1

In: Turgut, M., Yurttas, C, & Tubbs, R.S. (eds) (2018), Island of Reil (Insula) in
the Human Brain - Anatomical, Functional, Clinical and Surgical Aspects. New
York: Springer.

Chapter 10:

Participation of the Insula in Language

Alfredo Ardila
Florida International University
Miami, Florida, USA
ardilaa@fiu.edu

Abstract
Since the beginning of aphasia history it has been known that the insula
has a crucial role in language. The interest in the insula somehow
decreased during the 20th century, because it has not directly included in
the perisylvian language area proposed by Dejerine. During the late 20th
century it was proposed that the insula has a crucial role on speech
praxis. Since Wernicke it has been observed that insula damage is
frequently associated with aphasia; different types of aphasia have been
reported in cases of insula damage: motor planning and organization of
speech in Broca aphasia, repetition defects associated with conduction
aphasia, and the word-deafness component of Wernicke aphasia. Mutism
and oral apraxia have also been reported to be associated with left insula
pathology. Neuroimaging studies have corroborated that the insula has
extensive brain connection, including not only the frontal motor language
area, but also posterior language areas. Anatomical connections of the
insula point also to an important viscero-limbic role, and it may
accordingly be suggested that the insula is involved in verbal motivation.
 2

Key words: insula; language; apraxia of speech; language understanding;
fMRI; Broca aphasia; conduction aphasia; Wernicke aphasia;

`Unfortunately, the problem of insular aphasia, which would be so

important for our considerations, has not so far been clarified by
clinical observation. Meynert, de Boyer, Wernicke himself and others
maintain that the insula belongs to the speech area, while Bernard
and others, following Charcot, emphatically deny such a relation.
Nothing decisive concerning this problem emerged from Naunyn’s
survey. Although it seems highly probable that lesions of the insula
cause speech disorders (not only because of anatomical contiguity to
the so-called center), it is nevertheless impossible to state whether
the speech disorder is of a specific type and if so of what type.’ (Freud
1891, pp. 12-13).

Introduction
The controversies regarding the participation of the insula in language
has extended for over 100 years. During the 19th century it was assumed
that the insula had an evident participation in language. During the early
20th century Dejerine (1914) proposed the concept of “brain language
area”, including three cortical regions: (1) left frontal (posterior part of the
foot of F3, the frontal operculum, and the immediate surrounding zone,
including the foot of F2, and probably extending to the anterior insula), (2)
left temporal (encompassing the posterior first and second temporal gyri),
and (3) parietal (the angular gyrus) areas. This proposal was rapidly
accepted by virtually all researchers in the aphasia area (e.g. Head 1920;
Nielsen 1936; Luria 1966; Benson & Geschwind, 1971; Goodglass &
Kaplan, 1972; Albert et al. 1981). It was assumed that the brain language
zone corresponded in consequence to the perisylvian area of the left
 3

hemisphere; and the insula was forgotten. Until the end of the 20th century
the insula virtually disappeared as a language area. However, during the
last couple of decades the interest in the potential participation of the
insula in language has re-emerged.
The insula indeed is locate in the center of the language area. The
anterior segment of the insula extends to and interfaces with Broca’s area
while its posterior elements adjoin Wernicke’s area. Because of the
location in the epicentre of the human language area it is understandable
that the insula may play a significant role in language, as it was assumed
during the 19th century.

Involvement of the insula in aphasia

Brain pathology in cases of aphasia frequently includes an insular
extension (Gasquoine, 2014). Ever since Wernicke (1874), an extensive
body of clinical research has supported that the insula is frequently
involved in so-called “major” or perisylvian aphasic syndromes: Broca
aphasia, conduction aphasia, and Wernicke aphasia. Wernicke himself
directly related insula damage with conduction aphasia; noteworthy, the
first reported case of conduction aphasia presented an insular pathology.
Involvement of the anterior part of the insula in Broca aphasia was
reported many years ago, since the very beginning of the 20th century
(Bernheim, 1900; Dejerine, 1914). The word-deafness component of
Wernicke aphasia, on the other hand, has been since long time ago
associated with posterior insula damage (Liepmann & Storck, 1902).
Thus, the insula is frequently involved in these three major aphasia
syndromes.
Pathology involving only the insula, nonetheless, is rarely
reported. Alexander et al. (1987) presented two cases with CT evidence
of pathology limited to the left insula and subjacent extreme-external
capsules. An aphasia with mildly paraphasic production and agraphia was
 4

noted in both patients. Nielsen and Friedman (1942) reported several
cases from the literature with autopsy suggesting that the left insular
damage was associated with mild aphasia. Starkstein et al. (1988)
observed crossed aphemia associated with a right insular lesion. Fifer
(1993) reported a patient presenting a lesion involving the right insula as
well as the adjacent white matter; this patient presented with a unilateral
disorder in processing auditory information when speech materials were
presented to the left ear. Habib et al. (1995) found a case of bilateral
insula pathology, extending to a small part of the striatum on the left side,
and to the temporal pole on the right hemisphere. It was observed mutism
lasting several weeks; this patient did not respond to any auditory
stimulation, and made no effort to communicate. Baratelli et al. (2015)
reported a herpes simplex encephalitis patient with a left hemisphere
lesion limited to the left insula and to the left anterior parahippocampal
region. The aphasia type and language recovery were analyzed.
The language deficit was of fluent type, without phonological impairment,
and showed a good but incomplete recovery after four months.
In an extensive study Lemieux et al. (2012) reviewed the,
consecutive stroke database from two centers to identify patients insular
ischemic strokes during a four year period. Clinicoradiological correlation
was performed by distinguishing insular ischemic strokes involving the
anterior or posterior insular cortex. The authors identified 7 patients from
our institutions and 16 previously published cases. Infarcts were limited to
the anterior (n = 4) or posterior (n = 12) insula or affected both (n = 7).
The five most frequent symptoms were somatosensory deficits (n = 10),
aphasia (n = 10), dysarthria (n = 10), a vestibular-like syndrome (n = 8)
and motor deficits (n = 6). So, aphasia is no question a frequent clinical
manifestation of insular ischemic stroke.

Mutism has been sometimes reported in cases of insular pathology.

Transient mutism may be found in cases of left inferior motor cortex
 5

damage extending to the insula (Schiff et al. 1983, Alexander et al. 1989);
lasting mutism, on the other hand, appears to be associated with bilateral
lesions of the frontal operculum and anterior insula (Sussman et al., 1983;
Cappa et al., 1987; Groswaser et al., 1988, Pineda & Ardila, 1992). A
transient mutism and persistent auditory agnosia due to two successive
ischemic infarcts mainly involving the insular cortex on both hemispheres
were reported by Habib et al. (1995). Shuren (1993) observed a patient
who presented language initiation defects as a result of a left anterior
insular infarct. The author proposed that dominant hemisphere anterior
insular lesions impair the speech initiation loop. Alexander et al. (1989)
suggested that left cortical and sub-cortical opercular lesions frequently
result in a total speech loss associated with a right hemiparesis. Right
hemiparesis has a good recovery, but the oral apraxia responsible for the
mutism improves only slowly.
Regardless that most of the reports of insular aphasia refer to
vascular aphasia, other insular pathologies, such as insular tumor can
also be associated with aphasia (Duffau et al., 2001)

Apraxia of speech

The interest in the potential involvement of the insula in language re-

appeared during the late 20th; this renewed interest in the insula was
significantly associate with the publication of Dronkers’ paper “A new
brain region for coordinating speech articulation” (Dronkers, 1996). In this
paper it is reported that the left precentral gyrus of the insula is involved in
motor planning of speech. Twenty-five stroke patients with a disorder in
motor planning of articulatory movements, which Dronkers labelled as
`apraxia of speech’, were compared with 19 individuals without such
deficit. A robust double dissociation was observed. All patients with
articulatory planning impairments presented lesions including the anterior
 6

insula. This area was completely spared in all patients without these
articulatory defects. The conclusion was obvious: anterior insula
represents the crucial brain area in motor planning and organization of
speech. Verbal apraxia and agrammatism represent the two cardinal
characteristics of Broca aphasia, and hence, the anterior insula is directly
involved in Broca aphasia. Several reports have confirmed the
involvement of the anterior insula in speech praxis. During the following
years, several papers were published emphasizing the participation of the
insula in language (Ardila, 1999; Ardila, Benson & Flynn, 1997).For
instance, Nagao et al (1999) report a patient with apraxia of speech who
showed an acute infarct limited to the precentral gyrus of the insula.

In a critical study, 33 left hemisphere stroke patients with varying

degrees of speech impairment were asked to perform multiple repetitions
of single words; these words varied along three separate dimensions:
number of syllables, degree of articulatory travel (i.e., change between
places of articulation for consonants), and presence/absence of an initial
consonant cluster Baldo et al., 2011). The authors found that precentral
gyrus of the insula was critical for performance on the articulation task
across all three conditions. The authors concluded that the left precentral
gyrus of the insula is a critical area for intra- and inter-syllabic
coordination of complex articulatory movements, prior to end-stage
execution of speech commands.

Neuroimagining studies

Contemporary neuroimaging technique research has supported the active

involvement of the insula in linguistic processes. Activation of the insula
has been demonstrated in a diversity of linguistic tasks, including during
word generation task performance (Baker et al. 1997, McCarthy et al.
1993) and naming (Price et al. 1996). Lexical knowledge and word
retrieval evidently represent central linguistic processes frequently
 7

impaired in aphasia. Activation of the insula has also been reported
during phonological decision tasks (Rumsey et al. 1997). The insula has
also been found to be involved in the motor aspects of speech production
(Ackermann & Riecker, 2004) and naming (Owen et al., 2004).

Using fMRI it has been found that the insular cortex is sensitive to
phonological processing, particularly sublexical spelling-sound translation
(Borowsky, et al., 2006). It has been also reported that processing of
semantic violations relied primarily on the mid-portion of the superior
temporal region bilaterally and the insular cortex bilaterally (Friederici et
al., 2003)
Rousseaux et al. (1990), correlated linguistic impairment with
cerebral blood flow, and related verbal comprehension, naming, and
paraphasias to an asymmetry index of the insula and the lenticular
nucleus. The insula would accordingly not participate in some unique and
isolated language dimension, but rather would be active in different
linguistic aspects. It may therefore be concluded that contemporary
neuroimaging studies support to the assumption of a significant
participation of the insula in language. The insula would be involved not in
a single linguistic activity, but in several verbal processes simultaneously.
The anterior portion of the insula would be involved in the organization
and planning of language articulation, while the middle and posterior
portions would be involved with lexical knowledge, word retrieval,
language understanding, and phonological discrimination. By the same
token, functional imaging studies demonstrate that the insula participates
in several key auditory processes, such as allocating auditory attention
and tuning in to novel auditory stimuli, temporal processing, phonological
processing and visual-auditory integration (Bamiou, Musiek & Luxon,
2003). Zaccarella and Friederici (2015) using fMRi attempted to specify
the involvement of the insula during visual word processing using a sub-
regional parcellation approach. They found that word compared to letter
 8

string processing was strongly sub-regional sensitive within the anterior-
dorsal cluster only, and was left-lateralized.
Using graph theoretical analysis of functional MRI (fMRI) data in
healthy subjects, Fuertinger et al. 2015) quantified the speech network
topology by constructing functional brain networks of increasing
hierarchy. The authors identified a segregated network of highly
connected local neural communities (hubs) in the primary sensorimotor
and parietal regions. Compared to other tasks, speech production was
characterized by the formation of six distinct neural communities including
the prefrontal cortex, insula, putamen, and thalamus, which collectively
forged the formation of the functional speech connectome.

Contemporary meta-analytic studies

Recently, two major papers were published presenting meta-analyses of
insula fMRI activation, assessed using the Activation Likelihood Estimate
(ALE) method (Oh, Duerden, & Pang, 2014; Ardila, Bernal & Rosselli,
2014). In the first meta-analysis 42 fMRI studies including 639 healthy
adults were performed, comparing insula activation during performance of
language (expressive and receptive) and speech (production and
perception) tasks. Both tasks activated bilateral anterior insula. The
authors found that speech perception tasks preferentially activated the
left dorsal mid-insula, whereas expressive language tasks activated left
ventral mid-insula.
The second meta-analytic study used as well the Activation
Likelihood Estimation (ALE) technique was developed. Search conditions
were: (1) studies reporting insula activation; (2) studies using fMRI; (3)
context: normal subjects; (4) activations: activation only; (5) handedness:
right-handed subjects; (6) age 20–60 years; (7) domain: cognition,
subtype: language. By means of the BrainMap functional database, 26
 9

papers corresponding to 39 paradigms, and including 522 participants
were selected. Thirteen different activation clusters were found. Insula
connections included not only the classical areas involved in language
production (Broca’s area) and language understanding (Wernicke’s area)
but also areas involved in language repetition (supramarginal gyrus) and
other linguistic functions such as the left prefrontal lobe involved in
language control, and the Brodmann area (BA) 37, involved in lexico-
semantic associations. The first cluster includes the left claustrum (the
insular subcortical gray matter). The second cluster includes the anterior
cingulate gyrus (BA24) (involved in motor organization—motor
preparation/ planning, cognitive/motor inhibition—and language initiative)
and left BA6 (medial frontal gyrus). BA6 includes the supplementary
motor area (SMA). Cluster 3 included the right insula and the insular
subcortical gray matter (claustrum) and indicates an integrated activity of
the left and right insula. Cluster 4 referred to left BA7 (superior parietal
lobe). Therefore, the insula would also be part of the language system
related to some contextual and motor learning aspects of speech. The
following cluster involved BA37 (posterior inferior temporal gyrus, middle
temporal gyrus, and fusiform gyrus) and the cerebellar culmen. The
following two clusters (6 and 7) referred to two areas traditionally involved
in language: the left BA22 (superior temporal gyrus—part of Wernicke’s
area), and the left BA40 (supramarginal gyrus). Clusters 8 and 9 may
contribute to the motor aspects of speech and the attention control of
language. Cluster 10 on the other hand is similar to Cluster 7 and
includes theleftBA40 (inferior parietal lobe).
It was concluded that the insula represents a core area in
language processing and it is related, not just with language production
functions but also with language understanding processing. It could
consequently be conjectured that the insula is a core hub for language.
 10

Its strategic location between the anterior and posterior language areas
would be crucial to play a language coordinating function.

Language motivation and affect

Several reports have suggested that the insula may play an important role
in verbal motivation and affect. Habib et al. (1995) have proposed that
bilateral damage to the insula would disrupt the motivational mechanisms
that lead to the motoric production of human communication, by depriving
them of connections with various limbic structures. Hence, insular
damage would result in an impairment in linguistic motivation. Anatomical
studies suggest that the insula could well act as a cortical representation
of the limbic nervous system and, as such, may provide a direct input to
the brain emotional, that could in turn monitoring the affective tone of
language output. Within the same vein Beaty et al. (2017) have included
the right insula in an extended brain network involved in metaphor
production.
It can be considered that the insula is not only involved in
language, but also participates in emotional networks (Closet al., 2014)

Conclusion

Since the 19th century it has been known that the insula has a crucial role
in language processes. However, the interest in the insula somehow
decreased during the 20th century, because it has not directly included in
the perisylvian language area proposed by Dejerine. Since the last
decade of the 20th century the interest in understanding the participation
of the insula in language re-emerged. A crucial role on speech praxis was
initially postulated. It was also emphasized that the insula damage
frequently results in aphasia. Among the various language disturbances
associated with damage in the left insula are: motor planning and
 11

organization of speech in Broca aphasia, repetition defects associated
with conduction aphasia, and the word-deafness component of Wernicke
aphasia (Ardila, 1999; Ardila et al., 1997). Mutism and oral apraxia have
been reported to be associated with left insula pathology. Recent
neuroimaging studies have illustrated that the insula has extensive brain
connection, including not only the frontal motor language area, but also
posterior language areas, including the temporal and parietal lobes.
Anatomical connections of the insula point also to an important viscero-
limbic role, and it may accordingly be suggested that the insula is
involved in verbal motivation.

It can be concluded that the brain language are should be

reconsidered to include not only the perisylvian area of the left
hemisphere but also the insula.

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