Processing Speech and Thoughts during Silent Reading:

Processing Speech and Thoughts during Silent Reading:
Direct Reference Effects for Speech by Fictional
Characters in Voice-Selective Auditory Cortex
and a Theory-of-Mind Network

Ben Alderson-Day1, Jamie Moffatt1,2, Marco Bernini1, Kaja Mitrenga1
Bo Yao3, and Charles Fernyhough1

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Abstrait

■ Stories transport readers into vivid imaginative worlds, mais
understanding how readers create such worlds—populating
them with characters, objets, and events—presents serious
challenges across disciplines. Auditory imagery is thought to
play a prominent role in this process, especially when repre-
senting characters’ voices. Previous research has shown that
direct reference to speech in stories (par exemple., He said, “I’m over
here”) may prompt spontaneous activation of voice-selective
auditory cortex more than indirect speech [Yao, B., Belin,
P., & Scheepers, C. Silent reading of direct versus indirect
speech activates voice-selective areas in the auditory cortex.
Journal des neurosciences cognitives, 23, 3146–3152, 2011].
Cependant, it is unclear whether this effect reflects differential
processing of speech or differences in linguistic content,
source memory, or grammar. One way to test this is to com-
pare direct reference effects for characters speaking and

thinking in a story. Ici, we present a multidisciplinary
fMRI study of 21 readers’ responses to characters’ speech
and thoughts during silent reading of short fictional stories.
Activations relating to direct and indirect references were
compared for both speaking and thinking. Eye-tracking and
independent localizer tasks (auditory cortex and theory of
esprit [ToM]) established ROIs in which responses to stories
could be tracked for individuals. Evidence of elevated auditory
cortex responses to direct speech over indirect speech was
observed, replicating previously reported effects; no reference
effect was observed for thoughts. De plus, a direct reference
effect specific to speech was also evident in regions previously
associated with inferring intentions from communication.
Implications are discussed for the spontaneous representation
of fictional characters and the potential roles of inner speech
and ToM in this process. ■

INTRODUCTION

Stories can conjure complex imaginative worlds that offer
immersion and transportation for the reader (Vert,
2004; Vert, Brock, & Kaufman, 2004; Ryan, 1999;
Gerrig, 1993). Fictional characters in particular are some-
times experienced with a vividness and complexity,
which can linger beyond the page (Alderson-Day,
Bernini, & Fernyhough, 2017; Maslej, Oatley, & Mar,
2017). Understanding how these experiences are created
by the mind—often with apparent automaticity and
spontaneity—is a challenge for a wide range of disciplines
beyond psychology, including literary theory, narratology,
philosophy of mind, and cognitive neuroscience
(Herman, 2013). Far from passively “receiving” informa-
tion from the writer, readers actively and creatively en-
gage with fictional texts in a way that draws on multiple

1Durham University, 2University of Sussex, 3Université de
Manchester

psychological resources (Polvinen, 2016; Caracciolo,
2014; Kukkonen, 2014; Oatley, 2011; Bortolussi &
Dixon, 2003).

One approach to understanding the qualitative rich-
ness of the reading experience has been to study inner
speech (sometimes also referred to as inner monologue
or articulatory imagery; Alderson-Day & Fernyhough,
2015; Perrone-Bertolotti, Rapin, Lachaux, Baciu, &
Lœvenbruck, 2014). Intuitively, reading is often associated
with the sounding out of an “inner voice,” and the self-
reports of readers involve various kinds of auditory imag-
ery when engaged in a story (Vilhauer, 2016). Although
the reliability of readers’ introspective reports has been
questioned (Caracciolo & Hurlburt, 2016), empirical evi-
dence of inner speech involvement during silent reading
is well documented (Filik & Barber, 2011; Alexander &
Nygaard, 2008). De plus, silent reading appears to elicit
activity in perisylvian regions and auditory association
cortex (Magrassi, Aromataris, Cabrini, Annovazzi-Lodi, &
Moro, 2015; Perrone-Bertolotti et al., 2012), particularly
when characters’ voices and speech are being described

© 2020 Massachusetts Institute of Technology. Published under a
Creative Commons Attribution 4.0 International (CC PAR 4.0) Licence.

Journal des neurosciences cognitives 32:9, pp. 1637–1653
https://doi.org/10.1162/jocn_a_01571

(Brück, Kreifelts, Gößling-Arnold, Wertheimer, &
Wildgruber, 2014; Yao, Belin, & Scheepers, 2011). Tel
findings have been taken as evidence of the reading
experience—and its evocation of inner speech—being
almost akin to hearing external voices (Petkov & Belin,
2013).

A good example of this is provided by texts involving
direct speech. When direct reference is made to a char-
acter overtly speaking in a text (he said, “the cat is over
there”), it is thought to evoke a more vivid experience of
the storyworld than if the same overt speech is only indi-
rectly referred to (he said that the cat is over there). Il
has been suggested that the purpose of such construc-
tions is to demonstrate (and thus depict) a situation,
rather than merely describe it (Clark & Gerrig, 1990).
Evidence that this could resemble hearing an actual voice
is provided by Yao et al. (2011), who compared fMRI re-
sponses in auditory cortex for participants silently read-
ing short stories that contained either direct or indirect
reference to speech. Although both kinds of speech acti-
vated auditory cortex, direct speech was associated with a
greater response than indirect speech in voice-selective
regions of the right superior and middle temporal lobe
(as defined by a separate auditory localizer task; Belin,
Zatorre, Lafaille, Ahad, & Pike, 2000). As the stories were
very short (three to four sentences in total) and partici-
pants were not prompted to imagine the voices, charac-
ters, or stories in any specific way, this suggests that fairly
minimal textual markers for direct speech can elicit a re-
sponse in cortical regions that are selective for voice
perception.

If direct speech in text can prompt this kind of reac-
tion, a second question is why readers appear to respond
in this way. In a separate study, Yao, Belin, and Scheepers
(2012) observed similarly enhanced responses in voice-
selective regions for direct speech quotations when they
were being read by a monotonous voice. Building on
Barsalou’s theory of embodied cognition (Barsalou,
2008), they suggested that auditory cortex activation
may have a role in constructing a perceptual simulation
of the emotional prosody and intonation of the speaker’s
voice, given that such information is either absent or di-
minished in the case of both silent reading and monoto-
nous listening. This would not rule out perceptual
simulation during other kinds of silent reading but char-
acterizes direct reference as a cue to simulate supraseg-
mental and communicative properties of speech from
text ( Yao et al., 2011, 2012).

The effects of direct speech and its potential conse-
quences for simulation can be questioned, cependant. If di-
rect speech prompts more vivid imagery or provides
more communicative information (par exemple., tone or emotional
content), this would plausibly be reflected in reader com-
prehension. Cependant, in a series of behavioral experi-
ments, Eerland, Engelen, and Zwaan (2013) reported
inconsistent evidence for either perceptual or communi-
cative information being more available to readers after

direct speech quotations. Plutôt, they suggested that
the use of direct quotations prompts better memory for
the verbatim content of characters’ utterances, alors que
indirect speech assists the building of a situation model,
c'est, an overall “representation of the referential situa-
tion” (Eerland et al., 2013, p. 7; van Dijk & Kintsch, 1983).
Supporting this, source memory for characters’ utter-
ances is actually enhanced for indirect, not direct, speech
quotations (Eerland & Zwaan, 2018)—suggesting that the
potential vividness of direct speech is not used for track-
ing information about who said what (or could even ob-
struct such tracking, when compared to indirect speech).
Enfin, the typographical and grammatical differences be-
tween direct and indirect speech make it difficult to clearly
compare their specific consequences for mental simula-
tion. Along with potentially alerting the reader to pay
attention to text, direct sentences are typically shorter
than indirect sentences, are syntactically simpler, et
may be expected to prompt changes in reader perspective
(Köder, Maier, & Hendriks, 2015; Coulmas, 2011; Clark &
Gerrig, 1990). En tant que tel, the effect of direct speech on the
lecteur, and its potential function in the imaginative re-
sponse of reading, remains unclear.

One way to explore this topic—in a way that might be-
gin to address some of the above concerns—is to com-
pare references to characters’ speech with another kind
of representation that fictional narratives can involve:
characters’ thoughts. Although theories of mental simula-
tion during reading emphasize various forms of sensory
and embodied simulation (par exemple., Kurby & Zacks, 2013;
Zwaan, Madden, Yaxley, & Aveyard, 2004), fictional narra-
tives have been proposed to place specific sociocognitive
demands on the reader (Mar & Oatley, 2008; Zunshine,
2006). Typiquement, a reader must track the mental states of
multiple characters, following their beliefs, intentions,
and desires through a narrative, to make sense of actions,
decisions, and responses to events in the storyworld
(Spreng, Mar, & Kim, 2009; Herman, 2008; Palmer,
2004; Gerrig, Brennan, & Ohaeri, 2001),1 all of which im-
ply a central role for theory of mind (ToM) in the reading
processus.

How might this shed light on direct speech? First is be-
cause it provides a contrasting example of direct refer-
ence. Both indirect and direct references to thinking
are used in narratives. Indirect thought, which is usually
considered the representational norm (Leech & Short,
2007, p. 268), is more flexible and can be used to repre-
sent verbal, preverbal, and nonverbal mental processes
from the perspective of the character (par exemple., he thought
that X; he felt that Y; he was willing to do Z ). Direct
thought (also referred to as “quoted monologue”;
Cohn, 1978) is used to represent, verbatim, the linguistic
silent articulation of verbal thoughts (He thought “this is
so complicated!»).

The verbalized nature of depicting characters’ thoughts
is almost identical in form and complexity to direct speech
(c'est à dire., when used in a basic form; indirect thoughts in more

1638

Journal des neurosciences cognitives

Volume 32, Nombre 9

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extended narratives can be used in highly complex ways).
Contrasting these forms of speech and verbal thought can
therefore provide a test of Yao et al.’s (2011) interpreta-
tion of direct reference effects by assessing how specific
they might be to vocal information, while at the same
time controlling for typographic features. If Yao et al.’s
conjecture is correct, direct reference to speech—but
not necessarily thoughts—may be expected to elicit re-
cruitment of voice-selective regions of auditory cortex,
to specifically simulate the perceptual qualities of charac-
ters speaking out loud in the storyworld. In contrast, si
direct speech and direct thought elicit similar responses,
then a voice-specific account of direct reference would be
harder to maintain. It could be the case that both speech
and thoughts elicit some form of perceptual simulation
under direct reference, but they would be doing so de-
spite clear dissimilarities in the auditory scenario (one is
an external utterance, and the other is a form of internal
monologue). Plutôt, showing that direct reference to
speech and thoughts prompts a generally greater re-
sponse in auditory cortex could support alternative in-
terpretations: It may, Par exemple, simply reflect a
greater level of engagement that happens when quote
marks prompt the reader to pay attention to verbatim
content (Eerland et al., 2013). In that scenario, là
would be nothing special about speech for understand-
ing direct reference effects.

Deuxième, exploring sociocognitive processing and con-
trasting how this works for speaking and thinking are po-
tentially highly informative for understanding direct
speech effects. ToM has multiple components, each with
its own developmental trajectory (Fernyhough, 2008;
Tomasello, Carpenter, Call, Behne, & Moll, 2005).
Whereas some socio-cognitive skills are evident early in
infancy—such as the ability to follow others’ attentional
cues (Behne, Liszkowski, Carpenter, & Tomasello, 2012;
Woodward, 1998)—the ability to cognitively represent
others’ mental states when incorrect is thought to emerge
later in childhood (typically around 4 ans; Wellman,
Cross, & Watson, 2001). De la même manière, understanding prag-
matic information and speaker intention from prosody
shows a competence–performance gap, in which vocal
cues to emotion are recognized very early in infancy but
are only used consistently (in the face of, par exemple., conflicting
cues) in older children (Esteve-Gibert & Guellaï, 2018).
ToM and social cognition more generally are associated
with a canonical network of regions in the medial pFC
(mPFC), precuneus, and TPJ bilaterally (Molenberghs,
Johnson, Henry, & Mattingley, 2016; Schurz, Radua,
Aichhorn, Richlan, & Perner, 2014; Saxe & Kanwisher,
2003; Fletcher et al., 1995). Of these, representing the
thoughts and intentions of others in particular has been
argued to localize to regions of the TPJ and precuneus
(Schurz, Tholen, Perner, Mars, & Sallet, 2017; Saxe &
Powell, 2006), whereas mPFC has been linked to the pro-
cessing of more constant traits associated with self and
other (van Overwalle, 2009).

If direct speech prompts a detailed simulation of su-
prasegmental vocal information (such as emotional tone
or prosody), then this may also be reflected in social–
cognitive regions—specifically for areas associated with
interpreting or reasoning about a speaker’s communica-
tive intentions. Par exemple, using a nonverbal, cartoon-
based story task, Ciaramidaro et al. (2007) observed that
bilateral TPJ regions in particular are associated with
tracking different kinds of intent associated with socio-
communicative interactions. If direct speech prompted
similar activation, this would support an extension of
Yao et al.’s (2011) original theory to suggest that direct
reference involves constructing a broader, sociopercep-
tual simulation than merely how a voice sounds.
Contrastingly, if tracking characters and their intentions
is an ultimately separate process from simulating the per-
ceptual features of characters’ voices, then no direct ef-
fect for speech would necessarily be expected in ToM
régions. Plutôt, it is possible that references to charac-
ters’ mental states—but not their speech—would be
most likely to engage such regions, irrespective of any di-
rect reference effect.

To investigate this, we adapted Yao et al.’s (2011) par-
adigm to include direct and indirect references to charac-
ters’ verbal thoughts and speech in a 2 × 2 conception. Nous
used eye tracking and an auditory localizer task to study
cortical responses specific to each individual’s reading
times and voice-selective regions. To explore the broader
effect of direct speech in regions commonly associated
with inferring communicative intentions, we also included
a version of Ciaramidaro et al.’s (2007) story task as a sec-
ond localizer. Many standard ToM tasks use written short
stories in which characters’ false beliefs must be inferred
from textual information, but using such stories could be
expected to overlap considerably with other reading
tasks (in terms of both stimuli and task demands).
Plutôt, by using a wordless, cartoon-based ToM task,
we could avoid this potential confound with the demands
of our main direct/indirect story task. On the basis of the
original findings of Yao et al. (2011, 2012), we hypothe-
sized that (je) direct reference effects would be evident for
speech but not thoughts in auditory cortex. In accor-
dance with the claim that this facilitates prosodic and
communicative processing of the utterance, we also pre-
dicted that (ii) the voice-specific effect of direct reference
would extend to ToM-related regions. In contrast, Non
direct reference effects were expected for thoughts,
in either network.

MÉTHODES

Participants

An initial sample of 30 individuals took part in the full
MRI procedure, but nine participants did not produce a
full data set because of the following exclusions (one in-
cidental finding, one insufficient accuracy [<60%] on the Alderson-Day et al. 1639 D o w n l o a d e d l l / / / / j f / t t i t . : / / f r o m D o h w t t n p o : a / d / e m d i f r t o p m r c h . s p i l d v i r e e r c t c . m h a i e r d . u c o o m c n / j a o r t c i c n e / - a p r d t i 3 c 2 l 9 e 1 - 6 p 3 d 7 f 2 / 0 3 1 2 3 / 8 9 2 / 9 1 6 o 3 c 7 n _ / a 1 _ 8 0 6 1 2 5 0 7 1 1 7 p / d j o b c y n g _ u a e _ s 0 t 1 o 5 n 7 0 1 8 . p S d e f p e b m y b e g r u 2 e 0 s 2 t 3 / j t f . / o n 0 5 M a y 2 0 2 1 story task, two no clear voice-selective response on the auditory localizer task, five insufficient eye-tracking data; three male, six female). As such, analysis proceeded with a final sample of 21 (age: M = 23.49, SD = 6.63; three male, 18 female). All participants were right-handed, na- tive English speakers, with normal or corrected-to-normal vision. All procedures were approved by a university ethics subcommittee. Measures Story Task Following Yao et al. (2011), participants viewed a series of short stories containing two preparation sentences (Sentences 1 and 2) and a target sentence, containing a character either (i) speaking or thinking with (ii) direct or indirect reference. On each trial, participants viewed a fixation cross for 1–2 sec ( jittered at random), followed by one slide per sentence, presented sequentially (see Figure 1). Viewing times per slide were determined using the following formula: ( Words × 100 msec) + (Syllables × 50 msec) + 2000 msec. Mean presentation times were 5.61 and 5.72 sec for Sentences 1 and 2, respectively, and 5.95 and 6.22 sec for direct and indirect target sen- tences, respectively, reflecting the slightly longer length of indirect sentences on average (18.6 words per indirect sentence compared to 16.8 for direct sentences). To allow for sufficient trials in each condition, the number of stories was increased from the 90 trials used in Yao et al. (2011) to 120, split across two 20-min runs (additional stories were prepared by a narratologist, M. B., to follow the length, complexity, and style of the original stimuli and ensure balance across the four conditions). Each run also con- tained three 30-sec break periods, occurring every 20 trials. An attentional check (a simple comprehension question relating to factual content from the preceding story) was included after 25% of trials, with participants having 6 sec to respond.2 Four random orders of trials were generated, counterbalancing the combination of voice/thought and direct/indirect target sentences across participants. Eye- tracking timings were collected as an indicator of partici- pants’ reading responses for the two preparation sentences and the target sentence. Specifically, participants’ first D o w n l o a d e d l l / / / / j f / t t i t . : / / f r o m D o h w t t n p o : a / d / e m d i f r t o p m r c h . s p i l d v i r e e r c t c . m h a i e r d . u c o o m c n / j a o r t c i c n e / - a p r d t i 3 c 2 l 9 e 1 - 6 p 3 d 7 f 2 / 0 3 1 2 3 / 8 9 2 / 9 1 6 o 3 c 7 n _ / a 1 _ 8 0 6 1 2 5 0 7 1 1 7 p / d j o b c y n g _ u a e _ s 0 t 1 o 5 n 7 0 1 8 . p S d e f p e b m y b e g r u 2 e 0 s 2 t 3 / j / t . f o n 0 5 M a y 2 0 2 1 Figure 1. Adapted story task (A) with direct reference to voices and thoughts applied to auditory and ToM localizer regions (B). Figure 1B depicts left-sided sagittal view (rendered, p < .05, FWE); note that auditory and ToM regions were observed bilaterally (see Table 1). 1640 Journal of Cognitive Neuroscience Volume 32, Number 9 Table 1. Accuracy Rates, Reading Onsets, and Reading Times by Task Condition Direct Speech Indirect Speech Direct Thought Indirect Thought M SD Accuracy (%) 79.84 18.43 Reading onset (sec) Duration (sec) 0.56 4.02 0.32 0.54 M 79.21 0.57 4.10 SD 16.08 0.33 0.54 M 85.56 0.61 4.06 SD 15.87 0.36 0.44 M 86.35 0.67 4.11 SD 15.90 0.36 0.58 fixation (the beginning of the sentence) and last fixation (the final line of the target sentence) within the text area were used to define reading onsets and offsets of characters’ speaking and thinking in the target sentence. These were then directly included in the fMRI model to account for individual differences in the reading response. Auditory Localizer Task The auditory localizer task was identical to that used in Yao et al. (2011). Participants listened to 20 blocks of vo- cal stimuli and 20 blocks of nonvocal stimuli, along with 20 silent blocks that were used as a baseline. The blocks were presented randomly. Each block was 8 sec long, and the task lasted 10 min. The contrasting brain activity in response to the vocal and nonvocal stimuli reliably local- izes voice-selective areas of the auditory cortex ( Yao et al., 2011; Belin et al., 2000). ToM Task The cartoon-based ToM task was adapted from a task used by Walter et al. (2004) and Ciaramidaro et al. (2007). Participants viewed a sequence of three cartoon story vignettes (“story” phase) and were required to indi- cate a logical end of each story based on the three pre- sented images (“choice” phase). The story phase included either reasoning about characters’ intentions when communicating with others (e.g., a man indicating whether a seat is free on a train) or physical reasoning (e.g., a water pipe bursting). The images were displayed sequentially for 3 sec in the story phase and for 7 sec in the choice phase. The intertrial intervals lasted between 7 and 11 sec. In total, 10 ToM stories and 10 physical reasoning stories were presented in a random order. Participants answered (A, B, or C) by a button press. The task took 9 min to complete. The contrasting brain activity in response to the ToM reasoning stories compared to physical reasoning stories has been ob- served to prompt activity in brain regions often associated with ToM, including the right TPJ, precuneus, and anterior paracingulate cortex (Alderson-Day et al., 2016; Ciaramidaro et al., 2007; Walter et al., 2004). Data Acquisition fMRI data were acquired at Durham University Neuroimaging Centre using a 3-T Magnetom Trio MRI system (Siemens Medical Systems) with standard gradi- ents and a 32-channel head coil. T2*-weighted axial EPI scans were acquired with the following parameters: field of view = 212 mm, flip angle = 90°, repetition time = 2000 msec, echo time = 30 msec, number of slices = 32, slice thickness = 3.0 mm, interslice gap = 0.3 mm, and matrix size = 64 × 64. Story task data were collected across 2 × 20-min runs consisting of 600 volumes each; auditory and ToM tasks took roughly 10 min each and consisted of 300 and 281 volumes, re- spectively. The first three volumes of each EPI run were discarded to allow for equilibrium of the T2 response. For each participant, an anatomical scan was acquired using a high-resolution T1-weighted 3-D sequence (number of slices = 192, slice thickness = 1 mm, matrix size = 512 × 512, field of view = 256 mm, echo time = 2.52 msec, repetition time = 2250 msec, flip angle = 9°). Eye-tracking data were collected using a LiveTrack system (Cambridge Research Systems) with MATLAB 2016b (The Mathworks, Inc.). Data Analysis All MRI analyses were conducted using SPM Version 12 ( Wellcome Department of Cognitive Neurology) imple- mented in MATLAB. Images were slice-time corrected before being realigned to the first image to correct for head movement. Volumes were then normalized into standard stereotaxic anatomical Montreal Neurological Institute space using the transformation matrix calculated from the first EPI scan of each participant and the EPI template. The default settings for normalization in SPM12 and the standard EPI template supplied with SPM12 were used. The normalized data with a resliced voxel size of 2 × 2 × 2 mm were smoothed with an 8-mm FHWM isotropic Gaussian kernel to accommodate intersubject anatomical variation. The time-series data were high-pass filtered with a high-pass cutoff of 1/128 Hz, and first-order autocorrelations of the data were estimated and corrected for. Movement parameters from the re- alignment phase were visually inspected for outliers and included as regressors for single-participant (first-level) Alderson-Day et al. 1641 D o w n l o a d e d l l / / / / j f / t t i t . : / / f r o m D o h w t t n p o : a / d / e m d i f r t o p m r c h . s p i l d v i r e e r c t c . m h a i e r d . u c o o m c n / j a o r t c i c n e / - a p r d t i 3 c 2 l 9 e 1 - 6 p 3 d 7 f 2 / 0 3 1 2 3 / 8 9 2 / 9 1 6 o 3 c 7 n _ / a 1 _ 8 0 6 1 2 5 0 7 1 1 7 p / d j o b c y n g _ u a e _ s 0 t 1 o 5 n 7 0 1 8 . p S d e f p e b m y b e g r u 2 e 0 s 2 t 3 / j t . f / o n 0 5 M a y 2 0 2 1 Table 2. Whole-Brain Coordinates for (A) Auditory and (B) ToM Localizer Tasks k t z pFWE Location x y (A) Auditory (vocal > nonvocal)

L middle temporal gyrus

L superior temporal gyrus

L middle temporal gyrus

R superior temporal gyrus

R temporal pole

R superior temporal gyrus

−60

−58

−60

56

48

64

−14

−2

−36

−18

12

−4

(B) ToM (communicative inference > physical reasoning)

R middle cingulate cortex

L precuneus

WM

R middle temporal gyrus

R superior temporal gyrus

R middle temporal gyrus

WM

L temporal pole

L middle temporal gyrus

L superior medial frontal gyrus

R superior medial frontal gyrus

L posterior-medial frontal gyrus

L gyrus rectus

4

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12.76

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12.43

10.25

9.51

12.24

12.03

10.44

11.59

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10.35

10.68

9.21

7.92

7.76

7.34

6.34

6.29

6.58

6.39

6.08

5.95

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5.33

5.92

5.88

5.55

5.79

5.76

5.53

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5.26

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<.001 <.001 <.001 <.001 <.001 <.001 .015 −14 279 All results p < .05, FWE, at peak/cluster level. Minimum cluster size: k = 50. L = left; R = right; ToM = theory of mind; WM = white matter. analyses. ROI analyses were conducted using the Marsbar toolbox (Brett, Anton, Valabregue, & Poline, 2002). Individual ROIs were defined using p < .05 cor- rected for FWE at cluster level, in temporal cortical re- gions for the auditory localizer task and clusters in mPFC, precuneus, and TPJ regions for the ToM task. Where significant clusters were not evident for individ- ual participants at this level, a more liberal threshold of p < .001 (uncorrected) was used to maximize sensitiv- ity to individual differences; participants who showed no clusters in these regions even at the more liberal threshold were excluded from analyses (two auditory, six ToM). All whole-brain analyses are presented at p < .05, FWE, cluster-level corrected. All statistical anal- yses of mean beta values were conducted using R and jamovi; figures were generated using ggplot2 and MicroGL. Effect sizes are reported as Cohen’s d for pairwise comparisons and ηp 2 values for ANOVA main and interaction effects. ηp 2 values can be considered as small, moderate, and large effects with values of .099, .0588, and .1379, respectively (Richardson, 2011; Cohen, 1969). RESULTS Accuracy on the task was generally high (M = 82.5%, SD = 7.4%) indicating that participants maintained attention despite the 40-min duration of the task. Repeated- measures ANOVA with a 2 × 2 (Form × Reference) de- sign was used to compare behavioral responses for the four conditions (see Table 1). No main effects, interac- tion effects, or pairwise comparisons were significant for condition accuracy, although we observed a nonsig- nificant trend for participants to be slightly less accurate on speech trials compared with thought trials, F(1, 20) = 3.34, p = .082, ηp 2 = .14 ( p > .14 for all other effects
and comparisons). For the duration of reading times,
the only effect close to significance was for direct com-
pared with indirect reference, F(1, 20) = 3.57, p = .073,

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Tableau 3. Whole-Brain Coordinates for Speech and Thought Sentences vs. Baseline

Location

Direct speech

L middle temporal gyrus

L middle temporal gyrus

L middle temporal gyrus

R temporal pole

R temporal pole

L temporal pole

L inferior frontal gyrus

L inferior frontal gyrus

L superior medial frontal gyrus

L precentral gyrus

Indirect speech

L middle temporal gyrus

L middle temporal gyrus

L middle temporal gyrus

L temporal pole

L inferior frontal gyrus

L temporal pole

R medial temporal pole

R temporal pole

L middle temporal gyrus

L inferior frontal gyrus

L middle temporal gyrus

L middle temporal gyrus

L middle temporal gyrus

Direct thought

L middle temporal gyrus

L middle temporal gyrus

L middle temporal gyrus

R medial temporal pole

L superior medial frontal gyrus

L superior frontal gyrus

R middle temporal gyrus

L inferior frontal gyrus

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8.55

10.92

9.21

8.83

8.64

8.31

8.63

8.43

11.84

10.42

7.73

9.35

7.57

7.52

9.31

8.46

7.87

7.79

11.84

10.42

7.73

689

10.79

8.6

8.31

9.6

8.75

6.69

8.16

7.18

344

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5.83

5.49

6.17

5.69

5.58

5.51

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5.51

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5.74

5.14

5.12

5.73

5.45

5.25

5.22

6.38

6.04

5.2

6.13

5.5

5.41

5.81

5.55

4.79

5.35

4.99

<.001 <.001 <.001 <.001 <.001 <.001 <.001 <.001 <.001 <.001 <.001 <.001 <.001 <.001 <.001 <.001 Alderson-Day et al. 1643 D o w n l o a d e d l l / / / / j t t f / i t . : / / f r o m D o h w t t n p o : a / d / e m d i f r t o p m r c h . s p i l d v i r e e r c t c . m h a i e r d . u c o o m c n / j a o r t c i c n e / - a p r d t i 3 c 2 l 9 e 1 - 6 p 3 d 7 f 2 / 0 3 1 2 3 / 8 9 2 / 9 1 6 o 3 c 7 n _ / a 1 _ 8 0 6 1 2 5 0 7 1 1 7 p / d j o b c y n g _ u a e _ s 0 t 1 o 5 n 7 0 1 8 . p S d e f p e b m y b e g r u 2 e 0 s 2 t 3 / j f / t . o n 0 5 M a y 2 0 2 1 Table 3. (continued ) Location Indirect thought L middle temporal gyrus L middle temporal gyrus L middle temporal gyrus R medial temporal pole R temporal pole R middle temporal gyrus L superior medial gyrus L precentral gyrus L temporal pole L temporal pole L temporal pole x y z k t z pFWE −54 −50 −54 50 46 50 −10 −42 −52 −44 −46 −34 −26 −54 12 20 −38 54 −2 12 18 16 −2 −6 16 −24 −26 −2 28 56 −20 −16 −32 1048 343 56 76 70 206 11.2 11.07 9.74 10.78 10.4 8.91 8.89 8.52 8.42 8.42 6.69 6.23 6.2 5.85 6.13 6.03 5.6 5.59 5.48 5.44 5.44 4.79 <.001 <.001 <.001 <.001 <.001 <.001 All results p < .05, FWE, at cluster and peak levels. Minimum cluster size: k = 50. L = left; R = right; ToM = theory of mind; WM = white matter. ηp 2 = .15, which likely reflected the slightly longer lengths of indirect sentences. All other effects and comparisons for duration were also nonsignificant (all ps > .15).
Reading onsets, in contrast, showed a main effect of
Form, F(1, 20) = 7.10, p = .015, ηp
2 = .26, such that readers
were quicker to start reading speech trials; follow-up
pairwise comparisons indicated that this was only signifi-
cantly quicker for direct speech compared with indirect
thought, t = 2.20, df = 36.24, p = .035 (uncorrected), d =
0.4, all other ps > .10.

Whole-brain analyses—included here for descriptive
purposes—indicated that the vocal > nonvocal contrast
from the auditory localizer task was associated with sig-
nificantly greater activation in bilateral auditory cortices,
across the middle temporal gyrus (MTG) and superior
temporal gyrus (see Figure 1B and Table 2). Compared
with baseline, each of the four reading task conditions
was associated with temporal activation bilaterally, avec
the largest clusters being observed along the dorsal bank
of the left MTG (Tableau 3).

Responses to Characters’ Speech and Thoughts in
Voice-Selective Auditory Cortex

A repeated-measures ANOVA was used to compare mean
beta values in auditory ROIs for story passages containing
characters’ speech or thoughts (c'est à dire., Form) in direct or
indirect reference, in a 2 × 2 conception. No significant main
effect of Form was evident, F(1, 20) = 0.31, p = .584,
ηp
2 = .02, although a trend was observed for reference
in favor of direct quotation, F(1, 20) = 4.00, p = .059,
ηp
2 = .17. The interaction of Form and Reference was sig-
nificant, F(1, 20) = 7.08, p = .015, ηp
2 = .26. As displayed
in Figure 2, this was largely driven by a specific direct

reference effect for character’s speech, but not thoughts.
Pairwise comparisons indicated that mean beta values for
direct speech were significantly higher than those for in-
direct speech ( p = .006, d = 0.84, Bonferroni corrected),
but no other pairwise contrasts were significant (all ps > .25).

Responses to Speech and Thoughts in a
ToM Network

We then applied the same analyses to responses in a ToM
network identified via the cartoons task. As shown in

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Chiffre 2. Mean beta values for direct and indirect references to
characters’ speech and thoughts in voice-selective auditory cortex.
Error bars represent SEM.

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in voice-selective regions of the auditory cortex, no signif-
icant increase in signal was seen for this region when cor-
recting across the whole brain for direct versus indirect
speech (see Table 3). No regions were more active in
the reverse contrast (indirect > direct speech).

Other exploratory whole-brain comparisons indicated
few differences between conditions. Two exceptions
were direct speech versus direct thought and direct ref-
erence versus indirect reference (c'est à dire., with speech and
thought sentences combined). Direct speech compared
to direct thought was associated with greater activation
in the right insula and anterior and middle cingulate, dans-
cluding regions bordering on the pre-SMA (voir
Table 4B). Direct reference was observed to predomi-
nantly activate occipital and parietal regions more than
indirect reference (Table 4C). Their reverse contrasts (di-
rect thought > direct speech; indirect > direct) pro-
duced no significant clusters, even at an uncorrected

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Chiffre 4. Whole-brain activation for (UN) direct and indirect speech and
thoughts versus baseline, (B) direct > indirect speech (DS > IS), et
(C) DS > IS with group-level vocal > nonvocal contrast from auditory
localizer data. All activations significant at cluster level, p < .05, FWE, apart from C, which reflects p < .001, uncorrected (uncorr.). corr. = corrected. Alderson-Day et al. 1645 Figure 3. Mean beta values for direct and indirect references to characters’ speech and thoughts in ToM regions. Error bars represent SEM. Table 1, a range of typical regions were identified in the contrast between communicative inference reasoning and physical reasoning on the task, including the mPFC, precuneus, and TPJ bilaterally. Sixteen of the 21 individuals produced ToM networks with significant clus- ters in at least one of these regions, and their beta values were taken forward for ROI analysis (15/16, right TPJ; 12/16, left TPJ; 7/16, precuneus; 6/16, mPFC). When the mean beta values were compared in these areas in a repeated-measures ANOVA, no main effects of Form, F(1, 15) = 0.49, p = .493, ηp 2 = .03, or Reference, F(1, 15) = 1.74, p = .207, ηp 2 = .10, were observed, but a significant interaction was again evident, F(1, 15) = 9.39, p = .008, ηp 2 = .38.3 As Figure 3 shows, this too was driven by responses for direct speech (compared with indirect speech), and this was the only significant difference between the conditions ( p = .016, d = 0.90, Bonferroni corrected). We then conducted an exploratory whole-brain analysis to investigate any further potential differences for direct versus indirect speech. Significant increases in signal for direct over indirect speech were evident in three regions: right TPJ (encompassing right angular gyrus [AG] and MTG), left inferior frontal gyrus (IFG), and left superior parietal lobule (see Figure 4). Using the online meta- analytic tool Neurosynth ( Yarkoni, Poldrack, Nichols, Van Essen, & Wager, 2011), the most common functional terms associated with these regions were “network DMN” for the right AG (posterior probability = 0.73), “theory mind” for the right MTG ( p = .88), “semantic” for the left IFG ( p = .88), and “imagery” for the left SPL ( p = .78). Despite the apparent direct speech effect Table 4. Whole-Brain Activation Differences for (A) Direct vs. Indirect Speech, (B) Direct Speech vs. Direct Thought, and (C) Direct vs. Indirect Reference Location (A) Direct speech > indirect speech

R angular gyrus

R angular gyrus

R middle temporal gyrus

L inferior frontal gyrus

L inferior frontal gyrus

L inferior frontal gyrus

L superior parietal lobule

L middle occipital gyrus

L middle occipital gyrus

(B) Direct speech > direct thought

R middle cingulate cortex

R ACC

L ACC

Right insula

Right insula

(C) Direct > indirect

L superior parietal lobule

WM

L superior parietal lobule

L middle occipital gyrus

WM

R superior occipital gyrus

WM

R superior occipital gyrus

R middle occipital gyrus

R middle occipital gyrus

WM

R middle temporal gyrus

R middle temporal gyrus

R inferior temporal gyrus

L inferior occipital gyrus

L middle occipital gyrus

L inferior occipital gyrus

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3.50

<.001 <.001 .007 .002 .036 .00 .01 .00 .00 .01 .01 D o w n l o a d e d l l / / / / j f / t t i t . : / / f r o m D o h w t t n p o : a / d / e m d i f r t o p m r c h . s p i l d v i r e e r c t c . m h a i e r d . u c o o m c n / j a o r t c i c n e / - a p r d t i 3 c 2 l 9 e 1 - 6 p 3 d 7 f 2 / 0 3 1 2 3 / 8 9 2 / 9 1 6 o 3 c 7 n _ / a 1 _ 8 0 6 1 2 5 0 7 1 1 7 p / d j o b c y n g _ u a e _ s 0 t 1 o 5 n 7 0 1 8 . p S d e f p e b m y b e g r u 2 e 0 s 2 t 3 / j t f . / o n 0 5 M a y 2 0 2 1 All results p < .05, FWE, at cluster level, and p < .001, uncorrected, at peak level. Minimum cluster size: k = 50. WM = white matter. 1646 Journal of Cognitive Neuroscience Volume 32, Number 9 significance level ( p < .001, uncorrected, k > 50).
De la même manière, no whole-brain differences were observed be-
tween voices and thoughts overall or between indirect
forms of speech and thought, either at corrected or un-
corrected levels.

DISCUSSION

The aim of this study was to explore further the effect of
direct speech in the brains of readers. The main finding
of our results was to replicate the original effect reported
by Yao et al. (2011), namely, that direct speech in short
stories is accompanied by elevated responses in voice-
selective auditory regions of the brain, when compared
with indirect speech. Our findings go further than those
of Yao et al. in two key ways. D'abord, by comparing direct
and indirect references for speech and thoughts, our ROI
results demonstrate a specific effect of reference for char-
acters who are represented as speaking, but not when
they are represented as thinking. Deuxième, this direct
speech effect appears to extend beyond voice-selective
auditory cortex to also include regions that are used
when making inferences about communicative inten-
tion, based on a ToM localizer task (Ciaramidaro et al.,
2007). This pattern of results, donc, supports the ear-
lier observation that readers spontaneously engage sen-
sory cortices when faced with direct speech, but it also
implicates higher-order processes associated with gaug-
ing character intention and meaning.

Evidence of a direct speech effect in auditory cortex is
consistent with previous findings that such regions are
recruited during silent reading of characters’ speech
(Yao et al., 2011, 2012), which is in turn suggestive of au-
ditory verbal imagery being used during this process.
This aligns with behavioral evidence of phonologically
detailed imagery being involved in silent reading of vari-
ous kinds (Kurby & Zacks, 2013; Filik & Barber, 2011).
There is debate around how specific any such voice rep-
resentation would be: Kurby, Magliano, and Rapp (2009)
have argued that such effects are specific to familiar
voices only, whereas Petkov and Belin (2013) propose
that any kind of voice simulation is likely to reflect a ge-
neric speaking voice. Their argument for this is based on
phonological information specific to voice identity usually
being associated with anterior temporal cortex, alors que
those associated with direct speech in Yao et al. (2011),
Par exemple, are more focused on posterior temporal re-
gions (Petkov & Belin, 2013). Our findings cannot easily
arbitrate between these two possibilities (general vs. spe-
cific voices), as voice-selective auditory regions were
identified along the length of the superior temporal gyri
bilaterally. Cependant, we would speculate that any simula-
tion of a generic or specific voice is likely to vary consid-
erably across individuals. When asked, readers describe
drawing upon a wide range of active and creative strate-
gies to imagine the voices of characters, including other

familiar voices and their own voice (Alderson-Day et al.,
2017).

Perhaps more notable is the suggestion of direct
speech effects also being present in cortical regions often
associated with ToM in general and understanding
others’ intentions in particular.4 We chose a localizer
task that aimed to minimize superficial overlaps with
the primary task—using cartoons instead of a written
story format—and focused specifically on assessing un-
derstanding of communicative intentions over other
types of ToM reasoning, such as inferring false beliefs
(Ciaramidaro et al., 2007; Walter et al., 2004). This pro-
duced a network that, in our sample, primarily centered
around bilateral TPJ regions but also included precuneus
and mPFC in subsets of participants. Evidence of a direct
speech effect in these regions provides at least prima
facie support for the idea that text presented in this
way prompts engagement with what a character intends
to say (Yao et al., 2011, 2012), despite the mixed behavioral
evidence that direct reference primes any further commu-
nicative information about characters (Eerland et al., 2013).
De plus, our analysis suggests involvement of these re-
gions at a comparable level to responses in auditory net-
travaux, as indicated by the lack of any interaction effect
across the two ROIs.

Drawing strong conclusions about the role of these
regions in processing direct speech is fraught with dif-
ficulty. The areas highlighted by our ToM task are often
implicated in a range of attentional and cognitive pro-
cesses (Spreng et al., 2009; Mitchell, 2008), and making
broader claims based on the prior literature raises the risk
of reverse inference (Poldrack, 2006). Using Neurosynth
( Yarkoni et al., 2011), which provides at least a systematic
approach to informal reverse inference (Poldrack, 2011),
the strongest responses in the localizer task were in two
regions where the most common associations in the lit-
erature are with “mind tom” and “theory mind” (avec
posterior probabilities of .87–.90). De la même manière, in the ex-
ploratory whole-brain analysis, the right MTG peak in par-
ticular showed high z scores for tests of association (z =
12.00) and uniformity (z = 14.39) in a ToM meta-analytic
map of 181 études ( Yarkoni et al., 2011).

These regions have also been observed in similar work
examining sociocognitive responses to fiction reading by
Tamir, Bricker, Dodell-Feder, and Mitchell (2016), al-
though in their study, they observed preferential engage-
ment of the mPFC for social content in stories (describing
a person’s mental content), with medial temporal cortex
more closely indexing story vividness. In contrast, most
of our participants (15/16) activated the right TPJ on
our localizer task (compared with only six for mPFC),
and this was the only ToM region to be identified in
our whole-brain analysis comparing direct and indirect
speech. The right TPJ cluster that we observed in this
analysis included peaks in the right AG, extending dorsally
and caudally from areas that are often linked to represent-
ing others’ mental states (Bzdok et al., 2013). Both left

Alderson-Day et al.

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and right AG have been associated with support for the
default mode network, via the generation and processing
of transmodal information in the absence of stimulus in-
put (Murphy et al., 2018) and modality-independent con-
tributions to imagery (Daselaar, Porat, Huijbers, &
Pennartz, 2010). The right AG has also recently been im-
plicated in making valence judgments from nonverbal
cues: In a paradigm where participants were asked to
judge the intentions of musical alien “signals,” variations
in the consonance and dissonance of the stimuli (roughly
corresponding to positive and negative emotions) modu-
lated this region specifically (Bravo et al., 2017). Le
broader extension of this cluster, donc, may reflect
the generation and maintenance of intention-related im-
agery, rather than representing characters’ mental states,
or social content more generally. This being associated
with posterior ToM regions over mPFC would also be con-
sistent with van Overwalle’s (2009) distinction between a
posterior ToM subsystem supporting representation of
temporary and perceptually based intentions and goals,
versus an anterior pFC system that tracks and integrates
enduring social information over time.

When taken together, these findings broadly support
the interpretation of direct speech made by Yao et al.
(2011). Recall that, for Yao and colleagues, direct speech
prompts auditory imagery as a means of modeling
speaker prosody (et, finalement, communicative intent).
A counterhypothesis, provided by Eerland, Zwaan, et
colleagues, is that direct reference acts primarily as a
cue to simulate verbatim linguistic content—in other
words, emphasizing the words but, arguably, not the
conférencier (Eerland & Zwaan, 2018; Eerland et al., 2013).
Our data suggest that direct reference has a specific effect
for speech, and this extends to regions that would be
consistent with inferring communicative intentions.
De plus, this can be distinguished from the overall ef-
fect of direct reference, which primarily shows greater
engagement in visual areas of occipital and parietal cortex
(see Table 4C).

A curious characteristic of our data is the apparently
contradictory results for a direct speech effect in auditory
régions, which was evident in the ROI analysis, but not
for the whole-brain contrast. This likely reflects (je) indi-
vidual variability in the temporal voice area (Belin et al.,
2000), (ii) the effect of the more conservative statistical
correction required across the whole brain, et (iii) le
fact that both direct and indirect speech activate a range
of overlapping temporal regions, with any subsequent
difference in beta values being likely to be subtle.
Nevertheless, it should be noted that prominent differ-
ences across the cortex were observed in the right TPJ
(as discussed), left SPL, and left IFG, much more obviously
than for regions of the auditory cortex. The involvement of
the latter in particular is consistent with greater demand
being placed on inner speech production to support the
representation of direct speech, given the common asso-
ciation of Broca’s area with silent articulation (Alderson-

Day & Fernyhough, 2015; Kühn, Fernyhough, Alderson-
Day, & Hurlburt, 2014; Simons et al., 2010; Shergill et al.,
2001). Evidence from psycholinguistics research suggests
that greater involvement of articulatory processes in si-
lent speech results in more detailed acoustic properties
being represented in auditory imagery (Oppenheim &
Dell, 2010), and both external and internal speech have
been shown to consistently modulate auditory cortical re-
sponses (Okada, Matchin, & Hickok, 2018; Ylinen et al.,
2014; Shergill et al., 2002). En outre, two recent studies
of inner speech have highlighted how right-hemisphere
homologs of left-hemisphere language regions are re-
cruited when speech of others must be imagined
(Grandchamp et al., 2019; Alderson-Day et al., 2016). UN
potential model, alors, would be that a reader coming
across direct speech in a text is prompted to generate a
communicatively plausible perceptual simulation, via in-
ner speech, which involves the left IFG and right TPJ
working in concert to modulate voice-selective regions
of the auditory cortex. This is not to suggest that inner
speech (and other auditory imagery processes) would
not be evidenced in each of the task conditions (given
the widespread activation vs. baseline seen for all condi-
tion; see Table 2 and Figure 4A) but rather that direct
speech could place a specific demand on internal articu-
latory processes. In this scenario, direct reference effects
in auditory cortex would plausibly not be the primary
component of the reader’s response but a secondary
consequence of inner speech (and ToM) processes,
which may explain their relative prominence in our
whole-brain results.

Although the present results appear to have much to
say about how speech is treated by readers, they perhaps
say less about what is happening for characters’ thoughts.
Despite having received early theoretical attention in sty-
listics (Sharvit, 2008; Sotirova, 2004), until now, qualita-
tive differences between direct and indirect modes of
speech and thought representation have scarcely been
empirically investigated (for some exceptions using free
indirect discourse, see Fletcher & Monterosso, 2015;
Bray, 2007). A plausible assumption would be that
thought presentation (in direct or indirect reference)
would be more likely to engage ToM resources—that
est, a main effect of thoughts—compared with speech.
Why, alors, was this not seen? Insights from contempo-
rary cognitive narratology may be useful here, particularly
in relation to the problem of “accessibility” of others’
thoughts. Ordinarily, stories that are used to assess
ToM require the reader to make inferences about the
mental states of others; their actual beliefs are not made
explicit and may even conflict with the literal and imme-
diate content of what they say and how they act (par exemple., Saxe
& Kanwisher, 2003). Fictional narratives may sometimes
exploit this “accessibility gap” (par exemple., a suspect in a mystery
could have hidden motives), but they are also notable be-
cause they can give us apparent access to other minds via
direct and indirect references (Bernini, 2016; Cohn,

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1978). Although the stimuli used in our experiment con-
tained mental content, they did not necessarily make de-
mands in terms of mental state inference—in the thought
trials used in our experiment, the inner life of the charac-
ter is laid bare (par exemple., “He thought that he should go to the
shop”). Direct speech, in contrast, does not signal the in-
tonation, emotion, or intention of a character—they
must be simulated or otherwise inferred by the reader,
in a way that the ToM system is often considered to do
(Saxe & Kanwisher, 2003). En tant que tel, although counterin-
tuitive, our findings are in line with common views about
mental state inference (Spreng et al., 2009). It may also
be the case that direct speech in general is more vivid
and salient than direct reference to thinking, given the
whole-brain differences between speaking and thinking
seen in anterior insula and dorsal ACC (Uddin, 2016),
and the quicker orienting times we observed for speech
trials. Engagement with fictional storyworlds and charac-
ters is often argued to depend on the “experiential
traces” the reader brings from his or her own life
(Zwaan, 2008): The more we have access to an experi-
ence in the real world, the more it will be used to gener-
ate vivid and imaginative responses during reading. Quand
one considers the diminished, quasi-perceptual phenom-
enology that verbal thoughts are often claimed to possess
(Prinz, 2011; Jones & Fernyhough, 2007), it is perhaps no
surprise that characters’ thinking in a text did not provide
distinct patterns of activation that were as distinct as for
direct speech.

Another perspective—also provided by cognitive liter-
ary studies—is to consider how fictional minds may be
differently represented from the outside and the inside.
Kuzmičová (2013), Par exemple, has suggested that we
experience characters’ speech in literary texts as either
“outer reverberations” (when we read, as vicarious lis-
teners, about a character overtly speaking) and “inner re-
verberations” (when we voice a character’s words within
his or her perspective). In parallel, Caracciolo (2014) a
highlighted the contrast between attributing intentions
to characters and the direct, inner enactment of a charac-
ter’s thoughts and fictional consciousnesses more broadly.
These distinctions parallel the extensive literature on
perspective-taking and how this is instantiated in the
brain (par exemple., Ruby & Decety, 2001). It could be the case
that our different conditions prompted readers to adopt
d'abord- or third-person perspectives in response to speech
compared with thoughts or direct compared with indirect
reference. Cependant, the direction of these shifts is not
straightforward: Although it is sometimes assumed that
direct speech necessarily prompts adopting a first-person
perspective (speaking as the character), it is also under-
stood as focusing the reader on what it would be like to
hear the character speak to them (Clark & Gerrig, 1990).
De la même manière, thoughts could be seen to prime a first-person
perspective (thinking “from the inside”), but this will likely
depend on the position of the narrator, the reader’s iden-
tification with the character, and the wider context of the

narrative (Kuiken, Miall, & Sikora, 2004). En tant que tel, a key
area for further exploration is to systematically examine
how perspective shifts potentially interact with direct ref-
erence effects and speech/thought distinctions.

This study has a number of limitations. D'abord, it was
necessary to exclude some participants because of partial
data from eye-tracking or either of the independent loca-
lizer tasks, limiting the overall sample size. This also fur-
ther skewed our sex ratio, such that male participants are
underrepresented in our eventual sample (as can often
be the case for psychology studies recruited from univer-
sity populations, par exemple., Dickinson, Adelson, & Owen,
2012). Given the wide variability in individual differences
for reading responses, we chose to deploy these mea-
sures to be as specific as possible about both participants’
onset and offset times of reading target sentences and to
allow for the use of individually specific cortical networks.
This did not prohibit the recruitment of a larger sample
than the original study we sought to replicate ( Yao et al.,
2011), but for a topic (imagery) with typically small ef-
fects and potentially large variation, replication in larger
samples will be required for the exploration of individual
differences in imagery production across different kinds
of readers. Inner speech and imagery are highly suscep-
tible to individual differences in day-to-day use (Alderson-
Day et al., 2016), and effects of expertise (Borst, Niven, &
Logie, 2011) and variation across readers seem highly
likely.

Deuxième, our use of direct reference for thoughts (tel
as he thought “I should have finished this paper by
now”) could be questioned in terms of its relative famil-
iarity for readers. One of our aims for the study was to
use a stimulus that could act as a typographical and gram-
matical control comparison for direct and indirect
speech. Although use of quotation marks for thoughts
does feature in narratives, indirect references might be
thought of as many authors’ default option when refer-
ring to characters’ mental states (Leech & Short, 2007).
An alternative form of reference—such as using italics
to mark characters’ thoughts—may have been more fa-
miliar to readers but would also have added further typo-
graphical differences to the original contrast of interest:
direct versus indirect speech. The lack of any behavioral
differences (in terms of accuracy or reading time) être-
tween the thought conditions, and the lack of any pair-
wise or whole-brain differences, would suggest that this
had little effect on our participants. Cependant, further
careful behavioral (et, arguably, interdisciplinary)
work—incorporating the valuable insights of cognitive lit-
erary studies—is clearly required to elucidate how
readers interpret these kinds of text constructions when
depicting characters’ mental states.

Enfin, a related point about generalizability concerns
the fictional stories used in the experiment. For exper-
imental use, we used very minimal stories that were un-
likely to prompt extensive use of many of the processes
thought to be relevant to a reader’s experience of a

Alderson-Day et al.

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text, whether that involves identification with characters,
use of prior knowledge, management of expectations,
or feelings of transportation (Miall, 2011; Vert, 2004;
Kuiken et al., 2004). En tant que tel, this is still a very artificial
reading scenario for many participants. We cannot rule
out the possibility that there was something about this
situation in particular that may have posed unusual de-
mands or biased readers’ responses, such as encourag-
ing them to pay attention to or engage more with
specific aspects of the text (such as voices in particular).
Our attentional checks would adjudicate against this
interpretation—no significant differences in accuracy
were observed across the various task conditions—but,
in considering ecological validity, the experiential gap be-
tween full stories and these experimental sketches must
be borne in mind.

Notwithstanding these limitations, our findings have
important implications for future research on fiction,
reading, and imagination more generally. Our data broadly
support social cognitive approaches to fiction (Oatley,
2016; Tamir et al., 2016), but in a complex and unexpected
chemin. D'une part, the potential involvement of ToM
in simulating episodes of characters’ speech opens a new
avenue for research on fiction and mentalizing; on the
other hand, our findings for representing characters’
thoughts challenge the idea that engaging with the mental
states of others via fiction necessarily involves (or could
even enhance) ToM processes. Our findings also highlight
how readers likely draw on multiple perceptuomotor re-
sources to support a socially informed simulation of
speech, where prompted by the text. This is, arguably,
a creative and constructive process on the part of the
lecteur, which will be contingent on their own imaginative
skills and experience. Along with comparing individual
differences in this process, contrasting forms of reference
for speech and verbal thoughts offer a comparative meth-
odology for exploring how readers track speaking and
thinking through more complicated narratives. Free indi-
rect discourse, as seen in many modernist texts, demands
that the reader follow closely, or even make their own
inference, about exactly who is speaking or thinking in
a story (see Waugh, 2011, for a discussion of this topic).
Ici, reference or its absence could be considered as an
experimental tool to challenge the readers and place
them in situations of uncertainty about the speech and
thoughts in a narrative (as in Fletcher & Monterosso,
2015). A cet égard, more challenging texts offer an op-
portunity to push at the limits of readers’ creative and
imaginative capacities.

Conclusions

In conclusion, references to direct speech in fictional
stories are associated with the recruitment of not only
voice-selective auditory cortex but also regions that may
implicate gauging of characters’ communicative inten-
tion. De plus, this is a process that is apparently

specific to speech. We cannot conclude on the basis of
these findings that the function of this process is commu-
nicative inference per se, but we speculate that it goes
beyond a purely perceptual simulation of voice and re-
quires coordination between inner speech and ToM re-
sources. To experience a character’s voice in a story, dans
this sense, may be about not only what they say but also
how they say it and what they intend.

Remerciements

This work was supported by the Wellcome Trust ( WT098455
and WT108720). John Foxwell, Lucy May, and Anthony
Atkinson are thanked for their assistance with piloting and
eye-tracking; and David Smailes and the Hearing the Voice team
are thanked for their contributions to the early development of
the research question. For more information on this process,
please see Fernyhough (2015).

Reprint requests should be sent to Ben Alderson-Day,
Département de psychologie, Durham University, Science
Laboratories, South Road, Durham DH1 3LE, ROYAUME-UNI, ou par e-mail:
benjamin.alderson-day@durham.ac.uk.

Remarques

1. Exposure to literary fiction in particular has also been pro-
posed to enhance readers’ ToM skills (Oatley, 2016; Kidd &
Castano, 2013), although such claims have not always been sup-
ported in replication attempts (Kidd & Castano, 2018; Klein
et coll., 2018). For recent meta-analyses on this topic, voir
Dodell-Feder and Tamir (2018) and Mumper and Gerrig (2017).
2. A full list of the stories and questions used is available at
community.dur.ac.uk/benjamin.alderson-day/RVT_full_stim_
aldersonday.pdf.
3. This analysis was also run with ROIs that explicitly excluded
areas identified in the auditory localizer task, leading to almost
identical results: no main effects and a significant interaction,
F(1, 15) = 9.29, p = .008, ηp
2 = .38. On an individual level,
auditory and ToM ROIs overlapped in only 2 de 16 participants,
and this was to a minimal degree.
4. Although Yao et al.’s (2011) main analysis was ROI driven,
they also conducted an exploratory whole-brain analysis that
primarily identified regions of posterior temporal cortex and
occipital-fusiform regions associated with word reading.

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Alderson-Day et al.

1653Processing Speech and Thoughts during Silent Reading: image
Processing Speech and Thoughts during Silent Reading: image
Processing Speech and Thoughts during Silent Reading: image
Processing Speech and Thoughts during Silent Reading: image
Processing Speech and Thoughts during Silent Reading: image
Processing Speech and Thoughts during Silent Reading: image

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