Neuropsychol Rev (2014) 24:7787

DOI 10.1007/s11065-014-9246-9

REVIEW

Contributions of the Insula to Cognition and Emotion

Philip Gerard Gasquoine

Received: 31 October 2013 / Accepted: 8 January 2014 / Published online: 18 January 2014
# Springer Science+Business Media New York 2014

Abstract Historically, the insula was considered primary

gustatory cortex. Now it is known to play a more
comprehensive role in the processing of sensory information, including acting as primary cortex for interoceptive information, including autonomic nervous system mediated changes. As such, it is critical for emotional feeling in accord with the James-Lange theory, a
role previously ascribed to the limbic system.
Neuroimaged abnormal grey matter volumes or activity
levels in the insula have been associated with schizophrenia, eating disorders, anxiety and mood disorders,
conduct disorder, autism, addiction, and chronic pain.
The significance of these abnormal activity patterns
remains theoretical. Neuropsychological studies have
linked dominant insula injury with various symptoms
of aphasia, but its exact role in language processing
remains uncertain as most cases involve lesions that
extend into perisylvian language zones. Functional neuroimaging studies have found insula hyper-activations,
typically in conjunction with anterior cingulate cortex,
for all manner of experimental tasks including those
involving perception, intentional action, and consciousness. Such neuroimaged activity is unlikely to be taskspecific, but rather reflective of generic changes in
autonomic activity in response to salience, homeostatic
incongruence, or cognitive challenge.

Keywords Interoception . James-Lange theory of emotion .

Autonomic nervous system . Island of Reil . Insular cortex

P. G. Gasquoine (*)
Department of Psychology, University of TexasPan American,
1201 W. University Drive, Edinburg, TX 78541, USA
e-mail: drgdrg13@yahoo.com

Introduction
Many contemporary behavioral neuroscientists conceptualize
complex human behaviors as mediated by the flow of information (action potentials) through large scale distributed networks within the brain with modularization (specialized processing) taking place in various brain structures along the way
(e.g., Seeley et al. 2007). This may be little different from
previous conceptions of clinicians who followed in the tradition of Broca (1861) by investigating the effects on behavior
of naturally occurring brain lesions in humans. As illustration,
the influential Soviet neurologist Alexandra Luria wrote:
Each form of conscious activity is always a complex functional system and takes place through the combined working of
allbrain units, each of which makes its own contribution
(Luria 1973, p. 99: italics as in original).
Nowadays, behavioral neuroscientists have additional experimental techniques to supplement lesion/behavior change
analysis, especially the popular noninvasive neuroimaging
technique of functional magnetic resonance imaging (fMRI:
Ogawa et al. 1990). One benefit of the functional neuroimaging approach is that it has focused attention upon the brain
organization of human behaviors not assessed by traditional
neuropsychological tests of cognition that have dominated
lesion analysis studies. Such non-cognitive behaviors include
homeostatically incongruent states like hunger (e.g., Tataranni
et al. 1999) and self-generated emotional states (e.g., Damasio
et al. 2000). The role of brain structures that mediate such
behaviors have been poorly understood in the past. The insula
is one such structure.
A major reason for the historical lack of understanding of
the behavioral function of this enigmatic brain region is that
isolated lesions of the insula in humans are extremely rare. As
illustration, only 4 of 4,800 (0.0008 %) consecutive, first time
stroke patients recorded in the Lausanne Stroke Registry in
Switzerland between 1990 and 1999 had a lesion restricted to

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the insula as determined from static neuroimaging techniques

(Cereda et al. 2002). This has meant that lesion/behavior
change investigations of strokes affecting the insula are invariably single case studies, of very low N, and/or include
cases with injury extending beyond the insula. No other
variant of naturally occurring structural brain injury has been
found to affect the insula in isolation. There is a type of simple
partial seizure that is thought to emanate from the insula
(Isnard et al. 2004) and there have been studies of the symptoms accompanying this state along with electrical stimulation
studies of the insula in seizure patients (e.g., Afif et al. 2010;
Penfield and Faulk 1955; Stephani et al. 2011).
In the earliest conceptions the insula was chiefly known as
primary gustatory cortex (von Bechterew 1899). This attribution has survived to modern times (e.g., Ibaez et al. 2010), but
the insula has proved unlike the other primary sensory cortices
(visual, auditory, and somatosensory), for four main reasons:
(a) minimal neuronal response: primate studies have shown
that only 210 % of neurons in the insula respond to taste
stimuli (Sewards and Sewards 2001); (b) ipsilateral representation: taste information is conveyed to the ipsilateral insula
via the ventral posteromedial nucleus of the thalamus from the
Facial (VII), Glossopharngeal (IX), and Vagus (X) nerves; (c)
lack of chemotopic organization: there are five basic components of taste: sour; bitter; salty; sweet; and umami (savory).
There is little evidence that these basic components activate
different regions of the insula in humans (Schoenfeld et al.
2004); and (d) lack of specialization of sensory input: in
addition to taste information, the insula receives sensory information from the other four traditional senses (Sterzer and
Kleinschmidt 2010) in addition to interoceptive information
(Afif et al. 2010; Carleton et al. 2010; Cereda et al. 2002; Craig
et al. 2000; Damasio 2003; Penfield and Faulk 1955; Stephani
et al. 2011). Interoceptive is a generic term that includes
sensory information from the internal milieu including: temperature, pain, itch, tickle, sensual touch, muscular and visceral sensations, vasomotor flush, hunger, thirst, air hunger and
others (Paulus and Stein 2006, p. 60).
In more recent times, functional neuroimaging studies of
the insula have showed increased metabolic activity during a
wide range of experimental tasks: Activations in the AIC
(anterior insular cortex), along with anterior cingulate cortexareamong the most commonly reported foci in cognitive tasks, including those probing processes underlying perception and awareness (Sterzer and Kleinschmidt 2010, p.
611). Prominent coactivations of the insula (especially the
anterior subregion) and anterior cingulate cortex have also
been reported in many functional neuroimaging studies involving changes in autonomic reactivity mediated by salient
emotional or homeostatically incongruent stimuli including
thermal regulation, sexually explicit visual stimuli, pleasant
touch, hunger, breathlessness, and thermal pain (Gasquoine
2013). The chief question that these studies raised is what

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contribution the insula is making to these behaviors, a question that functional neuroimaging in isolation is poorly
equipped to address. There are general limitations in
interpreting variations in neuroimaged metabolic activity
levels in a given brain region in that they cannot be used to
determine exactly what contribution the brain region is making to the task (Coltheart 2006a, b). Neuroimaged metabolic
signals cannot distinguish between fundamental physiological
processes like excitatory/inhibitory or modulatory/
transmissive effects within a given neural region (Logothetis
2008). To circumvent such interpretational limitations, it has
been proposed that neuroimaging data be interpreted conservatively and combined with other sources of information to
better interpret brain-behavior relationships in a convergingmethods approach (e.g., Decety and Cacioppo 2010;
Gonsalves and Cohen 2010; Logothetis 2008; Stirling and
Elliott 2008). This review synthesizes information from brain
lesion/behavior change, stimulation, and functional neuroimaging studies on the insula with the goal of achieving a
parsimonious understanding of the contribution of the insula
to cognition and emotion. The preponderance of the evidence
suggests the insula acts as primary cortex for interoceptive
information that is critical for emotional feeling.

Neuroanatomy of the Insula

The insula (see Fig. 1) is that part of the cerebral cortex shaped
like an inverted triangle folded within, and forming the base
of, the lateral sulcus (sylvian fissure). It is completely obscured in the lateral view of the exposed cerebral cortex by
the overlying opercula of the temporal, parietal, and frontal
lobes. It was named by Johann-Christian Reil (1809), the
father of German psychiatry (Binder et al. 2007). Insula means
island in Latin. The insula is also known by the names island
of Reil, insular cortex, and intrasylvian cortex. It is a distinct
lobe of the cerebral cortex (Butti and Hof 2010), designated by
some as the fifth lobe (e.g., Augustine 1996), although all

Fig. 1 Insular cortex shown in a lateral view of the left cerebral hemisphere with the overlying opercula of the temporal, parietal, and frontal
lobes removed (Source: Grays Anatomy)

Neuropsychol Rev (2014) 24:7787

the other lobes of the cerebral cortex are named for bones of
the skull. It is the first region of cerebral cortex to develop and
differentiate, beginning at age 6 weeks of fetal life (Afif et al.
2007). In adults it comprises 2 % of the total cortical area
(Nieuwenhuys 2012).
The insula consists of five to six gyri arranged in a vertical
fan-like manner with the circular sulcus forming an upper
boundary. It has long been recognized that the number and
neuroanatomical arrangement of insula gyri varies naturally
among neurologically intact individuals, even between the
two sides of the same brain (Clark 1896; Craig 2010). Early
cytoarchitectonic studies on the human insula identified from
4 (areas 1316: Brodmann 1909) to 31 (Rose 1928) subregions. A frequently referenced subdivision of the insula based
on cytoarchitectonic study of Old World (rhesus) monkeys
defined concentric circular agranular (anterior/ventral),
dysgranular (medial), and granular (posterior/dorsal) subregions emanating from the lowest point (Mesulum and Mufson
1982). Variations in granularity pertain to the distribution of
neurons at layer IV, typically considered as having an input or
sensory function, of the cortical grey matter. The human insula
is much larger than that of the rhesus monkey and the boundaries of these subregions have not been precisely specified in
terms of sulci and gyri (Afif et al. 2007). Another oft used
subdivision is into anterior (three or four short gyri) and
posterior (two long gyri) subregions separated by the central
insular sulcus (Afif and Mertens 2010). The main branch of
the middle cerebral artery lies within this sulcus. Subdivision
of the human insula on the basis of cytoarchitecture remains
contentious (Nieuwenhuys 2012). Similarly, there is little
agreement on the functional specialization of subregions of
the insula from meta-analyses of functional neuroimaging
(Kurth et al. 2010; Mutschler et al. 2009) or electrical stimulation study (Stephani et al. 2011).
The insula has bidirectional connections with most of the
brain, but especially with orbitofrontal, anterior cingulate,
supplementary motor areas, parietal (primary and secondary
somatosensory areas), and temporal cortices and with subcortical structures that include especially the amygdala, globus
pallidus of the basal ganglia, and thalamus (Augustine 1985,
1996; Flynn et al. 1999: see Fig. 2). There is gradation of
connectivity throughout the region in that the anterior insula
has greater connectivity with the frontal lobe, while the posterior insula has greater connectivity with the parietal lobe.

Insula and Cognition

Insula Lesions and Sensory, Motor, Visual-perceptual,
Memory, and Executive Impairment
Isolated lesions of the insula, especially to the posterior subregion, following stroke have been shown to result in

79
Orbitofrontal cortex
Anterior cingulate
cortex
Supplementary motor
areas
Primary
somatosensory cortex
Insula
Secondary
somatosensory cortex

Temporal cortex

Thalamus

Amygdala

Globus pallidus

Fig. 2 Major bidirectional connections for the insula

impairment of somatosensory, gustatory, and vestibular processing (Cereda et al. 2002). Insula injury has also been
implicated in patients who do not respond to painful stimulation by withdrawal or negative emotional response (Berthier
et al. 1988). Aside from sensory impairments, insula injury
has also been linked with cognitive impairment across all
neuropsychological domains. Many of the cognitive symptoms ascribed to insula injury likely reflect diaschisis (von
Monakow 1914) or disconnection syndromes (Geschwind
1965). In diaschisis uninjured regions of the brain that connect
with injured regions experience reduced neural activity especially in the early stages post-injury. Modern functional neuroimaging of cerebral blood flow and cell metabolism has
confirmed that diaschisis occurs (Pearce 1994), although the
exact mechanism of action is unknown. Disconnection syndromes are a related concept implying that injury to a region
of the brain can permanently adversely impact the performance of connected regions by restricting information flow.
Reasons for suspecting that diaschisis and/or disconnection
syndromes are involved in the production of certain cognitive
symptoms following insula lesions include: (a) the rarity of
reports of the symptom following insula injury; (b) a previous
strong link between the symptom and injury to other brain
regions; and (c) the transient nature of the symptom following
sudden onset neurological conditions like stroke. Examples of
probable diaschisis/disconnection effects after left insula
stroke include: (a) truncal ataxia (Liou et al. 2010), a symptom
historically associated with cerebellar injury; and (b) declarative verbal memory (Manes et al. 1999b), a symptom historically associated with dominant hippocampal injury. In the
latter study, six patients with left insula infarction showed
poorer immediate and delayed recall of a story from the

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Wechsler Memory Scale Logical Memory test and poorer

delayed word-list recall than four patients with right insula
infarction. All patients were tested 48 weeks post, that is
within the accepted period of spontaneous recovery from
stroke (Cramer 2008) when diaschisis is especially prominent.
Examples of probable diaschisis/disconnection effects after
right insula stroke include: (a) anosagnosia for hemiplegia
(Karnath and Baier 2010) that also follows frontal, parietal,
and temporal lobe injuries (Appelros et al. 2007: Pia et al.
2004), injury that is limited to subcortical structures (House
and Hodges 1988), or the brainstem (Assenova et al. 2006). It
has been conceptualized as a disconnection syndrome whereby the injury to the brain results in a failure of the patient to
detect discrepancies between what s/he thinks should be happening (based on preinjury experiences) and the true state.
The mechanism for conscious awareness and self-correction
of errors is not triggered (Preston et al. 2010); (b) perseveration errors in pencil and paper target cancellation (Ronchi
et al. 2012), a symptom historically associated with dorsolateral frontal injury; and (c) multimodal neglect (Berthier, et al.
1987; Manes et al. 1999a), a symptom historically associated
with non-dominant parietal injury. In the last study, four
patients with right insula infarction showed poorer performance on tactile and auditory, but not visual, double simultaneous stimulation and poorer performance on two of three line
bisection tasks than five patients with left insula infarction. All
patients were tested 48 weeks post-stroke.
Insula Lesions and Language Impairment
Both historically and in terms of frequency the non-sensory
neuropsychological domain most associated with insula injury
is language, but the specific role that the insula plays in
language processing is far from clear. Impairments in sound
detection and auditory temporal processing have been associated with insula injury (Bamiou et al. 2006), but most of the
patients in this study had suffered infarctions that extended
into adjacent brain regions historically associated with auditory processing. Dysarthria, or impairment of the motoric
execution aspects of speech, has been: (a) associated with
insula injury following stroke in either hemisphere (Baier
et al. 2011; Cereda et al. 2002; Hiraga et al. 2010); (b)
observed to accompany the simple partial seizures that originate in the insula that can also cause laryngeal constriction and
mutism (Isnard et al. 2004); and (c) elicited by insular stimulation (Afif et al. 2010). Mutism has also been reported
following stroke that impacted the right insula and adjacent
brain regions (e.g., Berthier et al. 1987).
Traditional aphasic syndromes that have been linked with
dominant insula injury include: Brocas aphasia (Ardila
1999); apraxia of speech (Dronkers 1996); conduction aphasia
(Wernicke 1874); the word-deafness component of
Wernickes aphasia (Ardila 1999); and nonfluent progressive

Neuropsychol Rev (2014) 24:7787

aphasia (Gorno-Tempini et al. 2004). The last is one of three

variants of frontotemporal dementia, or Picks disease, characterized especially by apraxia of speech. All three variants of
frontotemporal dementia (the others are semantic dementia
and a behavioral variant) are thought to involve insula degeneration especially within the anterior subregion (Seeley 2010),
although this is not the only brain structure that is so affected.
Two primary areas of controversy concern the role of dominant insula injury in the production of: (a) Brocas aphasia and/
or apraxia of speech (to further complicate matters the distinction between these two concepts is very much disputed: e.g.,
Lebrun 1990) and (b) conduction aphasia. Dronkers (1996)
associated apraxia of speech, conceived chiefly as impairment
in motor planning of articulatory movements, with dominant
insula injury by comparing groups of language impaired stroke
patients with/without this condition. This methodology was later
found suspect as it does not completely rule out the possibility
that the impairment is associated with other injured regions
(Hillis et al. 2004). A recent study of 50 stroke patients linked
apraxia of speech with Brocas area (Richardson et al. 2012).
Conduction aphasia is characterized primarily by the presence of phonemic paraphasias accompanying impaired repetition and naming. It was originally postulated by Wernicke
(1874) who thought it would result from injury to the insula.
Later it became most associated with disruption of the arcuate
fasciculus that connects Brocas and Wernickes areas in the
dominant hemisphere (e.g., Anderson et al. 1999) and runs
partially beneath the insula (Nieuwenhuys 2012). In
discussing the anatomical basis of conduction aphasia,
Damasio and Damasio (1980) wrote: The exact functional
significance of damage to the insula may not be easy to settle
(p. 348). Isolated single case studies have linked conduction
aphasia with insula injury (e.g., Hyman and Tranel 1989;
Marshall et al. 1996). The proximity of Brocas area and the
arcuate fasciculus to the dominant insula suggests that the role
of diaschisis and/or disconnection syndromes in the production of these aphasia syndromes cannot be discounted. For
example, the single case of conduction aphasia reported by
Hyman and Tranel resolved completely after several months
and temporal lobe hypofusion was demonstrated by static
neuroimaging in the Marshall et al. case.
Although the insula is located within the core of the
perisylvian language regions in the dominant hemisphere, it is
not part of the traditional Wernicke-Geschwind model of language processing (Geschwind 1965, 1970). Neuropathology in
most of the cases linking aphasia with dominant insula injury
extended to adjacent structures generally associated with aphasic symptomatology. When the lesions were clearly restricted
to the dominant insula (Cereda et al. 2002), symptoms of
aphasia were inconsistent. In summary, the lesion data shows
injury to dominant insular cortex often produces symptoms of
aphasia, but the exact role of this region in language processing
remains unclear.

Neuropsychol Rev (2014) 24:7787

Insula and Emotion

Emotional Behavior and the Limbic System
The concept of a fundamental set of emotions originated from
Darwin (1872/1965) and his observations on the similarity
between human and animal emotional expression. One classification scheme of distinct and universally recognized facial
expressions in humans gives six fundamental emotions: (a)
fear; (b) anger; (c) sadness; (d) happiness; (e) disgust; and (f)
surprise (Ekman and Friesen 1971). Humans are thought to
differentiate among emotional states by interpreting external
cues associated with a generalized internal physiological response reflecting changes in autonomic nervous system activity (Dalgleish 2004), a concept that originated with the JamesLange theory of emotion (James 1884) and has persisted in
modified form to this day. In physiological studies of differing
emotional states the variance associated with generalized activation is far greater than the variance among specific emotional contexts (Lang 1994).
Neural control of emotional behavior is historically associated with the limbic system. Limbic derives from limbus
meaning border in Latin, a term introduced by Broca (1878)
to describe a neuroanatomical conception of a lobe of the
medial and basilar surfaces of the cerebral hemispheres that
formed the border around the brainstem and was thought to
play a primary role in olfaction. The insula was not part of this
lobe. Papez (1937) linked the cortex contained in Brocas
limbic lobe and its main subcortical connections with the
functions of central emotion (p. 743). In reference to the
same neuroanatomical area, MacLean (1952) applied the term
limbic system to describe a visceral brain that interprets
and gives expression to its incoming information in terms of
feeling (p. 415: italics as in original). Subsequently, the insula
was included within the limbic lobe on cytoarchitectonic
grounds (Yakovlev 1959). Later, Mesulum and Mufson
(1982) designated the insula as paralimbic cortex due to the
characteristic variation in granularity in layer IV that also
occurs in other cortical regions (e.g., cingulate cortex). The
functional specialization of paralimbic areas was conceived as
behaviors that integrate extrapersonal stimuli and the internal
milieu (p. 1). It is now seen as unlikely that the historical
conception of the limbic system as defined by Broca, Papez,
and MacLean functions as a singular unit (e.g., Devinsky et al.
1995; LeDoux 1996), given its well-established involvement
in cognitive functions (especially declarative memory) in
addition to emotional behavior.
Insula and the Autonomic Nervous System
Lesion, stimulation, and functional neuroimaging studies have
linked the insula with activity of the autonomic nervous
system including both sensory and motor (e.g., changes in

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respiration, blood pressure, and salivation: Augustine 1985)

components. There is often right side dominance for sympathetic effects. As illustration, strokes affecting the insula in
either hemisphere produce sympathetic hyper-activation that
depresses the immune system thereby increasing the chances
of post-stroke infections (Walter et al. 2013), but patients with
strokes involving the right insula have an increased likelihood
of death in the first 3 months post-injury (Christiansen et al.
2005) and an increased incidence of heart dysfunction
(Oppenheimer 2006). A caveat for interpreting these studies
it that it is difficult to factor out the contribution of preexisting
cardiac or other medical conditions to the post-stroke outcome. Stimulation of the right insula in five seizure disorder
patients produced changes in heart rate and blood pressure
(Oppenheimer et al. 1992). Functional neuroimaging has
shown that metabolism in the right insula correlates with
activities known to activate the sympathetic nervous system,
such as: bladder sensations (Griffiths et al. 2007); the
homeostatically incongruous state of breathlessness
(Banzett et al. 2000); and the magnitude of cardiac response to visually presented facial expressions of sadness,
anger, happiness, and disgust (Critchley et al. 2005). There
is limited evidence that the left insula mediates the activity
of the opponent parasympathetic nervous system (Craig
2005; Oppenheimer et al. 1996).
Insula and Emotional Feelings
Based on the role of the insula in encoding interoceptive
signals from the bodys internal milieu that reflect autonomic
activity, Damasio (2003) argued that the insula is the location
within the brain where subjective feelings of emotion are
generated. This role had previously been ascribed to the limbic
system (MacLean 1952). Functional neuroimaging studies on
emotional behavior have found both generic and emotionspecific effects. An example of the latter is the role of the
amygdala in fear (LeDoux 1996). Functional neuroimaging
studies (e.g., Hennenlotter et al. 2004; Phillips et al. 1997) and
a single case lesion study (Calder et al. 2000) have suggested
an emotion-specific role for the insula in disgust. This
emotion-specific role has not been confirmed by metaanalytical review of functional neuroimaging studies on emotion (Phan et al. 2004), where the data supported a generic role
for the insula in emotional behavior.
Functional Neuroimaging of the Insula in Mental Disorder
and Chronic Pain
Given the established role of the insula in encoding sensory
and motor aspects of autonomic system activity, it would be
expected that insula abnormalities should be linked to mental
disorders that involve emotional dysregulation. Functional
neuroimaging studies have implicated abnormal grey matter

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volumes or hyper-and hypo-activity in the insula with schizophrenia, eating disorders, anxiety and mood disorders, conduct disorder, autism, addiction, and chronic pain.
Grey Matter Volumes in the Insula
Meta-analytic review has shown that insula grey matter volume is decreased bilaterally, especially in the anterior subregion, in patients with schizophrenia (Shepherd et al. 2012),
stimulant drug dependence (Ersche et al. 2013; Mackey and
Paulus 2013), and bipolar disorder (Selvaraj et al. 2012). This
is not the only brain structure so affected in these patients and
the significance of these finding is unclear. Some specific
hypotheses have been generated, for example that insula dysfunction may lead to internally generated sensory information
being misattributed to an external source contributing to hallucinations in schizophrenia (Wylie and Tregellas 2010).
Alternatively, it is possible that the insula dysfunction, whatever its nature, is generic across psychiatric conditions. Grey
matter volume reductions in the left anterior insula have also
been found in major depression (Takahashi et al. 2010) and
bilateral reductions have been reported in posttraumatic stress
disorder (Chen et al. 2009) and conduct disorder (Sterzer et al.
2007). Grey matter volume enlargement in the anterior insula
bilaterally has been reported in obsessive-compulsive disorder
(Nishida et al. 2011).
Hyper-and Hypo-Neuroimaged Activity in the Insula
Several functional neuroimaging studies have shown increased
activity within the insula in either hemisphere during conscious
urges to take drugs of all types (see Naqvi and Bechara 2008
for review). None of these studies implicated the insula specifically and increased activation in regions of orbitofrontal,
anterior cingulate, and dorsolateral prefrontal cortex also occurred. These findings have been supplemented by the lesion
studies of Naqvi et al. (2007) and Gaznick et al. (2013) that
implicated injury to the insula or the insula plus basal ganglia,
respectively, in a sudden loss of the urge to smoke immediately
following brain injury onset in nicotine addicts. Lesions were
not confined to the insula in either study but extended into
adjacent structures including orbitofrontal cortex.
Increased blood flow and metabolism has also been found
in the insula of patients with: psychosis (during auditory
hallucinations: Sommer et al. 2008); major depression
(Drevets 2000); generalized social anxiety disorder (Klumpp
et al. 2013); bulimia (in response to pictures of food: Schienle
et al. 2009); and chronic back pain (Apkarian et al. 2005). In
chronic back pain patients, neuroimaged hyper-activity
reflected the chronicity (as opposed to the intensity) of the
pain (Baliki et al. 2006), but abnormalities in other brain
regions, such as reduced grey matter volumes in dorsolateral
frontal cortex, were also noted.

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Insula hyper-reactivity has been interpreted as evidence

that individuals with anxiety and depressive disorders overview interoceptive states as threatening or negative, respectively (Paulus and Stein 2006, 2010). In this regard, increased
metabolism in the insula has been linked with superior processing of interoceptive signals that have been associated with
anxiety specific arousal symptoms (Dunn et al. 2010).
Neuroimaged hypo-activity in the right insula has been
reported in autism and bilaterally in anorexia nervosa. In
autism it is thought to represent dysfunctional connectivity
patterns (Uddin and Menon 2009), while in anorexia nervosa
it is hypothesized to result from a disconnection between the
system responsible for adaptation to the external environment
and the system responsible for internal homeostasis (Nunn
et al. 2011).
Studies have linked neuroimaged activity levels in the
insula with patient response to treatment. For example, methamphetaminedependent patients who had just completed a
28-day rehabilitation program were more likely to relapse if
they had relatively less activity in multiple brain regions that
included the right insula during a simple two-choice task that
required the prediction of the location of a stimulus on a
computer screen (Paulus et al. 2005). Low levels of
neuroimaged activity in the anterior insula suggested a
better response to cognitive behavioral therapy in patients with major depression, while higher levels of
activity were associated with a better response to medication (McGrath et al. 2013).

Are Insula Contributions to Cognition Generic

or Task-Specific?
Table 1 lists behavioral functions (direct quotes) attributed to
the insula from 22 functional neuroimaging, lesion/behavior
change, electrical stimulation, and review studies. Although
not exhaustive, it demonstrates how such descriptions of the
behavioral function of insula have covered: (a) generic interoceptive reception/emotional feeling roles (seven studies); (b)
task-specific cognitive processing (ten studies); and (c) a
specific role in conscious awareness (five studies). The generic interoceptive reception/emotional feeling role pertains to
the encoding of changes in autonomic reactivity in response to
the salience (Seeley et al. 2007), homeostatic incongruence, or
challenging nature of the task. Examples of this generic interpretation of involvement in cognition include that: (a) insula
involvement in speech production, as indicated by increased
neuroimaged activity levels, likely reflects the response of the
autonomic nervous system to increased ventilation requirements (Ackermann and Riecker 2010); (b) anterior insula
cortex neuroimaged hyper-activations in perception studies
reflects increased sympathetic effects in response to the salience or challenging nature of the stimuli (Sterzer and

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83

Table 1 Behavioral functions ascribed to the insula from selected studies

Study

Function ascribed to the insula

Type of study

Ackermann and Riecker (2010)a posterior insularespiration-related metabolic (interoceptive) states rostral
componentsautonomic functions (p. 419)
part of the brain language area(and/or) verbal motivation and verbal affect (p. 79)
Ardila (1999)b
Baier et al. (2011)b
generation of speech motor execution (p. 1429)
b
Bamiou et al. (2003)
sound detectionallocating auditory attentionprocessing novel versus familiar
auditory stimuliauditory temporal processingmusic appreciation (p. 153)
tracking of risk (p. 645)
Bossaerts (2010)b
Brass and Haggard (2010)a
provide interoceptive signals that play an essential role in evaluating the
consequences of intentional action (p. 603: anterior insula)
neural correlate of consciousness (p. 59)
Craig (2009)c
Critchley et al. (2004)c
explicit awareness of, internal bodily processes (p. 193: right anterior
insula/operculum)
feelings of emotion (p. 260)
Damasio (2003)a
Dronkers (1996)b
motor planning of speech (p. 159)
Flynn et al. (1999)b
cardiovascular, gastrointestinal, vestibular, olfactory, gustatory, visual, auditory,
somatosensory, and motor modulationconditioned aversive learningpain
perception, stress induced immunosuppression, mood stability, sleep, and
language (p. 55)
Ibaez et al. (2010)a
coordination between internal and external information through emotional
subjective awareness (p. 397)
limb ownershipbeing involved in a movement (p. 411: right insula)
Karnath and Baier (2010)b
Manes et al. (1999b)b
verbal memory (p. 534: left insula)
c
Medford and Critchley (2010)
integrated awareness of cognitive, affective, and physical state (p. 540:
anterior insula)
interoceptive awareness, emotional responses,empathic processes
Menon and Uddin (2010)c
high-level cognitive control and attentional processes (p. 655).
conscious cue-induced urges (p. 61)
Naqvi and Bechara (2008)c
Penfield and Faulk (1955)a
gastric motility and stomach tone (p. 469)
Ronchi et al. (2012)b
perseveration (p. 588: right insula)
Seeley et al. (2007)a
personal salience, whether cognitive, homeostatic, or emotional, that requires
changes in sympathetic tone (p. 2349)
Small (2010)b
integrated oral sensory region that plays a role in feeding behavior (p. 551)
Stephani et al. (2011)a
somato-and viscerosensory cortex (p. 137)
a

Studies that implicate the insula in generic interoceptive reception and/or emotional feeling states

Studies that implicate the insula in specific cognitive tasks

Studies that implicate the insula in conscious awareness

Resting state connectivity shows temporally correlated activity in neural regions in the absence of a stimulus/task

Kleinschmidt 2010); and (c) anterior insula contributions to

intentional action reflect the provision of interoceptive information to be used in the evaluation of action consequences
(Brass and Haggard 2010).
Alternative explanations are that the insula plays a specific
role in each of these cognitive activities. Examples of other
task-specific behaviors attributed to the insula include feeding
behavior (Small 2010) and risk assessment (Bossaerts 2010).
Five studies in Table 1 implicated the insula in conscious
awareness including Craig (2009), Medford and Critchley
(2010), and Menon and Uddin (2010), but the issue remains
contentious. Cognitive behaviors like consciousness and intentional action have well-established relationships with other
regions of the cerebral cortex, especially prefrontal cortex.

Review
Review
Lesion analysis
Review
Review
Review
Review
Functional neuroimaging
Review
Lesion analysis
Review

Review
Review
Lesion analysis
Review
Review
Review (drug addiction)
Electrical stimulation
Lesion analysis
Resting state connectivityd
Review
Electrical stimulation

Changes in autonomic reactivity associated with the insula

typically occur independently of consciousness or volition. It
has been argued that on occasion some autonomic reactivity
can be represented in our consciousness. For example,
Critchley et al. (2004) used a heartbeat detection task whereby
participants had to judge if tonal feedback was synchronous (<
150 ms) with a heartbeat or was delayed (> 500 ms).
Participant accuracy was correlated with neuroimaged activity
levels in the right anterior insula. While this has been
interpreted as indicating conscious awareness of interoceptive
signals, such awareness is typically vague, especially in comparison with that of somatic sensations (Aziz et al. 2000). As
illustration, participants in heartbeat monitoring tasks only
identify about one third of the heartbeats recorded (Caseras

84

et al. 2013). Other aspects of autonomic nervous system

activity (e.g., changes in pupil dilation) are not represented
in conscious awareness. When interoceptive signals can vary
by location, as in pain, neuroimaged activity encoded in the
insula relates more to intensity than the spatial location of the
stimulus (Oshiro et al. 2009).

Summary
A synthesis of lesion/behavior change, stimulation, and functional neuroimaging studies of the insula suggests the following observations on its behavioral function:
1. The insula is commonly divided into anterior and posterior subregions. Some investigators also include a medial
subregion. There is not consistent agreement as to the
boundaries or the functional specialization of these subregions across studies. Divisions between anterior and
posterior subregions have been made on the basis of
cytoarchitechtonics or the central insular sulcus.
2. Historically conceived as a primary gustatory region, it is
now apparent that the insula processes a much wider
range of sensory stimuli. Most especially, it is the primary
reception area in the cerebral cortex for interoceptive
sensory information from the internal milieu associated
with changes in autonomic nervous system activity.
3. Lesion studies have implicated insula lesions with cognitive impairment across all neuropsychological domains,
but especially that of language. Although aphasic disturbances of various types frequently accompany dominant
insula injury, the exact role of the insula in language
processing remains unclear, as most cases involved lesions that extended into perisylvian language regions.
4. As the insula is the primary reception region of the brain
for interoceptive information reflecting changes in autonomic nervous system activity levels, it is considered as
critical for emotional feelings in accord with the JamesLange theory of emotion. This role was historically associated with the limbic system.
5. Lateralization effects in the insula reflect the wellestablished pattern of language processing in the dominant hemisphere, but also right sided processing of sympathetic effects and (possibly) left sided processing of
parasympathetic effects.
6. Neuroimaging has linked abnormal grey matter volumes
or hyper-and hypo-activity levels in the insula with
schizophrenia, eating disorders, anxiety and mood disorders, conduct disorder, autism, addiction, and chronic
pain. The significance of these abnormal activity patterns
remains theoretical.
7. Functional neuroimaging studies frequently show hyperactivations in the insula, typically in conjunction with

Neuropsychol Rev (2014) 24:7787

anterior cingulate cortex, for all manner of cognitive tasks,

including those involving perception, intentional action,
and consciousness. It is unlikely that the insula has a taskspecific function in so many cognitive activities, especially as they have well-established links with other brain
regions. The hyper-activations likely reflect generic
changes in autonomic nervous system activity in response
to the salient, homeostatically incongruent, or challenging
nature of the task. Autonomic nervous system activity
primarily occurs below the level of conscious awareness.
General recommendations for future studies on the insula
are: (a) to provide specific information as to how subregions
of the insula were determined; and (b) to account for the
generic autonomic nervous system effects during a given
experimental procedure, before invoking task-specific roles.

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