738392

research-article2017
AUT0010.1177/1362361317738392AutismHatfield et al.

Original Article

Autism

Autism spectrum disorder and 2019, Vol. 23(1) 212­–222

© The Author(s) 2017
Article reuse guidelines:
interoception: Abnormalities in sagepub.com/journals-permissions
DOI: 10.1177/1362361317738392
https://doi.org/10.1177/1362361317738392

global integration? journals.sagepub.com/home/aut

Timothy R Hatfield1 , Rhonda F Brown1,

Melita J Giummarra2 and Bigna Lenggenhager3

Abstract
Research over the past three decades has seen a revived interest in the way the human body—and the way in which it
is perceived—interacts with aspects of our experience. Consequently, interoception (i.e. the perception of physiological
feedback from the body) has recently been shown to be associated with a wide range of cognitive, emotional, and
affective functions, making it broadly relevant to the study of autism spectrum disorder. Although limited qualitative
accounts and empirical studies suggest that individuals with autism spectrum disorder encounter abnormalities when
perceiving and integrating physiological feedback from their bodies, other studies have suggested that people with/
without autism spectrum disorder do not differ in interoceptive ability after accounting for alexithymia. In this article,
we discuss the newly recognized importance of interoception in autism spectrum disorder with a focus on how deficits
in the perception of bodily feedback might relate to the core features and co-occuring psychopathology of autism
spectrum disorder. Finally, a new integrated theory is advanced which posits that people with autism spectrum disorder
may experience a reduced capacity to integrate interoceptive information that may result in a narrow attentional bodily
focus and reduced motivational and behavioral drives.

Keywords
alexithymia, autism spectrum disorder, body, interoception, sensory experiences, weak central coherence

Introduction
Sherrington’s (1906) classical taxonomy of the sensory
system originally described interoception as the process
of monitoring sensory inputs exclusively from the viscera,
such that it was functionally distinct from the propriocep-
tive (i.e. body and limb position) and exteroceptive (i.e.
touch, temperature, and pain) sensory modalities.
However, recent advances in neurofunctional anatomy
have provoked a substantial revision of the definition of
interoception to describe the sense of the physiological
condition of the entire body, not merely the viscera (Craig,
2002, 2003, 2011, 2014). Thus, the interoceptive sensory
system is now considered to be maintained by a homeo-
static afferent pathway that originates within the small-
diameter sensory afferent fibers that innervate all tissues
and organs, and which terminates in the posterior insula
(Craig, 2002, 2011). Specifically, information from the
small-diameter fibers is projected to pre-autonomic and
homeostatic sites in the caudal medulla via the lamina I
neurons in the spine, trigeminal dorsal horns, and the
nucleus of the solitary tract (NTS), and then on to the
hypothalamus and limbic sensory cortex (i.e. anterior
insula) and limbic motor cortex (i.e. anterior cingulate;
Craig, 2003). Following a posterior-to-anterior processing
gradient, the interoceptive activity that represents sensory
qualities (e.g. itch, pain, temperature) is processed in the
posterior insula, then re-represented and integrated in the
mid-insula, and finally integrated in the anterior insula
cortex (Craig, 2011). Consequently, it has been suggested
that the integration of inputs in the insula provides the
1The Australian National University, Australia
2Monash University, Australia
3University of Zurich, Switzerland
Corresponding author:
Timothy R Hatfield, Research School of Psychology, College of Health
& Medicine, The Australian National University, Canberra, ACT 2601,
Australia.
Email: timothy.hatfield@anu.edu.au
Hatfield et al.
template for a coherent representation of all feelings at
one moment or a “global emotional moment” (Craig,
2009, 2010, 2011). Importantly, functional neuroimaging
studies suggest that activation of the anterior insula is
uniquely associated with various subjective feelings
including internal bodily states (e.g. pain, temperature,
heart rate) and the experience of emotion (Craig, 2002,
2009, 2011; Kober et al., 2008; Zaki et al., 2012).
Despite these recent neuroanatomical advances in map-
ping the mechanisms and systems involved in interocep-
tion, there remain some inconsistencies in how the
definition and quantification of this multidimensional pro-
cess have been applied. Several terms that have been used
to describe different aspects of interoception (e.g. intero-
ceptive awareness, sensitivity, accuracy, and acuity) have
been inconsistently employed in the literature, and the dif-
ferences between objective and subjective modes of evalu-
ating interoception have generally not been acknowledged
(Garfinkel and Critchley, 2013). Recently, a tripartite
model of discrete interoceptive processes has been pro-
posed which makes a clear distinction between the objec-
tive, subjective, and metacognitive aspects of interoception,
and these have been shown to be dissociable in empirical
tests (Garfinkel et al., 2015). Thus, interoceptive accuracy
refers to a person’s behavioral performance on objective
tests of interoceptive ability, such as the heartbeat tracking
task (Schandry, 1981) and heartbeat discrimination task
(Katkin et al., 1983). Interoceptive sensibility refers to a
self-perceived interoceptive focus that is typically assessed
using subjective questionnaires, such as the Body
Perception Questionnaire (BPQ; Porges, 1993) and
Multidimensional Assessment of Interoceptive Awareness
(MAIA; Mehling et al., 2012). However, both these scales
make reference to “interoceptive awareness” rather than
“interoceptive sensibility.” Finally, interoceptive aware-
ness refers to the metacognitive awareness of interoceptive
accuracy which is the correspondence between objective
interoceptive accuracy and subjective self-reporting of
interoception (Garfinkel et al., 2015).
Many contemporary studies that have examined the
awareness of bodily sensations highlight the importance of
physiological feedback from the body to various cognitive,
motivational, emotional, and affective functions (Critchley
and Harrison, 2013). For example, awareness of physio-
logical changes in the body has been shown to be associ-
ated with emotional experience (Wiens, 2005). In
particular, Füstös et al. (2013) showed that the accurate
perception of bodily states can enable the more effective
regulation of emotion. Moreover, Fukushima et al. (2011)
showed that physiological feedback from the body permits
us to make inferences about the affective state of others
(e.g. empathy). Altogether, these findings suggest that the
perception of bodily sensations is an important part of sub-
jective and social human experience that provides us with
a sense of homeostatic instinct and self-awareness.
213
Furthermore, bodily feelings alert us to our physiological
state of well-being, prompting us to seek water if we are
thirsty or warmth when we are cold.
Together, these types of enquires are broadly relevant to
the understanding of autism spectrum disorder (ASD)
which is characterized by deficits in social communication
and social interaction across different contexts, difficulties
in socio-emotional reciprocity, presence of restricted inter-
ests and repetitive behavior, and atypical sensory process-
ing (American Psychiatric Association, 2013). While few
prior studies have directly examined interoception in peo-
ple with ASD, several authors have recently suggested that
people with ASD may experience sensory under-respon-
sivity to internal stimuli (Elwin et al., 2012; Fiene and
Brownlow, 2015). In addition, a novel theoretical perspec-
tive has suggested that dysfunctional interoception in ASD
may be the result of a disruption of the oxytocin system
(Quattrocki and Friston, 2014). Specifically, the authors
suggest that an oxytocin-mediated interoceptive dysfunc-
tion in early development prevents the contextualization of
interoceptive signals which are necessary to acquire asso-
ciations between internal and external cues. However, the
manner in which a putative dysfunction in the interocep-
tive system of people with ASD is related to the core fea-
tures and co-occurring symptomatology (e.g. anxiety and
depression) of the disorder is still unclear. Moreover, the
empirical findings to date have typically been viewed in
isolation, without consideration of the available neuropsy-
chological frameworks for cognitive and perceptual pro-
cessing in ASD, especially the local processing framework
(i.e. weak central coherence (WCC); Frith, 1989). At the
least, based on the scant empirical findings, a local pro-
cessing framework may offer insight into how interocep-
tion operates at the perceptual level in people with ASD.
For example, as detailed below, people with ASD may
preferentially process the local features of interoceptive
states but fail to integrate them together so that their global
significance can be ascertained. In this article, we critically
evaluate the existing literature examining interoception in
people with ASD and propose a new integrated theory of
interoception in ASD that attempts to incorporate intero-
ception into the WCC framework.
Autism and interoception
Historical traces
Dysfunctional bodily awareness in ASD is not a contem-
porary realization. Despite the relative paucity of intero-
ception research in the ASD sensory processing literature,
hints of interoceptive difficulties can be inferred from the
first descriptions of the disorder. For example, Kanner’s
(1943) original case studies of early infantile autism noted
a father’s description of his 5-year-old son with autism:
“He has never shown a normal appetite. Seeing children
214
eating candy and ice-cream has never been a temptation to
him” (p. 217). Similarly, another case describes an 8-year-
old girl: “At camp she slid into avitaminosis [i.e. vitamin
deficiency] and malnutrition but offered almost no verbal
complaints” (p. 229). In subsequent decades, Bettelheim
(1967) also explicitly made reference to childhood diffi-
culties in bodily awareness, commenting on a child’s per-
ceived lack of awareness during defecation: “If autistic
children show no reaction whatsoever to defecation, we
must assume that the process of alienation from their own
feelings has reached such proportions that they do not even
feel what goes on in their bodies” (p. 111). Thus, although
limited, this early clinical literature suggests that the pro-
cessing of interoceptive information may be impaired in
people with ASD.
Interoceptive brain structures and systems in
ASD
The brains of people with ASD are characterized by struc-
tural and functional abnormalities, with many theorists
suggesting that a difference in brain network connectivity
is responsible for many of the hallmark behavioral symp-
toms of the disorder (e.g. socio-emotional deficits). In
particular, functional neuroimaging studies have consist-
ently demonstrated significant low connectivity between
different brain regions in people with ASD (Just et al.,
2004, 2007, 2012; Koshino et al., 2008; Müller et al.,
2011), although some studies have reported higher con-
nectivity (Belmonte et al., 2004; Keown et al., 2013;
Wass, 2011). Importantly, substantial differences in the
functional connectivity of the networks subserving emo-
tion processing and interoception have been demonstrated
in people with ASD relative to controls. In particular,
Ebisch et al. (2011) showed disrupted functional connec-
tivity between the insula, amygdala, and somatosensory
cortices in people with high-functioning ASD. They sug-
gested that the results were consistent with a dysfunc-
tional interoceptive awareness network which may alter
subjective feelings and the ability to attribute emotional
valence to external events.
Recent neuroimaging studies have also suggested that
there are considerable group differences in the regions
that support interoceptive processing in people with ASD
relative to neurotypical controls (Barttfeld et al., 2012; Di
Martino et al., 2009). For example, in a disgust recogni-
tion task, people with ASD showed less activation in a
number of cortico-limbic brain regions including the left
insula (Ogai et al., 2003). Silani et al. (2008) also showed
that people with ASD exhibited reduced activation of the
anterior insula when they were required to introspect
about their feelings. However, the authors reported that it
was the degree of alexithymic traits that was predictive of
reductions in insula activation and that it was not attribut-
able to ASD, per se. In the same way, other authors have
Autism 23(1)
demonstrated that activation of the insula in people with
ASD is modulated by the level of alexithymic traits (Bird
et al., 2010), indicating that atypical activation of the
brain regions supporting interoception is not a necessary
feature ASD, but rather may be due to the frequently con-
comitant alexithymia. Nonetheless, a recent meta-analysis
of 24 functional neuroimaging studies that used social
paradigms has identified that the right anterior insula is
consistently hypoactive in people with ASD (Di Martino
et al., 2009). Thus, the results of these system-level neu-
roscience and functional neuroimaging studies provide a
persuasive starting point to assert that interoceptive pro-
cessing may be dysfunctional in people with ASD due to
fundamental dysregulation in the brain networks subserv-
ing interoception.
The body beneath the skin
Only in the past 5 years have researchers sought to exam-
ine the way in which individuals with ASD attend to and
process internally derived signals. Within this time, there
has been a surge in empirical work on interoception in peo-
ple with ASD, examining objective, subjective, and meta-
cognitive aspects of interoceptive awareness (DuBois
et al., 2016). However, much of this work has overlooked
previous neuropsychological frameworks of ASD, and
there has been little systematic examination of the extent
to which different methodologies and participant groups
have contributed to differences in the study results. As
such, the previous research findings on alterations of inter-
oception in ASD have yielded inconsistent and often con-
tradictory results.
Several studies have examined the subjective evalua-
tions of interoceptive ability in people with and without
ASD. For example, a qualitative study by Elwin et al.
(2012) examined the autobiographies of people with ASD
and reported that they experienced persistent interoceptive
abnormalities, especially hypo-sensitivities to internal
cues such as a difficulty in detecting and recognizing bod-
ily sensations (e.g. pain). However, as the participants had
written extensive autobiographies indicating normal or
greater intelligence (DuBois et al., 2016), the findings can-
not be generalized to larger heterogeneous ASD popula-
tions. Similar results were demonstrated in an online
survey assessing the subjective experience of functional
body and a thirst awareness in people with/without ASD.
People with ASD showed significantly lower body and
thirst awareness relative to controls (Fiene and Brownlow,
2015), using the Body Awareness Questionnaire (BAQ;
Shields et al., 1989) and a novel Thirst Awareness Scale
(TAS; Fiene and Brownlow, 2015), respectively. Although
again, most participants with ASD were assumed to be on
the higher functioning end of the spectrum.
Studies assessing behavioral performance (i.e. accu-
racy) on objective measures of interoception have tended
Hatfield et al.
to show mixed results. For example, Schauder et al. (2015)
assessed interoceptive accuracy using a heartbeat detec-
tion task in children with/without ASD and found that
although they performed equally in their ability to men-
tally track heartbeats, the children with ASD were better at
tracking their heartbeats over longer time intervals. While
it is not clear why the children with ASD demonstrated
superior heartbeat counting performance over longer inter-
vals, the authors proposed that people with ASD may dis-
proportionately allocate attentional resources to internal,
rather than external sensory information. However, others
have suggested that it may be due to non-specific task
adherence (Shah et al., 2016b). Alternately, Garfinkel et al.
(2016b) reported that adults with ASD showed impaired
interoceptive accuracy but an exaggerated subjective
awareness of bodily sensations relative to controls. They
suggested that this impaired ability to objectively detect
bodily signals accompanied by an inflated subjective per-
ception of bodily signals represents an interoceptive trait
prediction error (ITPE) that manifests as deficits in emo-
tional sensitivity and increased anxiety. However, although
the ASD participants completed a measure of autistic
traits, these scores were not directly assessed against the
measures of interoceptive accuracy, sensibility, or aware-
ness. Thus, a direct relationship between ASD status or
symptoms and interoception was not determined.
Several authors have argued that people with/without
ASD do not differ on measures of interoception after taking
account of alexithymic traits. In fact, the alexithymia
hypothesis suggests that where observed, the emotional
symptoms of autism will be due to alexithymia rather than
to the autism per se (Bird and Cook, 2013). Consistent with
this hypothesis, other researchers suggest that alexithymia
should be regarded as a reflection of a general impairment
in interoception (Brewer et al., 2016). However, again, the
empirical results to date have tended to be inconsistent. For
example, in a small study by David et al. (2013), autistic
traits but not alexithymic traits were positively correlated
with cardiac awareness score in a heartbeat counting task in
neurotypical adults. In contrast, other studies have empha-
sized the unique importance of alexithymia in explaining
the relationship between ASD diagnosis and interoception
(Shah et al., 2016a, 2016b). However, most prior studies
have typically used only a one-dimensional approach when
measuring interoception (Livingston and Livingston,
2016). Thus, although it is generally appreciated that inter-
oception is atypical in people with ASD, there is little clar-
ity as to the precise nature of the relationship between
interoceptive deficits, alexithymic traits, and ASD.
Local–global information processing
in ASD
The features of external sensory stimuli can be viewed as
having a hierarchical organizational structure such that
215
local features (e.g. specific details) are subordinate to
global features (e.g. complete picture). In neurotypical
individuals, the global features are typically processed
before the local features, an effect termed global prece-
dence (Navon, 1977). In contrast, people with ASD often
display a preference for and show superiority in the local
processing of sensory stimuli. For example, children with
ASD have been shown to respond faster and more accu-
rately on the Embedded Figures Task (EFT) relative to
controls (Jolliffe and Baron-Cohen, 1997; Shah and Frith,
1983). The EFT requires participants to search for and
detect a target figure that is hidden in a complex visual
pattern as quickly as possible (Gottschaldt, 1926). In a
recent meta-analysis, the pooled data demonstrated that
people with ASD show superior visuospatial performance
on tasks, such as the EFT and block design, and a stronger
local processing preference on Navon tasks (Muth et al.,
2014). Accordingly, several influential neurocognitive
models have been developed to better understand the per-
ceptual processing differences in people with ASD. In
particular, the WCC model (Frith, 1989) attempts to
explain the above privileged access to local parts and
details, and it has come to dominate the psychological
literature examining the local processing biases evident
in people with ASD.
Weak Central Coherence
The term “central coherence” was first used by Frith
(1989) to describe the propensity of neurotypical individu-
als to pull together large amounts of information with
potential global significance and to process it in context,
often at the expense of their memory for specific details.
As such, central coherence is the typical drive to pursue
meaning by extracting the gist or seeing the “big picture”
of the incoming information in everyday events. In con-
trast, WCC refers to a detail-focused local processing bias
that may lead to a failure to see the “big picture” (Happe
and Frith, 2006). Frith (1989) suggested that this central
processing deficit will manifest as a failure to detect global
meaning, and it may underlie many of the non-social defi-
cits and aptitudes that are known to characterize the ASD
experience, such as enhanced attention-to-detail and
greater sensory-perceptual discrimination.
The WCC account of ASD has received notable atten-
tion, and it has undergone considerable refinements since
its original formulation. Currently, WCC is conceptualized
as a cognitive style that is the result of two separate dimen-
sions: (1) a reduced global integration of information and
(2) a bias toward local processing (Happé and Booth,
2008). Consistent with this approach, local processing
biases have been observed experimentally in people with
ASD in visuospatial, auditory, and verbal domains, with
many of the observations related directly to perceptual
processing (Happe and Frith, 2006; Plaisted, 2001).
216
Mechanisms of local processing bias
Despite the prominence of the WCC model, the precise
neural mechanism(s) underlying the local processing
biases in people with ASD is unclear. Two possible mecha-
nisms have been proposed to posit abnormalities in either
the specialized brain regions or pathways used in a given
task, or diffuse changes in neural connectivity throughout
the brain (Happe and Frith, 2006). For instance, studies
that have investigated hemispheric asymmetry have con-
sistently shown that global integrative processing is pre-
dominantly lateralized to the right hemisphere, whereas
local processing is predominantly lateralized to the left
hemisphere (Boksem et al., 2012; Gable et al., 2013;
Volberg and Hübner, 2004; Volberg et al., 2009).
While functional abnormalities in the right hemisphere
may have potential implications for the WCC model, few
prior studies have shown evidence of right hemisphere
abnormalities in people with ASD (McKelvey et al., 1995;
Waiter et al., 2005), and some studies have suggested there
are left hemisphere abnormalities (Peterson et al., 2015).
In any case, two functional magnetic resonance imaging
(fMRI) studies have examined performance on a local–
global processing task in people with ASD, and they failed
to find clear evidence of right hemisphere dysfunction
(Damarla et al., 2010; Ring et al., 1999). Alternately,
abnormalities in the magnocellular pathway following the
dorsal stream of visual processing have been proposed to
account for the observed weak visuospatial coherence
(Milne et al., 2002; Pellicano et al., 2005). However, these
abnormalities cannot account for the auditory local pro-
cessing biases observed in people with ASD.
In contrast to a regional approach, other researchers
have suggested that the mechanism underlying WCC is
reduced anatomical and functional connectivity and syn-
chronization between distant cortical regions in the brain.
For example, in an early study using fMRI during a sen-
tence comprehension task, people with ASD showed con-
sistently reduced functional connectivity throughout the
cortical language system relative to controls (Just et al.,
2004). The results were interpreted to mean that there is
an underfunctioning of integrative neurocircuitry in peo-
ple with ASD which results in impoverished information
integration and is responsible for the occurrence of WCC.
Similarly, another early study proposed that the features
of ASD that are associated with WCC emerge due to an
impairment of temporal binding between specialized local
neural networks (Brock et al., 2002). More recently,
review papers examining structural and functional con-
nectivity studies in ASD have shown aberrant connectiv-
ity using diffusion tensor imaging (DTI; Travers et al.,
2012), magnetoencephalography (MEG; Lajiness et al.,
2014), and fMRI (Rane et al., 2015). Thus, although a
number of different explanatory frameworks have been
advanced, the mechanisms underlying WCC are still
unclear. Nonetheless, it is noteworthy that neural
Autism 23(1)
under-connectivity is purported to give rise to WCC and
that dysfunctional anterior insula connectivity is consist-
ently shown to play an important role in the social interac-
tion deficits of ASD (Anderson et al., 2010; Di Martino
et al., 2009; Uddin and Menon, 2009).
Local processing of interoceptive stimuli
Current concepts of interoception suggest that it is a home-
ostatic sensory capacity that represents the ongoing status
of all tissues and organs in the body (Craig, 2002, 2011).
Relying on specialized peripheral and central neural sub-
strates that represent all afferent activities, this system is
thought to generate distinct feelings from the body includ-
ing hunger, thirst, pain, temperature, itch, and all other bod-
ily sensations to assist us in meeting the challenges of the
environment. At the core of this conceptualization is an
appreciation that it provides a composite representation of
all salient body states at each moment of time which is
encoded as a feeling (Craig, 2010). Thus, it is as if a person
is presented with an ongoing interoceptive scene that is
built on the integration of local physiological features to
produce global features which may have homeostatic sig-
nificance, and which represents the entire physiological
condition of the body over time. This interoceptive scene is
analogous to Craig’s (2010) cinemascopic model of aware-
ness that is made up of a continuously updating series of
global emotional moments for each moment in time.
Importantly, we propose that this conceptualization of
interoception can be combined with the WCC model to
test specific hypotheses about ASD. This integrated the-
ory provides a novel framework for understanding how
interoception may be impoverished if local bodily stimuli
are not integrated into a composite whole. For instance,
thirst is a homeostatic drive state that arises from multiple
sensations and physiological signals such as mouth dry-
ness, viscous saliva, hypovolemia, hypotension, and
increased osmolality (Stevenson et al., 2015), and the
individual elements of the thirst experience can, therefore,
be viewed as local stimulus inputs into the interoceptive
scene, which when integrated together with other motiva-
tional and affective elements will result in a global feeling
state of thirst. However, a deficit in the integration of
these signals, consistent with WCC, may lead to an
impaired ability to perceive global features of homeo-
static significance. As a result, thirst might not be accu-
rately perceived consciously or subconsciously, resulting
in the failure to create appropriate motivational and
behavioral drive states to relieve the thirst. In short, a fail-
ure to properly integrate individual interoceptive inputs
into a homeostatic feeling or global emotional moment
(Craig, 2010) is analogous to not recognizing the prover-
bial forest for the trees. While the exact mechanism under-
lying this proposed dysfunction is not known, a failure
could occur at the levels of the anterior and posterior
Hatfield et al.
Figure 1. Illustration of the potential disturbance to the interoceptive system in ASD proposed in the integrated theory.
insula, whereby hypo-activity and/or under-connectivity
between these and other regions may reduce the likelihood
of the stimuli being integrated accurately. Such a proposal
is complementary to the Bayesian account of neurodevel-
opment in ASD which posits that an aberrant oxytocin
system in infancy may disrupt the precision of interocep-
tive contextualization and integration (Quattrocki and
Friston, 2014). Figure 1 shows an illustrated summary of
the potential failure to integrate local stimuli accurately
and the associated reduction in adaptive motivational and
behavioral drive states.
217
Subjective emotion can also be viewed as the unified
representation of physiological activity, and consequently,
both historical and contemporary models of emotional
processing have suggested that the subjective experience
of an emotion arises from the perception of physiological
feedback from the body (Damasio and Carvalho, 2013;
James, 1884; Schachter and Singer, 1962). Thus, when we
are afraid, we will experience symptoms associated with
autonomic arousal symptoms (e.g. elevated heart and
breathing rate, sweating, piloerection, vasoconstriction,
and pupil dilation; Kreibig, 2010; Levenson, 2003). Again,
218
each of these individual physiological responses can be
viewed as discrete local stimulus inputs that contribute to
the ongoing interoceptive scene and which contribute to
the global emotional feeling state of “fear.” Thus, if a per-
son with ASD has both a bias toward local interoceptive
information and a reduced capacity to integrate interocep-
tive information, then their interoceptive scene may be
fragmented, and this may prevent the accurate perception
of emotion.
Furthermore, a person’s failure to adequately integrate
the discrete interoceptive signals into a coherent whole
may lead to a narrowed attentional focus on local features
in the body, and this may lead to uncertainty in the inter-
pretation of emotions and feelings in the self and others
(i.e. alexithymia). Importantly, deficits in emotion recog-
nition are fundamental to the clinical understanding of
ASD (American Psychiatric Association, 2013) and alex-
ithymia (Brewer et al., 2016) and they are known to be
related to hypo-activity in the brain regions which support
interoceptive awareness (i.e. anterior insula; Silani et al.,
2008). Thus, the interoceptive differences experienced by
people with ASD could contribute to emotion recognition
and emotion regulation difficulties, social interaction
problems (e.g. failure to share perspectives and empa-
thize), and mental health problems (e.g. anxiety due to
uncertainty as to how to interpret and regulate autonomic
arousal symptoms).
General discussion
The objective measurement of interoceptive ability typi-
cally relies on behavioral evaluations to determine how
well a person perceives their bodily sensations. By con-
vention, methodological approaches to measuring intero-
ception have typically emphasized the detection of cardiac
activity (Kleckner et al., 2015). Other measures (e.g. res-
piratory, gastrointestinal, and genitourinary) have also
been used (Cameron, 2002), but the findings related to
interoceptive performance accuracy across different
modalities are mixed (Garfinkel et al., 2016a; Herbert
et al., 2012; Suschinsky and Lalumière, 2012). Two widely
used cardiac interoception paradigms are the heartbeat
counting (or mental tracking) method (Schandry, 1981)
and the heartbeat detection task (Katkin et al., 1983). Both
tasks provide a measure of cardiac interoceptive accuracy,
and they are often used interchangeably in the literature,
but the heartbeat counting task is the principle objective
method that has been used to assess interoception in peo-
ple with ASD (Garfinkel et al., 2016b; Schauder et al.,
2015; Shah et al., 2016a, 2016b).
Heartbeat detection is generally thought to provide an
indicator of general interoceptive ability, but from the
standpoint of WCC, cardiac activity may be considered as
a local feature in a person’s ongoing interoceptive scene.
In this case, WCC would estimate that people with ASD
Autism 23(1)
would perform better or at least the same, relative to typi-
cally developing individuals. Corroborating this assertion,
three prior studies that have compared interoceptive accu-
racy using the heartbeat counting task between people with
ASD and controls have shown no difference in their inter-
oceptive abilities (Schauder et al., 2015; Shah et al., 2016a,
2016b). Crucially, the study by Schauder et al. (2015)
showed that children with ASD were superior at mentally
tracking their heartbeats over longer time intervals.
Although the authors suggested that the results may be due
to an increased attentional focus on interoceptive cues and
a potential attentional trade-off between internal and exter-
nal cues, the results are also consistent with a bias for local
interoceptive information. On the contrary, Garfinkel et al.
(2016b) showed that while there was objective impairment
in the heartbeat counting task in adults with ASD, the ASD
group did not differ significantly from the control group
using heartbeat detection task. They suggested that because
accuracy on the heartbeat detection task requires simulta-
neous multimodal internal–external integration, this sen-
sory integrative process remains intact in people with
ASD. Although this is at odds with what would be pre-
dicted by a WCC model of interoceptive information pro-
cessing, it may be worth exploring performance differences
across these two tasks in larger samples of people with
ASD and in those with a wider range of severity in ASD
traits and features. In any case, if a local processing bias
exists for internally derived signals in people with ASD,
then it raises questions as to the utility of the heartbeat
detection task, unless it is used in combination with other
objective interoceptive measures.
Qualitative accounts and subjective measures of bodily
awareness may, therefore, offer the best means of disentan-
gling local and global interoceptive states. However, the
existing research examining subjective measures of intero-
ception has produced mixed results and/or the differences in
the measures do not permit direct comparison. For example,
one study showed that people with ASD scored significantly
higher on the BPQ (Porges, 1993) relative to controls
(Garfinkel et al., 2016b), whereas others showed that people
with ASD scored significantly lower on the TAS (Fiene and
Brownlow, 2015) and the BAQ (Shields et al., 1989) rela-
tive to controls (Fiene and Brownlow, 2015). Thus, the con-
flicting results may be due, in part, to the use of different
scale measures and the extent to which they assess local and
global sensory processes. For example, the BPQ mostly
asks about localized bodily symptoms (e.g. dry mouth,
facial twitches, heartbeat, and palms sweating; Porges,
1993), whereas the BAQ frames the questions in terms of
integrated global bodily states (e.g. “I know in advance if I
am getting the flu,” “I am aware of a cycle in my activity
level throughout the day,” and “I notice distinct body reac-
tions when I am fatigued”). Indeed, some items in the TAS
presume that a person already knows what the globally inte-
grated sensory experience of thirst is to answer the questions
Hatfield et al.
about specific sensations (e.g. “I notice distinct body reac-
tions when I am thirsty,” and “I notice distinct body reac-
tions when I am no longer thirsty”). Thus, it may be possible
to reconcile at least some of the conflicting results in the
literature by combining interoception with the WCC model
and examining the prior relevant literature from this stand-
point. Therefore, future research may benefit from separat-
ing local bodily symptoms and global bodily feelings when
examining interoceptive sensibility in ASD.
Specific predictions about emotion processing problems
in people with ASD can be tested by examining interocep-
tion from the perspective of the WCC. If there is a local
processing bias for interoceptive signals, then general tasks
of central coherence should generate results that are
inversely correlated with the results of emotion recognition
tasks in oneself and others. For example, the “Mind in the
Eyes Test” (Baron-Cohen et al., 2001), which has been
used to test the ability of individuals to read emotions and
other mental states from micro-expressions in the eyes of
other people, has been shown to be inversely correlated
with speed on the EFT in people with/without ASD (Baron-
Cohen and Hammer, 1997; Jarrold et al., 2000). This pat-
tern of results suggests that a perceptual bias toward the
local features of the environment will make the process of
recognizing emotions in others more difficult, as is the case
in people with ASD. Second, and by extension, feeling
states such as thirst which require the integration of multi-
ple interoceptive inputs should be more difficult to perceive
and interpret for people with ASD than simple feeling
states. Pertinently, Fiene and Brownlow (2015) reported
that individuals with ASD show significantly lower subjec-
tive thirst awareness relative to controls, which is consist-
ent with such a theorized local sensory processing bias.
The relationship between interoception and anxiety is
well established, with prior studies typically demonstrat-
ing positive associations between trait anxiety and meas-
ures of interoceptive accuracy (e.g. heartbeat detection;
Domschke et al., 2010). Indeed, anxiety is highly preva-
lent in people with an ASD diagnosis across all age
groups and affects about one-half (42%–56%) of people
with the disorder (Lai et al., 2014; Simonoff et al., 2008).
Importantly, Garfinkel et al. (2016a) showed that the
divergence between interoceptive accuracy and intero-
ceptive sensibility, or ITPE, was predictive of anxiety
symptoms in people with ASD beyond the severity of the
disorder. Along with others (Paulus and Stein, 2010),
they suggest that a person’s interoceptive structure may
represent a vulnerability factor for anxiety. A local pro-
cessing bias toward interoceptive stimuli may emphasize
the potential threat posed by autonomic reactivity (e.g.
heart rate) relative to other interoceptive and exterocep-
tive information that could be integrated into the intero-
ceptive scene to alleviate a person’s anxious feelings.
Such a hypothesis has yet to be empirically examined,
although a similar mechanism has been proposed to operate
219
in people with panic disorder such that their awareness of
fear-related physiological processes (e.g. cardiac activity)
is increased and their awareness of other processes is simul-
taneously neglected (Fairclough and Goodwin, 2007;
Limmer et al., 2015). Thus, while people prone to panic and
anxiety may have learned to bias their attention toward car-
diac activity due to its perceived potential for threat, people
with ASD may preferentially process cardiac information
as it represents a salient local stimulus.
Clinical implications
The use of a local–global approach in examining interocep-
tion in people with ASD and alexithymia may be informative
with regard to the use of therapy. For example, rather than
being a core deficit, WCC is conceptualized as a bias that can
be overcome with explicit task demands, and it is not consid-
ered to be a non-modifiable deficit (Happe and Frith, 2006).
Thus, global interoceptive processing might be improved in
people with ASD via the use of body-scan meditation tech-
niques that focus on all bodily regions. A recent study of an
unsupervised brief body-scan meditation intervention
showed improvement in somatosensory perceptual decision-
making in participants with medically unexplained symp-
toms (Mirams et al., 2013). While the authors suggested that
body-scan type meditation could reduce the misperception
of physical symptoms in individuals with medically unex-
plained symptom, people with ASD and alexithymia may
also experience benefits from increased attention and sensi-
tivity to their bodies. In fact, adults with ASD showed sig-
nificant reductions in depression, anxiety, and rumination
and an increase in positive affect in a randomized-controlled
trial of mindfulness-based therapy which included a body-
scan component (Spek et al., 2013). Importantly, other prior
studies have also noted the potentially therapeutic pathway
of interoception training and meditation in reducing distress
and anxiety (Schaefer et al., 2014; Serpa et al., 2014).
Applying clinical remediation or training approaches such as
these is likely to effectively enhance interoception in a range
of conditions in which local processing biases occur (e.g.
anorexia; Lopez et al., 2008).
Conclusion
This review discusses the potential importance of intero-
ception in better understanding the experiences and behav-
ior of people with ASD and also the potential mechanisms
underlying some of the socio-emotional and sensory fea-
tures of ASD. A picture of atypical interoceptive processing
in ASD is starting to emerge in the literature which incorpo-
rates recent advances in systems neuroscience and recent
empirical studies investigating objective and subjective
bodily awareness. However, the impairments appear to be
more strongly linked to alexithymia than to ASD, although
alexithymia frequently co-occurs with ASD. Furthermore,
220
by extending the WCC framework to include interoception,
the integrative theory that we have proposed may represent
a valuable starting point to better understand the core fea-
tures of ASD and the psychopathology associated with the
disorder. For example, the perceptual abnormalities out-
lined in such a framework may impair the perception of
internal sensory information in people with ASD and
potentially influence their experience of homeostatic and
emotional feeling states, and it may also contribute to com-
monly experienced affective symptoms (e.g. anxiety).
Current methodological limitations related to the
objective measurement of interoceptive processing pre-
sent a challenge in incorporating interoception into the
WCC framework. To fully understand the context of
interoception and awareness in ASD, it will be necessary
to appreciate the subjective evaluations of bodily percep-
tion. Ultimately, it will be necessary to investigate the
objective, subjective, and metacognitive aspects of inter-
oception in relation to both ASD and alexithymia before
we can fully understand the nature, quality, and extent of
the putative perceptual biases. Moreover, appreciation of
these experiences across the range of autism severity is
necessary given that much of the literature has focused
on high-functioning individuals with ASD who have a
greater capacity to comment on their internal sensations,
states, and emotions. Finally, the theoretical approach
that we have presented has potential implications for the
wider application of the WCC framework in ASD, which
has historically only accounted for non-social features of
ASD.
Funding
The author(s) disclosed receipt of the following financial support
for the research, authorship, and/or publication of this article:
MJG was supported by an Australian Research Council Discovery
Early Career Research Award (DE170100726). BL was funded
by the Swiss National Science Foundation.
ORCID iD
Timothy R Hatfield https://orcid.org/0000-0002-5223-5798
Melita J Giummarra https://orcid.org/0000-0001-7839-6058
Bigna Lenggenhager https://orcid.org/0000-0003-0418-9931
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