ORIGINAL RESEARCH ARTICLE

published: 28 June 2012

HUMAN NEUROSCIENCE doi: 10.3389/fnhum.2012.00192

The power of emotional valence—from cognitive to

affective processes in reading
Ulrike Altmann 1,2*, Isabel C. Bohrn 1,2 , Oliver Lubrich 2,3 , Winfried Menninghaus 2,4 and
Arthur M. Jacobs 1,2,5
1
Department of Education and Psychology, Freie Universität Berlin, Berlin, Germany
2
Languages of Emotion, Freie Universität Berlin, Berlin, Germany
3
German and Comparative Literature, Universität Bern, Switzerland
4
Department of Philosophy and Humanities, Freie Universität Berlin, Berlin, Germany
5
Dahlem Institute for Neuroimaging of Emotion, Freie Universität Berlin, Berlin, Germany

Edited by:
John J. Foxe, Albert Einstein College
of Medicine, USA
Reviewed by:
Eveline De Bruin, Unilever R&D
Vlaardingen, Netherlands
Ted Altschuler, Albert Einstein
College of Medicine, USA
*Correspondence:
Ulrike Altmann, Department of
Education and Psychology, Freie
Universität Berlin, Habelschwerdter
Allee 45, Berlin 14195, Germany.
e-mail: u.altmann@fu-berlin.de
The comprehension of stories requires the reader to imagine the cognitive and affective
states of the characters. The content of many stories is unpleasant, as they often deal with
conflict, disturbance or crisis. Nevertheless, unpleasant stories can be liked and enjoyed.
In this fMRI study, we used a parametric approach to examine (1) the capacity of increasing
negative valence of story contents to activate the mentalizing network (cognitive and
affective theory of mind, ToM), and (2) the neural substrate of liking negatively valenced
narratives. A set of 80 short narratives was compiled, ranging from neutral to negative
emotional valence. For each story mean rating values on valence and liking were obtained
from a group of 32 participants in a prestudy, and later included as parametric regressors
in the fMRI analysis. Another group of 24 participants passively read the narratives in a
three Tesla MRI scanner. Results revealed a stronger engagement of affective ToM-related
brain areas with increasingly negative story valence. Stories that were unpleasant, but
simultaneously liked, engaged the medial prefrontal cortex (mPFC), which might reflect
the moral exploration of the story content. Further analysis showed that the more the
mPFC becomes engaged during the reading of negatively valenced stories, the more
coactivation can be observed in other brain areas related to the neural processing of
affective ToM and empathy.
Keywords: emotion, empathy, fMRI, liking, literature, reading, theory of mind

INTRODUCTION mind (ToM) stories (Abu-Akel and Shamay-Tsoory, 2011; Brink

Humans are storytellers. They share daily experiences, tell each
other anecdotes, and exchange gossip (Baumeister et al., 2004;
Dunbar, 2004). Besides producing and performing live narratives
(McAdams Dan, 2001; Habermas and de Silveira, 2008), individ-
uals also extensively consume stories: we read them in newspapers
and magazines, in biographies and novels, via videotext or on
the internet (for the purpose of this paper, we will use the term
story in a broad sense synonymous with narrative). Yet even when
stories have a negative content, as they deal with conflicts or
crises, individuals not only understand, but also appreciate or
enjoy them. In a study by Berthoz et al. (2002), for example, sto-
ries with endings that were either embarrassing or violated social
norms were rated as funnier compared to stories whose endings
reaffirmed normative social behavior.
We have, however, only limited knowledge about the neural
processing of the emotional valence of stories. On the one hand,
research on the neural effects of valence focuses primarily on
the level of single words (Kuchinke et al., 2005; see Citron, 2012
for a review) or sentences (Willems et al., 2010; Kuchinke et al.,
2011). On the other hand, there is extensive literature on the
cognitive and emotional neural processing underlying theory of
et al., 2011; Schnell et al., 2011; Walter, 2012). In these studies,
individuals are often asked to make explicit cognitive and affec-
tive attributions to auditory or visually presented stories, and
the material is primarily selected for its capacity to invoke men-
tal state attributions. What has not been investigated so far is
the question to what extent the emotional valence of such sto-
ries might contribute to ToM related neural processes. Usually,
no data concerning the valence of ToM stories were included in
the analysis or reported as a selection criterion (but see Berthoz
et al., 2002; Brink et al., 2011). In the current study, we there-
fore took a reversed approach and investigated the contribution
of emotionally valenced story contents to ToM-related process-
ing in a passive reading task. In particular, we were interested
in whether increasing negative story valence would engage not
only more brain regions related to affective processing, but also
invite a stronger engagement of ToM-related regions. Story com-
prehension seems to be closely linked to ToM, as it presupposes
the understanding of actions and intentions of real or invented
protagonists (Ferstl et al., 2008).
In a recent meta-analysis, Mar (2011) reported a profound
overlap between text-based ToM studies and non-text-based ToM

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Altmann et al.
studies that used cartoons, pictures, animations or games as stim-
uli. Common activations comprised the dorsomedial prefrontal
cortex (dmPFC), bilateral posterior superior temporal sulcus
(pSTS), right temproparietal junction (TPJ), left inferior frontal
gyrus (IFG), bilateral medial temporal gyrus (MTG), and ante-
rior STS. Moreover, the analysis revealed large overlaps between
the ToM network and results from studies that primarily inves-
tigated comprehension of texts with a narrative structure rather
than ToM.
The literature distinguishes two components of ToM: cogni-
tive ToM and affective ToM (see Walter, 2012 for a recent review).
Cognitive ToM refers to mental state attribution in general (goals,
intentions and desires of others) and engages a network com-
prising the dmPFC, STS, and TPJ, as reflected in the meta-study
by Mar (2011). Affective ToM can be used almost synonymously
with cognitive empathy and relates to the capacity to understand
another’s affective state. Although the actual experience of a cor-
responding affective state is explicitly not assumed within the
framework of this concept, recent data suggest an interplay of
cognitive and affective processes (Schnell et al., 2011).
The first aim of the current study was to investigate the inter-
play of negative story content and ToM. We used a set of short
narrative texts that were either neutral or negative in valence.
The neutral stories deal with everyday events and actions. They
meet the definition of prototypical third-person narratives as they
“have a telic structure including an agent, a goal and a causal
sequence connecting the agent’s various actions with the achieve-
ment or nonachievement of the goal.” (Hogan, 2003, p. 205).
According to Bruner (1986, p. 35), good “storytelling, inevitably,
is about compelling human plights that are “accessible” to read-
ers.” (Bruner, 1986, p. 35). “Access” to a story presupposes the
comprehension of actions, intentions and goals of its protag-
onists, and should thus be closely linked with ToM processes
(Mason and Just, 2009; Mar, 2011). We therefore assume that,
compared to their neutral counterparts, the negative stories we
used were more effective in their potential to engage ToM, as their
negative valence is related to the above mentioned “plights”.
For the understanding of affective processes in reading, the
consideration of valence might be one aspect to consider. The
issue of liking could be another one. Highly interesting art-
work can be disturbing and unpleasant (Turner and Silvia, 2006),
and it has been known since Aristotle’s work on tragedy that
narrative contents do not have to be pleasurable in the sense
of positive valence in order to be liked. The second aim of
the current study was to investigate the neural substrate of
liking negatively valenced narratives. This at first glance para-
doxical tendency to like and enjoy unpleasant contents has
been investigated in media psychology regarding different narra-
tive contexts, including tragic television news and crime drama
(Zillmann et al., 1998; Raney, 2002; Raney and Bryant, 2002).
The enjoyment of unpleasant stories is not limited to a pos-
itive ending; in fact, a film without happy-end can also be
enjoyed (Schramm and Wirth, 2010). How can liking unpleas-
ant stories be explained? Disposition-based theories (Zillmann,
1994) postulate the involvement of two key factors: empathy
with the character and moral evaluation. Accordingly, the enjoy-
ment of unpleasant stories depends on the affective disposition
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Emotional valence of stories and ToM
and empathic reactions towards the characters as well as on
moral judgments of the outcomes the characters were confronted
with (deserved/undeserved). Correspondingly, increased sad film
enjoyment was reported for viewers with high empathy (de Wied
et al., 1994). In a study on crime drama, Raney (2002) found
that (a) the enjoyment of unpleasant contents was predicted by
moral judgments and that (b) moral judgments were predicted by
empathy. Parkinson and colleagues (2011) investigated the neural
processing during moral judgments of stories containing harm,
dishonesty, or disgust. The dmPFC was the only region that all
scenarios had in common and that therefore might represents
a general underlying evaluative processing. Together, these stud-
ies led us to the following assumptions: (1) If moral judgment is
associated with the enjoyment of unpleasant stories, as reported
for crime drama by Raney (2002), the dmPFC might be espe-
cially involved when negatively valenced (unpleasant) narratives
are simultaneously liked. (2) If moral judgments of narratives are
related to empathy, coactivation of empathy-related brain regions
and dmPFC can be expected.
Growing evidence suggests that reading (especially reading fic-
tion) has the capacity to modify personality traits (Djikic et al.,
2009) and is associated with better performance on scales of
empathy and social abilities (Mar et al., 2006). Therefore, further
insight in the affective processes in reading can help to inform
us about their contribution to ToM-related processes and their
potential capacity to enhance ToM development.
To sum up, we hypothesized that (1) passive reading and com-
prehension of both stories with neutral and negative valence
should engage the cognitive ToM network. (2) We further pre-
dicted that increasing negative story valence leads to (a) similar
activation patterns of affect-related brain regions as found in pre-
vious studies on single-word and sentence level. Moreover, we
expected to find (b) the affective ToM network to be involved, as
the negative valence is intertwined with the plights and conflicts
the protagonists are confronted with. Neutral narratives should
invite less ToM processes because the protagonists (and along
with them the reader) can follow their goals and intentions with-
out major disturbances or complications. (3) We expected to find
an interaction effect of negative valence and liking. In detail, read-
ing stories that are considered negative but simultaneously liked
should especially engage the mPFC, reflecting the moral moni-
toring of the characters and the plights they have to deal with.
(a) This region should show a functional coupling with brain
areas related to ToM and affective processing.
METHODS
MATERIAL
We used 80 short narratives (mean number of words: 48, range:
41–57), adopting half of them from the so called black stories,
a narrative-based game (© moses Verlag GmbH, 47906 Kempen,
www.moses-verlag.de). The plot of these stories was negatively
valenced (crimes, disasters, accidents), comparable to the con-
tent of daily news stories, but also of novels or crime stories. For
the other half of the material, we created 40 additional narratives
that had comparable content settings but were neutral in valence.
In these stories, protagonists pursued their goals smoothly and
were not confronted with plights. Negative and neutral narratives
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Altmann et al.
were matched on the number of protagonists, sentences, words,
and syllables, as well as on word frequency and comprehensi-
bility. We conducted a prestudy with 32 participants (16 female,
16 male) in order to make sure that content and style of the neg-
ative and the neutral narratives included only events that could
possibly occur, and that they could be read under two context-
label conditions (“real,” “invented”) which were used during the
task in the MRI scanner. A second group of 32 participants
(16 female, mean age = 24.6; SD = 4.3) rated the material on
valence (“How do you perceive the text?”, scale from −3-“very
negative” to 3-“very positive”) and liking (“Do you like the text?,”
scale from 1-“I do not like it at all” to 7-“I like it very much”). The
resulting mean rating values of each scale were then used in the
fMRI experiment to modulate parametric regressors for the effect
of valence and liking during reading (please refer to Table A7 in
the appendix for further details on the prestudy).
PARTICIPANTS
Twenty-four healthy, right-handed volunteers (12 female, mean
age = 26.5; SD = 6.7) took part in the study. Participants were
German native speakers and skilled readers (assessed with a
screening test that provides normdata for adults; SLS—Salzburger
Lesescreening, unpublished version). Only participants who did
not know the game black stories and were naive to its content were
included. All participants had normal or corrected-to-normal
vision and gave informed written consent in accordance with the
local research ethics committee.
TASK
A 2 × 2 repeated measures design was applied with one factor
varying the story-type (“negative,” “neutral”) and a second fac-
tor varying the context in which a story was presented (“real,”
“invented”). 20 stories were shown in each factor combina-
tion (Figure 1). We pseudo-randomized the order of conditions
FIGURE 1 | Experimental design and stimuli.
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Emotional valence of stories and ToM
as well as the presentation of the narratives across conditions,
but all participants read exactly the same set of black stories
and neutral narratives. During the fMRI experiment, a nar-
rative was presented for 20 s, displayed on five lines (shown
4 s each). Prior to the story, a context label (either “Real” or
“Invented”) was presented for 3 s. Participants were requested
to read the text silently and solve a verification task following
each text. By means of a cue (“Real?” or “Invented?”) partici-
pants were either asked, as an attention control task, whether
the story they just read was real, or they were asked whether it
was invented. Participants answered by pressing a button (“yes,”
“no”). The verification cues were presented in a pseudorandom-
ized order to avoid motor preparation during the reading phase
and to assure an equal assignment of question cues and required
responses with regard to each condition. Additionally, partici-
pants completed the Interpersonal Reactivity Index (IRI; Davis,
1983; German version: Paulus, 2009), which provides a four-
dimensional self-report estimate of empathy. In this study, we
focus on the “empathic concern” subscale, which assesses the
individual tendency to feel concern and compassion for other
people, because empathic concern was found to be associated
with increased interest in tragic television news (Hoffner et al.,
2009) as well as with the perception of a liked partner being
in pain (Singer et al., 2004). Empathic concern scores showed
a mean of 13.09 and a standard deviation of 1.79 (correspond-
ing mean of German population norms = 14.56, SD = 2.94;
norm data retrieved from Paulus, C., “Normtabellen des SPF,”
last modified November 21, 2011, http://bildungswissenschaften.
uni-saarland.de/personal/paulus/empathy/Normen.pdf).
fMRI DATA ACQUISITION
Functional data were acquired on a Siemens Tim Trio 3 T MR
imager at 3 T field strength. Four runs of 425 volumes were mea-
sured using a T2∗ weighted echo-planar sequence [slice thickness,
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Altmann et al.
3 mm; no gap; 37 slices; repetition time (TR), 2 s; echo time
(TE), 30 ms; flip angle, 90◦ ; matrix, 64 × 64; field of view (FOV),
192 mm; voxel-size 3.0 × 3.0 × 3.0 mm], and individual high-
resolution T1-weighted anatomical data (MPRAGE-sequence)
were acquired (176 slices; FOV, 256; TE, 2.52; TR, 1.9; matrix,
256 × 256; resolution 1.0 × 1.0 × 1.0 mm; sagittal plane; slice
thickness, 1 mm).
DATA ANALYSIS
Analysis of fMRI data was conducted with Brain Voyager QX [2.0]
(Brain Innovation, Maastricht, Netherlands; Goebel et al., 2006).
Functional Data were corrected for head-motion and for differ-
ent slice scan times using cubic spline interpolation. To remove
low-frequency signal drifts, a high-pass filter was applied with
a cutoff period three times the block length. Spatial smoothing
was performed using a Gaussian filter of 8 mm, full width at
half maximum. The functional maps of each participant were
then transformed into standard Talairach space (Talairach and
Tournoux, 1988).
Whole-brain statistical analysis was performed according to
the general linear model as implemented in Brain Voyager QX. On
the first level, the model was generated with two blocked regres-
sors for the reading of stories (negative, neutral) and two blocked
regressors for the instruction and attention task periods. Three
different models were constructed: (1) the first model included
an additional parametric regressor containing the mean valence
for each story/narrative, (2) the second model was added by a
parametric regressor containing the mean liking values for each
story/narrative (3) and in the third model we added the para-
metric interaction term of valence and liking (valence × liking).
For each model, individual contrast images from the first level
analysis were then applied to a second level random effects group
analysis, in which we tested for the parametric effects of (1) read-
ing negatively valenced stories (2) liking of texts, and (3) the
corresponding interaction (valence × liking). All parametric con-
trasts were reported whole brain corrected (p < 0.05) using false
discovery rate (FDR), and with an extend threshold of k = 20
voxels for the resulting clusters.
In correspondence with our initial hypotheses, we identified
the mPFC as involved in the interaction of negative valence ×
liking and selected it as the seed region for further psychophys-
iological interaction (PPI) analysis (Friston et al., 1997) with a
sphere of 10 mm around the peak voxel (−9, 41, 16). PPI analysis
provides a measure for task related functional connectivity—
it allows identifying brain regions that show a stronger co-
activation during one task (reading negative stories) as compared
to another (reading neutral stories). Particularly we were inter-
ested in whether valence-specific mPFC activation (i.e., reading
negative stories) is “coupled” with emotion and ToM-related
brain areas, especially the temporal poles (TP), the IFG, and the
temporoparietal junction (TPJ). The PPI regressor was calcu-
lated as the element-by-element product of the mean corrected
mPFC region of interest, and the task vector coding for the
valence-specific effect of reading black stories compared to read-
ing neutral narratives. To identify areas of the brain that showed
increased activity while reading black stories when mPFC activity
increased, individual PPI regressors where entered into a second
Frontiers in Human Neuroscience www.frontiersin.org
Emotional valence of stories and ToM
level random effects analysis [reported whole brain corrected
(p < 0.05) using FDR, k = 20 voxels]. Finally, we repeated the
analysis with individual scores on the empathic concern scale of
the IRI as covariate to examine areas of the brain that showed
a stronger coupling with mPFC depending on the individual ten-
dency to feel concern for other people [cluster level corrected (p <
0.05) using Monte Carlo simulations as implemented in Brain
Voyager, initial voxel level threshold p < 0.001 uncorrected].
RESULTS
CONJUNCTION OF NEGATIVE AND NEUTRAL NARRATIVES
Prior to the analysis that focused on emotional valence, we
applied a conjunction analysis in order to assess the com-
mon effects of neutral and negative narratives (Table A1 and
Figure 2A). This analysis revealed that neutral and negative nar-
ratives share extensive activation patterns including brain areas
that are regularly reported for cognitive ToM, namely the dmPFC
(BA 9), bilateral TP (TP: BA 38), and posterior superior temporal
gyrus (pSTG: BA 22/39).
PARAMETRIC EFFECTS
First, we analyzed the parametric effect of negative valence
(Table A2 and Figure 2B). This led to subcortical activations in
the left striatum (caudate body), left mediodorsal thalamus, and
left amygdala. Furthermore, the analyses revealed an extensive
fronto-temporal network including the left mPFC (mPFC: BA
8/9), the bilateral IFG (IFG: BA 45/47), and the TP (TP: BA 38),
the left fusiform/parahippocampal gyrus (BA 20/36), the bilateral
middle temporal gyrus (MTG: BA 21) and STG (BA 22), as well
as the bilateral posterior superior temporal gyrus extending to the
temporal parietal junction (STG/TPJ: BA 39).
Second, we analyzed the parametric effect of liking (Table A3).
Activations were found in the bilateral anterior STG/TP (BA 38),
anterior MTG (BA 21), IFG (IFG: BA 45/47), lingual/fusiform
gyrus (BA 18/19), and left posterior cingulate cortex (PCC:
BA 30).
Third, we analyzed the parametric interaction effect of valence
and liking (Table A4 and Figure 2C). Increasing negative valence
combined with increasing values of liking was associated with
activations in the bilateral mPFC (BA 9, 32), supramarginal
gyrus/TPJ (BA 39/40), and left dorsolateral prefrontal cortx
(DLPFC: BA 8).
PPI ANALYSIS
Next, we examined the results of the PPI analysis with the mPFC
as the seed region, as this region revealed an interaction effect of
valence × liking (Table A5 and Figure 3). Several regions showed
a stronger coactivation with the mPFC during the reading of
the negatively valenced stories as compared to the neutral narra-
tives (negative > neutral). Such coactivations could be observed
bilaterally in the IGF extending into the insula (BA 45/47/13),
the thalamus, the supramarginal gyrus (BA 40), and the vmPFC
(BA 10) as well as in the left amygdala, the right dorsal striatum
and the dorsal ACC (BA 32).
An additional PPI analysis was applied to take the individ-
ual tendency to feel concern for other people into account. It
was tested which areas of the brain showed higher functional
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Altmann et al.
FIGURE 2 | (A) Brain activation for story reading — conjunction
of negative stories and neutral stories (B) Parametric effect of
increasing negative valence (C) Brain activation for increasing
connectivity with mPFC during reading negative stories, depend-
ing on self-report scores at the empathic concern scale. The
bilateral anterior insula (BA 13) and the right posterior cingu-
late cortex (BA 31) (Table A5 and Figure 4) showed a stronger
coupling with mPFC for individuals who reported a stronger ten-
dency to feel concern for other people [cluster level corrected
(p < 0.05), initial voxel level threshold p < 0.001 uncorrected].
DISCUSSION
THE INTERPLAY OF STORY VALENCE AND ToM
This study aimed to investigate the hypothesis that narratives
with negatively valenced content invite for an increased engage-
ment of the affective mentalizing network (cognitive empathy).
The conjunction analysis between neutral and negatively valenced
narratives showed that they share the cognitive ToM network,
including dmPFC, bilateral TP, and left pSTS.
As hypothesized, these regions became more engaged with
increasing negative valence. The parametric analysis of valence
revealed activations in the bilateral posterior STS/TPJ and the
anterior subdivision of the mentalizing network, comprising
dmPFC, TP, and aSTS/MTG. This network appears to be espe-
cially involved when mental state reasoning requires the inter-
play of cognitive and affective components (cognitive empathy;
Preston and de Waal, 2002), e.g., for the inference of another
Frontiers in Human Neuroscience
Emotional valence of stories and ToM
negative story valence combined with increasing story
liking—Parametric interaction of valence × linking, whole brain
corrected (p < 0.05) using FDR.
person’s affective state. In a recent study, Schnell and colleagues
(Schnell et al., 2011) investigated cognitive empathy and found a
simultaneous activation of the anterior mentalizing network and
limbic structures, including the left amygdala, when affective as
compared to non-affective visuospatial states had to be inferred.
This network could also be observed when the participants made
affective judgments about social contexts from their own point
of view, without being explicitly asked to adopt a third-person
perspective. This fits the requirements of the task used in the cur-
rent experiment, as our participants were asked simply to read the
short narratives.
In line with those previous findings our results suggest a close
link between affective and cognitive components for mentaliz-
ing. Two key areas of cognitive ToM, the dmPFC and the TPJ,
were also related to valence. Particularly, the dmPFC and the TPJ
region seem to increase activation when updates of character-
related information and the processing of intentions are needed
(Saxe and Wexler, 2005; Mason and Just, 2009). In a study of
Hooker and colleagues (Hooker et al., 2008), participants had
to infer the emotional responses of characters in social scenes.
In order to predict correctly whether an emotional response of
a character might change or remain constant participants had
to update their character-related information. In line with our
results, the engagement of regions associated with mentalizing
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Altmann et al. Emotional valence of stories and ToM

FIGURE 3 | Brain regions showing positive connectivity with mPFC while reading negative stories compared to reading neutral stories, whole brain
corrected (p < 0.05) using FDR.

FIGURE 4 | Positive connectivity between mPFC and the anterior insula mPFC–connectivity while reading negative stories compared to reading
correlates with the individual tendency to feel concern for other people neutral stories), cluster level corrected (p < 0.05), initial voxel level
(Positive correlation between the empathic concern scale and threshold p < 0.001 uncorrected.

(STS/TPJ, mPFC, TP) and with emotion (IFG, thalamus) was
higher when the emotions of a character were likely to change
and unlikely to remain constant.
These observations are compatible with the negative sto-
ries in the current study: here, the intentions and emotions
of the characters often changed as they were confronted with
several “plights” (Bruner, 1986). In the neutral stories, on the
other hand, the characters could act upon their goals without
major disturbances. Apparently, the conflicts implicated in the
negative stories evoke more attributions of goals and thoughts
to the characters, as opposed to the neutral, everyday stories.
Such attributions and inferences are essential for understand-
ing the character and the story at large. And with increas-
ingly negative emotional valence, reading stories also engaged

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Altmann et al.
the bilateral IFG and additional subcortical structures com-
monly involved in emotion processing, namely the bilateral dorsal
striatum (caudate body), left mediodorsal thalamus, and left
amygdala.
The thalamus appears to be involved in general emotion
processing, independent of valence (Lane et al., 1997; Goldin
et al., 2005) or social/nonsocial content (Britton et al., 2006)
and has been shown to be involved when subjects empathize
with a protagonist suffering a threat (emotional empathy) com-
pared to empathizing with a protagonist in a neutral every-
day (cognitive empathy) situation (Nummenmaa et al., 2008).
Engagement of the amygdala has been reported for viewing pic-
tures of negative emotion (Lane et al., 1997) and watching sad
films (Goldin et al., 2005), but, similar to the thalamus, this
structure is presumably more sensitive to the salience than to
the valence of affective stimuli (Phan et al., 2004; Britton et al.,
2006).
In correspondence with our results, a recent meta analysis
(Mar, 2011) identified lefthemispheric amygdala activation to
be associated with story-based ToM, whereas righthemispheric
activation was found for nonstory-based ToM.
The IFG, which showed to be responsive to both emotional
valence and liking in the present study, is considered a possi-
ble human analogue to the mirror neuron system (Rizzolatti and
Craighero, 2004; Iacoboni, 2009). It was reported for the imita-
tion and observation of emotion in pictures (Carr et al., 2003),
imagery of emotional scripts (Sabatinelli et al., 2006), and might
also play a role in affective mentalizing (Schnell et al., 2011). In
correspondence with our results, Hynes et al. (2006) observed
the orbital part of the bilateral IFG (BA 47) and the left IFG
(BA 11) to be more strongly engaged in emotional perspective
taking than in cognitive perspective taking in a story-based men-
talizing task. Furthermore, potential mirror neuron activation in
bilateral IFG and STS was found to correlate with empathy scores
(Schulte-Rüther et al., 2007).
LIKING OF UNPLEASANT STORY CONTENTS
The parametric interaction analysis showed that when valence
and liking come together, the activation peak of the mPFC moves
inferiorly and closer to the anterior cingulate gyrus (−9, 41, 16),
compared to the activation peak resulting from the parametric
valence effect alone (−6, 44, 31). For reading negatively valenced
narratives (as compared to neutral ones) this region showed a
functional coupling not only with regions related to ToM, but also
with regions known to be involved in affective empathy (amyg-
dale, anterior insula, midcingulate cortex, and IFG; Walter, 2012).
These results were supported by the additional result showing a
stronger functional coupling between the mPFC seed region and
bilateral anterior insula engagement when the magnitude of the
participant’s empathic concern is taken into account. Similar cor-
relations between the anterior insula and self-reported empathy
have been observed for empathy of social pain (Masten et al.,
2011), and for the observation of other individuals receiving pain
(Singer et al., 2004, 2006).
These results support our initial hypotheses derived from
media psychology, which assumed that moral evaluations and
empathic reactions to characters in stories influence liking of
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Emotional valence of stories and ToM
unpleasant story contents (Raney, 2002; Parkinson et al., 2011).
Further studies will be needed to systematically investigate the
potential role of moral reasoning and its relationship to empa-
thy for affective and aesthetic processes in reading. Empathy and
moral reasoning about characters provide two possible factors
which influence enjoyment in reading. Others are very likely
involved as well, for instance stylistic features (Miall and Kuiken,
1994; Jacobs, 2011) and provided details (see Green et al., 2004
for an overview). A major challenge for future research will
be the implementation of paradigms that allow us to follow
the temporal dynamics of reading, particularly of longer, well-
crafted stories. As Green and Brock (2000) showed, well-crafted,
canonical stories were rated as more immersive. First steps in
that direction have already been made (e.g., Wallentin et al.,
2011).
LIMITATIONS OF THE STUDY
The values for valence and liking that were used for the paramet-
ric analysis of the fMRI study reflect mean data that were derived
from a prestudy and as such treated as characteristics of the sto-
ries. The fact that we did not use individual subject ratings of the
fMRI sample might limit the conclusions that can be drawn from
the results.
We decided for an implicit reading task as it was one pur-
pose of the study to assess the interplay of valence, liking and
ToM during (as much as possible) natural reading of stories. Post
scan data of individual subjects for valence and liking might have
strengthened the results. On the other hand, the usage of pretest
data allowed preventing subjects from fatigue effects, likely aris-
ing for rating the 80 stories again during an additional postscan
session. Moreover, it has been shown that especially liking judg-
ments can change with repeated exposure (Tan et al., 2006), which
might have distorted our data. The standard deviations for liking
judgments were comparable to those of valence (Table A7). Thus,
it could be excluded that the evaluation of liking implies lower
intersubject agreement than the evaluation of valence. Similar
observations have been made for judging the attractiveness of
faces (for an overview see Chatterjee et al., 2009) and for reading
poems (Martindale and Dailey, 1995). Given these potential lim-
itations, we consider a similar study using postscan rating data
desirable as to further inform us about individual effects in the
reader and to strengthen the present results.
CONCLUSION
To summarize, taking the emotional valence (from neutral to
negative) of the content of stories into account reveals the
full scale of ToM-related processing, ranging from cognitive
and affective ToM to components of affective empathy through
top-down processing during reading (Walter, 2012). Therefore,
it seems worthwhile to include valence for the investigation
of ToM and related processes either as a variable of inter-
est or as control variable (e.g., between conditions of inter-
est and control conditions). Given the extensive use of stories
in our daily life, their capacity to provide simulations of the
social world (Mar and Oatley, 2008), and to evoke even emo-
tional reactions as complex as liking unpleasant contents, we
should attempt to improve our understanding of the underlying
June 2012 | Volume 6 | Article 192 | 7
Altmann et al.
mechanisms and of how these processes might relate to learning
and development.
ACKNOWLEDGMENTS
This research was supported by the DFG-funded Cluster of
Excellence “Languages of Emotion” and the Dahlem Institute
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Conflict of Interest Statement: The
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Received: 12 March 2012; accepted: 09
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Citation: Altmann U, Bohrn IC,
Lubrich O, Menninghaus W and Jacobs
AM (2012) The power of emotional
valence—from cognitive to affective pro-
cesses in reading. Front. Hum. Neurosci.
6:192. doi: 10.3389/fnhum.2012.00192
Copyright © 2012 Altmann, Bohrn,
Lubrich, Menninghaus and Jacobs.
This is an open-access article dis-
tributed under the terms of the
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Commercial License, which permits
non-commercial use, distribution, and
reproduction in other forums, provided
the original authors and source are
credited.
June 2012 | Volume 6 | Article 192 | 9
Altmann et al. Emotional valence of stories and ToM

APPENDIX
Table A1 | Brain activation for story reading (negative stories AND neutral stories).

Cluster size Region BA Talairach coordinates Max. t-value

X Y Z

820 Inferior Frontal Gyrus 9 R 54.0 20.0 22.0 6.396.391

Postcentral Gyrus 3 R 60.0 −10.0 25.0 6.249.121
Precentral Gyrus 4 R 48.0 −13.0 52.0 5.192.021
Middle Frontal Gyrus 6 R 27.0 −13.0 61.0 4.339.577
Precentral Gyrus 6 R 39.0 −1.0 31.0 3.511.845
954 Middle Temporal Gyrus 21 R 48.0 −10.0 −11.0 9.289.865
Superior Temporal Gyrus 38 R 48.0 11.0 −14.0 8.176.331
posterior Insula 13 R 42.0 −19.0 −8.0 7.638.433
Superior Temporal Gyrus extending 22 R 42.0 −28.0 −2.0 6.806.289
into posterior Superior Temporal Gyrus

Superior Temporal Gyrus 38 R 33.0 26.0 −29.0 4.523.798

Middle Frontal Gyrus 11/47 R 27.0 38.0 −17.0 4.029.813
10940 Lingual Gyrus 18 R 12.0 −79.0 −5.0 41.358.112
Lingual Gyrus 18 L −12.0 −79.0 −5.0 21.272.240
Lingual Gyrus 17 L −21.0 −91.0 −2.0 19.262.949
Cerebellum * L −27.0 −73.0 −14.0 16.760.603
Precentral Gyrus 4 L −48.0 −7.0 43.0 13.336.107
Fusiform Gyrus 37 L −39.0 −61.0 −14.0 13.138.165
Cuneus 17 L −24.0 −82.0 13.0 12.455.202
Superior Temporal Gyrus extending into 22 L −54.0 −25.0 1.0 11.726.843
into posterior Superior Temporal Gyrus

Middle Temporal Gyrus 21 L −57.0 −10.0 −5.0 11.552.003

Superior Temporal Gyrus 38 L −51.0 8.0 −11.0 10.425.444
Lateral Geniculum Body L −21.0 −25.0 −2.0 9.180.390
Cerebellum * R 33.0 −55.0 −14.0 9.042.595
Thalamus * L −6.0 −28.0 −5.0 8.535.957
Middle Frontal Gyrus 9 L −39.0 11.0 22.0 6.862.788
Inferior Frontal Gyrus 47 L −39.0 26.0 −2.0 6.730.738
Parahippocampal Gyrus 35 L −21.0 −28.0 −14.0 5.937.679
Cerebellum * R 0.0 −49.0 −32.0 5.328.261
Sub-Gyral 20 L −36.0 −16.0 −17.0 4.988.610
Parahippocampal Gyrus 35 R 21.0 −28.0 −14.0 4.333.858
Parahippocampal Gyrus 34 L −18.0 −10.0 −14.0 4.179.035
Precentral Gyrus 4 L −27.0 −22.0 64.0 3.165.962
53 Medial Frontal Gyrus 9 L −6.0 53.0 34.0 3.812.754
Medial Frontal Gyrus 9 L −6.0 53.0 34.0 3.812.754
38 Caudate Body L −6.0 5.0 13.0 3.740.989
Putamen L −18.0 −1.0 10.0 3.450.212
167 Medial Frontal Gyrus 6 L −6.0 −1.0 55.0 6.942.378
38 Thalamus * L −9.0 −13.0 7.0 5.033.025

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Altmann et al. Emotional valence of stories and ToM

Table A2 | Brain activation for increasing negative story valence.

Cluster size Region BA Talairach coordinates Max. t-value

X Y Z

1556 Middle Temporal Gyrus 21 R 45.0 2.0 −23.0 −7.062.107

Superior Temporal Gyrus 22 R 45.0 −22.0 −5.0 −6.687.765
Superior Temporal Gyrus 22 R 57.0 −49.0 10.0 −5.981.107
Inferior Frontal Gyrus 11 R 24.0 32.0 −20.0 −5.366.148
Inferior Frontal Gyrus 47 R 48.0 29.0 −5.0 −5.282.379
Parahippocampal Gyrus 36 R 39.0 −22.0 −20.0 −5.018.050
Inferior Frontal Gyrus 45 R 48.0 23.0 10.0 −4.731.057
Superior Temporal Gyrus 22 R 48.0 −55.0 16.0 −4.456.529
Superior Temporal Gyrus 38 R 27.0 5.0 −38.0 −2.992.173
192 Cerebellum * R 21.0 −67.0 −29.0 −5.451.882
113 Caudate Body R 9.0 2.0 13.0 −5.290.515
22 Cerebellar Tonsil * R 6.0 −46.0 −35.0 −4.393.444
23 Medial Frontal Gyrus 11 L −3.0 47.0 −17.0 −5.042.429
713 Medial Frontal Gyrus 9 L −6.0 44.0 31.0 −7.031.443
Superior Frontal Gyrus 8 L −6.0 26.0 49.0 −5.530.106
Superior Frontal Gyrus 6 L −9.0 14.0 55.0 −4.287.555
28 Cerebellum * L −30.0 −88.0 −38.0 −5.304.901
3605 Middle Temporal Gyrus 21 L −54.0 −1.0 −23.0 −7.581.751
Inferior Frontal Gyrus 44 L −45.0 11.0 19.0 −7.456.018
Middle Temporal Gyrus 22 L −54.0 −43.0 1.0 −7.107.245
Superior Temporal Gyrus 38 L −42.0 17.0 −23.0 −6.966.726
Inferior Frontal Gyrus 11 L −24.0 32.0 −20.0 −6.805.636
Sub-Gyral 21 L −48.0 −28.0 −2.0 −6.792.051
Superior Temporal Gyrus 39 L −42.0 −58.0 19.0 −6.568.569
Inferior Frontal Gyrus 47 L −45.0 29.0 −5.0 −6.332.211
Caudate Body L −9.0 2.0 13.0 −5.455.053
Medial Dorsal Nucleus L −9.0 −16.0 10.0 −5.293.430
Amygdala L −18.0 −7.0 −8.0 −4.422.447
Fusiform Gyrus 20 L −39.0 −34.0 −17.0 −4.391.698
Uncus 28 L −27.0 2.0 −29.0 −4.382.841
Fusiform Gyrus 37 L −39.0 −43.0 −11.0 −4.220.282
Middle Frontal Gyrus 6 L −39.0 5.0 43.0 −4.129.689
Brainstem, Red Nucleus L −6.0 −25.0 −5.0 −3.936.012
Inferior Frontal Gyrus 47 L −57.0 32.0 −17.0 −3.860.281
Putamen L −27.0 −13.0 −8.0 −3.533.775
Middle Frontal Gyrus 8 L −33.0 20.0 40.0 −3.268.400

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Altmann et al. Emotional valence of stories and ToM

Table A3 | Brain activation for increasing story liking.

Cluster size Region BA Talairach coordinates Max. t-value

X Y Z

103 Inferior Frontal Gyrus 45 R 54.0 20.0 10.0 4.666.930

518 Superior Temporal Gyrus 38 R 42.0 14.0 −20.0 5.721.092
Superior Temporal Gyrus 38 R 54.0 20.0 −26.0 5.406.403
Middle Temporal Gyrus 21 R 57.0 −1.0 −14.0 5.036.968
Sub-Gyral 21 R 48.0 −10.0 −11.0 4.972.412
Superior Temporal Gyrus 22 R 45.0 −25.0 −5.0 4.305.095
30 Precuneus 31 R 24.0 −73.0 22.0 4.332.990
25 Medial Frontal Gyrus 6 L −6.0 2.0 61.0 4.623.124
52 Parahippocampal Gyrus 28 L −21.0 −25.0 −8.0 3.635.541
Cerebellum * L −3.0 −31.0 −5.0 3.600.545
3341 Middle Temporal Gyrus 22 L −54.0 −43.0 4.0 5.587.103
Superior Temporal Gyrus 38 L −48.0 11.0 −11.0 5.542.369
Superior Temporal Gyrus 22 L −48.0 −19.0 −5.0 5.364.425
Precentral Gyrus 6 L −45.0 −4.0 40.0 5.353.443
Superior Temporal Gyrus 22 L −54.0 −49.0 13.0 5.306.046
Cerebellum * R 21.0 −73.0 −23.0 4.993.320
Middle Occipital Gyrus 18 R 30.0 −79.0 −5.0 4.934.392
Cuneus 19 L −21.0 −88.0 28.0 4.894.963
Inferior Frontal Gyrus 45 L −57.0 14.0 19.0 4.853.875
Lingual Gyrus 17 R 12.0 −88.0 −2.0 4.847.248
Cuneus 18 L −18.0 −94.0 13.0 4.788.170
Middle Occipital Gyrus 18 R 24.0 −94.0 10.0 4.667.599
Cerebellum * L −33.0 −61.0 −8.0 4.616.873
Middle Occipital Gyrus 19 L −27.0 −88.0 10.0 4.599.266
Inferior Occipital Gyrus 18 R 30.0 −94.0 −5.0 4.459.953
Inferior Frontal Gyrus 45 L −51.0 26.0 4.0 4.417.239
Cuneus 30 R 6.0 −70.0 7.0 4.352.642
Middle Occipital Gyrus 18 L −36.0 −88.0 1.0 4.331.434
Cerebellum * L −15.0 −76.0 −11.0 4.291.461
Fusiform Gyrus 19 L −27.0 −82.0 −14.0 4.161.860
Fusiform Gyrus 37 L −39.0 −43.0 −11.0 4.049.331
Lingual Gyrus 19 R 15.0 −55.0 −2.0 4.028.108
Lingual Gyrus 18 L −18.0 −55.0 4.0 3.980.167
Cuneus 17 L −9.0 −76.0 10.0 3.888.908
Cerebellum * R 33.0 −58.0 −11.0 3.844.507
Inferior Occipital Gyrus 18 L −42.0 −88.0 −14.0 3.833.167
Inferior Frontal Gyrus 11 L −27.0 29.0 −23.0 3.559.359
Cerebellum * L −27.0 −37.0 −17.0
Parahippocampal Gyrus 30 R 15.0 −43.0 1.0 3.483.598

Frontiers in Human Neuroscience www.frontiersin.org June 2012 | Volume 6 | Article 192 | 12

Altmann et al. Emotional valence of stories and ToM

Table A4 | Brain activation for increasing negative story valence combined with increasing story liking (valence × linking).

Cluster size Region BA Talairach coordinates Max. t-value

X Y Z

266 Supramarginal Gyrus 40 L −57.0 −46.0 31.0 −5.475.258

Superior Occipital Gyrus 19 L −36.0 −79.0 31.0 −4.544.866
132 Medial Frontal Gyrus 9/32 L −9.0 41.0 16.0 −4.970.238
Medial Frontal Gyrus 9 L −9.0 41.0 28.0 −4.356.964
Superior Frontal Gyrus 9 L −18.0 44.0 28.0 −3.743.781
112 Middle Frontal Gyrus 8 L −36.0 23.0 40.0 −4.474.544
34 Middle Frontal Gyrus 10 L −42.0 56.0 7.0 −4.382.877
Middle Frontal Gyrus 10 L −33.0 47.0 −2.0 −3.886.662
38 Cerebellum * R 33.0 −70.0 −35.0 −3.703.044

Frontiers in Human Neuroscience www.frontiersin.org June 2012 | Volume 6 | Article 192 | 13

Altmann et al. Emotional valence of stories and ToM

Table A5 | Brain regions showing positive connectivity with mPFC while reading negative stories > reading neutral stories.

Cluster size Region BA Talairach coordinates Max. t-value

X Y Z

84 Middle Temporal Gyrus 21 R 48.0 −1.0 −17.0 5.344.678

Superior Temporal Gyrus 38 R 42.0 11.0 −20.0 5.268.341
941 Precentral Gyrus 6 R 42.0 −1.0 34.0 6.347.430
Middle Frontal Gyrus 9 R 33.0 17.0 31.0 5.649.367
Middle Frontal Gyrus 46 R 51.0 29.0 19.0 5.334.372
Putamen R 18.0 8.0 4.0 5.243.031
Putamen R 30.0 −1.0 7.0 5.216.119
Inferior Frontal Gyrus 10/46 R 48.0 35.0 1.0 5.191.519
Inferior Frontal Gyrus 10 R 39.0 47.0 1.0 5.183.762
Inferior Frontal Gyrus 9 R 36.0 8.0 22.0 5.183.069
Inferior Frontal Gyrus extending into Insula 45/47/13 R 48.0 20.0 4.0 5.168.429
Caudate Head R 18.0 17.0 1.0 4.853.752
Thalamus * R 9.0 −1.0 4.0 4.456.534
192 Supramarginal Gyrus 40 R 42.0 −43.0 34.0 6.638.423
Precentral Gyrus 4 R 45.0 −19.0 34.0 4.753.923
Inferior Parietal Lobule 40 R 57.0 −34.0 34.0 4.307.735
37 Fusiform Gyrus 37 R 42.0 −52.0 −11.0 4.580.812
299 Putamen R 33.0 −22.0 −2.0 5.305.676
Hippocampus R 27.0 −13.0 −14.0 5.082.986
Superior Temporal Gyrus 22 R 48.0 −37.0 7.0 4.951.488
Superior Temporal Gyrus 22 R 45.0 −22.0 −5.0 4.709.594
Caudate Tail R 36.0 −43.0 10.0 4.463.713
95 Superior Frontal Gyrus 10 R 12.0 56.0 −8.0 4.874.894
Superior Frontal Gyrus 10 R 27.0 65.0 1.0 4.775.339
Superior Frontal Gyrus 10 R 15.0 68.0 7.0 4.741.182
119 Cingulate Gyrus 32 R 6.0 14.0 37.0 5.477.295
70 Posterior Cingulate 29 L 0.0 −43.0 10.0 5.409.723
Posterior Cingulate 23 R 3.0 −37.0 22.0 4.375.017
49 Cerebellum * L −3.0 −58.0 1.0 4.554.553
104 Thalamus (ventral lateral nucleus) L −18.0 −16.0 10.0 4.994.545
Thalamus (anterior nucleus) L −6.0 −7.0 10.0 4.737.922
33 Middle Frontal Gyrus 9 L −24.0 29.0 34.0 4.578.666
728 Amygdala L −33.0 −7.0 −14.0 6.569.848
Superior Temporal Gyrus 38 L −45.0 20.0 −23.0 6.025.189
Middle Temporal Gyrus 21 L −48.0 −4.0 −14.0 5.425.207
Superior Temporal Gyrus 22 L −48.0 −22.0 −8.0 5.288.910
Inferior Frontal Gyrus extending into Insula 47/13 L −30.0 14.0 −17.0 5.172.719
Putamen L −21.0 11.0 4.0 4.972.111
Inferior Frontal Gyrus 9 L −45.0 5.0 34.0 4.660.889
Middle Frontal Gyrus 46 L −42.0 17.0 19.0 4.500.771
Inferior Frontal Gyrus extending into Insula 9/13 L −33.0 8.0 22.0 4.102.611
Precentral Gyrus 6 L −57.0 −4.0 37.0 3.943.306
1027 Inferior Parietal Lobule 40 L −48.0 −25.0 25.0 6.028.291
Inferior Parietal Lobule 40 L −39.0 −49.0 37.0 5.800.646
Inferior Parietal Lobule 40 L −51.0 −34.0 34.0 5.711.774
Postcentral Gyrus 2 L −36.0 −25.0 37.0 5.598.609
Precuneus 7 R 24.0 −67.0 28.0 4.901.719
Precuneus 7 R 6.0 −52.0 34.0 4.705.517
Superior Temporal Gyrus 13 L −51.0 −43.0 19.0 4.411.334
Precuneus 31 L −18.0 −73.0 28.0 4.256.221
Insula 13 L −30.0 −22.0 25.0 4.077.991
Precuneus 7 L −18.0 −64.0 40.0 3.988.875

Frontiers in Human Neuroscience www.frontiersin.org June 2012 | Volume 6 | Article 192 | 14

Altmann et al. Emotional valence of stories and ToM

Table A6 | Brain regions showing positive connectivity with mPFC depending on individual tendency to put feel concern for other people
(Positive correlation between the empathic concern scale and mPFC–connectivity while reading negative stories > reading neutral stories).

Cluster size Region BA Talairach coordinates Max. t-value

X Y Z

81 Inferior Frontal Gyrus/Anterior Insula 47/13 R 33 29 1 0.785256

Putamen R 24 8 −5 0.655270
17 Posterior Cingulate 31 R 18 −58 19 0.719141
16 Middle Frontal Gyrus 8 L −33 17 43 0.830213
53 Insula 13 L −36 14 −2 0.733469

Table A7 | Prestudy results for valence and liking.

Conditions Valence Liking

(Scale −3 to +3) (Scale 1–7)

Mean SD Mean SD

Negative stories −1.36 0.60 3.67 0.63

Neutral stories 0.52 0.38 3.60 0.51

Mean values for each story were used in the fMRI experiment to modulate parametric regressors for the effect of valence and liking during reading. A 2 × 2 repeated
measures ANOVA was conducted with the factors story type (negative, neutral) and context (real, invented) to examine whether participant’s judgments of valence
reflect a clear differentiation between stories that have been categorized as neutral or negative in earlier studies during the stimulus selection phase. The main
effect for story type [F(1, 31) = 72.83, p < 0.001] showed that this is the case. Results revealed no main effect of context [F(1, 31) < 1, p < 0.717] and no context ×
story type interaction [F(1, 31) = 3.67, p < 0.065]. We did not find any effect for liking judgments. Stories were equally liked, independently of valence [F(1, 31) < 1,
p < 0.741] or context labelling [F(1, 31) = 1.33, p < 0.258].
Descriptive data on the item level show a clear difference on average valence for negative stories (M = −1.36, SD = 0.60) and neutral stories (M = 0.52, SD = 0.38).
Average liking values are very similar for negative (M = 3.67, SD = 0.63) and neutral (M = 3.60, SD = 0.51) texts. The latter finding fits nicely to the observation
that an object must not provide positive valence in order to be liked.

Frontiers in Human Neuroscience www.frontiersin.org June 2012 | Volume 6 | Article 192 | 15

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Short- and Long-
Term Effects of a Novel on
Connectivity in the Brain
Gregory S. Berns ( , Kristina Blaine, Michael J. Prietula, and Brandon E. Pye

Published Online: 9 Dec 2013 https://doi.org/10.1089/brain.2013.0166 Information

Copyright 2013, Mary Ann

Abstract Liebert, Inc.

We sought to determine whether reading a novel causes measurable To cite this article:
changes in resting-state connectivity of the brain and how long these
Gregory S. Berns, Kristina Blaine,
changes persist. Incorporating a within-subjects design, participants Michael J. Prietula, and Brandon E.
received resting-state functional magnetic resonance imaging scans on Pye.
Short- and Long-Term Effects of a
19 consecutive days. First, baseline resting state data for a “washin”
Novel on Connectivity in the Brain.
period were taken for each participant for 5 days. For the next 9 days, Brain Connectivity. Dec 2013.
participants read 1/9th of a novel during the evening and resting-state 590-600.
http://doi.org/10.1089/brain.2013.0166
data were taken the next morning. Finally, resting-state data for a “wash-
out” period were taken for 5 days after the conclusion of the novel. On %
Published in
the days after the reading, signi^cant increases in connectivity were Volume: 3 Issue 6: December 9,
centered on hubs in the left angular/supramarginal gyri and right 2013
posterior temporal gyri. These hubs corresponded to regions previously Online Ahead of Print:
October 9, 2013
associated with perspective taking and story comprehension, and the Online Ahead of Editing:
changes exhibited a timecourse that decayed rapidly after the August 29, 2013
completionWeof use
the cookies
novel. Long-term
to give youchanges in connectivity,
a better experience which
on liebertpub.com. By continuing to use our site, you are agreeing to
persisted for several days after the reading, were observed in bilateral Keywords
somatosensory cortex, suggesting a potential mechanism for
“embodied semantics.” connectivity fMRI

“ reading resting state

A great book should leave you with many experiences, and slightly
 exhausted at the end. You live several lives while reading.– William
Styron, Conversations with William Styron.

”
Most people can identify books that have made great impressions on
them and, subjectively, changed the way they think. Some can even
point to a book that has changed their life. Stephen King, for example,
said that Lord of the Flies changed his life, “because it is both a story
with a message and because it is a great tale of adventure.” Joyce Carol
Oates pointed to Alice in Wonderland as “the book that most in_uenced
her imaginative life.” It seems plausible that if something as simple as a
book can leave the impression that one's life has been changed, then
perhaps it is powerful enough to cause changes in brain function and
structure. Here, we test this possibility by using functional magnetic
resonance imaging (fMRI) to track changes in resting-state brain activity
on a daily basis over a period of 3 weeks, during which individuals read
a complete novel.

Novels are stories, and stories are complicated objects of

communication (Abbott, 2008).* Although several linguistic and literary
theories describe what constitutes a story, neurobiological research has
just begun to elucidate brain networks that are active when processing
stories. To date, these studies have focused on the immediate response
to short stories (Mar, 2011). In other words, current neurobiological
theory of stories describes the network of brain regions that is active
and presumably responsible for cognitive processing of stories while
they are being consumed. While active tasks have traditionally been
used to identify functional networks within the brain, resting-state fMRI
has become a common tool to identify consistent patterns of correlated
activity, termed resting-state networks (RSNs) (Biswal et al., 1995, 2010;
Kelly et al., 2012; Raichle et al., 2001).

Cognitive and emotional interventions have been demonstrated to

cause transient changes in functional connectivity (Harrison et al.,
2008; Hasson et al., 2009; Mackey et al., 2013; Stevens et al., 2010), but
it is not known how long these changes last. Some changes appear to
be due to transient activation of speci^c regions, which persists for
minutes to hours (Hasson et al., 2009); while others may persist for
longer periods of time and may represent cortical reorganization
(Mackey et al., 2013). A limitation of these studies that makes it digcult
to determine what are short- and long-term changes is the small
number of resting-state scans actually performed.

To determine a timescale over which connectivity changes persist, we

measured changes in resting-state connectivity as a result of reading a
novel. We chose a novel over a short story because the length and
depth of the novel would afford a set of repeated engagements with
associated, unique stimuli (sections of the novel) set in a broader,
controlled stimulus context that could be consumed between several
scanning periods. A within-subjects design was selected for this pilot
study because of its substantive control of individual variability,
statistical power, and economic advantages in this type of study
(Anderson, 2001; Shadish et al., 2002).

Materials and Methods

Participants
A total of 21 participants were studied. Two were excluded from the
fMRI analyses: one for insugcient attendance, and the other for image
abnormalities. Before the experiment, participants were screened for
the presence of medical and psychiatric diagnoses, and none were
taking medications. There were 12 female and 9 male participants
between the ages of 19 and 27 (mean 21.5). Emory University's
Institutional Review Board approved all procedures, and all participants
gave written informed consent.

Reading material
Each participant was subject to 19 consecutive days (July 18, 2011–
August 5, 2011) of resting-state scans that consisted of a total
appointment time of less than 30 min at the same time each day. The
^rst 5 days and last 5 days were “wash-in” and “washout” sessions,
respectively. Each of the middle 9 scans was preceded by reading
approximately 1/9th of the novel (Pompeii: A Novel, by Robert Harris,
Fawcett, 2003). This novel was chosen because it was based on true
events but written as historical ^ction and conveyed in a classic
narrative arc (Freytag, 1900). During the “washin” and “wash-out”
sessions, the participants did not perform any other tasks except for the
resting-state scan (Fig. 1). For each of the other 9 days, the story days,
the participants performed the resting-state scan after taking a quiz and
self-report about the effect of the material presented in the portion of
the novel that was assigned for the previous night and included a ^ve-
point rating scale of how arousing the reading was (see Supplementary
Data for quizzes; Supplementary Data are available online at
www.liebertpub.com/brain). Through repeated scans, each participant
served as his or her own control to measure changes in resting-state
connectivity after the consumption of the novel.
FIG. 1.  Design of experiment (above). Participants underwent resting-
state functional magnetic resonance imaging scans on 19 consecutive
days (black arrows). On the evening before the middle 9 days of
scanning, participants read a portion of the novel, Pompeii. The mean
arousal rating across participants (below) showed a rising trend toward
the climax of the novel (error bars±1 standard error).

Scanning
The scanning was performed on a Siemens 3T Trio. Each participant
received only one T1-weighted structural image (TR=2600 ms, TE=3.93 
ms, _ip angle=8°, 224×256 matrix, 176 sagittal slices, and 1 mm cubic
voxel size) throughout the duration of the experiment. One functional
resting-state scan was acquired each day (223 volumes, TR=2000 ms,
TE=30 ms, _ip angle=73°, FOV=192 mm, 64×64 matrix, 33 axial slices,
and 3×3×3.5 mm resolution with an added 10% gap in the z-direction,
resulting in a resolution of 3×3×3.85 mm). Participants were instructed
to rest quietly with eyes closed.

Preprocessing
All of the preprocessing was performed using the 1000 Functional
Connectomes Scripts available from NITRC (www.nitrc.org). The only
modi^cation to these scripts was the addition of an iterative loop to
cycle through the 19days of data. The scripts performed the following
preprocessing procedures using FSL (Analysis Group, FMRIB) and AFNI
(NIMH). First, the anatomical image was deobliqued and reorientated to
the coordinate space that is compatible with FSL. Next, the image was
skull stripped.

The resting-state functional images were preprocessed through a multi-

step procedure. The images were deobliqued and reoriented similarly to
the anatomical images. A mean functional image was computed to
serve as a target for motion correction. Using 3dvolreg, the functional
images were then aligned to the mean image using two-pass Fourier
interpolation. To decrease edge artifacts from Fourier interpolation, a
zero pad of four voxels was added around the edges and stripped off
after motion correction. The images were then skull stripped to create a
mask that was then applied to the motion-corrected data. To allow for
full magnetization and settling on any startle responses from the onset
of the scanning, the eighth volume was used for registration to the
anatomical image. Spatial smoothing was performed using a 6-mm
Gaussian kernel. Grand mean scaling was performed with an intensity
normalization to 10,000. A low-pass ^lter of 0.1 Hz and a high-pass ^lter
of 0.005 Hz were applied for temporal ^ltering. The images were
detrended by calculating the mean of the temporally ^ltered image and
detrending with the addition of Legendre polynomials of an order up to
and including two. An image that was the addition of both the mean and
detrended image was created.

Three separate registration alignments were performed. The functional

(using the eighth image acquisition as a template) to anatomical
alignment was produced using a trilinear interpolation (six degrees of
freedom). The anatomical to standard brain (MNI152_T1) was created
again using a trilinear interpolation (12 degrees of freedom). The
transformation matrices of both of these steps were saved. A third
matrix, for the ability to transform between functional to standard, was
created by concatenating the matrices of the previous two steps. The
inverse of each of these matrices was also produced, providing the
option of registering freely between any combination of functional,
anatomical, or standard images.

Segmentation was performed to create individual images for each

tissue type and individual probability maps. The tissue types recognized
as cerebrospinal _uid (CSF) and white matter (WM) were masked.
These masks were used to control for nuisance signals. We utilized the
global signal, WM and CSF segmentation masks, and the six motion
parameters to adjust the functional signals for the effects of
physiological noise and motion (Yan et al., 2013). Although adjustment
for global signals is controversial, we opted to take a conservative
approach and control for physiological noise (Fox et al., 2009). This
approach may protect against false positives but may introduce
spurious negative correlations (Murphy et al., 2013; Weissenbacher et
al., 2009), so our analysis focused only on changes in positive
correlations.

Analysis
Two of the participants were not present on the ^rst day of scanning,
and a third was absent on the last day of scanning. Therefore, the 19
participants were analyzed over 17 consecutive days of scanning (days
2–18).

Using a prede^ned network of 160 regions of interest (ROI) (Dosenbach

et al., 2010), we extracted the time series of each ROI for spheres of 6 
mm radius for each person on each day. We chose this set of ROIs
because the number of ROIs strikes a balance between a reasonable
number and good cortical coverage. Because the original paper was a
developmental neuroscience study, our results on reading are
particularly relevant to the study of cognitive development. This yielded
a four-dimensional matrix of ROI×volume×person×day
(160×223×19×17). Next, we computed the pairwise cross-correlation
between ROIs for each scan session (Mackey et al., 2013), yielding a
matrix with dimensions 160×160×19×17. We then applied the Fisher z-
transformation to normalize the correlation coegcients, which are
bounded by±1.

Statistics were performed using the Network-Based Statistics (NBS)

Connectome v1.2 (Zalesky et al., 2010). All connections in the z-
transformed correlation matrices were submitted to a one-sided t-test
to see which individual connections were signi^cantly different based
on the speci^ed contrast. The design matrix for these tests included a
column for each day and dummy variables for each subject to control
for subjectwise differences in mean correlations. Thus, there were 36
columns (17 days + 19 subjects). Contrasts were speci^ed as vectors of
differences across the 17 day-columns (see below). Due to the large
number of elements in the correlation matrix (12,720 unique elements),
connections that surpassed p<0.001 (t=3.32) in signi^cance were then
submitted to a permutation test to control for familywise error rate. NBS
is more sensitive to detecting networks of topologically connected
nodes, while the related approach—false discovery rate—is more
sensitive to strong, focal connections. Since we assume that a novel
engages many regions, its effects are likely to be extended over a
network of connections rather than a small number of connections. For
this reason, we used the NBS intensity statistic. For each contrast, 5000
permutations were performed. Permutations were restricted to be done
within subjects only.

Three sets of contrasts were speci^ed. First, to determine whether there

were any signi^cant changes in connectivity between the beginning and
the end of the story, we examined the contrasts: [washout−washin] and
[washin−washout]. Second, to determine short-term changes in
connectivity as a result of the story and potential reactivation due to the
daily quiz, we contrasted: [story−washin−washout], with appropriate
weightings for the different number of days. Finally, to determine long-
term changes that were related to the story but persisted beyond the
reading days, we contrasted: [story+washout−washin], again with
appropriate weightings for the different number of days.

Networks were visualized by displaying nodes and connections in

BrainNet Viewer (www.nitrc.org/projects/bnv/). Timeseries for each
network were computed by averaging the correlation coegcients in
each connection of the network for each subject on each day and then
computing the mean and standard error across subjects for each day.
Results
Consistent with theories of plot structure, the mean arousal ratings of
the story rose consistently throughout the story and culminated with the
climax—the eruption of the volcano and the destruction of Pompeii (Fig.
1).

For the ^rst set of contrasts, [washout−washin] showed positive

correlations that changed signi^cantly between the beginning and the
end of the story. This was a small network of eight nodes and eight
connections, all in the cerebellum (Table 1 and Fig. 2). The timeseries of
this network showed both a monotonic trend throughout most days, but
interestingly, the largest increase in correlation was after the ^rst night's
reading. During the story days, the correlation _uctuated, but not below
pre-story levels, and rose on the last story day with a continued rise
after the story. The opposite contrast, [washin−washout] revealed a
slow decline in correlations within a network between the left
cerebellum and left pre/post central gyrus. These correlations were
generally low (<0.1) and decreased to ∼0.05 by the end of the
experiment. Given these monotonic trends and preponderance of
connectivity changes within cerebellar regions, we do not consider
these changes related to the story.

FIG. 2.  Networks associated with increased connectivity after the

novel [p=0.022 corrected for familywise error rate (FWER)] and
decreased connectivity (p=0.003 corrected for FWER). Both networks
showed generally monotonic changes in correlation strength with time,
suggesting that these changes may not be related directly to the novel
itself. It is noteworthy that both networks have strong hubs in the
cerebellum.

Table 1. Washout Versus Washin Connections

Node MNI Label Node MNI Label

Washout>washin (p=0.022)

  −6 L cerebellum 131 −34 L

120 −60 −67 cerebellum
−15 −29
  1 Cerebellar 131 −34 L
130 −66 vermis −67 cerebellum
−24 −29

  −24 L cerebellum 140 33 R

109 −54 −73 cerebellum
−21 −30

  −24 L cerebellum 150 −21 L

109 −54 −79 cerebellum
−21 −33

  −6 L cerebellum 150 −21 L

120 −60 −79 cerebellum
−15 −33

  −6 L cerebellum 151 −6 L
120 −60 −79 cerebellum
−15 −33

  1 Cerebellar 151 −6 L
130 −66 vermis −79 cerebellum
−24 −33

  −34 L cerebellum 155 18 R

131 −67 −81 cerebellum
−29 −33

Washout<washin (p=0.003)

   −6 L mid 43 0 SMA
30 17 cingulate −1
34 52

   −2 ACC 103 −59 L middle

19 30 −47 temporal
27 11
   −6 L mid 103 −59 L middle
30 17 cingulate −47 temporal
34 11

   −12 L thalamus 103 −59 L middle

57 −12 −47 temporal
6 11

   11 R thalamus 103 −59 L middle

58 −12 −47 temporal
6 11

   −30 L putamen 103 −59 L middle

61 −14 −47 temporal
1 11

   0 SMA 109 −24 L

43 −1 −54 cerebellum
52 −21

   −42 L insula 109 −24 L

48 −3 −54 cerebellum
11 −21

   −38 L pre/post 110 −37 L

62 −15 central g −54 cerebellum
59 −37

   −42 L insula 113 −34 L

48 3 −57 cerebellum
11 −24

   −38 L pre/post 113 −34 L

75 −27 central g −57 cerebellum
60 −24

   −42 L insula 120 −6 L

48 −3 −60 cerebellum
11 −15

   −38 L pre/post 120 −6 L

62 −15 central g −60 cerebellum
59 −15

   −38 L pre/post 120 −6 L

75 −27 central g −60 cerebellum
60 −15

   −42 L insula 121 −25 L

48 −3 −60 cerebellum
11 −34

   0 SMA 128 21 R
31 15 −64 cerebellum
45 −22

   0 SMA 128 21 R
43 −1 −64 cerebellum
52 −22

   −42 L insula 130 1 Cerebellar

48 −3 −66 vermis
11 −24

   −38 L pre/post 130 1 Cerebellar

75 −27 central g −66 vermis
60 −24

   −42 L insula 131 −34 L

48 −3 −67 cerebellum
11 −29

   −55 L 136 −9 L
97 −44 supramarginal −72 precuneus
 30 g 41

  −59 L middle 136 −9 L

103 −47 temporal −72 precuneus
11 41

Node is the node number based on the sorting in table S6 by

Dosenbach et al. (2010). Network signi^cance is based on 5000
permutations, correcting for FWER. MNI are x, y, z coordinates. Label
is based on AFNI “whereami” function and CA_ML_18_MNIA atlas.

FWER, familywise error rate; MNI, montreal neurologic institute; AFNI,

analysis of functional neuroimage.

To isolate the short-term changes associated with reading the story, we

combined the washin and washout periods and contrasted them with
the story days. This contrast identi^ed three independent networks that
had signi^cant increases in connectivity during the story days (Table 2).
Network 1 had a prominent hub around the left angular and
supramarginal gyri with connections to both the precuneus and medial
frontal lobe (Fig. 3). There was also a signi^cant connection to the left
lingual gyrus in the vicinity of the hippocampus. The timecourse of
correlations within this network showed a striking pattern of a sharp
rise on the ^rst story day, reaching its peak on the last story day,
followed by a nonlinear decay. Network 2 was a bilaterally distributed
network without prominent hubs (Fig. 4). Signi^cant connections were
mostly posterior and located in the superior temporal gyri and cuneus
(Table 2). The timecourse was not as clearly related to the story, with
correlations peaking on the second story day and then declining. The
correlations were also lower than in Network 1. Finally, Network 3 had
signi^cant connections between a hub in the right middle temporal
gyrus and the left pre/post central gyrus and left superior temporal
gyrus (Fig. 5). The timecourse of correlations of this network showed
the same striking increase with the onset of the story. Unlike Network 1,
this network did not have a nonlinear decay. The magnitude of
correlations were similar to that of Network 1.

FIG. 3.  Network 1 (p=0.009 corrected for FWER) of nodes and

connections with signi^cantly increased correlation during story versus
nonstory days. This network was concentrated in a hub around the left
angular and supramarginal gyri, with connections to medial prefrontal
cortex. The timecourse of correlations across days showed a sharp rise
beginning on the ^rst post-story day and a decay after the end of the
novel.

FIG. 4.  Network 2 (p=0.012 corrected for FWER) of nodes and

connections with signi^cantly increased correlation during story versus
nonstory days. This sparse network was located in posterior temporal
gyri with connections to the cuneus. The timecourse of correlations
across days showed a rise beginning on the ^rst post-story and peaking
on the second story day, followed by a decline throughout.

FIG. 5.  Network 3 (p=0.023 corrected for FWER) of nodes and

connections with signi^cantly increased correlation during story versus
nonstory days. This sparse network was located in posterior temporal
gyri with connections to the central sulcus. The timecourse of
correlations across days showed a sharp rise beginning on the ^rst
post-story that was sustained at a relatively constant level throughout
the story, followed by a sharp decline post-story.

Table 2. Story Versus Nonstory Connections

Node MNI Label Node MNI

Network 1 (p=0.009)

 90 −8 L lingual 97 −55 L

−41 g/hippocampus −44 supramarginal
3 30 g

 11 −11 L sup medial g 104 −53 L angular g

45 −50
17 39

 13 8 R ACC 104 −53 L angular g

42 −50
−5 39

 17 23 R sup frontal g 104 −53 L angular g

33 −50
47 39
 17 23 R sup frontal g 107 44 R angular g
33 −52
47 47

 97 −55 L supramarginal g 108 −5 L precuneus

−44 −52
30 17

  −53 L angular g 108 −5 L precuneus

104 −50 −52
39 17

 97 −55 L supramarginal g 111 10 R precuneus

−44 −55
30 17

 97 −55 L supramarginal g 115 −11 L precuneus

−44 −58
30 17

  −53 L angular g 134 −36 L angular g

104 −50 −69
39 40

 97 −55 L supramarginal g 136 −9 L precuneus

−44 −72
30 41

  −53 L angular g 136 −9 L precuneus

104 −50 −72
39 41

 97 −55 L supramarginal g 141 −2 L cuneus

−44 −75
30 32
 97 −55 L supramarginal g 155 18 R cerebellum
−44 −81
30 −33

Network 2 (p=0.012)

 43 0 SMA 82 −41 L sup temp g

−1 −37
52 16

 70 42 R sup temp g 86 34 R post central

−24 −39 g
17 65

 82 −41 L sup temp g 129 19 R lingual g

−37 −66
16 −1

 64 −47 L post central g 145 −16 L sup occipital

−18 −76 g/cuneus
50 33

 82 −41 L sup temp g 145 −16 L sup occipital

−37 −76 g/cuneus
16 33

 89 58 R sup 145 −16 L sup occipital

−41 temp/supramarginal −76 g/cuneus
20 g 33

 51 46 R post central 148 15 R cuneus

−8 g/insula −77
24 32

 70 42 R sup temp g/insula 148 15 R cuneus

−24 −77
17 32
 89 58 R sup 148 15 R cuneus
−41 temp/supramarginal −77
20 g 32

 82 −41 L sup temp g 156 −37 L inf occipital

−37 −83 g
16 −2

Network 3 (p=0.023)

 60 59 R sup temp g 68 −54 L sup temp g

−13 −22
8 9

 68 −54 L sup temp g 77 −24 L post central

−22 −30 g
9 64

 49 −44 L post central g 123 46 R middle temp

−6 −62 g
49 5

 62 −38 L precentral g 123 46 R middle temp

−15 −62 g
59 5

 68 −54 L sup temp g 123 46 R middle temp

−22 −62 g
9 5

 69 41 R pre/post central g 123 46 R middle temp

−23 −62 g
55 5

 75 −38 L pre/post central g 123 46 R middle temp

−27 −62 g
60 5
  75 −38 L pre/post central g 135 39 R middle
−27 −71 temp/occipital
60 13 g

Node is the node number based on the sorting in table S6 by

Dosenbach et al. (2010). Network signi^cance is based on 5000
permutations, correcting for FWER. MNI are x, y, z coordinates. Label
is based on AFNI “whereami” function and CA_ML_18_MNIA atlas.

To identify potential long-term changes in connectivity, we contrasted

[story+washout−washin]. This contrast identi^ed increases in
correlation during story days and persisted during the washout period.
One network was identi^ed with bilateral connections between pre/post
central gyri, middle and superior temporal gyri, and insula (Table 3 and
Fig. 6). The timecourse of correlations showed the increase occurring
with the onset of the story, peaking on the sixth or eighth story day, and
declining slightly afterward. All the correlations during the washout days
were higher than the washin period (Fig. 6).

FIG. 6.  Network (p=0.001 corrected for FWER) of nodes and

connections with signi^cantly increased correlation during story days
and that persisted beyond the story. This network was located
bilaterally around the central sulcus with sparse connections to the
insula and occipital regions. The timecourse of correlations across days
showed a gradual rise beginning on the ^rst post-story day that was
sustained beyond the end of the novel.

Table 3. Story+Washout Versus Washin Connections

Node MNI Label Node MNI Label

Story+washout>washin (p=0.001)

 18 34 R inf 42 43 R insula

32 frontal 1
7 g/insula 12

 53 44 R 62 −38 L
−11 pre/post −15 precentral
 38 central g 59 g

 42 43 R insula 69 41 R
1 −23 pre/post
12 55 central g

 43 0 SMA 69 41 R
−1 −23 pre/post
52 55 central g

 49 −44 L post 70 42 R sup

−6 central g −24 temp g
49 17

 52 −54 L 74 18 R
−9 pre/post −27 pre/post
23 central g 62 central g

 51 46 R 75 −38 L
−8 pre/post −27 pre/post
24 central g 60 central g

 70 42 R sup 75 −38 L

−24 temp g −27 pre/post
17 60 central g

 54 −47 L 76 −30 L insula

−12 pre/post −28
36 central g 9

 74 18 R 76 −30 L insula

−27 pre/post −28
62 central g 9

 62 −38 L 77 −24 L post

−15 precentral −30 central g
59 g 64
 64 −47 L post 77 −24 L post
−18 central g −30 central g
50 64

 69 41 R 77 −24 L post

−23 pre/post −30 central g
55 central g 64

 62 −38 L 78 51 R
−15 precentral −30 sup/mid
59 g 5 temp g

 70 42 R sup 78 51 R
−24 temp g −30 sup/mid
17 5 temp g

 74 18 R 78 51 R
−27 pre/post −30 sup/mid
62 central g 5 temp g

 76 −30 L insula 82 −41 L sup

−28 −37 temp g
9 16

 64 −47 L post 83 −53 L sup/mid

−18 central g −37 temp g
50 13

 42 43 R insula 95 43 R
1 −43 sup/mid
12 8 temp g

 82 −41 L sup 118 −34 L inf

−37 temp g −60 occipital
16 −5 g

 49 −44 L 123 46 R middle

 −6 pre/post −62 temp g
49 central g 5

 52 −54 L 123 46 R middle

−9 pre/post −62 temp g
23 central g 5

 65 46 R 123 46 R middle

−20 pre/post −62 temp g
45 central g 5

 69 41 R 123 46 R middle

−23 pre/post −62 temp g
55 central g 5

 64 −47 L post 145 −16 L sup

−18 central g −76 occipital
50 33 g/cuneus

 65 46 R 145 −16 L sup

−20 pre/post −76 occipital
45 central g 33 g/cuneus

 95 43 R 145 −16 L sup

−43 sup/mid −76 occipital
8 temp g 33 g/cuneus

 82 −41 L sup 156 −37 L inf

−37 temp g −83 occipital
16 −2 g

Node is the node number based on the sorting in table S6 by

Dosenbach et al. (2010). Network signi^cance is based on 5000
permutations, correcting for FWER. MNI are x, y, z coordinates. Label
is based on AFNI “whereami” function and CA_ML_18_MNIA atlas.

Discussion
 Before interpreting the changes in RSNs, it is worth examining the
repeatability of resting-state scans. Previous work has shown that three
resting-state scans—two within an hour and one 5–16 months later—
demonstrated a modest to high degree of repeatability in the spatial
components identi^ed through temporal concatenation independent
component analysis (ICA) as well as targeted ROIs (Damoiseaux et al.,
2006; Shehzad et al., 2009; Zuo et al., 2010), especially if signals from
the CSF were regressed out as nuisance variables (Chang and Glover,
2009; Li et al., 2012). Similar test–retest reliability from temporal
concatenation ICA was obtained in older adults scanned twice, 1 year
apart (Guo et al., 2012). Measures of theoretical graph connectivity
showed moderate test–retest reliability, depending on temporal ^lter
parameters (Braun et al., 2012). Therefore, RSNs are a viable measure
of brain network reorganization due to a salient experience, as these
networks appear relatively stable and reliable across time in the
absence of signi^cant events. This raises the question of whether
reading a novel is sugciently powerful to cause a detectable
reorganization of cortical networks.

The timescale of the effect of a novel may be both short and long term.
Short-term effects might be observed immediately after reading. For
example, RSNs are known to be altered by recent language
comprehension tasks (Hasson et al., 2009) as well as visual
categorization tasks (Stevens et al., 2010). Although the chapter
readings were performed during the evenings before scans, the quizzes
occurred just before the scan. The quizzes, therefore, might be
responsible for such immediate changes in resting state, though the
tasks differ in their orientation. The primary (evening) task involved

Pactive consumption of the story, while the next morning the quiz task
S involved re_ection of the story, where the latter task would likely engage
in areas associated with autobiographical recall. The magnitude of the
arousal score was not signi^cantly correlated with the changes in
connectivity. Even so, recent evidence suggests that resting-state
changes persist for a day after a cognitive intervention, such as
neurofeedback with a “focusing” effect on loci of activity (Harmelech et
al., 2013). Thus, although the reading quiz may be partly responsible for
short-term changes in RSNs, there is now evidence that these changes
may also be due to carryover from the previous evening.

Considering both the evening carryover and the quiz reactivation as

short-term effects, three independent cortical networks demonstrated
increases in connectivity as a result of the novel. Network 1 (Fig. 3)
displayed the strongest localization to a hub centered around the left
angular and supramarginal gyrus with increases in connectivity to both
the medial prefrontal cortex (MPFC) and cuneus. The timecourse of
correlation within this network displayed a nonlinear decay during the
washout period that is consistent with a lingering, but decaying effect.
This decay suggests that the changes were not solely due to the quiz,
which would have had more of an on–off effect as seen in Network 3
(Fig. 5). The nodal center around the left angular gyrus in Network 1 is
consistent with this region's well-known role in language
comprehension. A recent meta-analysis of theory of mind studies
identi^ed the left angular gyrus as the third most likely region to be
activated if the task was story based (behind right angular gyrus and
MPFC—both of which also appear in Network 1) (Mar, 2011). Thus, the
implication is that the activation of these regions during the evening
reading carried over to the next morning as changes in connectivity.

One explanation expands on the concept of resting state as a dynamic

organizational construct (Deco et al., 2011). These “resting-state”
networks marshal the brain's earlier functional engagement of the novel,
particularly the multimodal associative regions around the
temporoparietal junction (TPJ) (angular and supramarginal gyri, and
middle and superior temporal gyri). This interpretation rests on the
principle that the brain is a prediction engine. That is, the resting state
of the brain is best viewed as being in a “constant inner state of
exploration, in which the brain generates predictions about the likely
network con^guration that would be optimal for a given impending
input” (Deco et al., 2011). Accordingly, since earlier cognitive
experiences may modulate the resting-state connectivity maps, the task
of reading Pompeii conditionally altered the resting state of our
participants with a bias toward a hybrid mentalizing-narrative network
con^guration even though they were not actively engaged in a task.

Could these speci^c neural effects be mere consequences not related

to story consumption? Variance attributed to the story consumption
“treatment” certainly contains an error term, but the resultant network
components identi^ed are distinctly task speci^c and revealed across
participants rising above the heterogeneous in_uences of their “off-
task” (reading days) as well as “pre-task” (washin period) experiential
milieu. It is, however, possible, that the observed changes in
connectivity could be due to the overall experimental context of being
scanned after reading chapters from a novel—the experiment itself
triggers an active process of remembering the previous night's reading,
which was further primed by a quiz. Even so, the fact that reading a
novel caused changes in cortical connectivity places a bound on the
stability of RSNs. While largely stable, the resting state should properly
be conceived of as quasi-static and subject to both short- and long-term
dynamic recon^gurations.
Longer-term changes in connectivity were identi^ed by the contrast of
[story+washout−washin]. This contrast identi^ed connections that
increased in strength during the story days and remained elevated after
the novel (Fig. 6). This network was heavily concentrated around the
central sulcus bilaterally with connections to bilateral posterior
temporal gyri and insula. This network corresponds closely to a
previously identi^ed RSN comprising somatosensory and motor regions
(De Luca et al., 2006). Correlated _uctuations in motor cortex are well
known and may not be due to a speci^c cognitive process (Biswal et al.,
1995; Xiong et al., 1999). Thus, we are left with the question as to why
these correlations increased with the onset of the novel.

One possibility for increases in somatosensory cortex connectivity is

that reading a novel invokes neural activity that is associated with bodily
sensations. This is called the theory of “embodied semantics” (Aziz-
Zadeh and Damasio, 2008). Somatosensory cortex activation has been
previously demonstrated by the reading of metaphors, especially if they
are tactile metaphors (Lacey et al., 2012). It is plausible that the act of
reading a novel places the reader in the body of the protagonist, which
may alter somatosensory and motor cortex connectivity. It is interesting
to note, however, that the regions previously called the “protagonist
network”—dorsomedial PFC and right TPJ (Mason and Just, 2009)—
constitute a very different network than the somatosensory regions and
bear more similarity to the networks we identi^ed as having short-term
changes. This network can be compared with the predominately
cerebellar network identi^ed by the [washout−washin] contrast (Fig. 2).
Although the latter represents the most obvious change from before
and after the novel, the timecourse is largely monotonic and not clearly
related to the novel per se. However, it is possible that both the
cerebellar changes and the somatosensory changes re_ect changes in
motor control related to the act of reading. Such processes might relate
to oculomotor coordination and attention, for example, and have
nothing to do with the content of the novel.

In summary, we have demonstrated that across the likely array of

diverse experiences encountered by our participants, there was a
detectable and signi^cant common alteration of their RSN associated
with reading sections of a novel the previous evening. Moreover, these
changes could be segregated into networks associated with short-term
changes originating near the left angular gyrus and long-term changes
dispersed bilaterally in somatosensory cortex. It remains an open
question for further study as to how lasting these effects are, but our
results suggest a potential mechanism by which reading stories not
only strengthen language processing regions but also affect the
individual through embodied semantics in sensorimotor regions.
Acknowledgments
This research was supported by a grant from DARPA (D11AP00289).
The views, opinions, and/or ^ndings contained in this article are those
of the authors and should not be interpreted as representing the ogcial
views or policies, either expressed or implied, of the Defense Advanced
Research Projects Agency or the Department of Defense. This work was
approved for Public Release, Distribution Unlimited. The authors are
grateful for the input of Andrew Brooks, Lisa LaViers, W. Gavin Ekins,
and Ting Li.

Author Disclosure Statement

No competing ^nancial interests exist.

*“Story” and “narrative” are often used interchangeably. Technically,

narrative (vs. expository text) is the representation of events, consisting
of a story (as a sequence of events, participating characters, and causal
associations linking acts) and discourse (how the story is physically
conveyed). Thus, all conveyed stories are mediated by a narrative
discourse (e.g., voice, text, video, actors).4

) * +
© 2023 Mary Ann Liebert, Inc., publishers. All rights reserved, USA and
worldwide.
Call us toll free at (800) M-LIEBERT (800-654-3237).
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Think abouteveryone
who buys books for
the fun of it a big
corporations thatfeed
heseaddictions

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 crazy

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me

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 don'tknow if I'll reread books
but some
give me
the vibe

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 If extintrins
motivat

I do
this

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 Research Report
Psychological Science

Becoming a Vampire Without Being Bitten: 22(8) 990–994

© The Author(s) 2011
Reprints and permission:
The Narrative Collective-Assimilation sagepub.com/journalsPermissions.nav
DOI: 10.1177/0956797611415541

Hypothesis http://pss.sagepub.com

Shira Gabriel and Ariana F. Young

University at Buffalo, State University of New York

Abstract
We propose the narrative collective-assimilation hypothesis—that experiencing a narrative leads one to psychologically become a
part of the collective described within the narrative. In a test of this hypothesis, participants read passages from either a book
about wizards (from the Harry Potter series) or a book about vampires (from the Twilight series). Both implicit and explicit
measures revealed that participants who read about wizards psychologically became wizards, whereas those who read about
vampires psychologically became vampires. The results also suggested that narrative collective assimilation is psychologically
meaningful and relates to the basic human need for connection. Specifically, the tendency to fulfill belongingness needs through
group affiliation moderated the extent to which narrative collective assimilation occurred, and narrative collective assimilation
led to increases in life satisfaction and positive mood, two primary outcomes of belonging. The implications for the importance
of narratives, the need to belong to groups, and social surrogacy are discussed.

Keywords
need to belong, collective identity, reading, social identity
Received 2/2/11; Revision accepted 4/8/11

We read to know we are not alone.
—C. S. Lewis in Shadowlands (Attenborough, 1993)
Humans are driven by a need for social connection (Baumeis-
ter & Leary, 1995; Maslow, 1968), which propels them to
affiliate with collectives (i.e., groups).1 Purportedly, the sur-
vival value of collective life for humans’ evolutionary ances-
tors (Caporael & Brewer, 1995; Wilson, 1978) led to the
evolution of internal mechanisms that propel modern humans
to form and join collectives (Stevens & Fiske, 1995). These
mechanisms predispose people to experience pleasure, such as
Narratives have previously been linked to fulfillment of
belongingness needs, albeit in a less direct manner. For exam-
ple, correlational data suggest that narratives increase social
skills by enabling people to learn the rules of human interac- mimic
tion and empathy (Mar & Oatley, 2008; Mar, Oatley, Hirsch,
dela Paz, & Peterson, 2006; Oatley, 1999). In addition, identi-
fying with characters while reading a narrative leads to a
merging ofbecome
you self withprotagonist
the characters (Sestir & Green, 2010).
Further, narratives play a role in alleviating loneliness, such
that people are drawn to familiar narratives when they feel
lonely, feel less lonely after engaging in familiar narratives,

increased life satisfaction and positive affect, from collective
affiliations (Baumeister & Leary, 1995; Myers, 1992). The
potency of the desire for collective bonds leads people to eas-
ily assimilate collective identities, even on the basis of the
most minimal criteria (Tajfel, 1970). People may even assimi-
ope
tending late a collective to which they do not belong, adopting its
fitin behaviors, attitudes, and traits (DeMarree, Wheeler, & Petty,
2005; Kawakami, Young, & Dovidio, 2002). Here, we propose
that narratives provide the positive experience of connection
to a collective. Specifically, we propose and examine the nar-
rative collective-assimilation hypothesis, the hypothesis that
experiencing a narrative leads to psychological assimilation of
the collective described within the narrative.
r
and engage in narratives to decrease the detrimental effects of
social rejection (Derrick, Gabriel, & Hugenberg, 2009).
Finally, many of the neural regions that are activated when
people read about activities overlap substantially with the
same
regions that are activated when people imagine and actuallyactivation
engage in the activities (Speer, Reynolds, Swallow, & Zacks,officio
realactin
2009).
Corresponding Author:
Shira Gabriel, Department of Psychology, University at Buffalo, SUNY, Park
Hall, Buffalo, NY 14260
E-mail: sgabriel@buffalo.edu
Narrative Collective Assimilation
In summary, previous research demonstrates that there is a
strong human desire to belong to collectives and that people
easily assimilate collective identities. Further, people are
drawn to narratives when they feel lonely, and the brain
regions activated during reading of narratives are highly
similar to the regions involved in actual behavior. Thus, dispa-
rate branches of research imply the narrative collective-
assimilation hypothesis. In the experiment reported here, we
tested this hypothesis by examining whether reading a passage
from Harry Potter and the Sorcerer’s Stone (Rowling, 1999)
would lead participants to “become” wizards, and reading a
passage from Twilight (Meyer, 2005) would lead participants
to “become” vampires. We also tested the hypothesis that
narrative collective assimilation would be moderated by level
of collective self-construal; that is, we expected that the more
participants tended to fulfill their social needs through identi-
fication with collectives, the more they would exhibit narra-
tive collective assimilation. Finally, we reasoned that narrative
collective assimilation would lead to the same effects as sati-
ated belongingness needs—life satisfaction and positive
mood—and we tested this hypothesis as well.
Method
Participants were 140 undergraduates (72 men, 68 women;
79% White; mean age = 19 years) at the University at Buffalo,
State University of New York. They had completed the Collec-
tive and Relational Self-Construal Scales (Cross, Bacon, &
Morris, 2000; Gabriel & Gardner, 1999), measures of the ten-
dency to fulfill the need to belong through collective and rela-
tional bonds, during a mass-testing session earlier in the
semester. An example item from the Collective Self-Construal
Scale is, “When I join a group, I usually develop a strong sense
of identification with that group.”
Participants were told that the purpose of the study was to
examine people’s responses to books and movies. First, par-
ticipants read a passage from either Twilight (Meyer, 2005) or
Harry Potter and the Sorcerer’s Stone (Rowling, 1999), with
the direction that they should read as they would normally do
when reading for their own pleasure. Participants in the Twi-
light condition read chapter 13 (“Confessions”), in which
Edward (a vampire) describes what it is like to be a vampire to
his romantic interest, Bella. Participants in the Harry Potter
condition read chapters 7 (“The Sorting Hat”) and 8 (“The
Potions Master”), in which Harry and his friends (all wizards)
are sorted into “houses” and Harry first encounters Severus
Snape. Participants advanced to the next part of the experi-
ment when they finished the assigned passage or when 30 min
had elapsed. Participants read for an average of 25.5 min.
After participants finished reading, they completed an iden-
tity Implicit Association Test (identity IAT; Gabriel, Kawakami,
Bartak, Kang, & Mann, 2010; Nosek, Banaji, & Greenwald,
2002), the scores from which were our main dependent vari-
able. This response latency task assessed participants’ implicit
991
identification with vampires relative to wizards. Participants
completed two critical blocks, each consisting of 40 trials. In
one critical block, participants were instructed to categorize
“me” words (myself, mine) and “wizard” words (wand, broom-
stick, spells, potions) using the same response key and to cate-
gorize “not me” words (they, theirs) and “vampire” words
(blood, undead, fangs, bitten) using another response key. In the
other critical block, these pairings were reversed. Participants
were instructed to respond as quickly and accurately as possi-
ble. The underlying rationale for the IAT is that the speed with
which participants respond to two stimuli using the same key is
an indication of implicit associations between the two catego-
ries. In identity IATs, it is assumed that people are quicker to
respond to trials in which the self is indicated with the same key
as the assimilated collective than to trials in which the self is
indicated with the same key as some other group (Gabriel
et al., 2010). Thus, we predicted that participants who read the
Harry Potter chapters would respond more quickly when “me”
words and “wizard” words were categorized using the same key
rather than different keys, whereas those who read the Twilight
chapter would respond more quickly when “me” words and
“vampire” words were categorized using the same key rather
than different keys.
We next administered an explicit (albeit somewhat indirect)
measure of collective assimilation, which we call the Twilight/
Harry Potter Narrative Collective-Assimilation Scale. Embed-
ded among filler questions were three items designed to mea-
sure collective assimilation of Twilight vampires (“Compared
to the average person, how high do you think you could jump?”
“How long could you go without sleep?” and “How sharp are
your teeth?”) and three items designed to measure collective
assimilation of Harry Potter wizards (“How British do you
feel?” “Do you think, if you tried really hard, you might be
able to make an object move just using the power of your
mind?” and “Do you think you might ever be able to make
yourself disappear and reappear somewhere else?”).
Participants then completed the Transportation Scale, a
measure of level of absorption into a story (Green & Brock,
2000); a five-item version of the Positive and Negative Affect
Schedule (Watson, Clark, & Tellegen, 1988), a measure of
mood; a single-item life-satisfaction measure (“Right now, in
most ways my life is close to my ideal”; Schimmack & Oishi,
2005); and questions about the books and participants’ reading
habits.
Results
IAT scores were calculated by subtracting the mean response
latencies for trials that paired “me” words with “vampire”
words and “not me” words with “wizard” words from the
mean response latencies for trials that paired “me” words with
“wizard” words and “not me” words with “vampire” words.
Thus, higher (positive) IAT scores indicated stronger associa-
tions between the self and vampires, whereas lower (negative)
992
IAT scores reflected stronger associations between the self and
wizards. Five participants were excluded from analysis
because more than 10% of their response latencies were less
than 300 ms.2 Scores on the explicit measure, the Twilight/
Harry Potter Narrative Collective-Assimilation Scale, were
computed by averaging responses to the wizard items (α = .43)
and responses to the vampire items (α = .60) separately, con-
verting these averages to z scores, and then subtracting the
z score for wizard items from the z score for vampire items.
Mood was calculated by reverse-scoring the negative-mood
items and averaging these scores with the scores for positive
items (α = .61).
Scores on the identity IAT were examined using multiple
regression analyses recommended by Aiken and West (1991).
The predictor variables were collective self-construal, book
read (represented as a dichotomous variable: 0 = Harry Potter;
1 = Twilight), and the interaction of collective self-construal
and book read. All predictors were centered at their means (as
were all predictors in subsequent analyses). All regression
terms were entered into the model and interpreted simultane-
ously. The overall regression was significant, F(3, 128) =
20.04, p < .001. The narrative collective-assimilation hypoth-
esis was confirmed: Participants who read the Harry Potter
chapters self-identified as wizards (M = −0.27), whereas par-
ticipants who read the Twilight chapter self-identified as vam-
pires (M = 0.30), t(131) = 7.54, p < .001; semipartial r2 = .30.
Our second hypothesis was that narrative collective
assimilation would be moderated by level of collective self-
construal. Indeed, analysis of the identity IAT scores revealed
that the Collective Self-Construal × Book Read interaction
was significant, β = −0.17, t(128) = 2.34, p = .02, semipartial
r2 = .03 (see Fig. 1). Although participants low in collective
self-construal (1 SD below the mean) became members of the
0.5 High CSC
0.4
Association→
Low CSC
Vampire-Self
0.3
0.2
IAT Score
0.1
0
–0.1
←Wizard-Self
Association
–0.2
–0.3
–0.4
–0.5 Harry Potter Twilight
Passage Passage
Fig. 1. Score on the implicit measure of narrative collective assimilation (the
identity Implicit Association Test, or IAT) as a function of book read and level
of collective self-construal (CSC). Positive values indicate identification with
vampires, whereas negative values indicate identification with wizards. “High”
CSC was defined as 1 standard deviation above the mean, and “low” CSC was
defined as 1 standard deviation below the mean.
Gabriel, Young
salient in-group, β = 0.38, t(128) = 3.67, p < .001, semipartial
r2 = .07, participants high in collective self-construal (1 SD
above the mean) identified even more strongly with that group,
β = 0.72, t(128) = 6.83, p < .001, semipartial r2 = .25. In other
words, the more participants tended to fulfill their social needs
through collective identification, the more narrative collective
assimilation they exhibited.3
Next, we examined the explicit measure, scores on the
Twilight/Harry Potter Narrative Collective-Assimilation Scale.
The overall regression was significant, F(3, 132) = 3.43, p = .02.
The narrative collective-assimilation hypothesis was again
confirmed: Participants who read the Harry Potter chapters
self-identified as wizards (M = −0.14), whereas participants
who read the Twilight chapter self-identified as vampires (M =
0.15), t(132) = 2.17, p = .03, semipartial r2 = .07. The hypothesis
that collective assimilation would be moderated by collective
self-construal was again supported, β = 0.27, t(132) = 2.17,
p = .002, semipartial r2 = .03 (see Fig. 2). Participants who
were high in collective self-construal exhibited narrative
collective assimilation, β = 0.033, t(132) = 2.75, p = .007,
semipartial r2 = .05, whereas those low in collective self-construal
did not, p = .75.4
To test the third hypothesis, that narrative collective assimi-
lation leads to the same effects as satiating the need to belong
(i.e., increased life satisfaction and improved mood), we com-
puted overall narrative collective assimilation by reversing the
IAT scores for participants in the Harry Potter condition. Thus,
for people in both conditions, higher, positive collective-
assimilation scores meant greater collective assimilation. As
expected, greater narrative collective assimilation predicted
both increased life satisfaction (β = 0.215, p = .012) and
increased mood (β = 0.18, p = .04). Increased life satisfaction
also predicted increased mood (β = 0.275, p = .001). To test for
Narrative Collective-Assimilation Score
0.3
Association→
Vampire-Self
High CSC Low CSC
0.2
0.1
0
–0.1
–0.2
←Wizard-Self
Association
–0.3
–0.4
–0.5
–0.6
–0.7 Harry Potter Twilight
Passage Passage
Fig. 2. Score on the explicit measure of narrative collective assimilation (the
Twilight/Harry Potter Narrative Collective-Assimilation Scale) as a function of
book read and level of collective self-construal (CSC). Higher values indicate
identification with vampires, whereas lower values indicate identification with
wizards. “High” CSC was defined as 1 standard deviation above the mean, and
“low” CSC was defined as 1 standard deviation below the mean.
Narrative Collective Assimilation
mediation, we regressed positive mood on overall narrative
collective assimilation and life satisfaction simultaneously.
Increased life satisfaction still predicted mood (β = 0.275, p =
.001), but the relationship between narrative collective assimi-
lation and mood was no longer significant (β = 0.121, p = .153;
see Fig. 3). A Sobel test confirmed that the mediation was sig-
nificant (z = 1.95, p = .05).5 In other words, narrative collec-
tive assimilation put participants in a good mood because it
made them feel better about their lives.
Discussion
The proposed narrative collective-assimilation hypothesis was
supported by results for both explicit and implicit measures;
participants who read the Harry Potter chapters associated
themselves with wizards, whereas those who read the Twilight
chapter associated themselves with vampires. Furthermore,
for both measures, the effects were moderated by the degree to
which participants tended to fulfill their belongingness needs
through collectives. This finding supports our argument that
narrative collective assimilation is related to the desire to
belong to groups. Also supportive of the link between narra-
tive collective assimilation and belongingness needs is our
finding that greater narrative collective assimilation predicted
increased life satisfaction and positive affect, two common
outcomes of a satiated need to belong.
This research complements and extends previous research
on narratives and fulfillment of social needs (e.g., Mar & Oat-
ley, 2008). For example, although previous research found that
narratives alleviate loneliness (Derrick et al., 2009), no spe-
cific mechanism for that action was identified. The current
research suggests that narratives alleviate loneliness by pro-
viding a collective identity that is easily assumed and psycho-
logically rewarding.
This research also increases understanding of how social sur-
rogates, or symbolic means of fulfilling relationships, function.
Previous research demonstrated that social surrogates can
provide symbolic relationship partners (i.e., affiliation with
Narrative 0.18* (0.12)
Positive
Collective
Mood
Assimilation
0.215** 0.275** (0.267**)
Increased
Life
Satisfaction
Fig. 3. Results of the analysis of increased life satisfaction as a mediator
of the relationship between narrative collective assimilation and positive
mood. The figure shows standardized regression coefficients; values outside
parentheses were obtained when each predictor was entered into the model
separately, and values inside parentheses were obtained when the predictors
were entered simultaneously. Asterisks indicate significant coefficients (*p
.05; **p .01).
993
celebrities; Derrick, Gabriel, & Tippin, 2008) and remind one of
existing relationships (i.e., foods can have cognitive associa-
tions with primary relationship partners; Troisi & Gabriel, 2011).
The current research suggests another mechanism for social sur-
rogacy: affiliation with symbolic groups.
Finally, the current research increases what is known about
the ability of collective connections to fulfill belongingness
needs. Although connections to groups have long been identi-
fied as a means of social connection, most research on the
need to belong has tended to focus on relational bonds (Bau-
meister & Leary, 1995). This is surprising given the preva-
lence of group identifications in society (e.g., team fandom,
family groups, ethnic-group affiliation). The present research
adds to what is known about the utility and availability of col-
lective identities by demonstrating the ease and consequences
of collective assimilation.
The pleasure of immersing oneself in narratives is not sur-
prising or novel to anyone who has ever been lucky enough to
get lost in a good book. However, the current research sug-
gests that books give readers more than an opportunity to tune
out and submerge themselves in fantasy worlds. Books pro-
vide the opportunity for social connection and the blissful
calm that comes from becoming a part of something larger
than oneself for a precious, fleeting moment.
Acknowledgments
The authors would like to thank Amanda Arnst, Jessica Egles, Alyssa
Geisler, Kelly Law, Michael Neste, Nicole Wilcocks, and Rachel
Wollenberg for their help with data collection.
Declaration of Conflicting Interests
The authors declared that they had no conflicts of interest with
respect to their authorship or the publication of this article.
Notes
1. The other component of the need to belong is the need to have
intimate relationships (Brewer & Gardner, 1996).
2. Seven participants indicated suspicion when probed about the
experiment (i.e., they thought the study was examining whether
reading a book would change one’s personality). Analyses exclud-
ing these participants yielded results indistinguishable from those
obtained when these participants were included, so they were not
removed from the sample.
3. Liking of the book, transportation, relational self-construal, and
previous exposure to the book also interacted with book read. How-
ever, when all of the interactions and main effects were entered into
the model concurrently, only the two-way interactions of book read
with collective self-construal and liking of the book remained. The
more participants liked the book, the stronger the collective assim-
ilation. More details on this and other analyses are available upon
request from the first author.
4. No other variables, including relational self-construal, were sig-
nificant moderators of the effect of book read on the explicit measure.
5. The alternate mediation model, with mood as a mediator of
the relationship between narrative collective assimilation and life
994
satisfaction, was not supported. The relationship between narrative
collective assimilation and life satisfaction remained significant
when mood was entered into the model, and the Sobel test was not
significant.
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 Social Cognitive and Affective Neuroscience, 2016, 215–224

doi: 10.1093/scan/nsv114
Advance Access Publication Date: 4 September 2015
Original Article

Reading fiction and reading minds: the role of

simulation in the default network
Diana I. Tamir,1 Andrew B. Bricker,2 David Dodell-Feder,3 and
Jason P. Mitchell3
1
Department of Psychology, Princeton University, Princeton, NJ, USA, 2Department of English, McGill
University, Montreal, Quebec, Canada, and 3Department of Psychology, Harvard University, Cambridge, MA,
USA
Correspondence should be addressed to Diana I. Tamir, Peretsman-Scully Hall, Washington Street, Princeton, NJ 08544, USA.
E-mail: dtamir@princeton.edu

Abstract
Research in psychology has suggested that reading fiction can improve individuals’ social-cognitive abilities. Findings from
neuroscience show that reading and social cognition both recruit the default network, a network which is known to support
our capacity to simulate hypothetical scenes, spaces and mental states. The current research tests the hypothesis that
fiction reading enhances social cognition because it serves to exercise the default subnetwork involved in theory of mind.
While undergoing functional neuroimaging, participants read literary passages that differed along two dimensions: (i) vivid
vs abstract and (ii) social vs non-social. Analyses revealed distinct subnetworks of the default network respond to the two
dimensions of interest: the medial temporal lobe subnetwork responded preferentially to vivid passages, with or without
social content; the dorsomedial prefrontal cortex (dmPFC) subnetwork responded preferentially to passages with social and
abstract content. Analyses also demonstrated that participants who read fiction most often also showed the strongest social
cognition performance. Finally, mediation analysis showed that activity in the dmPFC subnetwork in response to the social
content mediated this relation, suggesting that the simulation of social content in fiction plays a role in fiction’s ability to
enhance readers’ social cognition.

Key words: fiction; reading; theory of mind; default network; simulation; functional MRI

Introduction who demonstrate greater civic engagement, including higher

Readers of fiction can transcend the here-and-now to experi-
ence worlds, people and mental states that differ vastly from
their local reality. The consequences of reading, however, ex-
tend far beyond the subjective experience of any one individual.
Researchers from fields as diverse as evolutionary psychology,
literary studies and anthropology have independently credited
literacy as a possible explanation for such fundamental societal
shifts as the decline in human violence over the past few centu-
ries, the development of desire-based over rule-based social
interactions, and the advent of ‘modern subjectivity’ (Lukacs,
1920; Watt, 1957; Ong, 1982; McKeon, 1987; Habermas, 1991;
Pinker, 2011). Such large-scale societal impacts may neverthe-
less begin with small behavioral changes in individual readers,
levels of volunteering, donating and voting, than non-readers
(Katz, 2006). How might reading effect its influence on these
individuals?
Recent research in psychology suggests that readers make
good citizens because reading may improve one’s ability to em-
pathize with and understand the thoughts and feelings of other
people. Readers of fiction score higher on measures of empathy
and theory of mind (ToM)—the ability to think about others’
thoughts and feelings—than non-readers, even after controlling
for age, gender, intelligence and personality factors (Mar et al.,
2006, 2009, 2010). Developmental work has likewise shown a
correlation between reading and social cognition. Children be-
tween the ages of four and six who were exposed to more

Received: 25 May 2015; Revised: 1 September 2015; Accepted: 1 September 2015

C The Author (2015). Published by Oxford University Press. For Permissions, please email: journals.permissions@oup.com
V

215
 216 | Social Cognitive and Affective Neuroscience, 2016, Vol. 11, No. 2
juvenile fiction performed better at ToM tasks than children
with less exposure, again controlling for such potentially con-
founding factors as age, gender, vocabulary and parental in-
come (Mar et al., 2010). Other developmental work has similarly
demonstrated that the frequency of parent–child picture book
reading and parents’ use of mental state terms predict false-
belief task performance (Adrian et al., 2005), and that the use of
stories that contain more emotional, social and psychologically
convincing content predicts empathy and socioemotional ad-
justment (Aram and Aviram, 2009). Recent experimental re-
search has further shown that fiction reading plays a causal
rather than just correlational role in the development of social-
cognitive skills, such that among adults, fiction reading en-
hances ToM performance (Kidd and Castano, 2013) and em-
pathy (Bal and Veltkamp, 2013).
However, not all reading improves social cognition. One
study found that after controlling for demographic factors, per-
sonality traits and exposure to non-fiction and other fiction
genres, only exposure to Romance significantly predicted ToM
performance (Fong et al., 2013). In another series of studies,
though high-quality ‘literary’ fiction consistently improved so-
cial cognition, lower-quality fiction and non-fiction did not
(Kidd and Castano, 2013). Indeed, people who regularly read
non-fiction do not have better social abilities and may have
interesting
worse social abilities than more infrequent readers of non-
fiction (Mar et al., 2006, 2009). However, at least one study found
that people randomly assigned to read either literary fiction or
literary non-fiction did not differ in empathy change pre- to
post-reading; only when taking participants’ openness into ac-
count did the expected difference between fiction and non-
fiction emerge (Djikic et al., 2013).
Developmental research further suggests that quality and
genre may not be the only features that moderate reading’s abil-
ity to improve social cognition; content, and the kinds of cogni-
tive demands that a piece makes on readers, may also play an
world important role. In one experiment, children who read books
that required them to construct their own social interpretations
building performed better on social-cognition tasks than children
exposed to stories that explicitly provided such metacognitive
language (Peskin and Astington, 2004). In a similar vein, adults
assigned to read fiction over a 1-week period demonstrated
positive changes in empathy only when they reported high
emotional transportation into the story (Bal and Veltkamp,
2013), suggesting that immersion into and simulation of the
mental and emotional lives of the characters may be the mech-
anism of change.
Taken together, these findings suggest that the effectiveness
with which literature improves social cognition may depend on
how well it demands attention to others’ mental states. That is,
perhaps literary fiction improves social cognition to the extent
that it requires readers to mentally construct social contexts.
Such high-quality practice in simulation—or the capacity to ex-
perience realities outside of the ‘here-and-now’, including
hypothetical events, distant worlds, and other people’s subject-
ive experience—then translates into real-world consequences
for readers’ social cognition (Zunshine, 2006).
Because we now know a great deal about the neural networks
involved in such simulation processes, work in neuroimaging
presents a unique way to test this prediction. In particular, our
brain’s default network, which comprises the medial prefrontal
cortex (mPFC), posterior cingulate cortex (PCC), posterior superior
temporal sulcus (pSTS), temporal parietal junction (TPJ), anterior
medial temporal gyrus and medial temporal lobes (MTLs), is re-
sponsible for supporting our capacity for simulation (Raichle
et al., 2001; Buckner and Carroll, 2007; Schacter and Addis, 2007;
Spreng et al., 2009). The default network is recruited whenever
people conjure up experiences outside of their local experiences,
such as thinking about the future or the past, mentally construct-
ing places and spaces, imagining hypothetical events and think-
ing about another’s perspective (Okuda et al., 2003; Addis et al.,
2007; Buckner and Carroll, 2007; Hassabis et al., 2007; Schacter
et al., 2007; Szpunar et al., 2007; Botzung et al., 2008; Hassabis and
Maguire, 2009; Tamir and Mitchell, 2011). The entirety of the de-
fault network has been associated with simulation in general but
research has also demonstrated that two distinct subnetworks of
the default network are recruited differentially when simulating
vivid spatial content and mental content, respectively (Andrews-
Hanna et al., 2010). More specifically, scene construction and the
simulation of vivid physical spaces rely on neural structures
within more ventral aspects of the default network, such as the
ventromedial prefrontal cortex (vmPFC), hippocampal and para-
hippocampal gyri and retrosplenial cortex (Hassabis et al., 2007;
Hassabis and Maguire, 2009), structures that comprise the default
network’s MTL subnetwork. Conversely, studies of human social
cognition suggest that thinking about people and mental states
recruits a separate set of regions, including the dorsomedial pre-
frontal cortex (dmPFC), anterior temporal poles and TPJ (Mitchell
et al., 2002; Saxe and Wexler, 2005; Mitchell, 2008), structures that
comprise the default mode’s dmPFC subnetwork. Researchers
have noted the overlap between the dmPFC subnetwork and the
network of brain regions associated with ToM, suggesting that
ToM may rely on simulation processes (Andrews-Hanna et al., processing
2010; Spreng and Grady, 2010; Mars et al., 2012).
Fiction reading, which engenders simulations of vivid and
blu
social content, also recruits the default network (Mar, 2004,
interacti
2011). Thus, the overlap between reading and simulation is per-
haps unsurprising, given that narratives often invoke vivid de-
scriptive language to transport readers to far-off places, and
engage readers with characters’ actions, interactions and men-
tal states. Both the simulation of physical spaces and of mental
entities provides a plausible explanation for why reading reli-
ably activates the default network (Mar, 2004; Yarkoni et al.,
2008, 2011; Mason and Just, 2009; Speer et al., 2009). However,
the relation between default network activity and the simula-
tion of scenes and minds, respectively, during fiction reading
has yet to be explicitly tested. This study first empirically tests
the hypothesis that reading recruits the default network be-
cause it evokes both of these types of simulation.
In addition, we capitalize on this hypothesized overlap be-
tween simulation and fiction reading to test the hypothesis
that, by recruiting the default network while reading, readers
may practice the types of simulation necessary for solving so- reading
cial tasks. However, not all types of simulation should play a is plastic
commensurate role in the relation between reading and for real
enhanced social cognition. As suggested by previous research world
(Peskin and Astington, 2004), only simulations of social content scenarios
should provide relevant social practice, whereas simulations of
non-social scenes, events or hypothetical scenarios should fail
to provide relevant social practice. This proposed dynamic rela-
tionship between neural function and experience is supported
by neuroplasticity literature, which has demonstrated that re-
peated engagement in cognitive processes can lead to positive
changes in the neural networks supporting those cognitive
processes (Draganski and May, 2008; Klingberg, 2010; Anguera
et al., 2013; Lovden et al., 2013; Merzenich et al., 2014). Thus, here,
repeated engagement in social simulation vis-à-vis fiction read-
ing may lead to beneficial changes in the default network,
which may carry concomitant benefits for social ability. Said

otherwise, fiction reading may impact social ability through its
effect on the neural system supporting social simulation. We
note the possibility that individuals who have greater ToM abil-
ity may read more fiction, in which case greater ToM ability
may predispose individuals to engage in more simulation,
which may make reading fiction more enjoyable. However, cur-
rent empirical work suggests a causal effect of fiction reading
on ToM (Peskin and Astington, 2004; Mar et al., 2010; Bal and
Veltkamp, 2013; Kidd and Castano, 2013), rather than the re-
verse. Thus, taken together, we further hypothesize that neural
activity while simulating social content, but not vivid physical
scenes, should mediate the relation between fiction reading
and ToM.
To test these hypotheses, participants in this study under-
went functional neuroimaging scanning while they read literary
excerpts designed to engender both simulations of vivid phys-
ical scenes and simulations of social content. Outside of the
scanner, participants provided measures of their reading behav-
ior and ToM ability. We expect that reading, which engenders
simulation, should preferentially recruit the default network.
More specifically, simulations of vivid scenes should evoke ac-
tivity in the MTL subnetwork of the default network whereas
simulations of social content should evoke activity in the
dmPFC subnetwork of the default network. Further, we expect
that the extent of fiction reading should predict ToM perform-
ance, replicating previous research on the relation between
reading and social cognition. Finally, to the extent that simula-
tion of social content provides the practice necessary for im-
provements in social cognition, we expect that neural activity
specific to social simulations should mediate this relation be-
tween fiction reading and ToM.
Method
Participants
Twenty-six (16 female) right-handed, native English speakers
with no history of neurological problems participated in this
study (M age ¼ 21.2 years; range ¼ 19–26 years). All participants
provided consent in a manner approved by the Committee on
the Use of Human Subjects at Harvard University.
fMRI reading task
While undergoing fMRI scanning, participants read a series
of literary passages, excerpted from a wide variety of sources,
including novels, biographies, magazines, newspapers and self-
help books (Supplementary Materials A). Each passage came
from a unique source. For each trial, participants were pre-
sented with one passage to read (M length ¼ 85 words;
range ¼ 56–106). Instructions emphasized that participants
should pay full attention as they read each passage and that
they did not need to finish reading the passage within the time
allotted. Passages remained on screen for up to 30 s.
Participants pressed a button under their index finger if they
finished the passage before 30 s after which the passage was
replaced by a fixation cross for the remainder of the 30 s period.
Four seconds of fixation followed each 30 s reading period.
Passages varied systematically along two orthogonal dimen-
sions: (i) the vividness with which they described physical
scenes (Vivid vs Abstract) and (ii) whether or not they described
a person or a person’s mental content (Social vs Non-social).
Passages were selected and categorized based on the pretest
ratings of a separate set of participants across a variety of
D. I. Tamir et al. | 217
features: Social passages were selected to be highly social and
personal, and contain either one person or groups of people;
Vivid passages were selected to be high on vividness and move-
ment and low on abstractness. There were a total of four pas-
sage types. Vivid/Social passages describe vivid scenes or
events that include references to mental states, individuals or
groups of people. Vivid/Non-social passages describe vivid
physical scenes or events but lack references to mental states,
individuals or groups of people. Abstract/Social passages use
abstract language and thus lack easily imagined physical scenes
but include references to mental states, individuals or groups of
people. Finally, Abstract/Non-social passages use abstract lan-
guage and lack imaginable physical scenes, people and mental
states. Importantly, the four types of passages did not differ in
their pretest ratings of boringness or wordiness or in average
reading time (Supplementary Materials B).
During scanning, participants read 13 passages of each type,
for a total of 52 passages. In addition, 13 fixation periods (each
lasting 30 s) were included. The 52 passages and fixation periods
were presented in a random order for each participant, divided
among five consecutive runs of 442 s each.
fMRI data acquisition and analysis
Functional data were acquired using a gradient-echo echo-pla-
nar pulse sequence (TR ¼ 2 s; TE ¼ 30 ms) on a 3T Siemens Trio.
Images were acquired using 36 axial, interleaved slices with a
thickness of 3 mm (0.5 mm skip) and 3 " 3 in-plane resolution
and online motion correction. Functional images were prepro-
cessed and analyzed using SPM8 (Wellcome Department of
Cognitive Neurology, London, UK; http://www.fil.ion.ucl.ac.uk/
spm/). Data were first spatially realigned to correct for head
movement and then unwarped to reduce image distortions.
Images were then normalized to a standard anatomical space
(3 mm isotropic voxels) based on the ICBM 152 brain template
(Montreal Neurological Institute). Normalized images were then
spatially smoothed using an 8 mm FWHM Gaussian kernel.
Preprocessed images were analyzed using a general linear
model in which the events were modeled using a canonical
hemodynamic response function and covariates of no interest
(session mean, linear trends and six motion realignment par-
ameters). Events began at the onset of the presentation of the
passage and lasted for a duration of either 30 s, or until the par-
ticipant indicated that he or she had finished reading the pas-
sage by pressing a button. Trials were conditionalized based on
the type of passage presented, resulting in four conditions of
interest: Vivid/Social, Vivid/Non-social, Abstract/Social and
Abstract/Non-social. To test whether reading recruited the de-
fault network, primary analyses identified voxels in which
BOLD response differed along the two dimensions of interest;
that is, vividness (Vivid/Social þ Vivid/Non-social) > (Abstract/
Social þ Abstract/Non-social) and sociality (Vivid/
Social þ Abstract/Social) > (Vivid/Non—social). Analyses were
performed individually for each participant, and contrast
images generated within each participant were subsequently
entered into a second-level analysis treating participants as a
random effect. Group level whole-brain contrasts employed an
experiment-wise threshold of P < 0.05 corrected for multiple
comparisons per Slotnick and Schacter’s (2004) specifications;
Monte Carlo simulations indicated use of a statistical criterion
of 54 or more contiguous voxels at a voxel-wise threshold of
P < 0.01.
To test the hypothesis that the MTL subnetwork would pref-
erentially respond to the vividness of passages and the dmPFC
 218 | Social Cognitive and Affective Neuroscience, 2016, Vol. 11, No. 2
subnetwork to the socialness of passages, we assessed neural
responses to the reading task in independently defined regions-
of-interest (ROIs). ROIs were defined as 8 mm spheres centered
on the coordinates for each of the 11 regions independently
identified by Andrews-Hanna (2010). Using functional connect-
ivity analyses, Andrews-Hanna et al. (2010) identified 11 default
network regions, divided into three functionally and anatomic-
ally distinct subnetworks: (i) a MTL subnetwork that comprises
the hippocampal formation, parahippocampal cortex, retrosple-
nial cortex, posterior intraparietal lobe and vmPFC; (ii) a dmPFC
subnetwork that comprises the dmPFC, temporal pole, lateral
temporal cortex and temporal-parietal junction; and (iii) a ‘core’
subnetwork that comprises the PCC and mPFC (Andrews-Hanna
et al., 2010). Parameter estimates for the Social > Non-social and
Vivid > Abstract contrasts were extracted from these ROIs to
examine how each subnetwork responds to simulating physical
and mental events during reading. Activity within each subnet-
work was then calculated as the average parameter among the
regions composing that network. Follow-up analyses evaluated
individual ROI response within each network. Four outliers
(>2.5 s.d. of the mean) were identified and Winsorized by
replacing them with the next highest non-outlying value and
adding 10% of that value to maintain variance. Of note, the dir-
ection and significance of the findings did not change when
using the Winsorized values.
Behavioral measures
In addition to the reading task, participants also completed be-
havioral surveys outside of the scanner. We measured partici-
pants’ exposure to both fiction and non-fiction and their social-
cognitive abilities to assess whether participants showed the
expected relation between fiction reading and ToM (Mar et al.,
2006, 2009).
Fiction reading. Participants completed the author recognition
test (ART) to assess the extent to which they read fiction and
non-fiction in their daily lives. This measure was originally de-
veloped by Stanovich and West (1989) but participants in this
study saw the most recently updated and validated version of
the ART developed by Mar et al. (2006). For this test, participants
were presented with the names of fiction authors (50 names),
non-fiction authors (50 names) and 40 foils, and were asked to
place checkmarks next to the names that they recognized as au-
thors. Participants needed only to recognize a name as that of
an author but were not required to have read any of the author’s
work. Participants were told that some of the names were of
people who are not writers. In this way the ART discourages
guessing and overcomes potential issues of self-report bias.
Following Stanovich and West (1989), fiction and non-fiction
ART scores were calculated separately as the number of fiction
or non-fiction author names a participant recognized, respect-
ively, minus the number of foils they reported recognizing.
ToM. Participants also completed a ToM task that assessed the
extent to which they spontaneously think about intentions
when judging an individual’s behavior. For this task, partici-
pants read 48 vignettes in which an actor engages in a behavior
with either a negative or neutral outcome, on the basis of either
a negative or a neutral intention (Young et al., 2010a). The nega-
tive and neutral intentions were fully counterbalanced with the
negative and neutral outcomes across 48 stimuli, resulting in 12
vignettes of four types: (i) no harm—neutral intention/neutral
outcome, (ii) intentional harm—negative intention/negative
outcome, (iii) accidental harm—neutral intention/negative out-
come and (iv) attempted harm—negative intention/neutral out-
come. After reading each vignette, participants judged the
permissibility of the actor’s behavior on a scale from 1 (forbid-
den) to 5 (permissible). Participants read and answered all moral
judgment questions at their own pace. Given that the intention
differs from the outcome in the accidental and attempted harm
scenarios, judgments of moral permissibility reflect the extent
to which participants take into account the actor’s intention as
opposed to the outcome. Thus, to the extent that participants
consider the actor’s intention, participants should judge the ac-
tion in accidental harm scenarios as more permissible and judge
the action in attempted harm scenarios as less permissible (see
Supplementary Materials C for more information). Using this
task, previous researchers have shown that responses to these
two scenario types provide a sensitive measure of ToM (Young
et al., 2010a,b; Moran et al., 2011).
fMRI reading task memory assessment. Finally, participants com-
pleted a surprise memory assessment for the stimuli presented
during the reading task. This task measured the extent to which
they had remained attentive during the scanning session based
on their ability to recognize a series of sentences that either had
been presented during scanning (‘old’) or had not been seen
(‘new’). Performance was assessed by calculating d-prime for
each participant (Supplementary Materials D).
Brain-behavior analysis
One outlier was identified in the moral judgment data and was
Winsorized. We evaluated the relationship between the behav-
ioral measures with bivariate Pearson correlations. These val-
ues are accompanied by bias-corrected and accelerated 95% CIs
generated from 5000 bootstrap samples in SPSS. To evaluate the
hypothesis that fiction reading impacts ToM ability, in part,
through its effect on the neural bases of social simulation, we
tested a mediation model with fiction reading (fiction ART
score) as the predictor variable, neural activity for social simula-
tion as the mediator and as an index of ToM ability, perform-
ance on the moral judgment task as the outcome variable. We
used a non-parametric bootstrapping procedure to estimate the
indirect effect; that is, the path from the predictor to the out-
come variable through the mediator (fiction reading ! neural
basis of social simulation ! ToM ability). Estimates of the indir-
ect effect are accompanied by bias-corrected and accelerated
95% CIs derived from 5000 bootstrap samples. As measures of
effect size, we provide the proportion of variance accounted for
by the mediated effect (R2med) and, as recommended by
Preacher and Kelley (2011), j2, which represents the ratio of in-
direct effect observed relative to the maximum possible indirect
effect. This analysis was implemented in SPSS with the
PROCESS macro (Hayes, 2013).
Results
fMRI results
Our primary analyses identified brain regions that responded to
the two features of interest: (i) the vividness with which pas-
sages described physical scenes and events and (ii) whether or
not they described a person or a person’s mental content.
Consistent with earlier research, both vivid passages and social
passages recruited regions of the default network significantly
more than abstract and non-social passages. A whole-brain

random-effects contrast of Social>Non-social passages revealed
activity in dmPFC, vmPFC, lateral temporal cortex from the
temporal pole to the TPJ bilaterally, bilateral hippocampi and bi-
lateral IFG (Figure 1A; Table 1). A whole-brain random-effects
contrasts of Vivid>Abstract passages revealed robust activity
in MTL structures, including hippocampus and parahippocam-
pus bilaterally, retrosplenial cortex and precuneus (Figure 1B;
Table 1).
To test the hypothesis that subnetworks of the default net-
work respond differentially to each of these two features of the
passages, we assessed neural responses to the four passage
types within the three subnetworks identified by Andrews-
Hanna et al. (2010) (Figure 1C). Consistent with the whole-brain
analysis demonstrating that the subnetworks differentially re-
spond to the social content and vividness of the passages, a 3
Subnetwork (Core, MTL, dmPFC) " 2 Vividness (Vivid,
Abstract) " 2 Sociality (Social, Non-Social) repeated-measures
ANOVA revealed an interaction between Subnetwork and
Sociality, F(2, 50) ¼ 3.51, P ¼ 0.04, partial g2 ¼ 0.12, Subnetwork
and Vividness, F(2, 50) ¼ 19.64, P < 0.001, partial g2 ¼ 0.44, and a
three-way interaction between Subnetwork, Sociality and
Vividness, F(2, 50) ¼ 4.96, P ¼ 0.01, partial g2 ¼ 0.17.
Follow-up 2 " 2 repeated-measures ANOVA within subnet-
works revealed that the vividness of the passages significantly
affected activity in the MTL subnetwork, F(1, 25) ¼ 4.38, P < 0.05,
Cohen’s d ¼ 0.42, but the presence of people or mental content
in the passages had no effect on the MTL subnetwork, F(1,
25) ¼ 0.21, P ¼ 0.65, d ¼ 0.09. This suggests that the MTL network
responded most robustly to passages that vividly described
scenes but did not differentiate between passages with or
without people. No interaction effects between the social and
vivid factors were observed, F(1, 25) ¼ 2.66, P ¼ 0.12, partial
g2 ¼ 0.10.
In contrast, a 2 " 2 repeated-measures ANOVA over activity
in the dmPFC subnetwork revealed two main effects. First, the
presence of people in the passages significantly affected activity
in the dmPFC subnetwork, F(1, 25) ¼ 8.21, P < 0.01, d ¼ 0.57, such
that this subnetwork responded more robustly to the presence
of people and mental states in literary passages. Unexpectedly,
we also observed a second main effect: abstract passages eli-
cited more activity in the dmPFC subnetwork than vivid pas-
sages, F(1, 25) ¼ 28.72, P ¼ 0.001, d ¼ 1.07, suggesting that the
dmPFC subnetwork responds robustly to abstract content. No
Fig 1. BOLD differences for main effect of (A) Social > Non-social (B) Vivid > Abstract, (C) and results of ROI analysis of both contrasts. Both whole-brain and ROI analyses
show that the dmPFC subnetwork of the default network responded most robustly to literary passages containing people or mental content, whereas the MTL subnet-
work responded most robustly to vivid physical descriptions.
D. I. Tamir et al. | 219
interaction effects between the social and vivid factors were
observed, F(1, 25) ¼ 0.01, P ¼ 0.92, partial g2 ¼ 0.00.
The core subnetwork did not show differential responses to
Social vs Non-social passages, F(1, 25) ¼ 0.48, P ¼ 0.49, d ¼ 0.14 or
Vivid vs Abstract passages, F(1, 25) ¼ 1.50, P ¼ 0.23, d ¼ $0.24. No
interaction effects between the social and vivid factors were
observed, F(1, 25) ¼ 0.18, P ¼ 0.67, partial g2 ¼ 0.01. Responses in
individual regions within the network are presented in
Supplementary Materials E.
Thus, both the whole-brain and ROI analyses suggest that
the default network does indeed respond differentially to liter-
ary passages depending on their content. Different subnetworks
of the default network distinguished between the vividness of a
passage and the social content of a passage. The MTL network
responded most robustly to passages designed to be easy to
simulate because they are rich in vivid details, whereas the
dmPFC subnetwork responds most robustly to passages de-
signed to be easy to simulate because they contain references to
people or mental states.
Behavioral results
Performance on the scanner task and behavioral measures
are presented in Supplementary Materials D and F, respect-
ively. We found that people who read more fiction were more
likely to take intentions into account when judging attempted
harm scenarios (i.e. negative intention/neutral outcomes).
Specifically, participants’ fiction ART scores were
significantly correlated with their ratings on the moral judg-
ment task, r(24) ¼ $0.44, P ¼ 0.02, 95% CI [$0.66, $0.12] (Figure
2C), such that greater fiction reading was associated with
judging actions as less permissible on attempted harm
scenarios.
This positive association between reading and ToM was
specific to fiction reading. Non-fiction ART scores among par-
ticipants did not correlate with moral judgments of failed
harm, r(24) ¼ $0.15, P ¼ 0.48, 95% CI [$0.48, 0.22], even though
the extent to which participants read fiction and non-fiction
was highly correlated, r(24) ¼ 0.80, P < 0.001, 95% CI [0.64, 0.91].
Such findings replicate numerous previous studies that dem-
onstrate that exposure to fiction, but not non-fiction, predicts
enhanced ToM (Mar et al., 2006, 2009, 2010; Kidd and Castano,
2013).
 220 | Social Cognitive and Affective Neuroscience, 2016, Vol. 11, No. 2

Table 1. Peak voxel and cluster size for all regions obtained from a Table 1. (continued)
contrast of Social > Non-social and Vivid > Abstract (cluster-level cor- Anatomic label x y z Volume Max t
rected P < 0.05)
Cerebellum 30 $83 $26 2659 6.12
Anatomic label x y z Volume Max t
$42 $73 $22 167 3.99
Middle frontal gyrus $46 11 52 1294 5.78
40 59 $4 814 4.22
Social > non-social
Inferior parietal lobule 44 $57 40 1132 4.69
Anterior temporal pole 34 31 $42 5362 6.99
Occipital cortex 46 $83 $4 272 4.43
$44 9 $36 6189 6.96
$4 $91 $20 71 3.51
Primary motor cortex $42 $15 72 1159 6.94
36 $83 18 78 3.40
54 1 58 591 5.74
PCC $2 $43 32 191 3.90
22 $5 84 202 4.49
Inferior frontal gyrus 60 27 6 502 5.21
46 29 $10 88 3.39
Brain-behavior results
Cerebellum 26 $83 $34 586 5.01
$26 $79 $38 146 4.21 Based on the extant literature, we hypothesized that the neural
0 $51 $36 77 3.12 basis of social simulation would explain the link between fiction
Dorsomedial prefrontal cortex $10 59 46 1741 4.75 reading and ToM ability. That is, fiction reading improves ToM,
Ventromedial prefrontal cortex 6 49 $14 302 4.38 in part, through its effect on the neural basis of social simula-
Occipital cortex 16 $107 24 55 3.98 tion. Because reading mental and vivid physical passages differ-
$8 $81 10 139 3.57 entially recruited distinct subnetworks of the default network,
$50 $97 6 55 3.43 we were able to address this question using dmPFC subnetwork
16 $79 14 61 2.86 activity to social passages as an index of social simulation. If fic-
Non-social > social tion reading enhances ToM because doing so activates or trains
Inferior temporal gyrus $58 $55 $12 2554 7.45 the neural networks involved in ToM (i.e. the dmPFC subnet-
Middle frontal gyrus 48 37 24 16836 7.02 work), then we would expect the dmPFC subnetwork response
$30 35 $12 6.85
to the mental simulations to mediate the relation between fic-
Inferior parietal lobule $50 $49 52 12634 6.78
tion reading and ToM.
52 $41 50 6.10
These questions were addressed with mediation analysis.
Middle temporal gyrus 60 $47 $10 1028 6.08
The paths between the predictor (fiction ART), mediator (dmPFC
Superior temporal gyrus 52 1 $6 1186 4.68
subnetwork activity for Social > Non-social) and outcome vari-
$52 $7 $2 142 4.29
Parahippocampal gyrus 24 $37 $4 113 4.40
able (moral judgments on the attempted harm scenarios) were
$32 $43 $10 589 3.85 significant in the predicted directions (Figure 2): fiction reading
34 $21 $32 60 3.26 was positively associated with considering actors’ intention
Cerebellum 56 $67 $42 348 4.14 when making moral judgments, and with dmPFC subnetwork
Insula $36 13 4 451 3.98 activity; dmPFC subnetwork activity was positively associated
Occipital cortex 44 $75 4 98 3.48 with considering actors’ intention when making moral judg-
Vivid > abstract ments. Importantly, the direct effect of fiction reading on moral
Parahippocampal gyrus 18 $13 $20 1038 6.82 judgments was no longer significant when controlling for
Inferior parietal lobule $66 $35 38 1098 6.71 dmPFC subnetwork activity. Bootstrap analysis of the indirect
62 $39 44 58 3.01 effect (coefficient ¼ $0.02, SE ¼ 0.01) generated a CI that did not
Fusiform gyrus $34 $35 $20 1556 6.63 encompass zero, 95% CI [$0.05, $0.001], indicating that dmPFC
Angular gyrus $36 $87 36 809 6.37 subnetwork activity mediated the relationship between fiction
44 $75 32 640 4.77 reading and moral judgments (R2med ¼ 0.16, 95% CI [0.02, 0.37];
Middle temporal gyrus $52 $63 $2 679 5.54 j2 ¼ 0.21, 95% CI [0.03, 0.46]).
PCC 10 $53 14 1743 4.94 Since the dmPFC subsystem also responded preferentially to
Middle frontal gyrus $38 35 18 931 4.88
abstract vs vivid passages, one possibility is that simulation of
48 51 24 66 3.21
abstract and non-social features of fiction in this subsystem
Inferior frontal gyrus 48 31 6 326 3.98
contributed to the mediated effect. We evaluated this possibility
22 27 $12 162 3.38
by running an additional mediation model controlling for the
Precuneus $8 $35 46 378 3.60
dmPFC subnetwork’s response to Abstract > Vivid passages. The
STS $40 $1 $20 87 3.29
Superior frontal gyrus 22 13 48 76 3.01
findings remain unchanged (Supplementary Materials G).
Abstract > vivid Another possibility is that fiction reading may impact ToM
Superior temporal gyrus $50 11 $24 10531 9.10 through its effect on the neural system selective for non-social
Inferior frontal gyrus $50 27 $12 8.25 simulation of vivid scenes. To evaluate this idea, we tested one
64 19 22 61 3.36 additional model using neural activity in the MTL subnetwork
62 9 38 166 3.26 for Vivid > Abstract as the mediator. Bootstrap analysis of the
Middle temporal gyrus $60 $23 $6 10531 7.96 indirect effect revealed that non-social simulation of vivid
64 $41 $12 2546 6.31 scenes also did not mediate the relation between fiction and
Dorsomedial prefrontal cortex $10 49 46 4539 8.08 ToM ability (Supplementary Materials H).
Ventromedial prefrontal cortex 0 43 $20 1478 6.71
Temporo-parietal junction $52 $63 30 1929 6.70 Discussion
Insula $32 3 8 591 6.54
The link between fiction reading and ToM occurs at multiple
continued levels of analysis. Psychologically, fiction readers possess
 D. I. Tamir et al. | 221

Fig 2. Depiction of significant correlations between (A) fiction reading scores on the fiction ART and dmPFC subnetwork activity during Social > Non-social passages, (B)
between dmPFC subnetwork activity during Social > Non-social passages default activity and ToM task performance and (C) between fiction reading and ToM task per-
formance. (D) The effect of fiction reading on ToM task performance through dmPFC subnetwork activity for social passages. Bootstrap analysis of the indirect effect
indicated that dmPFC subnetwork activity mediated the relationship between fiction reading and ToM task performance. Unstandardized path coefficients shown with
SE in parentheses for each path. The dotted line represents the direct effect of fiction reading on ToM task performance (i.e. controlling for the effect of dmPFC subnet-
work activity). Note that the ‘More Intention’ and ‘Less Intention’ anchors on plots B and C are for visualization purposes only; participants rated each story on the
Moral Judgment Task from 1 (forbidden) to 5 (permissible). *P < 0.05.

stronger social-cognitive abilities than both non-readers and
non-fiction readers (Mar et al., 2006, 2009, 2010). Historically,
highly literate societies, especially societies that produced psy-
chologically rich literature, function more empathically and less
violently than less literate societies (Lukacs, 1920; Watt, 1957;
Ong, 1982; McKeon, 1987; Habermas, 1991; Pinker, 2011). And
neurally, fiction reading and social cognition recruit an overlap-
ping neural network (i.e. the default network) (Mar, 2004, 2011).
This study not only replicates previous findings that fiction
reading both enhances social cognition and recruits the default
network but also draws together these findings to test two
hypotheses about the nature of this relation between reading
fiction and ToM.
First, this study demonstrates that fiction reading recruits
the default network because it elicits at least two distinct types
of simulation: the simulation of vivid physical scenes and the
simulation of people and minds. Each type of simulation re-
cruited distinct subnetworks of the default network. Consistent
with prior work evaluating non-social vs social scene construc-
tion (Hassabis et al., 2014), simulations of physical scenes pri-
marily recruited the MTL subnetwork of the default network,
while simulations of people and minds primarily recruited the
dmPFC subnetwork of the default network. Interestingly, and
unexpectedly, this study also found that the dmPFC subnetwork
was significantly more responsive to abstract content than vivid
physical content. This finding may be consistent with prior lit-
erature on semantic and conceptual processing. For example,
prior work has found dmPFC and left TPJ to be preferentially
engaged during abstract or high-level construal tasks (e.g. gen-
erating semantic categories) vs low-level construal tasks (e.g.
describing visual characteristics) regardless of the social con-
tent (Baetens et al., 2014), a finding that has been further sub-
stantiated with meta-analytic data (Binder et al., 2009).
Similarly, focusing on the abstract features of personal memo-
ries preferentially recruits dmPFC and left TPJ, whereas focusing
on concrete features of personal memories recruits aspects of
the MTL subsystem (D’Argembeau et al., 2014). Thus, our find-
ings regarding the preferential response of the dmPFC subnet-
work to abstract vs vivid passages converge with other findings
on the role of this network in abstract processing.
Second, this study capitalized on these neural findings to
test the hypothesis that fiction reading improves ToM by pro-
viding readers with the opportunity to exercise or practice men-
tal simulation capacities that are also recruited during social-
cognitive tasks. Said otherwise, fiction reading may impact ToM
through its influence on the neural basis of social simulation.
Mediation analysis was consistent with this idea. Specifically,
we found that dmPFC subnetwork response to simulating
 222 | Social Cognitive and Affective Neuroscience, 2016, Vol. 11, No. 2
people and minds mediated the relation between fiction read-
ing and ToM. This effect was not changed when controlling for
the dmPFC subnetwork’s response to non-social abstract infor-
mation. Furthermore, using MTL subnetwork activity to vivid vs
abstract scenes, we further ruled out the possibility that fiction
reading impacts ToM through its effect on non-social simula-
tion. Together, the results suggest that any positive effect of
reading fiction on social-cognitive abilities might be due to the
influence of reading on neural networks involved in simulating
social content and not non-social vivid scenes. This finding is
consistent with that of behavioral work demonstrating that fic-
tion reading over a 1-week period was associated with an in-
crease in empathy only when readers reported a high level of
transportation (i.e. simulation) of the characters’ mental lives
and story events (Bal and Veltkamp, 2013). We note that re-
search in this field is nascent, and as such alternative models
may be viable. For example, a reverse mediation model whereby
ToM causally impacts the amount of fiction reading through its
impact on social simulation. However, the model tested here, in
which fiction reading impacts ToM through its effect on social
simulation specifically, is most consistent with the extant lit-
erature regarding the nature and direction of the relations be-
tween the variables.
These findings also suggest that future research should
focus on the content of literature to understand the relation be-
tween reading and ToM. Literature that effectively engages a
reader in social content should be most likely to improve ToM;
literature that does not successfully engage a reader in social
thought, or that taxes a reader’s imagination only with hypo-
thetical events and places, should not. Previous researchers
have studied how genre (fiction vs non-fiction) or the quality
(literary vs non-literary) of such works improves ToM (e.g. Fong
et al., 2013; Kidd and Castano, 2013). The current research
manipulated content irrespective of genre. As such, these find-
ings suggest a need to reinterpret previous findings in terms of
content, and the kinds of cognitive demands that content
makes on readers. For example, literary fiction may just more
effectively depict social content than low-quality literature, and
fiction may more often traffic in social content than non-
fiction.
Nevertheless, the fact that the social content of a passage
may play an important role in shaping social cognition raises
important consequences for future research in both psychology
and literary studies alike. For instance, our study does not dis-
cretely define and test every kind of social or mental inter-
action. That is, our ‘social’ passages sometimes contain either
one person or groups of people; they depict either the appear-
ance of an individual or describe a character’s abstract mental
content; or they describe purely social interactions among
groups of individuals. For both the psychologist and the literary
scholar, a closer analysis of the content of the ‘social’ passages
might reveal which aspects of social interaction and mental
content fiction readers respond to most robustly, both neurally
and psychologically. Future investigations into reading and so-
cial cognition would benefit significantly from a more in-depth
understanding of which specific aspects of this range of social
content most effectively drive the relation between fiction read-
ing and enhanced ToM.
In a similar way, our passages range over a host of broad lit-
erary techniques associated with fiction without studying
which techniques in particular provide the most relevant
opportunities for simulation. Our ‘social’ passages include dif-
ferent techniques to supply readers with psychological informa-
tion about their characters: direct representations of mental
content; multi-level psychological inferences (‘she believed that
he believed . . . ’); free-indirect discourse (or the rendering of
first-person thoughts into third-person narration); and physical
actions linked to psychological states, thoughts and emotions
(e.g. facial expressions and postures). Further narrative elem-
ents, such as issues related to first- and third-person narration,
also present opportunities for further research into the relation
among fiction, social-cognitive abilities and simulation.
The current findings must be interpreted in the context of
several limitations. For one, given our sample size, the brain-
behavior correlations and mediation findings may be overesti-
mates of the true population effect (Button et al., 2013).
Additionally, though it is tempting to draw a causal inference
from the mediation findings, the current data are cross-
sectional and cannot definitively speak to a causal relationship
between the variables. As such, future longitudinal research
should endeavor to establish that fiction reading enhances so-
cial-cognitive abilities. Such a connection would hold extremely
important real-world implications, perhaps guiding the direc-
tion of higher education and social initiatives more broadly, as
well as potentially providing a tolerable and cost-effective inter-
vention for social cognitive deficits in clinical populations.
However, this is not to preclude the possibility that future re-
search might establish an inverse relation: that social-cognitive
abilities instead cause people to read fiction. In either case,
there is still a great deal to be learned about the nature of social
cognition, its relation to fiction reading, and their impacts on
both personal choices and behavior.
Acknowledgments
The authors thank John Bender, Susanna Carmona, Juan
Manuel Contreras, Eshin Jolly, Joe Moran, Brandi Newell,
Kenneth Parreno, Amitai Shenhav, Emma Templeton,
Blakey Vermeule, Adam Waytz and Jamil Zaki for helpful
advice and assistance.
Funding
Andrew Bricker was supported by fellowships from the
Social Sciences and Humanities Research Council of Canada
and the Andrew W. Mellon Foundation.
Supplementary data
Supplementary data are available at SCAN online.
Conflict of interest. None declared.
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