Neuropsychologia 171 (2022) 108240

Contents lists available at ScienceDirect

Neuropsychologia
journal homepage: www.elsevier.com/locate/neuropsychologia

Neural differences in social and figurative language processing on the

autism spectrum
William W. Graves a, *, Hillary Levinson a, Linsah Coulanges b, Shannon Cahalan a, Daniel Cruz c,
Catherine Sancimino d, Vanessa H. Bal e, Miriam Rosenberg-Lee a
a
Department of Psychology, Rutgers University – Newark, Newark, NJ, USA
b
Department of Psychology, University of Pittsburgh, Pittsburgh, PA, USA
c
Psychology, Hackensack Meridian Health – Mountainside Medical Center, Montclair, NJ, USA
d
Department of Pediatric Neuroscience, Albert Einstein College of Medicine, Bronx, NY, USA
e
Graduate School of Applied and Professional Psychology, Rutgers University – New Brunswick, Piscataway, NJ, USA

A R T I C L E I N F O A B S T R A C T

Keywords: Individuals on the autism spectrum often have trouble with social and figurative language. As social language is
MeSH terms) often figurative, it can be challenging to disentangle the cognitive and neural sources of these difficulties. Neural
Humans systems for social cognition and language comprehension overlap in areas involved in retrieving linguistic
Language
meaning (semantics), such as the anterior temporal lobe (ATL), ventro-medial prefrontal cortex (vmPFC), pos­
Autism spectrum disorder
Brain
terior cingulate cortex (PCC), and posterior middle temporal gyrus (pMTG). Using adjective-noun phrases, we
Magnetic resonance imaging manipulated social/nonsocial and figurative/literal dimensions, which we expected to activate distinct but
Semantics overlapping regions. We hypothesized that activation differences in the group with autism (AUT) would be
Social cognition greater for more social and figurative stimuli. During fMRI, participants in the AUT group (N = 19) and those in
the non-autistic comparison (NAC) group (N = 22) made familiarity judgments to 192 phrases in a balanced 2 ×
2 (social/nonsocial x figurative/literal) design. Social phrases activated the PCC in all participants, but only the
NAC group activated the vmPFC. Figurative phrases were rated as more literal by the AUT group, with the
figurative-literal phrase contrast showing no activation in the AUT group, but activating the PCC and right pMTG
in the NAC group. The one significant group-level neural difference was for the social-figurative condition
predicted to be most different between groups: greater activation for the AUT group in the right ATL. Differences
in the right ATL and pMTG in the AUT group suggest altered engagement of right homologues of the canonical
semantic network being recruited for processing combined social and figurative language.

1. Introduction
Individuals on the autism spectrum, including those without a his­
tory of language delays, have been found to exhibit difficulties com­
prehending some kinds of abstract language such as metaphor (Vicente
and Falkum, 2021; Vulchanova et al., 2015). Whether these difficulties
arise primarily from a linguistic source, or relate to other core features of
autism, such as difficulties with social cognition, has been challenging to
pinpoint. This challenge arises from the fact that abstract language and
metaphor often contain social content. Like difficulties with social
processing, abstract language comprehension difficulties in autism have
been found both in terms of behavioral performance (Kalandadze et al.,
2018; Vulchanova et al., 2015) and differences in neural systems (Groen
et al., 2008; Phan et al., 2021). Abstract language contains both social
content and varying degrees of figurative or literal content (Borghi et al.,
2017). Notably, the neural systems found to modulate with different
degrees of social content are largely the same as those that respond to
different degrees of semantic content during language comprehension
(Smallwood et al., 2021). In this study we sought to better understand
the neural underpinnings of language comprehension differences in
autism by determining whether there are distinct neuro-cognitive
mechanisms behind social and figurative language processing.
1.1. Social language processing
While there are wide-ranging individual differences, in general,

* Corresponding author.
E-mail address: william.graves@rutgers.edu (W.W. Graves).

https://doi.org/10.1016/j.neuropsychologia.2022.108240
Received 2 March 2022; Received in revised form 13 April 2022; Accepted 18 April 2022
Available online 21 April 2022
0028-3932/© 2022 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-
nc-nd/4.0/).
W.W. Graves et al.
individuals with autism are often considered to have challenges in
comprehending social content, including social language (Groen et al.,
2008; Kalandadze et al., 2018; Levy and Perry, 2011; Tager-Flusberg
et al., 2005). Those without a history of measurable adverse effects on
language development or verbal intelligence can have quite strong
general language and reading skills. Yet these same individuals may
stumble when it comes to socially focused language such as irony,
sarcasm, or pragmatics that depend on inferring the mental states of
others (Landa and Goldberg, 2005; Williams et al., 2015).
The neural underpinnings of challenges with social language in
autism point to differences largely within the same midline and lateral
temporo-parietal areas that are typically activated for processing social
language. This network of areas includes anterior midline structures
such as the medial prefrontal cortex (mPFC), posterior midline struc­
tures such as the posterior cingulate cortex (PCC) and precuneus, the
anterior temporal lobe (ATL), and lateral structures such as the posterior
middle temporal gyrus (pMTG) and angular gyrus (AG) (Denny et al.,
2012; Molenberghs et al., 2016; Olson et al., 2007; Van Overwalle,
2009). Largely overlapping with these areas of the mentalizing network
are areas consistently found to activate for language comprehension
(semantic processing), including the mPFC, PCC, ATL, pMTG, and AG
(Binder and Desai, 2011; Binder et al., 1999, 2009; Mattheiss et al.,
2018). The goals of the current study were twofold. One was to deter­
mine whether neural areas responding to social or figurative content will
differ when those factors are orthogonally manipulated. The other was
to determine if neural areas responding to manipulation of social or
figurative content will differ for participants with compared to those
without autism.
1.2. Figurative language processing
Pursued largely independently from research on the neural basis of
social comprehension differences in autism, there is also a considerable
body of empirical research on the neural basis of general language
comprehension differences in autism (Harris et al., 2006; Kana et al.,
2006; Knaus et al., 2008; Phan et al., 2021; Sahyoun et al., 2010), with
fewer studies focusing on neural differences in figurative language such
as metaphor (Chouinard et al., 2017) or puns (Kana and Wadsworth,
2012). The central research question of the current study is to determine
whether there is an interaction between processing social content and
processing figurative content. More specifically, it may be the case that,
for autistic adults, social processing is mediated by different neural re­
gions than in non-autistic adults, particularly when social information
occurs in figurative contexts. Such a finding would help clarify the na­
ture of differences in abstract language comprehension that have been
reported for individuals with autism.
In terms of predictions for figurative compared to literal content,
studies have demonstrated a great deal of variation with regard to the
neural loci identified, even at the level of the hemisphere (Brownell
et al., 1990; Eviatar and Just, 2006; Gagnon et al., 2003). One source of
this variation may relate to the different types of non-literal language
used in different studies. This possibility is reinforced by two different
large-scale meta-analyses of functional neuroimaging studies in
non-autistic participants processing figurative (non-literal) language.
One (Bohrn et al., 2012) revealed that areas of the semantic network
such as the medial and lateral parietal and lateral temporal areas are
consistently activated for processing literal language, while areas acti­
vated for different kinds of non-literal language such as metaphor, irony,
and sarcasm were more heterogenous, and in turn less consistent, across
the cortex. There were, however, a few areas activated for figurative
compared to literal language processing that included the bilateral
inferior frontal and anterior temporal cortices. The other meta-analysis
(Rapp et al., 2012) similarly combined the various types of non-literal
language and contrasted them with literal language. Their findings
were similar to Bohrn et al. in revealing bilateral (but greater on the left)
activation across studies for figurative compared to literal language in
2
Neuropsychologia 171 (2022) 108240
inferior frontal cortex. However, in their meta-analysis, the left but not
right ATL was activated for figurative language (Rapp et al., 2012).
Considered together with studies pointing to a prominent role of the ATL
in social (Olson et al., 2007; Ross and Olson, 2010) and semantic
cognition (Patterson et al., 2007; Visser et al., 2010), particularly on the
left when language is involved (Binder et al., 2009; Rice et al., 2015;
Wang et al., 2010), we can predict a divergence from activation in the
left ATL if there is divergent processing of social and figurative language
in individuals with autism.
Regarding the typical neural patterns related to semantic processing,
it is notable that these regions reside in higher-order association
cortices, rather than more primary sensory or motor cortices. By
contrast, a pattern seen in the handful of studies of semantic processing
in autism is that participants on the spectrum activate sensory-related
areas such as the lateral occipital cortex. For the same semantic pro­
cessing conditions, activation for non-autistic participants is restricted
to areas of association cortex that have been independently related to
semantics (Gaffrey et al., 2007; Harris et al., 2006).
More generally, a recent meta-analysis of studies examining the
neural basis of semantic processing in autism suggested only limited
findings to be consistent across studies (Phan et al., 2021). Whether their
paucity of results was due to a lack of relevant studies (i.e., N = 11 in the
meta-analysis), or spatial heterogeneity of functional neural organiza­
tion in autism, was not clear. Nevertheless, a finding that did consis­
tently emerge was greater activation for the autistic than non-autistic
groups in the left pMTG bordering on the posterior superior temporal
gyrus (pSTG). This region is at a nexus point between the semantic
network that includes pMTG and the language network that includes
pSTG associated with phonological (speech sound or auditory word
form) processing (Graves et al., 2008; Vigneau et al., 2006). The pSTG is
also among the areas consistently under-activated for readers with
dyslexia compared to typical readers (Richlan et al., 2009), suggesting
that the autistic group may have been relying more than the non-autistic
group on phonology as a route to semantic processing. Altogether, these
findings raise the possibility that the typical neural networks underlying
semantics are at least partly distinct from the neural substrates involved
in semantic processing for individuals with autism.
1.3. Social and figurative language processing
Social and figurative language use are generally folded into the label
of language pragmatics, yet pragmatics extends beyond the simple
adjective-noun stimuli considered here to also include wider linguistic
constructs such as sentence-level syntax, multi-sentence discourse, and
prosody (Bach, 2004; Pylkkänen and McElree, 2006; Tager-Flusberg
et al., 2005). In an effort to distinguish the neuro-cognitive effects of
social and figurative content, we use relatively simple stimuli that we
can carefully control to orthogonalize the social and figurative factors.
Indeed, disentangling the distinct contributions of correlated factors is a
well-established approach in the cognitive neuroscience of language.
For a related example, cognitive and computational models of reading
are broken down into visual (orthographic), auditory (phonological),
and concept (semantic) components (Harm and Seidenberg, 2004; Plaut
et al., 1996; Seidenberg and McClelland, 1989). Even though orthog­
raphy and phonology are highly correlated in alphabetic languages,
examining their separate mechanisms has been critically helpful in
teasing apart the corresponding neural systems involved in reading
(Graves et al., 2010; Taylor et al., 2013). Here we take a similar
approach to neural systems related to pragmatic language in autism by
carefully constructing and selecting stimuli to test for potentially distinct
neural processing of its social and figurative aspects.
In studying the possibly separate and intersecting neural effects of
varying social and figurative content, we note that there have been
relatively few studies examining neural responses to manipulating both
of these factors. One of the few related studies of which we are aware
used photographic stimuli displaying social situations in which speech
W.W. Graves et al.
bubbles (as seen, for example, in comic books) contained either ironic
statements or statements that were only sensible if interpreted idio­
matically (Wakusawa et al., 2007). These stimuli were presented in fMRI
alternating between blocks where the phrases were to be interpreted
based on the pictured situational context or interpreted literally. The
idiom condition activated the vmPFC only in the social context. The
irony condition activated the dorso-medial PFC (dmPFC) and vmPFC in
the social context but not the literal context, and the right ATL in both
contexts. This study suggests that activation in the dmPFC, vmPFC, and
right ATL may relate to interactions between degrees of social and
figurative context.
Another relevant study considered predictions for separate neural
activations to a social compared to a semantic manipulation for the same
single words or (two-word) phrases. Specifically, Lin and colleagues
crossed amount of social content with a semantics-related factor such as
degree of either imageability (i.e., subjective ratings of the degree to
which words call to mind an image) or meaning plausibility. They found
adjacent and partially overlapping activation for social and semantic
factors in regions of bilateral inferior parietal lobule and pMTG, and
activation for social content alone in bilateral ATL (Lin et al., 2018,
2020). These findings are relevant to the current study in that both use
short, carefully controlled phrases to enable manipulation of social
content separately from a potentially related linguistic factor. Critically
however, the current study, aside from using English instead of Chinese
phrases, manipulates the degree of figurativeness. In this way, we can
investigate the effects of two factors that are both relevant to compre­
hension difficulties in autism, socialness and figurativeness. The po­
tential neural differences in responses to these factors among autistic
individuals have until now been largely unexplored.
1.4. Current study
Issues with pragmatic language that involve aspects of higher-level,
inferential meaning are typically studied using longer stimuli, as in
passage comprehension. However, use of such stimuli in the functional
magnetic resonance imaging environment can be challenging for a
number of reasons. For example, small differences in perceptual span of
stimuli, or uncontrolled differences in form or meaning at the individual
word, sentence, or discourse level can lead to unexpected neural effects
and difficulties interpreting contrasts between conditions (Vigliocco
et al., 2012), as our own pilot investigations also revealed. The challenge
then is to elicit social and figurative language processing in a way that
also allows for exerting experimental control over the factors being
investigated.
We address this challenge by using two-word, adjective-nouns
phrases. Studies using short phrases and even single words have been
shown to elicit differences in semantic processing between groups of
individuals with and without autism (Kamio et al., 2007; Norbury and
Nation, 2011). Specifically, we use two-word (adjective-noun) phrases
rather than single-word or discourse-level stimuli. We chose this
approach because single words are unlikely to provide sufficient context
to enable separate manipulation of both social and figurative content.
Conversely, discourse-level stimuli afford less control, and the typical
length of these stimuli would make it difficult to include sufficient
numbers of stimuli during fMRI to be sensitive to the relatively subtle
manipulations used here. Moreover, making judgments to two-word
phrases is a well-established task we have used in previous studies for
probing the neural basis of combinatorial semantic processing, or con­
ceptual combination (Graves et al., 2010; Graves et al., 2013). Within
the more extensive semantic network identified in previous studies
(Binder et al., 2009), semantic processing of two-word phrases has been
particularly shown to activate areas of ventro-medial prefrontal cortex
(vmPFC) and anterior temporal lobe (ATL), along with both left and
right-sided posterior middle temporal and angular gyri (pMTG and AG)
(Boylan et al., 2017; Graessner et al., 2021; Graves et al., 2010;
Pylkkänen, 2019).
3
Neuropsychologia 171 (2022) 108240
We designed the current study to bring additional insight into how
social and figurative language domains are typically organized in the
brain, and how they respond in a population with known but under­
specified processing differences. Building on the theoretical and
empirical background described above, we predict anterior and poste­
rior midline and lateral temporo-parietal areas previously found to be
related to social and semantic cognition to differ in the group with
autism, particularly for areas such as the ATL and pMTG related to
combining concepts in these domains. Maximal differences are expected
when processing phrases that are both social and figurative, as these
domains are often challenging for individuals with autism.
2. Material and methods
2.1. Participants
Interested participants who self-reported to be on the autism spec­
trum (AUT group) were recruited by advertisement and word-of-mouth
contacts from communities in central and northern New Jersey. These
included disability service offices in colleges and universities. Non-
autistic comparison (NAC group) participants were recruited for the
study through advertisement in the local university community and the
psychology research participant pool. All participants provided written,
informed consent as approved by the Rutgers University Institutional
Review Board (Rutgers IRB). Participants were compensated $15 per
hour for behavior-only sessions and $30 per hour for MRI sessions.
Participants consisted of 41 individuals either with AUT (N = 19,
mean age = 21.3) or without (N = 22, mean age = 21.3). Participants in
the NAC group met the following criteria: 1) no documented history of
psychiatric or neurologic disorders, 2) right-handed, 3) English learned
as a primary language no later than five years old, 4) no metal embedded
in soft tissue or other contraindications to MRI, and 5) minimum full-
scale IQ score of 85 on the Wechsler Abbreviated Scale of Intelligence
– II, as estimated by the vocabulary and matrix reasoning sub-tests
(FSIQ-2 from the WASI-II; Wechsler, 2011).
Participants in the AUT group met the same study criteria as the NAC
group, except that they had a primary diagnosis of autism spectrum
disorder. Diagnostic status was confirmed with objective psychometric
instruments and evaluation, including the Autism Diagnostic Observa­
tion Schedule – Second Edition (ADOS-2) Module 4 (Hus and Lord,
2014), administered either by a licensed clinical psychologist who was
research-reliable or a doctoral student supervised by such a clinician.
To verify that the groups were matched on the relevant factors
(summarized in Table 1), we performed t-tests of mean differences be­
tween groups. Results did not reveal any reliable differences (all p > 0.1)
between the groups with regard to age (t (39) = 0.72) or FSIQ2 (t (39) =
1.11). The verbal comprehension index (VCI) and perceptual reasoning
index (PRI) were calculated for all participants but one in the AUT group
who only completed the vocabulary and matrix reasoning subtests. The
VCI (t (38) = 1.65) and PRI (t (38) = 1.32) also did not significantly
differ between groups (p > 0.1). Gender ratios were also found to not be
reliably different between groups by χ2 testing (χ2 = 0.40, p > 0.1).
Table 1
Characteristics of participants in the two study groups. Numbers in parentheses
are standard deviations. FSIQ2 was available for all participants and is based on
the vocabulary and matrix reasoning subtests of the WASI. VIQ and PIQ were
available for all NAC participants and 18 participants in the AUT group. Groups
did not reliably differ on any of these measures (ps > 0.1).
AUT NAC
N 19 22
Gender 15 M, 4 F 19 M, 3 F
Age 20.8 (1.9) 21.3 (2.3)
FSIQ2 114.2 (12.8) 110.2 (10.4)
VIQ 109.8 (12.4) 104.5 (8.2)
PIQ 111.3 (13.0) 106.2 (11.2)
W.W. Graves et al. Neuropsychologia 171 (2022) 108240

2.2. Experiment stimuli and design or concreteness (for matching phrases on a general semantic factor, as
described below) using a 1–7 Likert-type scale. Examples of phrases not
The stimuli consisted of two-word, adjective-noun phrases. Each included for rating were given with anchor points illustrating that
phrase was rated as high or low on socialness and high and low on higher numbers corresponded to greater levels figurativeness, social­
figurativeness, and selected to create a fully crossed 2 × 2 design. Social ness, or concreteness. Data integrity was checked by comparing each
phrases were defined as using language that “relates to interactions of participant’s ratings to the mean rating from the remaining participants
people, groups of people, or society as well as reflections on interactions for the same items. Participants whose ratings were more than 2.5 SD
with other people.” Figurative phrases were defined as “departing from from the mean were excluded. This resulted in each phrase being rated
the usual or most basic sense of the words.” There were 48 phrases per by at least 27 participants.
cell (social figurative, social literal, nonsocial figurative, nonsocial Using these stimulus characteristics and ratings, we selected phrases
literal) for a total of 192 phrases. Characteristics of these phrases, that clearly and reliably differed in figurativeness and socialness across
including examples and relevant factors on which they are matched, are the four conditions. These conditions were also statistically matched for
shown in Table 2. surface characteristics (number of letters) along with factors related to
To generate these stimuli, we first conducted a norming study. Using familiarity (frequency of adjective, noun, and whole-phrase occurrence)
the SUBTLEX-US database, we considered words with concreteness and semantics (concreteness). Selection of phrase stimuli to meet these
ratings (Brysbaert et al., 2014) and whose dominant part of speech was criteria was performed using the Stochastic Optimization of Stimuli
either adjective or noun. We then selected the 100 most frequent ad­ software (Armstrong et al., 2012). Characteristics for the final set of 192
jectives and 400 most frequent nouns, and paired them in order to derive phrases are summarized in Table 2. Phrases that were bimodally
40,000 possible pairs. Initial winnowing of this list was performed by distributed to reliably differ on socialness (t (190) = 34.1, p < 0.001) did
cross-referencing with the Google bigrams database (Brants and Franz, not reliably differ in figurativeness (t (190) = 0.7, p > 0.5), and phrases
2006). Here bigrams refers to two-word combinations, from which we bimodally distributed for figurativeness (t (190) = 36.9, p < 0.001) did
selected only those from our list that appeared more than 1000 times on not reliably differ in socialness (t (190) = 0.6, p > 0.5). The other factors
the Internet by the year 2006. This resulted in 5000 pairs, which were being controlled for (Table 2, below the double line) did not reliably
manually inspected for likelihood of cleanly fitting into one of the cells differ across conditions (all ps > 0.1). The full set of stimuli and their
in our 2 × 2 design. Use of this database-driven approach made clear relevant characteristics are provided as a comma-separated-values file in
that meaningful figurative phrases are unlikely to emerge by this supplementary material.
method. We therefore supplemented figurative phrases by consulting
the free online Farlex Idioms and Slang Dictionary (2017). This yielded a 2.3. Image acquisition
set of 1084 candidate phrases that were submitted for independent
rating as described next. All brain scans were acquired at the Rutgers University Brain Im­
To select the final set of phrase stimuli for the fMRI experiment, we aging Center (Newark, NJ) using a 3 T S TIM Trio with a 12-channel
obtained ratings from a separate group of participants recruited using head coil. Following acquisition of initial localizer slices in each of the
Amazon Mechanical Turk. To ensure high-quality ratings, we restricted axial, sagittal, and coronal planes, a high-resolution 3D anatomical
participants to those in the United States who had a record of completing (MPRAGE) scan was acquired using the following parameters: TR =
10,000 other assignments with a 99% acceptance rate from other re­ 1900 ms, TE = 2.52 ms, flip angle 9◦ , image matrix = 256 × 256, field of
questors. Potential raters were told to accept the assignment only if they view = 256 mm, number of slices = 176, slice thickness = 1 mm, voxel
considered English their native or first language. Raters then completed size = 1 mm isotropic. Next fMRI data were acquired as T2*-weighted
a short demographic questionnaire, and anyone reporting an age below gradient-echo single-shot echo planar images (EPI) with the following
18 years or a language other than English as their first/native language parameters: TR = 2000 ms, TE = 25 ms, flip angle = 77◦ , image matrix
was excluded from providing ratings. Participants gave written = 64 × 64, field of view = 195 mm, number of slices = 35, slice
informed consent in accordance with the Rutgers IRB and were thickness = 3.0 mm with interleaved acquisition, voxel size = 3 mm
compensated $4.00 per half hour. Each participant was given a list of isotropic. These were acquired during four runs of 3 min 28 s each,
360 unique phrases and asked to rate them on figurativeness, socialness, during which participants performed the task described next. Stimuli
were viewed through a prism mirror mounted to the head coil, and
finger-press responses were recorded using an MRI-compatible button
Table 2
box.
Rows show each factor either being manipulated (social and figurative, above
the double line, with examples in parentheses) or being controlled (remaining
rows below the double line). Columns show each cell in the completely crossed 2.4. In-scanner task
2 × 2 factorial design. Cells contain means (with standard deviations) for each
value. The values for social, figurative, and concrete come from subjective rat­ As the experiment was meant to probe neural responses to orthog­
ings and are therefore unitless. Frequencies are log-transformed number of onally manipulated social and figurative content of phrases, we chose
occurrences. the familiarity judgment task as a way of prompting participants to
Social Nonsocial attend to the meaning of the phrases without directing them to focus
specifically on the social or figurative aspects of their meaning. We also
Figurative Literal Figurative Literal
(blue blood) (gentle (sharp taste) (sore assumed the groups would have roughly equal access to their own fa­
hug) throat) miliarity judgments, thereby eliciting phrase comprehension while
Social 4.95 (0.66) 4.96 (0.86) 2.15 (0.28) 1.90 (0.39)
minimizing the involvement of potentially domain-general performance
Figurative 5.73 (0.46) 2.75 (0.58) 5.62 (0.41) 2.54 (0.57) differences such as attention or time-on-task.
Concrete 3.98 (1.74) 4.41 (1.54) 3.87 (1.43) 4.06 (1.43) Participants were instructed to press a button on a handheld MRI
Number of 9.77 (2.41) 10.77 9.98 (2.27) 10.04 compatible two-button box to indicate whether each phrase they were
letters (2.73) (2.32)
seeing was familiar or unfamiliar. Familiar phrases were defined in the
Phrase 10.52 (1.62) 10.64 10.56 (1.81) 10.87
frequency (1.81) (1.59) instructions as ones they “use or hear often”. Following an initial 8 s of
Adjective 7.94 (1.70) 8.06 (1.69) 7.74 (2.03) 7.71 (1.17) fixation at the beginning of each run to allow EPI saturation effects to
frequency stabilize, either a fixation or a trial would appear. During trials, the
Noun 8.05 (1.50) 8.56 (1.76) 8.05 (1.67) 8.08 (1.76) stimulus (phrase) was displayed for 1.5 s, followed by a fixation cross for
frequency
0.5 s. The screen would then return to a fixation cross for a randomly

4
W.W. Graves et al.
jittered inter-trial interval (mean: 2.5 s). Reaction time (RT) was
recorded as the time from stimulus onset to a button press.
A potential concern about using familiarity judgment as the in-
scanner task is that it does not explicitly probe the degree to which
participants comprehended the phrases in social or figurative terms. To
address this, a subset of participants in the AUT (N = 14) and NAC (N =
15) groups rated the 192 phrases seen during scanning in terms of so­
cial/nonsocial and figurative/literal content. This outside-the-scanner
task followed the same procedure as described above for Amazon Me­
chanical Turk.
2.5. Data analysis
Analysis of behavioral data from the in-scanner task was conducted
as a 2 × 2 (within-subjects for social and figurative) x 2 (between
groups) ANOVA to test for potential group differences in rate of
“familiar” responses, with subjects as the random factor. The same
ANOVA setup was also performed with RT as the dependent variable.
Functional neuroimaging data were analyzed using AFNI software
(Cox, 1996; Cox and Hyde, 1997) within custom Linux shell scripts
(tc-shell under Ubuntu 18.04). The reconstructed data were converted
from DICOM to AFNI HEAD/BRIK format. Motion correction and
anatomical alignment to the functional data were performed using the
AFNI script, align_epi_anat.py (Saad et al., 2009). TRs with motion
displacement greater than 10% from the mean were determined using
3dToutcount and censored from further analysis. A nuisance regressor
was also created to model non-physiological signal fluctuation using the
first principal component (calculated using the AFNI program 3 dpc) of
the signal from the lateral ventricles.
To ensure that any neural differences found between groups could
not be due to differences in amount of motion during fMRI, we per­
formed motion correction and analyzed the resulting motion parameters
as part of the standard AFNI processing pipeline (Saad et al., 2009).
Specifically, six motion parameters were calculated when registering
each image volume in the fMRI time series to a central volume: Three
directions of rotation, and three directions of displacement. The tem­
poral derivative of this 6-element vector for a given time point relative
to the previous one was calculated to determine the amount of motion
from one time point to the next. The derivative vectors were then
combined into a single term by taking the Euclidean (L2) norm, yielding
one motion value per time point. The mean across time points was the
mean motion for each participant. As these values were quite small
overall (AUT: 0.08 mm, NAC: 0.09 mm), and no participant in either
group exceeded 0.22 mm of displacement from one frame to the next, we
did not apply a cutoff threshold for participant inclusion. Motion did not
significantly differ between groups (t39 = 0.74, p = 0.47).
First-level regression analysis of individual participant data was
performed using 3dDeconvolve, with seven nuisance covariates and six
explanatory variables of interest. Of the seven nuisance covariates, 6
were motion parameters, 3 for rotation in the pitch, roll, and yaw di­
rections, and 3 for displacement in those directions, along with the
physiological noise regressor from the lateral ventricles as described
above. The six explanatory variables were 4 for the cells in the 2 × 2
design: social/figurative, social/literal, nonsocial/figurative, nonsocial/
literal, along with a term modeling any null responses, and a continuous,
z-score normalized regressor for trial RT. The resulting output images
were then nonlinearly warped to Talairach space (the TT_N27 brain
space; Lancaster et al., 2000) using diffeomorphic techniques in
3dQwarp and smoothed with a 5 mm FWHM kernel using the AFNI
program 3dmerge.
Second-level analysis of the NAC and AUT groups was performed
with the AFNI program 3dttest++ to compare first-level contrasts be­
tween groups at each voxel. The “-Clustsim” option was used to imple­
ment current best-practice approaches to map-wise cluster corrections
for multiple comparisons that also take into account the smoothness of
the data (Cox et al., 2017). For all contrasts reported, voxel-level
5
Neuropsychologia 171 (2022) 108240
thresholds were set to p < 0.005, and mapwise cluster corrected to p
< 0.05.
3. Results
3.1. Behavioral
Task responses during scanning were not reliably different between
the groups. Participants in the AUT group had slightly higher rates of
judging the phrases to be familiar (81%) compared to the NAC group
(78%), but this was not a statistically significant difference (F1,39 = 1.16,
p = 0.29). The 2 × 2 × 2 ANOVAS (Fig. 1A) did reveal that during
scanning, literal phrases were judged to be familiar (F1,39 = 215.7, p <
0.001) more often (85.9% familiar) than were figurative phrases (72.3%
familiar). However, there was no main effect of group, and these ratings
did not interact with group status. Similarly, social phrases were judged
to be familiar more often (84.9%) than were nonsocial phrases (73.3%,
F1,39 = 206.9, p < 0.001). Here too, there was no main effect of group,
nor was there an interaction with group status. There was, however, an
interaction between the social and figurative factors (F1,39 = 34.3, p <
0.001), such that phrases that were both nonsocial and figurative were
judged on average to be less familiar (64% familiar) than were the other
three conditions (ranging from 81% familiar for social/figurative
phrases to 89% familiar for social/literal phrases, with nonsocial/literal
phrases falling in between at 83% familiar).
Regarding response times for making these judgments, the AUT
group averaged 990 ms (ms), while the NAC group averaged 970 ms.
These means were also not reliably different between the groups (F1,39
= 0.29, p = 0.60). Overall, the RT data followed a pattern similar to the
one described above for the rate at which phrases were judged to be
familiar. That is, participants were faster overall to make judgments to
literal phrases (961 ms) than to figurative ones (994 ms). This main
effect of figurativeness on RT was statistically significant (F1,39 = 26.3,
p < 0.001), but did not interact with group (Fig. 1B). Similarly, social
phrases were judged more quickly on average (958 ms) than were
nonsocial phrases (997 ms). This main effect of phrase socialness was
statistically reliable (F1,39 = 58.4, p < 0.001), but did not interact with
group. Unlike the pattern above for rate of familiarity, the RT for making
those ratings did not show an interaction between levels of socialness
and figurativeness.
To determine whether the groups were comprehending the phrases
differently in terms of social or figurative content, a subset (71%, AUT N
= 14, NAC N = 15) of participants who underwent scanning also
completed post-scan ratings of the phrases on Likert-type scales of
figurative/literal and social/nonsocial. There was a significant interac­
tion of figurative/literal ratings with group (Fig. 2), such that the AUT
group rated the figurative phrases as more literal than the NAC group
did (F1,29 = 9.06, p = 0.003). There were no group differences in the
social/nonsocial ratings (F1,29 = 0.01, p = 0.937).
3.2. Functional neuroimaging
We tested fMRI data for a main effect of social content by contrasting
neural responses to social minus nonsocial phrases. Both groups showed
reliable activation for this contrast. Social phrases showed activation
relative to nonsocial phrases primarily in midline structures, including
bilateral PCC, precuneus, and dmPFC (Fig. 3, Table 3). Additionally,
only the NAC group showed activation for social phrases in the vmPFC.
Although this apparent difference between groups did not reach statis­
tical significance in the whole brain analysis, a plot of the relative ac­
tivations in the vmPFC (Fig. 3C) revealed a numerically larger difference
between activation for social and nonsocial phrases for the NAC
compared to the AUT group.
We also tested for main effects in the contrast between figurative and
literal phrases. The NAC group showed activation for literal phrases
primarily in the bilateral precuneus and PCC, and right pMTG (Fig. 4A,
W.W. Graves et al.
Fig. 1. Performance on the in-scanner task. There were no differences between groups in terms of proportion of phrases judged to be familiar (A) or response times
(RT) to those make those judgments (B). Significant effects of phrase type are described in the text of the Results section. AUT: autism, NAC: non-autism comparison.
Fig. 2. Post-scan ratings in terms of figurative/literal on a Likert-type scale for
the 192 phrases seen during scanning. The ratings were obtained from a subset
of the scanned participants in the autism (AUT, N = 14) and non-autism
comparison (NAC, N = 15) groups. The AUT group rated figurative phrases
as significantly more literal than did the NAC group.
6
Neuropsychologia 171 (2022) 108240
Table 3). No brain areas showed significant activation for this contrast in
the AUT group (Fig. 4B, Table 3). To illustrate the directions of activa­
tion for the conditions making up this contrast, BOLD signal parameter
estimates are plotted separately for the figurative and literal conditions
in each group (Fig. 4C). As shown in the plot, the difference between the
figurative and literal conditions is greater for the NAC group than the
AUT group. Although this apparent group difference does not reach
statistical significance when correcting for multiple comparisons at the
whole brain level, it is notable that activation differences for figurative
compared to literal phrases appear robustly in the NAC group and not at
all in the AUT group. The full 2 × 2 × 2 ANOVA for the neural results did
not yield any significant between-group effects.
Rounding out the planned comparisons, we considered each of the
four cells in the 2 × 2 design. Reliable group differences were found for
activation in the social-figurative cell in the regions of the bilateral mid-
cingulate extending to supplementary motor area, and the right ATL
(Fig. 5A, Table 4). The ATL activation was of particular interest due to its
known role in semantics (as summarized in the Introduction). Plotting
the activation in this region for each group revealed deactivation for the
NAC relative to AUT group (Fig. 5B).
To better understand the nature of the right ATL group result, we
performed an exploratory follow-up analysis of social/nonsocial and
figurative/literal main effects in which we kept the voxel-level threshold
at p < 0.005, but relaxed the extent threshold so that it was no longer
mapwise corrected to p < 0.05. This revealed a trend toward a main
effect of socialness in which social phrases showed numerically greater
activation than non-social phrases in the left ATL for the NAC group only
(Fig. 6). No such trend was found for main effects of figurativeness in the
left ATL.
4. Discussion
The overall aim of this study was to determine the neural basis of
differences in abstract language processing among individuals with
autism, separating components related to degree of figurative or social
language content. Main effects of the figurative-literal contrast were
found in several semantics-related areas such as PCC and pMTG.
W.W. Graves et al. Neuropsychologia 171 (2022) 108240

Table 3
Peak coordinates for the main effects of the contrast between literal and figu­
rative phrases (upper rows), and the contrast between social and nonsocial
phrases (lower rows), separated by group. L = left, R = right.
Location of extreme point Cluster size X Y Z z-
(mm3) score

Social > nonsocial

NAC group
Bilateral medial prefrontal 9260
L medial superior frontal gyrus 0 51 37 4.56
L gyrus rectus 0 57 − 6 4.45
Bilateral medial parietal 6719
L posterior cingulate − 5 − 51 26 5.28
AUT group
Bilateral medial parietal 6002
L posterior cingulate 0 − 50 27 4.62
Bilateral medial prefrontal 4156
R medial superior frontal gyrus 2 59 17 4.20
Literal > figurative
NAC group
Bilateral medial parieto- 30,216
occipital
R precuneus 7 − 54 32 5.14
L cuneus − 1 − 84 31 4.34
R superior parietal lobule 20 − 64 54 4.17
L superior parietal lobule − 28 − 74 37 4.12
L occipital pole 0 − 89 − 6 4.06
Right posterior temporal and 6577
inferior parietal
R posterior middle temporal 57 − 49 5 4.87
gyrus
AUT group
None
Figurative > literal
None
R medial superior frontal gyrus 2 43 42 3.77

vmPFC, was activated in the NAC group only. Critically, the social and
figurative cell of the 2 × 2 factorial design that was expected to show the
greatest group differences indeed showed a reliable difference in group
activation in the right ATL.

4.1. Social-nonsocial contrast

As social impairments constitute a critical criterion for AUT diag­

Fig. 3. Social minus nonsocial contrast for the NAC group (A), AUT group (B),
and parameter estimates in vmPFC (C) illustrating the relative effects in the
contrast for both groups.
Strikingly, this pattern was detected for the NAC group only. This result
aligns with the findings that participants in the AUT group rated figu­
rative phrases as more literal than did the NAC group. Main effects of the
social-nonsocial contrast, however, were found for both groups in
bilateral PCC and dmPFC, consistent with previous studies of social
stimulus processing. Another region expected for this contrast, the
nosis, we expected neural differences in the processing of social
compared to non-social phrases. While there were indeed numerical
differences between the groups in results of this social-nonsocial
contrast in the vmPFC, it is also striking that both groups showed acti­
vation in areas of the PCC and dmPFC that have been found in previous
studies of social cognition (Denny et al., 2012). This pattern of results,
along with the fact that both groups endorsed similar proportions of the
social and nonsocial phrases as being familiar, suggests that group dif­
ferences due to overall social content may be minimal for these rela­
tively simple stimuli.
Of note, however, is the marginally significant group differences for
the social-nonsocial contrast in the vmPFC. While the vmPFC is
considered part of the brain network for social cognition, recent studies
have also aimed to determine whether different parts of that network
play somewhat different roles in social cognition. For example, a recent
large-scale meta-analysis of neural components of social cognition found
the vmPFC to be related to verbally mediated judgments of peoples
beliefs or traits (Schurz et al., 2021). A separate line of research with
brain lesion-deficit analyses has linked the vmPFC to valuation of
reward-related signals, potentially in relation to signals from other parts
of the body, such as signals that are detected from autonomic skin
conductance responses (Bechara et al., 1996). This interpretation is
compatible with previous lesion-deficit studies showing that damage to
the vmPFC can lead to atypical social decisions (Koenigs and Tranel,
2007; Koenigs et al., 2007). There is also a separate line of research on

7
W.W. Graves et al. Neuropsychologia 171 (2022) 108240

Fig. 5. Significant group differences in the social, figurative condition in the

right ATL (A), along with plots of size and direction of effects (B).

Table 4
Peak coordinates of the between-group activation differences for social, figu­
rative phrases.
Fig. 4. Figurative minus literal contrast for the NAC group (A), AUT group (B), Location of extreme point Cluster size X Y Z z-
and parameter estimates in right pMTG (C) illustrating the relative effects in the (mm3) score
contrast for both groups. AUT > NAC
Bilateral cingulate and 3190
aspects of AUT related to atypical awareness or perception of one’s own supplementary motor
L middle cingulate − 11 3 36 4.32
body (Schauder et al., 2015). To the extent that autonomic or other R supplementary motor area 5 − 9 50 4.05
sensory signals are integrated with ongoing cognitive processes during R anterior temporal lobe 2199
tasks such as the one used here, it is relevant to note that the vmPFC has R temporal pole 34 8 − 23 4.89
been hypothesized to be related to multisensory integration issues in
autism (Martínez-Sanchis, 2014).

8
W.W. Graves et al.
Fig. 6. Exploratory follow-up analysis showing trends in the left ATL (A) for
both groups toward a main effect of socialness (B). Although it did not survive
extent thresholding, there was a clear trend toward greater activation for social
compared to non-social phrases in the NAC group compared to the AUT group.
4.2. Figurative-literal contrast
The overall pattern of results for this contrast was that the NAC group
showed greater activation for the literal condition in the figurative
minus literal contrast in the right pMTG, extending inferiorly into the
inferior temporal gyrus and superiorly into the AG, along with bilateral
activation of the PCC. These results largely replicate those of a previous
study from one of us (WG). In that study, straightforwardly meaningful
two-word phrases (e.g., “ski jacket”) were contrasted with the reverse of
each phrase (e.g., “jacket ski”) that were independently judged to be
largely meaningless. If the literal phrases in the current study are
considered to be more straightforwardly meaningful than the figurative
phrases (e.g., “cold air” compared to “cold turkey”), then the current
study can be seen as at least partially replicating our previous finding in
which more meaningful phrases activated the right pMTG, extending to
AG, and bilateral PCC (Graves et al., 2010). Similar findings have been
reported for conventional (easily interpretable) compared to novel (less
interpretable) metaphors in German noun-noun phrases that also acti­
vated the bilateral PCC, right pMTG and AG (Forgács et al., 2012).
While the above pattern of straightforward, meaningful two-word
phrases activating the right pMTG and bilateral PCC may be repli­
cable, other studies report the left ATL to be prominently involved in
combining verbal concepts (Baron and Osherson, 2011; Baron et al.,
2010; Bemis and Pylkkänen, 2011; Li and Pylkkänen, 2021; Pylkkänen,
2019; Westerlund and Pylkkänen, 2014). Studies have also found both
the left ATL and bilateral AG to be modulated by conceptual combina­
tion (Boylan et al., 2017), while still others have reported activation for
meaningful two-word conceptual combination in the AG bilaterally but
not ATL (Graessner et al., 2021; Lin et al., 2020). These varied findings
leave open the question of the relative contributions of the ATL and AG
to conceptual combination in minimal phrase units such as those used
here.
9
Neuropsychologia 171 (2022) 108240
To distinguish the roles of the ATL and AG in conceptual combina­
tion, one theory emphasizes the distinction between taxonomic and
thematic relations (Mirman et al., 2017). Taxonomic relations share
category-related features (e.g., dog—bear, or coffee—tea), whereas
thematic relations co-occur in similar events, contexts, or scenarios (e.g.,
dog—leash, coffee—spoon). Reviews have summarized evidence
pointing to the ATL as more associated with taxonomic relations, and the
AG as more associated with thematic relations (Davis and Yee, 2019;
Mirman et al., 2017). For example, survivors of stable, focal lesions to
the left ATL exhibit deficits primarily in processing taxonomically
related items, while participants with lesions to the left AG have deficits
in processing thematically related items (Schwartz et al., 2011). In terms
of the current study, stimuli consisted of adjective-noun phrases that
were not selected in terms of taxonomic or thematic relations. Of the
two, however, adjective-noun phrases would seem to be more analogous
to thematic relations, as the constituent words combine to form an
overall concept where the first word is descriptive and has no require­
ment to be taxonomically related to the second. Therefore, it may be that
the right AG activation found here relates to processing phrases con­
taining especially clear, meaningful descriptions.
Of particular importance to the aims of the current study is the lack
of any significant differences between figurative and literal phrases in
the AUT group. While the magnitude of this contrast did not formally
survive whole-brain correction for multiple comparisons, the fact that
the NAC group activated numerous expected areas for this contrast and
the AUT group significantly activated none is nonetheless striking.
While we are not aware of functional neuroimaging studies specifically
examining responses in participants with AUT to figurative compared to
literal phrases, two previous studies are relevant. One fMRI study con­
trasted neural responses in AUT and NAC participants to metaphor
processing. They found greater activation in the AUT group in visual
association cortex, thalamus, and cerebellum (Chouinard et al., 2017). A
second fMRI study compared neural responses to verbal humor (puns)
between AUT and NAC groups, finding greater activation in the AUT
group for puns in the bilateral inferior parietal lobule, right hippocam­
pus and middle frontal gyrus (Kana and Wadsworth, 2012). Although
these studies did find activation for their groups of AUT participants in
their respective non-literal language conditions, they were in entirely
non-overlapping areas, making it difficult to draw specific
between-study conclusions about what brain areas to expect to be
activated. A salient potential source of this variability could be the de­
gree of matching between groups. Participants in the AUT group from
Kana and Wadsworth (2012) had significantly lower verbal IQ overall
compared to the NAC group (103.7 compared to 114.5), while Choui­
nard et al. (2017) matched the groups in their study for non-verbal IQ
but did not report verbal IQ. We suggest that careful matching of groups
for both verbal and non-verbal IQ, as in the current study, may help
increase consistency of findings by reducing uncontrolled variability.
Relative to neural activation studies, figurative language processing
in AUT has been studied more extensively in behavioral studies. Such
studies were compiled in a recent meta-analysis, which concluded that
figurative language processing is indeed atypical in AUT (Kalandadze
et al., 2018). Relatedly, participants with AUT have been shown to have
difficulty calling to mind the multiple meanings of homographs as a
function of context (e.g., “In her eye/dress, there was a big tear.”; Happé,
1997). While such issues with lexical ambiguity are not always found in
AUT groups (Norbury, 2005), focusing on a single meaning, rather than
modulating meaning activation by context, is broadly consistent with
the “weak coherence” theory of cognition in autism (Happé and Frith,
2006). This theory would presumably predict retrieval of a single,
usually literal, meaning regardless of small differences in linguistic
context. In the current adjective-noun phrase task, this may have led the
AUT group to treat literal and figurative phrases in much the same way,
thereby attenuating the neural differences between them. Indeed, rat­
ings from a large subset of participants in the current study showed that
the AUT group interpreted the figurative phrases as more being literal
W.W. Graves et al.
than did the NAC group.
Before turning to the between-groups result, we note that the within-
group results for the social-nonsocial contrast and the NAC result for the
figurative-literal contrast appear to partially overlap the putative default
mode network, particularly in the PCC. The default mode network is so
called because it is more active at rest than during task performance.
Whether the function of that network relates to particular cognitive
processes or a metabolically determined “baseline activation” is highly
controversial (Buckner et al., 2008; Raichle, 2015; Smallwood et al.,
2021). As there is no true baseline in fMRI (Gusnard and Raichle, 2001),
studies often report inter-trial intervals or blocks of no task as an implicit
“resting” baseline. While we have followed that convention in the bar
graphs to illustrate the relative pattern of activation, the activation maps
themselves are based on direct contrasts between conditions. For
example, Fig. 3 shows activation greater for social than nonsocial, and
Fig. 4 shows activation greater for literal than figurative. Which side of
these contrasts should a priori be considered baseline or default is not
obvious. Instead, we have focused on the separate but related question
of what kinds of cognitive processes are typically associated with acti­
vations in this set of brain areas. There is a large body of evidence
showing that these areas process semantics (Binder et al., 2009; Lanzoni
et al., 2020; Mattheiss et al., 2018; Murphy et al., 2018; Wirth et al.,
2011), and our stimuli were designed to modulate processing within the
semantic system. Viewed this way, the activation patterns shown in
Figs. 3 and 4 comport with the interpretation that the stimuli and
experimental design did indeed succeed in modulating the target neural
and cognitive systems.
4.3. Social and figurative phrases
Greater activation in the AUT group for phrases that were both social
and figurative was found in a cluster spanning the mid-cingulate cortex
and supplementary motor area (SMA), along with a separate cluster in
the right ATL. The mid-cingulate and SMA are functionally associated
with a larger network of fronto-parietal regions found to activate across
domains for tasks requiring attention to external stimuli relative to rest
(Duncan, 2010; Fedorenko et al., 2013). While such tasks have been
found to include those relevant to social cognition, such as making social
preference decisions (Chen et al., 2010), the broadly domain-general
nature of responses found in these regions makes it unlikely to be spe­
cific to the social and figurative stimuli considered here. Additionally,
the proportion of phrases judged to be familiar, and the RT for those
responses, were not reliably different between groups. That plus the fact
that trial-level RT was included in the fMRI analyses suggests that the
results do not simply reflect domain-general effects such as time-on-task,
executive functioning, or working memory demands.
The social and figurative condition was the cell of our 2 × 2 factorial
design predicted to differ most between the groups. By placing demands
simultaneously on social and linguistic comprehension, we expected
brain response in this condition to reveal areas where the AUT group
was processing these stimuli differently from the NAC group, presum­
ably reflecting a difference in social and semantic networks. Greater
activation in the AUT group for the right rather than left ATL is
consistent with this expectation. Meta-analyses of semantic processing
have found the left more than right ATL to be typically activated (Visser
et al., 2010; Wang et al., 2010), particularly for tasks involving language
stimuli (Binder et al., 2009; Rice et al., 2015). The left/right distinction
in these meta-analyses is relative, however, as activation in both ATLs
was found, but with greater activation in left ATL (Binder et al., 2009;
Rice et al., 2015; Visser et al., 2010). Only one of these meta-analyses
separately analyzed neural responses to social stimuli. They found
bilateral ATL activation across studies with stimuli aimed at social
concepts (Rice et al., 2015). Therefore, activation of the right ATL,
rather than left or bilateral, by the AUT group for social and figurative
phrases suggests that this group recruited key areas of the semantic
processing network in a relatively atypical fashion.
10
Neuropsychologia 171 (2022) 108240
Further support for the interpretation that greater activation of the
right ATL for social-figurative phrases in the AUT group represents
atypical recruitment of semantics comes from an exploratory follow-up
analysis. By keeping the voxel-level threshold constant but relaxing the
spatial extent threshold, we identified a set of voxels in the left ATL
(Fig. 6A) that showed a trend toward greater activation for social
compared to non-social phrases in the NAC group only (Fig. 6B). This is
consistent with previous studies showing neurotypical left ATL
involvement in social semantics (Lin et al., 2018, 2020; Simmons et al.,
2010). While this follow-up analysis was strictly exploratory, it lends
qualitative support to the interpretation that the AUT group atypically
recruited ATL nodes of the semantic network. The novel finding from the
current study is that processing for social concepts in the ATL shifts
rightward in the AUT group, particularly when those concepts also
contain figurative content.
Another, non-exclusive interpretation of the greater activation of the
right ATL by the AUT group for the social/figurative phrases relates to
the putative role of the ATL in processing taxonomic, rather than the­
matic, relations. While the set of phrase stimuli used here were not
selected in terms of taxonomic or thematic relations, a preference for
taxonomic interpretations, whether implicit or explicit, could poten­
tially explain the greater recruitment of the ATL by the AUT group.
Indeed, directly testing this possibility could be a fruitful direction for
future study, as we are aware of no existing studies to specifically
compare neural responses to taxonomic and thematic processing across
AUT and NAC groups.
4.4. Potential limitations
There is an inherent trade-off between careful sample selection and
generalization to wider populations. In this the current study is no
exception. Testing a sample where most participants were un­
dergraduates helped to ensure that participants would be generally
familiar with the kinds of tasks they would be asked to perform for the
study, as well as more generally being able to schedule and keep ap­
pointments, and tolerate the MRI environment for an extended time.
While these abilities were necessary for study completion, our choice of
participants also means that the results may not generalize to samples
who could not perform these tasks. Conversely, most participants in both
groups tested here seem to be doing well in college overall, at least in
terms of mainlining successful enrollment. This raises the possibility that
the greater activation seen in the AUT group in the right ATL for the
social/figurative condition could relate to the process of managing to
deal with a world that is not particularly attuned to neuro-diversity.
A second concern that arises with many studies involving special
populations is sample size. While it is true that more participants is
generally better, note that the current group of participants with autism
(N = 19) and the overall sample size (N = 41) is larger than the two most
relevant functional neuroimaging studies of non-literal language in AUT
that we were able to identify in the literature, with Ns of 16 (Kana and
Wadsworth, 2012) and 12 (Chouinard et al., 2017) in their AUT groups.
To address the related concern of potential false-positive results, we took
care to use current best practices to minimize potential false-positives in
our whole-brain analyses (Cox et al., 2017; Eklund et al., 2016). This
may have come with the trade-off of limiting sensitivity to the poten­
tially subtle differences that might be expected between groups that are
as well matched as those in the current study. Ultimately, however, we
acknowledge that this relatively limited sample size warrants caution in
interpreting how generalizable the findings may be to new samples.
Additionally, we choose to use adjective-noun pairs as stimuli (full
set provided as supplementary material) instead of more linguistically
complex sentences or even discourse-level materials. In doing so, we
emphasized experimental control at the necessary expense of ecological
richness. It is therefore unknown at this point whether the patterns seen
in the current results would obtain with more complex or less tightly
controlled stimuli.
W.W. Graves et al.
Finally, while we made every effort to ensure that levels of social and
figurative content were orthogonally manipulated in our stimuli, this
does not change the fact that these types of content typically co-occur in
everyday language. Such co-occurrences may lead participants to make
automatic associations between social and figurative content. Viewed
another way, however, these possible automatic associations can actu­
ally be seen as a potential source of the observed effects. That is, stan­
dard conceptual networks activated by social or figurative concepts are
expected to spread activation to other related social or figurative con­
cepts, as in classically conceived spreading activation networks
(reviewed by Ober and Shenaut, 2006). However, in the current case the
AUT group was interpreting the figurative phrases more literally, and
neurally activating areas not typically engaged in verbal semantic pro­
cessing. Viewing this as resulting from a difference in the spread of
activation among nodes of a semantic network organized through a
lifetime of associations that may be subtly different in the AUT
compared to NAC groups may be a helpful way to interpret the source of
the differences reported here.
4.5. Conclusion
This study addressed the question of whether the apparent diffi­
culties individuals with AUT can have in processing pragmatic aspects of
language, such as social and figurative content, is related to differences
in activation of brain areas related to processing social and figurative
content. Participants with AUT showed neural activations broadly
similar to the NAC group when processing phrases in terms of overall
social or figurative content. There were, however, clear trends toward
differences in key areas of the social cognition network, where only the
NAC participants activated vmPFC for more social phrases, and only the
NAC group showed differences in activation between figurative and
literal phrases. Critically, there was a significant difference in the right
ATL between the groups for phrases that were expected to be processed
with maximal difference between groups – those that were both social
and figurative. We suggest that the partially distinct sets of brain areas
shown here to respond to social and figurative content are atypically
organized, and that activation in the right ATL in particular may be
related to some of the challenges individuals with AUT often report in
processing social and figurative language. The greater activation in the
AUT group for social/figurative phrases in the ATL also raises the pos­
sibility that the AUT group may be processing the phrases in a taxonomic
fashion. An implication of this for future exploration is that school-age
individuals with autism may be more readily taught social and figura­
tive content if it is presented in a taxonomic rather than thematic
fashion.
Credit author statement
William W. Graves: Conceptualization, Methodology, Formal
analysis, Resources, Writing – original draft, Writing – review & editing,
Visualization, Supervision, Project administration, Funding acquisition.
Hillary Levinson: Conceptualization, Methodology, Formal analysis,
Investigation, Data curation, Writing – original draft, Project adminis­
tration. Linsah Coulanges: Investigation, Data curation, Writing – re­
view & editing, Project administration. Shannon Cahalan:
Investigation, Data curation, Writing – review & editing, Project
administration. Daniel Cruz: Investigation, Writing – review & editing.
Catherine Sancimino: Investigation, Writing – review & editing.
Vanessa H. Bal: Investigation, Writing – review & editing, Supervision.
Miriam Rosenberg-Lee: Conceptualization, Methodology, Writing –
review & editing, Supervision, Project administration, Funding
acquisition.
Acknowledgements
We gratefully acknowledge the study participants, and the numerous
11
Neuropsychologia 171 (2022) 108240
research assistants who helped with recruitment and data collection.
Funding for this study was provided by a Rutgers University Busch
Biomedical Grant to MRL and WWG.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.neuropsychologia.2022.108240.
References
Armstrong, B.C., Watson, C.E., Plaut, D.C., 2012. SOS! an algorithm and software for the
stochastic optimization of stimuli. Behav. Res. Methods 44 (3), 675–705.
Bach, K., 2004. Pragmatics and the philosophy of language. In: Horn, L.R., Ward, G.
(Eds.), The Handbook of Pragmatics. John Wiley & Sons, Incorporated, pp. 463–487.
Baron, S.G., Osherson, D., 2011. Evidence for conceptual combination in the left anterior
temporal lobe. Neuroimage 55, 1847–1852.
Baron, S.G., Thompson-Schill, S.L., Weber, M., Osherson, D., 2010. An early stage of
conceptual combination: superimposition of constituent concepts in left
anterolateral temporal lobe. Cognit. Neurosci. 1 (1), 44–51.
Bechara, A., Tranel, D., Damasio, H., Damasio, A.R., 1996. Failure to respond
autonomically to anticipated future outcomes following damage to prefrontal cortex.
Cerebr. Cortex 6 (2), 215–225.
Bemis, D.K., Pylkkänen, L., 2011. Simple composition: a Magnetoencephalography
investigation into the comprehension of minimal linguistic phrases. J. Neurosci. 31
(8), 2801–2814.
Binder, J.R., Desai, R.H., 2011. The neurobiology of semantic memory. Trends Cognit.
Sci. 15 (11), 527–536.
Binder, J.R., Desai, R.H., Graves, W.W., Conant, L.L., 2009. Where is the semantic
system? A critical review and meta-analysis of 120 functional neuroimaging studies.
Cerebr. Cortex 19, 2767–2796. https://doi.org/10.1093/cercor/bhp055.
Binder, J.R., Frost, J.A., Hammeke, T.A., Bellgowan, P.S.F., Rao, S.M., Cox, R.W., 1999.
Conceptual processing during the conscious resting state: a functional MRI study.
J. Cognit. Neurosci. 11 (1), 80–93.
Bohrn, I.C., Altmann, U., Jacobs, A.M., 2012. Looking at the brains behind figurative
language—a quantitative meta-analysis of neuroimaging studies on metaphor,
idiom, and irony processing. Neuropsychologia 50 (11), 2669–2683.
Borghi, A.M., Binkofski, F., Castelfranchi, C., Cimatti, F., Scorolli, C., Tummolini, L.,
2017. The challenge of abstract concepts. Psychol. Bull. 143 (3), 263.
Boylan, C., Trueswell, J.C., Thompson-Schill, S.L., 2017. Relational vs. attributive
interpretation of nominal compounds differentially engages angular gyrus and
anterior temporal lobe. Brain Lang. 169, 8–21.
Brants, T., Franz, A., 2006. Web 1T 5-gram Version 1. Linguistic Data Consortium,
Philadelphia.
Brownell, H.H., Simpson, T.L., Bihrle, A.M., Potter, H.H., Gardner, H., 1990.
Appreciation of metaphoric alternative word meanings by left and right brain-
damaged patients. Neuropsychologia 28 (4), 375–383.
Brysbaert, M., Warriner, A.B., Kuperman, V., 2014. Concreteness ratings for 40 thousand
generally known English word lemmas. Behav. Res. Methods 46 (3), 904–911.
Buckner, R.L., Andrews-Hanna, J.R., Schacter, D.L., 2008. The brain’s default network:
anatomy, function, and relevance to disease. Ann. N. Y. Acad. Sci. 1124, 1–38.
Chen, A.C., Welsh, R.C., Liberzon, I., Taylor, S.F., 2010. ‘Do I like this person?’A network
analysis of midline cortex during a social preference task. Neuroimage 51 (2),
930–939.
Chouinard, B., Volden, J., Cribben, I., Cummine, J., 2017. Neurological evaluation of the
selection stage of metaphor comprehension in individuals with and without autism
spectrum disorder. Neuroscience 361, 19–33.
Cox, R.W., 1996. AFNI: software for analysis and visualization of functional magnetic
resonance neuroimages. Comput. Biomed. Res. 29, 162–173.
Cox, R.W., Chen, G., Glen, D.R., Reynolds, R.C., Taylor, P.A., 2017. FMRI clustering in
AFNI: false-positive rates redux. Brain Connect. 7 (3), 152–171. https://doi.org/
10.1089/brain.2016.0475.
Cox, R.W., Hyde, J.S., 1997. Software tools for analysis and visualization of fMRI data.
NMR Biomed.: An International Journal Devoted to the Development and
Application of Magnetic Resonance In Vivo 10 (4-5), 171–178.
Davis, C.P., Yee, E., 2019. Features, labels, space, and time: factors supporting taxonomic
relationships in the anterior temporal lobe and thematic relationships in the angular
gyrus. Language, Cognition and Neuroscience 34 (10), 1347–1357.
Denny, B.T., Kober, H., Wager, T.D., Ochsner, K.N., 2012. A meta-analysis of functional
neuroimaging studies of self-and other judgments reveals a spatial gradient for
mentalizing in medial prefrontal cortex. J. Cognit. Neurosci. 24 (8), 1742–1752.
Duncan, J., 2010. The multiple-demand (MD) system of the primate brain: mental
programs for intelligent behaviour. Trends Cognit. Sci. 14 (4), 172–179.
Eklund, A., Nichols, T.E., Knutsson, H., 2016. Cluster failure: why fMRI inferences for
spatial extent have inflated false-positive rates. Proc. Natl. Acad. Sci. U.S.A. 113
(33), 7900–7905.
Eviatar, Z., Just, M.A., 2006. Brain correlates of discourse processing: an fMRI
investigation of irony and conventional metaphor comprehension. Neuropsychologia
44 (12), 2348–2359.
Fedorenko, E., Duncan, J., Kanwisher, N., 2013. Broad domain generality in focal regions
of frontal and parietal cortex. Proc. Natl. Acad. Sci. U.S.A. 110 (41), 16616–16621.
W.W. Graves et al.
Forgács, B., Bohrn, I., Baudewig, J., Hofmann, M.J., Pléh, C., Jacobs, A.M., 2012. Neural
correlates of combinatorial semantic processing of literal and figurative noun noun
compound words. Neuroimage 63 (3), 1432–1442.
Gaffrey, M.S., Kleinhans, N.M., Haist, F., Akshoomoff, N., Campbell, A., Courchesne, E.,
Müller, R.-A., 2007. A typical participation of visual cortex during word processing
in autism: an fMRI study of semantic decision. Neuropsychologia 45, 1672–1684.
Gagnon, L., Goulet, P., Giroux, F., Joanette, Y., 2003. Processing of metaphoric and non-
metaphoric alternative meanings of words after right-and left-hemispheric lesion.
Brain Lang. 87 (2), 217–226.
Graessner, A., Zaccarella, E., Hartwigsen, G., 2021. Differential contributions of left-
hemispheric language regions to basic semantic composition. Brain Struct. Funct.
226 (2), 501–518.
Graves, W.W., Binder, J.R., Desai, R.H., Conant, L.L., Seidenberg, M.S., 2010a. Neural
correlates of implicit and explicit combinatorial semantic processing. Neuroimage
53, 638–646. https://doi.org/10.1016/j.neuroimage.2010.06.055.
Graves, W.W., Binder, J.R., Seidenberg, M.S., 2013. Noun-noun combination:
meaningfulness ratings and lexical statistics for 2,160 word pairs. Behav. Res.
Methods 45 (2), 463–469.
Graves, W.W., Grabowski, T.J., Mehta, S., Gupta, P., 2008. Left posterior superior
temporal gyrus participates specifically in accessing lexical phonology. J. Cognit.
Neurosci. 20, 1698–1710.
Groen, W.B., Zwiers, M.P., van der Gaag, R.-J., Buitelaar, J.K., 2008. The phenotype and
neural correlates of language in autism: an integrative review. Neurosci. Biobehav.
Rev. 32 (8), 1416–1425.
Gusnard, D.A., Raichle, M.E., 2001. Searching for a baseline: functional imaging and the
resting brain. Nat. Rev. Neurosci. 2, 685–694.
Happé, F., 1997. Central coherence and theory of mind in autism: reading homographs in
context. Br. J. Dev. Psychol. 15 (1), 1–12.
Happé, F., Frith, U., 2006. The weak coherence account: detail-focused cognitive style in
autism spectrum disorders. J. Autism Dev. Disord. 36 (1), 5–25.
Harm, M.W., Seidenberg, M.S., 2004. Computing the meanings of words in reading:
cooperative division of labor between visual and phonological processes. Psychol.
Rev. 111 (3), 662–720.
Harris, G.J., Chabris, C.F., Clark, J., Urban, T., Aharon, I., Steele, S., McGrath, L.,
Condouris, K., Tager-Flusberg, H., 2006. Brain activation during semantic processing
in autism spectrum disorders via functional magnetic resonance imaging. Brain
Cognit. 61 (1), 54–68.
Hus, V., Lord, C., 2014. The autism diagnostic observation schedule, Module 4: revised
algorithm and standardized severity scores. J. Autism Dev. Disord. 44 (8),
1996–2012. https://doi.org/10.1007/s10803-014-2080-3.
International, F., 2017. The Farlex Idioms and Slang Dictionary. CreateSpace
Independent Publishing Platform. https://books.google.com/books?id=N3Z
stAEACAAJ.
Kalandadze, T., Norbury, C., Nærland, T., Næss, K.-A.B., 2018. Figurative language
comprehension in individuals with autism spectrum disorder: a meta-analytic
review. Autism 22 (2), 99–117.
Kamio, Y., Robins, D., Kelley, E., Swainson, B., Fein, D., 2007. Atypical lexical/semantic
processing in high-functioning autism spectrum disorders without early language
delay. J. Autism Dev. Disord. 37, 1116–1122.
Kana, R.K., Keller, T.A., Cherkassky, V.L., Minshew, N.J., Just, M.A., 2006. Sentence
comprehension in autism: thinking in pictures with decreased functional
connectivity. Brain 129, 2484–2493.
Kana, R.K., Wadsworth, H.M., 2012. The archeologist’s career ended in ruins”:
hemispheric differences in pun comprehension in autism. Neuroimage 62 (1), 77–86.
Knaus, T.A., Silver, A.M., Lindgren, K.A., Hadjikhani, N., Tager-Flusberg, H., 2008. fMRI
activation during a language task in adolescents with ASD. J. Int. Neuropsychol. Soc.
14 (6), 967–979.
Koenigs, M., Tranel, D., 2007. Irrational economic decision-making after ventromedial
prefrontal damage: evidence from the Ultimatum Game. J. Neurosci. 27 (4),
951–956.
Koenigs, M., Young, L., Adolphs, R., Tranel, D., Cushman, F., Hauser, M., Damasio, A.,
2007. Damage to the prefrontal cortex increases utilitarian moral judgements.
Nature 446 (7138), 908–911.
Lancaster, J.L., Woldorff, M.G., Parsons, L.M., Liotti, M., Freitas, C.S., Rainey, L.,
Kochunov, P.V., Nickerson, D., Mikiten, S.A., Fox, P.T., 2000. Automated Talairach
atlas labels for functional brain mapping. Hum. Brain Mapp. 10, 120–131.
Landa, R.J., Goldberg, M.C., 2005. Language, social, and executive functions in high
functioning autism: a continuum of performance. J. Autism Dev. Disord. 35 (5),
557–573. https://doi.org/10.1007/s10803-005-0001-1.
Lanzoni, L., Ravasio, D., Thompson, H., Vatansever, D., Margulies, D., Smallwood, J.,
Jefferies, E., 2020. The role of default mode network in semantic cue integration.
Neuroimage 219, 117019.
Levy, A., Perry, A., 2011. Outcomes in adolescents and adults wiht autism: a review of
the literature. Research in Autism Spectrum Disorders 5, 1271–1282.
Li, J., Pylkkänen, L., 2021. Disentangling semantic composition and semantic association
in the left temporal lobe. J. Neurosci. 41, 6526–6538.
Lin, N., Wang, X., Xu, Y., Wang, X., Hua, H., Zhao, Y., Li, X., 2018. Fine Subdivisions of
the semantic network supporting social and sensory–motor semantic processing.
Cerebr. Cortex 28 (8), 2699–2710. https://doi.org/10.1093/cercor/bhx148.
Lin, N., Xu, Y., Yang, H., Zhang, G., Zhang, M., Wang, S., Hua, H., Li, X., 2020.
Dissociating the neural correlates of the sociality and plausibility effects in simple
conceptual combination. Brain Struct. Funct. 225 (3), 995–1008.
Martínez-Sanchis, S., 2014. Neurobiological foundations of multisensory integration in
people with autism spectrum disorders: the role of the medial prefrontal cortex.
Front. Hum. Neurosci. 8, 970.
12
Neuropsychologia 171 (2022) 108240
Mattheiss, S.R., Levinson, H., Graves, W.W., 2018. Duality of function: activation for
meaningless nonwords and semantic codes in the same brain areas. Cerebr. Cortex
28 (7), 2516–2524. https://doi.org/10.1093/cercor/bhy053.
Mirman, D., Landrigan, J.-F., Britt, A.E., 2017. Taxonomic and thematic semantic
systems. Psychol. Bull. 143 (5), 499.
Molenberghs, P., Johnson, H., Henry, J.D., Mattingley, J.B., 2016. Understanding the
minds of others: a neuroimaging meta-analysis. Neurosci. Biobehav. Rev. 65,
276–291.
Murphy, C., Jefferies, E., Rueschemeyer, S.-A., Sormaz, M., Wang, H.-t., Margulies, D.S.,
Smallwood, J., 2018. Distant from input: evidence of regions within the default
mode network supporting perceptually-decoupled and conceptually-guided
cognition. Neuroimage 171, 393–401.
Norbury, C., 2005. Barking up the wrong tree? Lexical ambiguity resolution in children
with language impairments and autistic spectrum disorders. J. Exp. Child Psychol.
90 (2), 142–171.
Norbury, C., Nation, K., 2011. Understanding variability in reading comprehension in
adolescents with autism spectrum disorders: interactions with language status and
decoding skill. Sci. Stud. Read. 15 (3), 191–210.
Ober, B.A., Shenaut, G.K., 2006. Semantic memory. In: Handbook of Psycholinguistics.
Elsevier, pp. 403–453.
Olson, I.R., Plotzker, A., Ezzyat, Y., 2007. The enigmatic temporal pole: a review of
findings on social and emotional processing. Brain 130, 1718–1731.
Patterson, K., Nestor, P.J., Rogers, T.T., 2007. Where do you know what you know? The
representation of semantic knowledge in the human brain. Nat. Rev. Neurosci. 8,
976–987.
Phan, L., Tariq, A., Lam, G., Pang, E.W., Alain, C., 2021. The neurobiology of semantic
processing in autism spectrum disorder: an activation likelihood estimation analysis.
J. Autism Dev. Disord. 51 (9), 3266–3279.
Plaut, D.C., McClelland, J.L., Seidenberg, M.S., Patterson, K., 1996. Understanding
normal and impaired word reading: computational principles in quasi-regular
domains. Psychol. Rev. 103 (1), 56–115.
Pylkkänen, L., 2019. The neural basis of combinatory syntax and semantics. Science 366
(6461), 62–66. https://doi.org/10.1126/science.aax0050.
Pylkkänen, L., McElree, B., 2006. The syntax-semantics interface: on-line composition of
sentence meaning. In: Handbook of Psycholinguistics. Elsevier, pp. 539–579.
Raichle, M.E., 2015. The brain’s default mode network. Annu. Rev. Neurosci. 38,
433–447.
Rapp, A.M., Mutschler, D.E., Erb, M., 2012. Where in the brain is nonliteral language? A
coordinate-based meta-analysis of functional magnetic resonance imaging studies.
Neuroimage 63 (1), 600–610.
Rice, G.E., Lambon Ralph, M.A., Hoffman, P., 2015. The roles of left versus right anterior
temporal lobes in conceptual knowledge: an ALE meta-analysis of 97 functional
neuroimaging studies. Cerebr. Cortex 25 (11), 4374–4391.
Richlan, F., Kronbichler, M., Wimmer, H., 2009. Functional abnormalities in the dyslexic
brain: a quantitative meta-analysis of neuroimaging studies. Hum. Brain Mapp. 30
(10), 3299–3308.
Ross, L.A., Olson, I.R., 2010. Social cognition and the anterior temporal lobes.
Neuroimage 49, 3452–3462.
Saad, Z.S., Glen, D.R., Chen, G., Beauchamp, M.S., Desai, R., Cox, R.W., 2009. A new
method for improving functional-to-structural MRI alignment using local Pearson
correlation. Neuroimage 44, 839–848.
Sahyoun, C.P., Belliveau, J.W., Soulières, I., Schwartz, S., Mody, M., 2010. Neuroimaging
of the functional and structural networks underlying visuospatial vs. linguistic
reasoning in high-functioning autism. Neuropsychologia 48 (1), 86–95.
Schauder, K.B., Mash, L.E., Bryant, L.K., Cascio, C.J., 2015. Interoceptive ability and
body awareness in autism spectrum disorder. J. Exp. Child Psychol. 131, 193–200.
Schurz, M., Radua, J., Tholen, M.G., Maliske, L., Margulies, D.S., Mars, R.B., Sallet, J.,
Kanske, P., 2021. Toward a hierarchical model of social cognition: a neuroimaging
meta-analysis and integrative review of empathy and theory of mind. Psychol. Bull.
147 (3), 293.
Schwartz, M.F., Kimberg, D.Y., Walker, G.M., Brecher, A., Faseyitan, O.K., Dell, G.S.,
Mirman, D., Coslett, H.B., 2011. Neuroanatomical dissociation for taxonomic and
thematic knowledge in the human brain. Proc. Natl. Acad. Sci. U.S.A. 108 (20),
8520–8524.
Seidenberg, M.S., McClelland, J.L., 1989. A distributed, developmental model of word
recognition and naming. Psychol. Rev. 96 (4), 523–568.
Simmons, W.K., Reddish, M., Bellgowan, P.S., Martin, A., 2010. The selectivity and
functional connectivity of the anterior temporal lobes. Cerebr. Cortex 20, 813–825.
Smallwood, J., Bernhardt, B.C., Leech, R., Bzdok, D., Jefferies, E., Margulies, D.S., 2021.
The default mode network in cognition: a topographical perspective. Nat. Rev.
Neurosci. 1–11.
Tager-Flusberg, H., Paul, R., Lord, C., 2005. language and communication in autism. In:
Volkmar, F.R., Paul, R., Klin, A., Cohen, D. (Eds.), Handbook of Autism and
Pervasive Developmental Disorders: Diagnosis, Development, Neurobiology, and
Behavior. John Wiley & Sons Inc, pp. 335–364. https://doi.org/10.1002/
9780470939345.ch12.
Taylor, J.S.H., Rastle, K., Davis, M.H., 2013. Can cognitive models explain brain
activation during word and pseudoword reading? A meta-analysis of 36
neuroimaging studies. Psychol. Bull. 139 (4), 766–791. https://doi.org/10.1037/
a0030266.
Van Overwalle, F., 2009. Social cognition and the brain: a meta-analysis. Hum. Brain
Mapp. 30, 829–858.
Vicente, A., Falkum, I.L., 2021. Accounting for the Preference for Literal Meanings in
Autism Spectrum Conditions. Mind & Language.
Vigliocco, G., Tranel, D., Druks, J., 2012. Language Production. The Cambridge
handbook of psycholinguistics, p. 443.
W.W. Graves et al.
Vigneau, M., Beaucousin, V., Hervé, P.Y., Duffau, H., Crivello, F., Houdé, O., Mazoyer, B.,
Tzourio-Mazoyer, N., 2006. Meta-analyzing left hemisphere language areas:
phonology, semantics, and sentence processing. Neuroimage 30, 1414–1432.
Visser, M., Jefferies, E., Lambon Ralph, M.A., 2010. Semantic processing in the anterior
temporal lobes: a meta-analysis of the functional neuroimaging literature. J. Cognit.
Neurosci. 22 (6), 1083–1094.
Vulchanova, M., Saldaña, D., Chahboun, S., Vulchanov, V., 2015. Figurative language
processing in atypical populations: the ASD perspective. Front. Hum. Neurosci. 9, 24.
Wakusawa, K., Sugiura, M., Sassa, Y., Jeong, H., Horie, K., Sato, S., Yokoyama, H.,
Tsuchiya, S., Inuma, K., Kawashima, R., 2007. Comprehension of implicit meanings
in social situations involving irony: a functional MRI study. Neuroimage 37 (4),
1417–1426.
Wang, J., Conder, J.A., Blitzer, D.N., Shinkareva, S.V., 2010. Neural representation of
abstract and concrete concepts: a meta-analysis of neuroimaging studies. Hum. Brain
Mapp. 31, 1459–1468.
Wechsler, D., 2011. Wechsler Abbreviated Scale of Intelligence, second ed. The
Psychological Corporation.
13
Neuropsychologia 171 (2022) 108240
Westerlund, M., Pylkkänen, L., 2014. The role of the left anterior temporal lobe in
semantic composition vs. semantic memory. Neuropsychologia 57, 59–70.
Williams, D.L., Minshew, N.J., Goldstein, G., 2015. Further understanding of complex
information processing in verbal adolescents and adults with autism spectrum
disorders. Autism 19 (7), 859–867.
Wirth, M., Jann, K., Dierks, T., Federspiel, A., Wiest, R., Horn, H., 2011. Semantic
memory involvement in the default mode network: a functional neuroimaging study
using independent component analysis. Neuroimage 54 (4), 3057–3066.
Further reading
Graves, W.W., Desai, R., Humphries, C., Seidenberg, M.S., Binder, J.R., 2010b. Neural
systems for reading aloud: a multiparametric approach. Cerebr. Cortex 20,
1799–1815. https://doi.org/10.1093/cercor/bhp245.