al., 2011;
Kong et al., 2013; Versace et al., 2010; Wang et al., 2012). Similarly, in schizophrenia, another
disorder that includes the symptom of anhedonia, group differences in functional connectivity
have been reported during experiences such as working memory and social processing (e.g.,
Eack et al., 2013; Mukherjee et al., 2013; Straube et al., 2013). Aside from categorical group
differences, studies of mental health have also employed functional connectivity to examine
function in neural circuitry that varies with a dimensional characteristic, such as neuroticism or
symptom severity (Cremers et al., 2010; Davey et al., 2012; Doucet et al., 2013; Servaas et al.,
2013; Yue et al., 2013). Several studies have now used both approaches, describing functional
connectivity as a correlate of both between-group differences and continuous, within-sample
variability (e.g., Davey et al., 2012). Most relevant to our questions in the current study, a study
of adults recently reported that trait anhedonia is associated with altered response and altered
functional connectivity in contexts involving pleasant musical stimuli (Keller et al., 2013).
Research with adolescents introduces important considerations. One is the definition of
adolescence itself, which has been debated in the fields of psychology, anthropology, and
pediatrics, among others. Adolescence is defined as the period between the end of puberty and
the attainment of adult-level status and competence. Specifying an age range for this
developmental period requires consideration of a variety of factors (e.g., psychological and
biological processes), as well as consideration of the ongoing developmental tasks. For research
purposes, it is particularly important to consider the ongoing developmental tasks that are
relevant to a research question when defining an adolescent population. Given that the
developmental tasks of adolescence are postulated to include impulse control, accurate
assessment of risk vs. reward, and affect regulation during challenges (Hazen et al., 2008) and
that the neural circuitry underlying these cognitive and behavioral functions continues
Mutual Liking and Social Anhedonia 6
development throughout the teen years and into the 20s (Lenroot & Giedd, 2006), we defined
adolescence as occurring through the early 20s. That is, our deliberate focus on a population in
which the processes of interest (i.e., function in reward circuitry, social processing) have not yet
reached adult levels led us to consider this a study of adolescent development. Other approaches
might classify participants over age 18 as adults based on legal or cultural changes in status at
that age (e.g., attaining the right to vote or perform military service). It is also notable that many
psychology studies conducted with undergraduate samples describe their participants as adults
rather than late adolescents, and this practice has been identified as a key limitation for
interpreting findings on constructs that involve self and social cognition (Sears, 1987). Other
terms have also been used for the late adolescent developmental period, such as “emerging
adulthood” (Conger & Little, 2010). Based upon our focus on neural reward circuitry and the
emergence of a symptom relevant to several forms of psychopathology that have onset during
the later stages of brain development, we studied a population from age 18-21 years, termed
hereafter as late adolescents.
In addition, investigating function in reward circuitry during adolescence requires
sufficient context. First, reward circuitry is undergoing substantial development during
adolescence (Spear, 2000). The function of the striatum and mPFC in response to reward
changes with puberty (Forbes, 2009) and with adolescent development (Bjork et al., 2004;
Ernst et al., 2005; Galván et al., 2007). Thus, these brain regions are especially malleable and,
consequently, potentially vulnerable to developing dysfunction during this period (Davey et al.,
2008). Second, the value and salience of social contexts increases as adolescents spend more
time with peers, experience more intense emotions in peer contexts, and pursue newly valued
abstract social rewards (e.g., initiating a romantic relationship; Furman & Simon, 1999; Larson
et al., 1999; Steinberg, 2008). Hence, a class of rewards that is perhaps most important to
anhedonia, to the function of frontostriatal reward circuitry, and to adolescent development is
social reward. In particular, adolescents’ reward circuitry and decision-making behavior are
Mutual Liking and Social Anhedonia 7
sensitive to peer context (Albert et al., 2013; Chein et al., 2011), and peer experiences are
postulated to influence the development of reward circuitry (Albert et al., 2013). Thus, peer
social reward is likely to be salient during this developmental period.
The model presented by Davey et al. (2008) may be further elaborated to emphasize the
role of being accepted by peers when peers’ opinions are highly valued, as opposed to being
minimally valued or ambiguous. Accordingly, Davey and colleagues (2010) developed a social
reward paradigm in which participants view images of unfamiliar peers and rate how much they
thought they would like the peer. Participants are led to believe that their own images will also
be evaluated by these unfamiliar peers. In a separate fMRI assessment session, participants
receive feedback about whether these unfamiliar peers liked them (i.e., social reward) or
whether unfamiliar peers were unable to provide ratings (i.e., ambiguous outcome). A key
feature of this paradigm is the participants’ own ratings of the unfamiliar peers, which allows
the neural response to both acceptance in general and to mutual acceptance to be investigated.
Indeed, Davey et al. (2010) found that being liked activates primary reward regions, such as the
nucleus accumbens and vmPFC, for 15- to 24-year-old adolescents and young adults.
Furthermore, mutual acceptance, or being liked by highly regarded peers, activated the mPFC
more strongly in that study than being liked by less-liked peers. Response to social reward could
be particularly relevant to social anhedonia, or difficulty experiencing pleasure from social
contact (Blanchard et al., 2000). Whereas previous studies have focused on anhedonia as
broadly related to hedonic capacity—a construct that can include physical, affective, and social
components—the importance of social reward to adolescents and the disruption of social
functioning in depression together point to the importance of understanding neural response to
social reward in relation to social anhedonia.
Using an fMRI paradigm involving feedback about peer social acceptance—specifically,
presenting whether unfamiliar peers the participant had rated for their likeability also rated the
participant as likeable—we examined the association of social anhedonia with adolescents’
Mutual Liking and Social Anhedonia 8
mPFC response and functional connectivity during social reward contexts. Specifically, we
examined functional connectivity between the ventral striatum and the mPFC, two key regions
in reward circuitry that are also relevant to processing social reward and likely to respond in
tandem to social reward. We hypothesized that late adolescents’ social anhedonia would be
associated with higher mPFC response during the social reward of mutual acceptance, as
opposed to acceptance from peers whom participants liked less. In addition, to confirm that
mutual liking—rather than all positive feedback—was particularly associated with altered mPFC
response, we examined whether mPFC response to social reward in general was associated with
Participants were 27 late adolescents, aged 18-21 (M=20.4, SD=.80; 51.9% female; 78%
European American, 7% African American, 4% Asian and 11% other). An additional 7
participants took part in the study but were excluded from analyses (and, therefore, did not
contribute data to the current paper) for the following reasons: not completing the
questionnaires (n=1), not completing the fMRI task (n=5), or excessive movement during fMRI
data acquisition (n=1). These 7 participants had higher severity of both anhedonia and
depression than the 27 participants in the final data set (F = 5.81, p = .02 and F = 5.65, p = .02,
respectively). Sample characteristics and results described below all refer to the sample of 27
The sample was recruited in the Pittsburgh metropolitan area by flyers, online ads, and
from a previous large, psychometric study of depression and temperament in young adults (n =
6) (Olino et al., 2013). To obtain variability in anhedonia, recruitment materials targeted both
healthy adolescents and adolescents with depression. Participants were allowed to have current
Major Depressive Disorder (n=9) and anxiety disorders other than Obsessive-Compulsive
Disorder (Generalized Anxiety Disorder (GAD) n=2, Social Phobia n=1). Participants were
Mutual Liking and Social Anhedonia 9
excluded if they met criteria for Bipolar Disorder or lifetime Substance Dependence because of
those disorders’ association with altered reward function (Koob & Le Moal, 2008; Nusslock et
al., 2012). Diagnoses were made by interview with the KSADS-PL (Kaufman et al., 2000). All
participants were free of lifetime psychotropic medications and free of illicit drugs on the day of
the scan. Written informed consent was obtained after a complete description of the study and
study procedures according to the guidelines of the University of Pittsburgh Institutional Review
Participants completed the Revised Chapman Social Anhedonia Scale (RSAS; Eckblad et
al., 1983), a widely used, 40-item true/false questionnaire that focuses on anhedonia
experienced in social contexts (e.g., “having close friends is not as important as many people
say”). We used the total score of the RSAS measure (M=7.17, SD=7.22, Range 0-26), for which
internal consistency was high in this sample (α=.91). Anhedonia was sufficiently normally
distributed, as indicated by its acceptable level of skewness (1.49, SE=.45, p>.001).
The Center for Epidemiologic Studies Depression Scale (CES-D; Radloff, 1977) is a 20-
item questionnaire that assesses depressive symptom severity, and contains 4 items related to
anhedonia (e.g., “I felt I was just as good as other people”). We used the total score of the CES-D
(M=12.63, SD=14.52, Range 0-45), for which internal consistency was high in this sample
(α=.97). Depression severity was sufficiently normally distributed, as indicated by its acceptable
level of skewness (.94, SE=.45, p>.001).
2.2.3 Social reward task
We adapted the likeability task developed by Davey and colleagues (Davey et al., 2010).
Participants were asked to rate photos of other late adolescents (40 photos; 50% female) based
on how much they thought that they would like the people in the photos. They were told that
Mutual Liking and Social Anhedonia 10
they would, in turn, be rated by other participants. Approximately 3 weeks before the scan,
participants had their photo taken and rated the photo stimuli. Ratings were made on a 1-9
scale, with 1 being “not at all” liked and 9 being ”very much” liked. Photo stimuli were drawn
from a database of face stimuli (Martinez & Benavente, 1998;
http://www.ece.osu.edu/~aleix/ARdatabase.html). As ratings of likeability may have been
influenced by sexual attraction, participants were asked about their sexual orientation. One
participant reported being “100% homosexual,” and the remaining participants reported being
either “mostly heterosexual” or “100% heterosexual.” As a result, we did not include sexual
identity in our analyses (Corliss et al., 2009).
Ratings of the photo stimuli were classified into 3 sets of stimuli for each participant:
most-liked stimuli (participant’s 8 highest-rated faces, 4 of each sex), least-liked stimuli
(participant’s 8 lowest-rated faces, 4 of each sex), and neutral stimuli (16 faces per participant,
based on rank order 7-14 of each sex). These classifications were used to create the stimulus sets
for the 3 types of blocks in the task: mutual liking blocks contained favorable feedback from the
8 stimuli rated by the participant as most highly liked, received liking blocks contained
favorable feedback from the 8 stimuli rated by the participant as least liked, and ambiguous
blocks contained no feedback from the neutral stimuli. Stimuli ranked 5, 6, 15, and 16 for each
sex (i.e., 8 total stimulus faces) were discarded to increase the differential value between mutual
liking, received liking, and ambiguous stimuli. For both mutual and received-liking blocks, the
participant was told that the individuals in the photos had selected the participant as a highly
liked peer. As a comparison, the participant was told that individuals in the ambiguous block did
not have a chance to rate them. We term this type of feedback ambiguous because participants
could interpret the no-feedback information in several possible ways.
The fMRI portion of the task was administered in a block design. Stimuli for positive-
feedback blocks were displayed on a green background, and stimuli for ambiguous blocks were
displayed on a white background (Fig 1). The task contained positive-feedback blocks and
Mutual Liking and Social Anhedonia 11
ambiguous-feedback blocks. Each of the 32 stimulus photographs included in the final fMRI
paradigm (16 positive-feedback faces [8 high-rated and 8 low-rated] and 16 ambiguous-
feedback faces) were presented 3 times over 8 blocks. Each block consisted of 12 stimuli for a
total of 84s, for a total task time of 12 minutes and 8 seconds. Of the 8 blocks, 2 were primarily
mutual liking, 2 were primarily received liking, and 4 were primarily ambiguous feedback. Each
block also contained 2 stimuli of the other type (e.g., an ambiguous block contained 2 positive-
feedback faces). A proportion (16.67%) of non-block-category stimuli was included within each
block in order to reduce the predictability and habituation that can accompany the repeated
viewing of stimuli from a single stimulus category within a block. Each stimulus photograph was
presented for 3s, with a jittered inter-trial fixation screen between stimuli (1, 3, 5, or 7 s). The
inter-block interval (IBI) was 8 seconds. The instructions for each block were presented with
either a green (positive feedback) or white (ambiguous feedback) background to signal the block
During the scan, the participants were instructed to pay attention to the faces in the
photos and try to remember who liked them and who had not rated them. Participants were
instructed to press any button when they saw a face, to confirm that they were awake and
attending to the task. After the scan, participants completed ratings in which they recalled how
good they felt when they saw each of the stimulus photos; the order of stimuli shown (faces from
ambiguous blocks vs. faces from positive-feedback blocks) was counterbalanced across
participants. At the end of the scan session, participants were debriefed on the deception in the
task and informed that their photo had not been rated by others.
2.3 fMRI acquisition and preprocessing
Each participant was scanned using a Siemens 3T Trio scanner. BOLD functional images
were acquired with a gradient echo planar imaging (EPI) sequence and covered 39 axial slices,
3.1 mm thick, beginning at the cerebral vertex and encompassing the entire cerebrum and the
majority of the cerebellum (TR/TE=2000/25ms, FOV=20cm, matrix=64×64, flip angle=90°).
Mutual Liking and Social Anhedonia 12
All scanning parameters were selected to optimize the quality of the BOLD signal while
maintaining a sufficient number of slices to acquire whole-brain data. Before the collection of
fMRI data for each participant, we acquired a reference EPI scan that we visually inspected for
artifacts (e.g., ghosting) and for good signal across the entire volume of acquisition. The fMRI
data from all included participants were cleared of such problems.
Preprocessing and image analysis were completed using SPM8
(http://www.fil.ion.ucl.ac.uk/spm). For every scan, images for each participant were segmented
and then realigned to correct for head motion. Data sets were then selected for quality based on
our standard small-motion correction (<3mm). Realigned images were spatially normalized into
standard stereotactic space (Montreal Neurological Institute template) using a 12-parameter
affine model. Normalized images were smoothed with a 6mm full-width at half-maximum
Gaussian filter. Voxel-wise signal intensities were ratio normalized to the whole-brain global
mean. Voxels were resampled during preprocessing to be 2mm3.
2.4 Data Analytic Strategy
We conducted two sets of analyses: one testing mPFC response to social reward and the
other testing functional connectivity between the nucleus accumbens and the mPFC. The first
set of analyses addressed whether mPFC response to reward varied with anhedonia. The second
set of analyses addressed whether anhedonia was associated with the coordination of two
important regions in reward circuitry during the processing of social reward. For both sets of
analyses, we used widely applied, conventional strategies as implemented through SPM8:
general linear models for mPFC response, and psychophysiologic interaction for functional
In preparation for both sets of analyses, for each participant and scan, first-level
predetermined condition effects (i.e., main effects of task) at each voxel were calculated using a
t-statistic, producing a statistical image for the 2 contrasts of interest: (a) mutual liking >
received liking and (b) all positive feedback (i.e., mutual liking + received liking) > ambiguous
Mutual Liking and Social Anhedonia 13
feedback. We were primarily interested in neural response to mutual liking > received liking, as
this contrast reflects being reciprocally liked by an unfamiliar peer vs. being liked by an
unfamiliar peer whom you had not rated as highly likeable. This contrast is proposed to capture
both receiving social reward and weighing one’s own preferences in social stimuli. We included
the all positive feedback > ambiguous feedback contrast to examine whether social anhedonia
was also associated with disrupted response to social reward in general, regardless of the
participant’s own preferences.
2.4.1 Analyses of mPFC Response to social reward
To test the association of social anhedonia with mPFC response to social reward,
preprocessed data sets were analyzed using second-level random effect models that account for
both scan-to-scan and participant-to-participant variability to determine task-specific regional
responses. Individual contrast images were then included in the group-level (i.e., second-level)
mPFC region of interest (ROI) analysis to test our hypotheses. The ROI was constructed using
the WFU PickAtlas Tool (v2.4) and defined as a 25 mm-radius sphere including medial
Brodmann Area (BA) 10 and BA32, which are part of the anterior rostral mPFC region
implicated in social cognition and self-systems (Forbes et al., 2010; Forbes et al., 2012).
Regression analyses were used to test associations between task-related mPFC reactivity and
social anhedonia using the RSAS total score as a regressor in SPM. We also conducted
exploratory, whole-brain analyses for these same models to confirm that the mPFC was one of
the main regions engaged. To test whether social anhedonia was related to mPFC response on
its own or because of its relation to depression, we then conducted regression analyses with
RSAS but included CES-D score as a covariate to adjust for the influence of depressive severity.
2.4.2. Functional connectivity analyses
To test whether mPFC response detected in our first set of analyses might reflect the
disruption of coordination of reward-circuit regions in relation to social anhedonia, we then
conducted functional connectivity analysis. We conducted a single, fairly popular type of
Mutual Liking and Social Anhedonia 14
functional connectivity analysis: task-based PPI. The purpose of PPI analyses was to examine
task-specific changes in the association between mPFC and ventral striatum connectivity in
relation to level of social anhedonia. Functional connectivity allows the testing of functional
interaction of separate regions within neural circuitry (Friston, 1994, 2011; O’Reilly et al., 2012).
In the case of PPI, this functional interaction is related to a specific context (see O’Reilly et al.,
2012 for an excellent tutorial on PPI). In PPI, a region is selected as a seed – in the current
study, the bilateral nucleus accumbens – and the strength of its association to other regions is
examined during a psychological context – in the current study, the receipt of mutual vs.
received liking. Specifically, the time course of the seed region’s response to the contrast of
interest is modeled, entered into a GLM regression, and then used to create an interaction term
between task-related time course and seed-region time course. This interaction is then used to
identify voxels whose activity is associated with the task-seed interaction. Because of our focus
on the potential role of the mPFC in social reward and anhedonia, we masked our PPI analyses
using the same mPFC ROI described above.
Since we were interested not only in whether mPFC and VS had associated responses to
social reward but whether this association was related to individual differences in social
anhedonia, we conducted second-level regression analyses with social anhedonia score
predicting PPI. We masked these results for mPFC response and also conducted exploratory
whole-brain analyses to confirm that mPFC was among the areas whose functional connectivity
with VS varied with level of social anhedonia. This approach has now been used in many studies
to investigate whether psychopathology is related to functional coordination among regions in
affective circuitry (e.g., Keller et al., 2013). In fact, given that neural response and PPI-based
functional connectivity address different research questions—that is, task responding in specific
brain regions vs. functional coordination between regions during a task context – it is now often
the case that studies include both techniques in a single paper (e.g., Eack et al., 2013).
Mutual Liking and Social Anhedonia 15
PPI analyses included 26 participants because one participant was removed from
analyses due to incorrect file format. Given our hypotheses, the bilateral nucleus accumbens was
selected as our seed region of interest and we evaluated correlations of activation between the
nucleus accumbens and the mPFC. We examined whether social anhedonia was associated with
fronto-striatal connectivity by regressing PPI on social anhedonia. We also conducted
exploratory whole-brain analyses for our PPI hypothesis tests, to confirm that the mPFC was
one of the main regions interacting with the accumbens during social reward.
For analyses testing our hypotheses, simulations in the AlphaSim program in AFNI were
used to estimate the minimum number of contiguous voxels in each cluster required to avoid
Type I error (corrected cluster level threshold of p<.05) for the ROI masks. These simulations
resulted in the following minimum extent for significant clusters: 136 voxels for mPFC and 580
voxels for whole-brain findings.
As in previous studies using this task, female faces were rated higher than male faces
(t=3.01, df=2, 25, p<0.01). Reaction times did not differ significantly between male and female
faces. There was no correlation between post-scan ratings and mean reaction time for all stimuli
(r=-.02, p=.92). Participant sex was unrelated to reaction time (F=0.02, p=.91), stimulus ratings
(F=.737, p=.34), and post-scan ratings (F=2.60, p=.12). There was no significant difference
between participants’ post-scan ratings of their response to positive-feedback and ambiguous-
feedback stimuli. Age was associated with higher RSAS (r=.45, p<.02) and CES-D scores (r=.52,
p<.01). Sex was unrelated to RSAS (F=1.56, p=0.22) or CES-D scores (F=0.20, p=0.51).
3.2 Neural response
3.2.1 Effects of task
Analyses with a within-sample t-test indicated that, as expected, participants exhibited
mPFC response to mutual liking > received liking. ROI analysis yielded 2 clusters (dorsal
Mutual Liking and Social Anhedonia 16
anterior cingulate, 132 voxels, [peak voxel (Talairach): -14, 35, 31], t=2.47, p<.05; pregenual
anterior cingulate; 60 voxels, [-14, 40, 20], t=2.35, p<.05), both of which also emerged in whole-
brain analyses for the effects of this contrast (Table 1). Whole-brain analyses for mutual liking >
received liking also yielded response in several other mPFC clusters, as well as in reward
processing regions such as the caudate, social processing areas such as the precuneus, and
affective regions such as the insula. As expected, participants also exhibited mPFC response to
all positive > ambiguous feedback. ROI analyses yielded 4 clusters (perigenual anterior
cingulate, 206 voxels, [-8,41,10], t=2.72, pcorrected<.05; rostral mPFC/dorsal mPFC/perigenual
anterior cingulate, 161 voxels, [6,52,15], t=2.37, pcorrected<.05). Whole brain analyses for all
positive > ambiguous feedback also yielded response in several mPFC clusters, as well as in
social/self-processing regions (e.g., temporoparietal junction, precuneus, posterior cingulate),
reward regions (e.g., caudate, putamen, globus pallidus), and affective regions (e.g., insula,
ventrolateral PFC). As participants varied in anhedonia and depressive symptoms, however,
these findings do not represent the typical adolescent response to this task and thus are not
suitable as a mask for hypothesis tests (see Davey et al., 2010, for a thorough exploration of the
effects of this task in a healthy sample).
3.2.2 Social anhedonia and neural response to social reward
Higher social anhedonia was associated with heightened response to mutual liking >
received liking in a large cluster in the mPFC (4826 voxels, [1,44,41], t=4.01, pcorrected<.05; Fig.
2). This cluster included the following subregions of mPFC, as described by Etkin et al. (2011):
dorsal mPFC, pregenual anterior cingulate, and rostral mPFC.
Whole-brain analyses yielded a large cluster with a peak in the mPFC (see Table 2 for all
whole-brain results), confirming that the mPFC was one of the key brain regions associated with
social anhedonia. Whole-brain analyses also indicated that the striatum (ventral and dorsal),
precuneus, dorsolateral prefrontal cortex, and insula were among the other regions whose
response to mutual liking was associated with level of social anhedonia (see Table 2 for all
Mutual Liking and Social Anhedonia 17
whole-brain results for hypothesis tests). Social anhedonia was unrelated to mPFC response to
all positive > ambiguous feedback.
3.2.3 Specificity of association between social anhedonia and neural response to social reward
To examine whether the association between social anhedonia and response in reward-
circuit regions is evident after adjusting for depression, we re-computed regressions of brain
function on social anhedonia, including CES-D score as a covariate. The association of social
anhedonia remained in two somewhat smaller clusters of activation in the mPFC (peak in BA8,
814 voxels, [15,36,42], t=3.29, pcorrected<.05; peak in BA9, 908 voxels, [6,46,18], t=2.97,
pcorrected<.05) in response to mutual liking > received liking. These clusters included the
following regions: dorsal mPFC, pregenual anterior cingulate, and anterodorsal anterior
3.2.4 Depression and neural response to social reward
Higher depression severity was associated with heightened response to mutual liking >
received liking in a very large cluster centered in dorsal mPFC (peak in BA9, 2971 voxels,
[14,1,44], t=3.36, pcorrected<.05). This cluster included the following subregions: dorsal mPFC and
anterodorsal anterior cingulate. Whole-brain analyses confirmed the association of the mPFC
with depression, and they also indicated that function in the dorsolateral prefrontal cortex and
precuneus was greater in late adolescents with higher depressive symptoms (Table 2).
Depression was unrelated to mPFC response to all positive > ambiguous feedback.
3.2.5 Social anhedonia and frontostriatal connectivity
Higher social anhedonia was associated with greater positive connectivity between the
bilateral nucleus accumbens and the mPFC (peak in BA9, 1110 voxels, [-12,41,33], t=3.44,
pcorrected<.05; peak in BA32, 292 voxels, [-12,29,36], t=3.26, pcorrected<.05) in response to mutual
liking > received liking. This large cluster contained the following subregions: dorsal mPFC,
pregenual anterior cingulate, rostral mPFC, and anterodorsal anterior cingulate. Whole brain
analyses confirmed that the mPFC was a primary region whose function varied positively with
Mutual Liking and Social Anhedonia 18
that of the nucleus accumbens in late adolescents with higher social anhedonia. The large cluster
containing the mPFC in our whole-brain findings also included the VS, orbitofrontal cortex,
temporoparietal junction, and insula (Table 1).
3.2.6 Specificity of association between social anhedonia and frontostriatal connectivity
As with analyses to examine neural response above, we examined the contribution of
social anhedonia over and above that of depression severity by re-computing regressions of
frontostriatal connectivity on social anhedonia, including CES-D score as a covariate and
masking with the results of the original regression. The association of social anhedonia with
greater positive connectivity between the bilateral nucleus accumbens and the mPFC remained
(peak in BA9, 656 voxels, [6,54,32], t=3.90, pcorrected<.05; Figure 3) in response to mutual liking
> received liking. This large cluster was located in the dorsal subregion of the mPFC.
In a sample of late adolescents with varying levels of anhedonia, we found that neural
response and frontostriatal connectivity to social reward were related to anhedonia and
depression. Specifically, late adolescents with higher levels of social anhedonia and depression
showed greater mPFC response and stronger positive connectivity between the mPFC and the
nucleus accumbens to receiving positive peer social feedback. Notably, anhedonia and
depression were associated with altered reward-circuit function to feedback about mutual
liking, in which late adolescents learned that peers whom they liked also liked them. The mPFC
response to social reward was specific to anhedonia, as it remained even when depressive
symptoms were covaried. Thus, anhedonia itself, above the effects of depression, was associated
with disrupted neural responding to peer social feedback. These findings are among the first to
focus on the neural substrates of adolescent anhedonia and to investigate social anhedonia in
relation to social reward.
Interestingly, depression and social anhedonia were only associated with neural
response to mutual liking relative to received (non-mutual) liking, not simply to all positive peer
Mutual Liking and Social Anhedonia 19
social feedback. Both mutual liking relative to received liking and positive feedback relative to
ambiguous feedback elicited response in mPFC and other reward- and social-processing regions
in this sample, but only the former was related to social anhedonia. Late adolescents higher in
social anhedonia did not respond differently to positive feedback in general, but they showed
greater mPFC response and greater positive functional connectivity between the nucleus
accumbens and mPFC in response to mutual compared to received feedback about being liked.
Because fMRI analyses rely on difference scores (i.e., contrasts of conditions), this pattern could
reflect greater mPFC response (or VS-mPFC functional connectivity) to mutual liking, less
mPFC response to received liking, or both. Given the putative functions of dorsal mPFC, these
results could reflect altered social processing involving judgments of others (Denny et al., 2012)
or over-regulation of response to mutual liking (Ochsner et al., 2012; Phillips et al., 2008). Our
results for functional connectivity indicate that the mPFC might engage more intensely when the
VS responds in pleasant social contexts, potentially dampening the VS response and resulting in
a positive correlation of function between these two regions during mutual liking. Alternatively,
the mPFC could be signaling to elicit greater or more sustained response from the VS, which is
not responding with typical magnitude or duration. Results for dorsal mPFC response could also
indicate that late adolescents with anhedonia might respond to mutual liking as if it were
aversive or, alternatively, to received liking as if it were less salient. The former possibility is
consistent with previous findings on greater mPFC response to pleasant self-relevant stimuli in
people with depression or anhedonia, which is the opposite pattern found in healthy adults, who
exhibit greater mPFC response to unpleasant stimuli (Keedwell et al., 2005a, 2005b).
Given the location of our findings in the mPFC in relation to meta-analytic findings
about self-relevant judgments (Denny et al., 2012), our results suggest that late adolescents with
higher levels of anhedonia could possibly experience mutual liking as reflecting negatively on
their identity or self, or, alternatively, could experience received liking as less meaningful for
their identity or self. These late adolescents might thus be less motivated by mutual liking or less
Mutual Liking and Social Anhedonia 20
sensitive to its value. A possible cognitive consequence for such a response in reward circuitry
could be that late adolescents with anhedonia second-guess or discount their own preferences.
Then, instead of being more drawn to people they like, they could defer to others’ preferences,
potentially allowing others to make decisions about social behavior for them. Thus, they might
have weaker social preferences or might be less guided by their social preferences, so that they
may not distinguish between liking and not liking others. Another possibility is that late
adolescents with high levels of anhedonia respond to mutual liking with a kind of dread, in
which they expect the other person to end up disliking or rejecting them, thereby dealing a blow
to their identity and social status. These speculative interpretations are based upon the putative
functions of the mPFC and the content of our task condition, and it will be compelling for future
studies to examine the nuances of anhedonia’s influence on late adolescents’ social experience
The pattern of greater mPFC response we found could be related specifically to social
anhedonia or to the context of social reward. Notably, Wacker et al. (2009), despite
hypothesizing that mPFC function would be related to anhedonia, did not find such an
association in adults. That study used a monetary reward fMRI task, however, and defined
anhedonia more generally. The recent study by Keller et al. (2013) reported findings in a set of
affect-related regions that did not include mPFC, but they employed musical stimuli and focused
on psychiatrically healthy adults. However, as with that study, we also found that anhedonia was
associated with altered functional connectivity of VS and other reward-related regions. In the
current study, although we focused specifically on the mPFC, we also found that VS response—
occurring as part of two large clusters, one of which also contained the mPFC and OFC, and one
of which also contained dorsal striatum and thalamus—was related to higher social anhedonia.
The positive correlation between response in this large cluster and social anhedonia could reflect
the role of reward-circuit regions in processing both rewarding and potentially aversive stimuli.
Whole-brain results also indicated that late adolescents’ social anhedonia was associated with
Mutual Liking and Social Anhedonia 21
greater response or greater functional connectivity with the nucleus accumbens in regions
implicated in social processing (i.e., the precuneus and temporoparietal junction; Mar, 2011;
Van Overwalle et al., 2009), affective processing (i.e., insula, orbitofrontal cortex; Phan et al.,
2002; Phillips et al., 2008), and effortful affect regulation (dorsolateral prefrontal cortex;
Phillips et al., 2008). In addition, the dorsolateral prefrontal cortex and precuneus exhibited
greater response in those with higher depression. Together, these findings support the role of
the mPFC and suggest that other regions in reward and affective circuitry also contribute to
Similar to our results with social anhedonia, we found that depression severity was
related to greater mPFC response to mutual liking relative to received liking. This is consistent
with our previous findings on the association between adolescent depression and neural
response to monetary reward (Forbes et al., 2009; Morgan et al., 2013), as well as with some
other researchers’ findings on mPFC function in adult anhedonia and depression (Keedwell et
al., 2005a, 2005b; Knutson et al., 2008). However, unlike other previous findings in adolescent
depression (Forbes et al., 2009; Forbes et al., 2006), our whole-brain findings did not indicate
that depressive severity was related to response in the VS. One possibility related to our choice
of fMRI task is that the mPFC is especially sensitive to social rewards, and depression
particularly disrupts responding to a social reward in that region rather than in the VS. Also,
unlike a previous study using this social reward task with late adolescents (Davey et al., 2011),
we did not find that depression was related to amygdala response to social reward. That study
took a group-differences approach, however, comparing a group of clinically depressed young
people with a group of healthy young people. In addition, that study used an event-related
version of the social reward task.
Our study is the first to examine reward-circuit response to social reward in relation to
adolescent anhedonia. In contrast, most extant findings on reward in adolescents with
depression or at risk for depression have used monetary or other non-social classes of reward
Mutual Liking and Social Anhedonia 22
(e.g., Forbes et al., 2009; Gotlib et al., 2010; McCabe et al., 2012; Monk et al., 2008; Olino et al.,
2011). Despite the importance of the social context to adolescent depression and to adolescents’
reward processing (e.g., Chein et al., 2011), the study of social reward and its association with
affective disorders has been minimal in adolescent research. In addition, much of the research
on personally relevant stimuli has focused on social loss or exclusion rather than social reward.
Social stimuli are especially salient if they are self-relevant, and previous studies either have
used stimuli such as images, videos, or audio clips of loved ones (e.g., Hooley et al., 2005;
Leibenluft et al., 2004; Whittle et al., 2012) or have employed paradigms that mimic situations
participants encounter in the real world (e.g., playing a computer game with peers, as in
Eisenberger et al., 2003). Thus, it is valuable to note that our mPFC results echo findings for
adolescents’ response to social evaluation paradigms, such as those reported with the social-
exclusion cyberball task (Bolling et al., 2011) and with a simulated chat room task (Silk et al.,
2012). It is not surprising that mPFC function is sensitive to social reward in various forms, such
as being liked, being included, and being selected for interaction.
Furthermore, this is the first study to link late adolescents’ social anhedonia to functional
connectivity in reward circuitry, with greater coordination between the nucleus accumbens and
mPFC during the experience of mutual relative to received liking. Thus, it appears that the
mPFC is not simply responding differently to social reward but is also coordinating its response
more closely with the nucleus accumbens in those with higher levels of social anhedonia.
Admittedly, the application of functional connectivity to answer questions about whether the
coordination among regions in affective or social circuitry varies with psychopathology is a
somewhat new approach. Nonetheless, this approach is also becoming widely applied in clinical
neuroscience (e.g., Almeida et al., 2009; Davey et al., 2012; Mukherjee et al., 2013). The current
findings thus build on an important emerging literature in the neural bases of affective and
Mutual Liking and Social Anhedonia 23
In terms of adolescent brain development, our findings indicate that low subjective
response to social reward is associated with disrupted function in reward circuitry. This
association might be especially pronounced during late adolescence, given the dramatic changes
in social context, the pursuit of high-intensity rewarding experiences, and the enhanced
intensity of social goals during this developmental period (Davey et al., 2008). In addition,
disrupted function in reward-relevant regions or in frontostriatal connectivity could be
particularly evident in response to acceptance from peers, rather than to feedback involving
people of other ages. One question we are now investigating is whether personally relevant peer
social reward—for example, video stimuli of a close friend displaying positive affect during a
conversation with the adolescent in the scanner—is particularly evocative of differences in
neural circuitry that are linked to reward-related problems.
Clinically, the implications of our findings are that disrupted function of neural reward
circuitry could play a role in the development of depression. Anhedonia is a strong prognostic
factor for the onset and clinical course of depression in adolescents (McMakin et al. 2012; Pine
et al., 1999; Spijker et al., 2001; Wilcox & Anthony, 2004) and its association with depression
could be mediated by neural response to reward. Social anhedonia, which was the focus of the
current study, might be addressed by preventive interventions that target adolescents who
experience that symptom. In adolescents who are experiencing high levels of anhedonia in the
context of depression, encouraging the pursuit and experience of social rewards in treatments
such as behavioral activation therapy (Dimidjian et al., 2011) or savoring (McMakin et al., 2011)
could be fruitful. Given that late adolescents with high social anhedonia were less responsive to
mutual liking in the current study, adolescents with depression might be taught to be aware of
or enhance their own responses to others. For example, mindfulness exercises could strengthen
their experience of their social preferences and possibly facilitate a more intense response to
feedback from people who are important to them.
Mutual Liking and Social Anhedonia 24
While the present study represents an important step forward, it has limitations worth
noting, including its relatively small sample, cross-sectional design, and focus on late
adolescents. To fully address the range of variability in a continuous characteristic such as
anhedonia, it will be valuable for future studies to include larger samples. We acknowledge that
our data might reflect a restricted range of anhedonia severity, as the participants who could not
be included in fMRI analyses tended to have higher severity of anhedonia than those who were
included. Thus, it will be important to include broad range of anhedonia in future studies. If
anything, this difference might have led to Type II error, as restricted range attenuates
statistical power to detect associations. In addition, our strict adjustment for multiple
comparisons resulted in very large clusters of BOLD response that encompassed many regions, a
challenge that commonly occurs when using extent-based thresholds. This approach can also
increase the risk of Type II error, since it can limit the ability to detect response in small regions.
As a result of our cross-sectional design, we are unable to address anhedonia as a precursor to
depression or the role of neural reward circuitry in the development of depression in late
adolescents who are high in anhedonia. We describe our participants as late adolescents, which
is consistent with evidence of continued development in reward- and affect-related neural
circuitry into the early 20s (Lenroot & Giedd, 2006). We chose this developmental focus because
of the importance of social reward and the ability to pursue that reward more freely during this
period. While people in this developmental period may be assuming some adult-like status and
roles in their lives, leading some researchers to describe this period as emerging adulthood (e.g.,
Conger & Little, 2010), they are likely to have quite a bit in common with younger adolescents in
terms of their reward-related behavior and anhedonia. In addition, because our participants are
past the typical age of onset of depression (Kessler et al., 2001; Lewinsohn et al., 1994), and
many have already developed depression, our focus on neural response to reward in this age
group could capture some effects of depression.
Mutual Liking and Social Anhedonia 25
It is critical to examine the role of anhedonia across the range of development, and thus
the inclusion of children, younger adolescents, and adults in future studies will be valuable for
addressing potential developmental influence in the association between anhedonia and neural
response to social reward. In addition, we focused on social reward and social anhedonia, which
is only one type of anhedonia. It will be important to investigate other types of anhedonia in
relation to neural response to various classes of reward stimuli. Furthermore, the instrument we
used, the RSAS, assesses trait-like rather than state-like anhedonia, and it will be worthwhile for
future work to consider episode-relevant anhedonia as a symptom. Finally, the examination of
other types of psychopathology (e.g., substance use, psychosis, eating disorders) in future
studies will also allow the examination of the role of anhedonia across other types of reward-
The current study is the first to focus on the neural correlates of adolescent anhedonia.
This characteristic, which is a cardinal symptom in depression, a prognostic factor for the
development of depression, and a potential endophenotype of depression, is an important
dimension of psychopathology that reflects alterations in positive valence systems. In addition,
this study is one of few to examine neural response to social reward, a class of reward that is
especially salient to adolescents and may have meaning for the development of depression
(Forbes, 2009). Our findings of high responding in neural regions implicated in the regulation
of reward responding, in combination with our finding of heightened positive connectivity
between the VS and mPFC, provide preliminary evidence on the pathophysiology of adolescent
anhedonia. These findings could thus contribute to our understanding of the role of anhedonia
in the developmental psychopathology of reward-related problems.
Mutual Liking and Social Anhedonia 26
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Table 1. Whole-brain Results for Neural Response to Social Reward
Mutual Liking > Received Liking
Superior Parietal Cortex
Middle Temporal Gyrus
Inferior Parietal Cortex
Inferior Parietal Cortex
mPFC/Supplementary Motor Area
All Positive Feedback > Ambiguous Feedback
Precentral Gyrus/mPFC/Postcentral Gyrus/Superior
Gyrus/Thalamus/Globus Pallidus/Lingual Gyrus
Lateral Temporal Gyrus/Temporoparietal
Junction/Superior Lateral Gyrus
Precentral Gyrus/Postcentral Gyrus
Inferior Parietal Cortex
Superior Temporal Gyrus
Middle Temporal Gyrus
2228 4.29 34 -8 44
12909 4.15 -26 -90 21
Mutual Liking and Social Anhedonia 33
Note: Analyses were conducted within the entire sample (n=27, df=26), using the contrasts
indicated. All analyses were thresholded at were puncorrected<.05 and cluster size>30 voxels.
DLPFC: dorsolateral prefrontal cortex. mPFC: medial prefrontal cortex. PFC: prefrontal cortex.
Mutual Liking and Social Anhedonia 34
Table 2. Whole-brain Results for the Association of Social Anhedonia with Neural Response
and Functional Connectivity
Regions in Cluster
Neural Response and Social Anhedonia
Thalamus/Caudate Tail/Posterior Cingulate/VS
Precuneus/Superior Parietal Cortex
Neural Response and Depressive Symptom Severity
Precuneus/Inferior Parietal Cortex
Functional Connectivity and Social Anhedonia
Note: Analyses involved the association of (1) neural response to reward and social anhedonia
(n=27, df=26); (2) neural response to social reward and depressive symptom severity (n=27,
df=26); and (3) functional connectivity with the bilateral nucleus accumbens and social
anhedonia (n=26, df=1,24). The contrast of interest was mutual liking > received liking. Social
anhedonia was assessed with the Revised Social Anhedonia Scales (Eckblad et al., 1983).
Depressive symptoms were assessed with the Center for Epidemiologic Studies Depression Scale
(Radloff, 1977). All analyses were thresholded at pcorrected<0.05 using extent computed with
AlphaSim to adjust for Type I error. mPFC: medial prefrontal cortex. VS: ventral striatum.
DLPFC: dorsolateral prefrontal cortex. The first region listed for each cluster is the location of
the peak voxel.
Mutual Liking and Social Anhedonia 35 Download full-text
Figure 1. Block-design social reward fMRI task included in the current study. The task was
adapted from the work of Davey et al. (2010). Participants rated 20 female and 20 male face
stimuli on how much they imagined that they would like each person. Ratings were used to
create personalized stimulus sets for the 3 block types: mutual liking (positive feedback, from
most-liked faces), received liking (positive feedback, from least-liked faces), and ambiguous (no
feedback, from neutral faces).
Figure 2. Late adolescents’ social anhedonia (RSAS; Eckblad et al., 1983) in relation to medial
prefrontal cortex (mPFC) response (in arbitrary units, extracted from cluster mean) to mutual
liking relative to received liking. The large cluster yielded by this analysis includes the following
subregions of the mPFC: dorsal mPFC, pregenual anterior cingulate cortex, and rostral mPFC.
Scatterplots depict the association between social anhedonia score and mPFC response
(extracted as the mean across the entire cluster), for illustration purposes.
Figure 3. Late adolescents’ social anhedonia (RSAS; Eckblad et al., 1983) in relation to positive
functional connectivity between the bilateral nucleus accumbens and the mPFC during mutual
liking relative to received liking. Two large clusters yielded by this analysis include the following
subregions of the mPFC: (1) dorsal mPFC and (2) pregenual anterior cingulate, rostral mPFC,
and anterodorsal mPFC. Scatterplots depict psychophysiologic interaction (PPI) values,
(extracted as the mean across the entire larger cluster), for illustration purposes. R2 value
reflects the effect size of the association between social anhedonia and the mean PPI value
across the depicted cluster.