NeuroRehabilitation 32 (2013) 437–443 437

DOI:10.3233/NRE-130866
IOS Press

Review Article

Anatomy of the vestibular system: A review

Sarah Khana,∗ and Richard Changb
a JFK Johnson Institute of Rehabilitation, Edison, NJ, USA
b Mount Sinai Medical Center, New York, NY, USA

Abstract.
INTRODUCTION: A sense of proper sensory processing of head motion and the coordination of visual and postural movements
to maintain equilibrium is critical to everyday function. The vestibular system is an intricate organization that involves multiple
levels of sensory processing to achieve this goal.
PURPOSE: This chapter provides an overview of the anatomical structures and pathways of the vestibular system.
SUMMARY: The five major vestibular structures are located in the inner ear and include: the utricle, the saccule, and the lateral,
superior, and posterior semicircular canals. Hair cells on the neuroepithelium of the peripheral vestibular organs carry sensory
impulses to primary processing centers in the brainstem and the cerebellum. These areas send input via ascending and descending
projections to coordinate vital reflexes, such as the vestibuloocular reflex and the vestibulospinal reflex, which allow for the
proper orientation of the eyes and body in response to head motion. Specific connections regarding higher level cortical vestibular
structures are poorly understood.
CONCLUSION: Vestibular centers in the brainstem, cerebellum, and cerebral cortex function to integrate sensory information
from the peripheral vestibular organs, visual system, and proprioceptive system to allow for proper balance and orientation of the
body in its environment.

Keywords: Vestibular apparatus, kinocilium, stereocilia, utricle, saccule, semicircular ducts, macula, cupula, striola, vestibular
nuclear complex, vestibuloocular reflex, vestibular spinal reflex

1. Introduction forces on the body. This information is processed by

vestibular centers in the brain to allow the body to
The vestibular system is a complex sensory organi- maintain balance and proper spatial orientation during
zation which involves the communication between the movement, as well as the correct processing of visual
peripheral vestibular apparatus, the ocular system, pos- images during motion.
tural muscles, the brainstem, cerebellum and the cortex.
Small structures in the inner ear make up the vestibu-
lar apparatus and detect head motion and gravitational 2. Bony and membranous labyrinth

∗ Address for correspondence: Sarah Khan, JFK Johnson Institute

The peripheral vestibular system is located in the
of Rehabilitation, 65 James Street, Edison, NJ 08818, USA. Tel.: +1 inner ear and consists of a bony labyrinth and a mem-
732 321 7000; E-mail: dr.s.khan22@gmail.com. branous labyrinth. This system sits in the otic capsule

1053-8135/13/$27.50 © 2013 – IOS Press and the authors. All rights reserved
438 S. Khan and R. Chang / Anatomy of the vestibular system

Fig. 1. Bony & Membranous Labyrinth. The bony labyrinth consists of the cochlea, an oval cavity called the vestibule, and the semicircular
canals. The membranous labyrinth is contained within the bony labyrinth and consists of the utricle, the saccule, and the lateral, superior and
posterior semicircular ducts. The semicircular ducts end in a dilated area called the ampulla that contains the hair cell receptors.

in the petrous portion of the temporal bone. The bony tors called hair cells. These receptor cells are embedded
labyrinth consists of: the cochlea, an oval cavity called in a membrane of neuroepithelium. The basic structure
the vestibule, and the semicircular canals. The cochlea of the hair cell includes a single large kinocilium and
is a snail shaped structure containing the Organ of approximately 70–100 stereocilia on its apical end [10].
Corti, which is the receptor for hearing. The struc- The stereocilia are organized in rows that range from
tures of the bony labyrinth are filled with a fluid that the tallest, which are the closest to the kinocilium, and
is continuous with and similar in composition to cere- progressively decrease in size to the shortest stereocilia,
bral spinal fluid, known as perilymph [5]. This fluid is which are the farthest from the kinocilium. The kinocil-
drained by the perilymphatic duct into the adjacent sub- ium resembles a true cilium, but is non-motile and has
arachnoid space [9]. The membranous labyrinth houses a 9 + 2 microtubule doublet arrangement [10] (a central
the sensory epithelium and structures of the vestibular pair surrounded by 9 pairs). In contrast, the stereocilia
apparatus, and is suspended in the perilymph within the are made up of actin rich parallel filaments coated with
bony labyrinth. various isoforms of myosin [2].
Endolymph flows throughout the structures of the “Tip links” connect the tips of shorter stereocilia
membranous labyrinth and is similar in composition to to the body of their adjacent taller stereocilia [2].
intracellular fluid [5]. This fluid is produced by capillar- When head motion results in tilting of the stereocilia
ies in the stria vascularis [9] in the wall of the cochlear toward the kinocilium, shifting of the “tip links” causes
duct and absorbed by the endolymphatic sac [9]. The mechanical opening of the transduction channels result-
vestibular apparatus is composed of 5 organs: the utri- ing in an influx of K + . This results in a depolarization
cle, the saccule, and the lateral, superior, and posterior of the hair cell and opens Ca++ channels at the base of
semicircular ducts. The utricle and saccule are located the hair cell. Ca++ influx stimulates neurotransmitter
in the vestibule. The semicircular ducts are contained release into the synapses with afferent vestibular nerve
in the bony semicircular canals (Fig. 1). fibers increasing their firing rate. The bending of the
stereocilia away from the kinocilium decreases the “tip
3. Hair cells link” tension and results in mechanical closure of the
channel. This causes hyperpolarization of the hair cell
The vestibular system has two types of sensory neu- which closes the Ca++ channels and decreases the neu-
roepthelium, the macula and crista ampullaris. Both rotransmitter release [9], reducing the firing rate of the
structures contain rod-shaped sensory mechanorecep- vestibular nerve fibers (Fig. 2).
 S. Khan and R. Chang / Anatomy of the vestibular system 439

Fig. 2. Hair Cell “tip link” function. When head motion results in tilt-
ing of the stereocilia toward the kinocilium, shifting of the “tip links” Fig. 4. Utricle and Saccule. The stereocilia in the macula are oriented
causes mechanical opening of the transduction channels resulting in relation to a curvilinear line called the striola. This line is an area
in an influx of K and depolarization of the hair cells. The bend- of thinning in the utricle and an area of thickening in the saccule.
ing of the stereocilia away from the kinocilium decreases the “tip The kinocilia and stereocilia of the hair cells are oriented toward the
link” tension and results in mechanical closure of the channel causing striola in the utricle and away from the striola in the saccule.
hyperpolarization of the hair cell.

The interior of the macula is coated with a gelatinous

membrane embedded with small calcium carbonate
particles called otoliths, or otoconia. Vestibular recep-
tor hair cells project through this otolitic membrane.
Because the otoliths are denser than endolymph, grav-
ity is able to deflect the stereocilia of the hair cells when
the head is stationary [5]. Linear movement or tilting of
the head causes inertial drag and shearing force between
the otolitic membrane and the macular surface which
causes bending of the hair cells [10].
The stereocilia in the macula are oriented in relation
Fig. 3. Two Types of Hair Cells. Type I has a rounded base surrounded
by a nerve calyx of the afferent nerve fiber. Type II hair cells are
to a curvilinear line called the striola [13]. This line is an
columnar with bouton synaptic connections to their afferent fibers. area of thinning in the utricle and an area of thickening
in the saccule [10]. The kinocilia and stereocilia of the
There are two structural types of hair cells. Type I hair cells are oriented toward the striola in the utricle and
has a rounded base surrounded by a nerve calyx of the away from the striola in the saccule. This distribution
afferent nerve fiber. The majority of hair cells have a of hair cells in different directions means that various
Type II structure. They are columnar cells with bouton patterns of hair cell stimulation may occur based on the
synaptic connections to their afferent fibers. Type I hair degree to which head tilt occurs. Motion will stimulate
cells are associated with irregular afferents that have a one group of hair cells while inhibiting another group
high variability of resting discharge. Type II hair cells and yet, also have no effect on another group of hair
usually synapse on regular afferents with a low vari- cell receptors [13]. This intricate pattern of response is
ability of resting discharge. Both hair cell types also critical to relating accurate information regarding head
have efferent connections from vestibular nuclei that position to the central nervous system (Fig. 4).
modulate their sensitivity (Fig. 3) [9]. Another important physiologic property of the mac-
ula is that of adaptation. When the head tilt stimulus
4. Utricle and saccule remains beyond a few seconds, the bent hair cells and
the depolarized membrane potentials begin to return to
The utricle and saccule are structures of the static normal. This allows the hair cells to be responsive to
labyrinth that sense the orientation of the head in further positional changes [10].
space. They respond to linear acceleration, gravitational
forces, and tilting of the head. Each contains a sensory
neuroepithelium called the macula. The macula of the 5. Semicircular ducts
utricle senses motion in the horizontal plane, while the
macula of the saccule senses motion in the vertical plane The semicircular ducts have the same basic struc-
[2]. ture as the bony semicircular canals in which they are
440 S. Khan and R. Chang / Anatomy of the vestibular system

contained. They make up the kinetic labyrinth that

senses angular acceleration or rotation of the head and
are oriented at right angles to one another. The superior
and posterior ducts are aligned in a 45 degree angle to
the sagittal plane, and the lateral canals are aligned in
a 30 degree angle in the axial plane [6]. Contralateral
semicircular ducts oriented in the same plane are paired
as follows:

Semicircular Duct Pairs [12] Fig. 5. Ampulla of the Semicircular Ducts. The ampulla is a dilation
at the end of each of the semicircular ducts. It contains the sensory
Right Superior Left Posterior neuroepithelium, the crista ampullaris and is coated by the cupula,
Left Superior Right Posterior which is a gelatinous substance through which hair cells are embed-
Left Horizontal Right Horizontal ded. Rotational acceleration causes endolymph motion that displaces
the cupula, and bends the hair cells in the opposite direction of the
rotation [10].
This arrangement allows for a 3-dimensional vector
representation of rotational acceleration. Each duct is
sensitive to movement in its specific plane. rejection.” This means that simultaneous firing of both
The semicircular ducts open into the utricle. At the semicircular ducts of the pair is ignored by the central
end of each of the ducts there is a dilation called the nervous system. This situation may occur with eleva-
ampulla which contains the sensory neuroepithelium, tion of body temperature, as in a fever, and is not related
the crista ampullaris. It is coated by the cupula, which to motion. Finally, this physiologic arrangement also
is a gelatinous substance through which hair cells are allows for compensation for sensory overload [5].
embedded [9]. The crista ampullaris is histologically
similar to the macula. The cupula is thicker than the 6. Vestibular ganglion
gelatinous membrane of the macula, and also does not
contain otoliths [9]. The kinocilia of the hair cells in The vestibular ganglion, also known as Scarpa’s Gan-
the lateral ducts are oriented toward the utricle, and the glion, is located in the lateral portion of the internal
kinocilia of the superior and posterior duct hair cells are auditory meatus [11]. It is composed of close to 20,000
oriented toward the duct. bipolar cell bodies that receive afferent impulses from
Rotational acceleration causes endolymph motion the hair cells of the crista ampullaris and the maculae.
that displaces the cupula, and therefore bends the hair The vestibular ganglion is divided into a superior and
cells in the opposite direction of the rotation [2]. This an inferior division which are connected by an isthmus
results in opening of the ion channels and depolariza- [13]. The peripheral fibers of the superior division of the
tion of the hair cell which increases the firing of its vestibular ganglion terminate in the crista ampullaris of
afferent fibers. When the rotational velocity of the head the superior and lateral semicircular ducts, as well as
becomes constant, the cupula returns to an upright posi- the macula of the utricle. The macula of the saccule and
tion, and the membrane potential of the cell normalizes. the crista ampullaris of the posterior semicircular ducts
Rotational deceleration of the head results in cupula receive peripheral vestibular branches from the inferior
displacement in the same direction as the head move- section of the vestibular ganglion [13].
ment [2]. This closes the ion channels of the hair cell
which causes it to become hyperpolarized and results
in a reduction in afferent nerve firing (Fig. 5). 7. Vestibular nerve
Endolymph flow that causes excitation in one
semicircular duct will inhibit the hair cells of the con- Axons from the superior and inferior divisions of
tralateral duct it is paired with. There are a few main the vestibular ganglion merge to form the vestibular
advantages of this system. First, it allows for sensory nerve. It combines with the cochlear nerve to become
redundancy in the event there is pathology of one semi- the vestibulocochlear nerve. This nerve travels with the
circular duct. In this instance, vestibular input regarding facial nerve, nevus intermedius, and the labyrinth artery
movement in its plane will be received from the other through the internal auditory canal, which transverses
duct in its pair. Another advantage is “common mode through the petrous temporal bone to the posterior fossa
 S. Khan and R. Chang / Anatomy of the vestibular system 441

[5]. The nerve fibers travel past the cerebellopontine 9. Vestibulocerebellum

angle and enter the brainstem at the pontomedullary
junction. At this point, the vestibular nerve separates The role of the cerebellum in the vestibular sys-
from the cochlear nerve. The majority of the affer- tem is that it functions as an adaptive processor.
ent vestibular fibers project to the ipsilateral vestibular It monitors vestibular performance and will readjust
nuclear complex in the pons. Some of the nerve fibers vestibular input through inhibitory input as necessary
project to the flocculo-nodular lobe of the cerebellum [5]. The “vestibulocerebellum” is composed of the floc-
and the adjacent vermian cortex [11]. conodular lobe and the vermian cortex. The ipsilateral
cerebellum can send efferent information to bilateral
vestibular nuclei. It has projection fibers that go directly
8. Vestibular nuclear complex to the ipsilateral vestibular nuclei, and to the ipsilat-
eral fastigial nucleus. Axons from the fastigial nucleus
This complex is the primary processor of vestibular project to the contralateral vestibular nuclei via the
input and consists of four major nuclei: medial, supe- juxarestiform body [11]. This area has an important
rior, lateral, and inferior [13]. These nuclei are also role in the generation of postural reflexes and orienting
known as: Schwalbe, Bechterew, Deiter, and descend- behaviors.
ing, respectively [6, 14]. They are located under the The cerebellar flocculus adjusts the gain of the
floor of the fourth ventricle and extend from the rostral vestibuloocular reflex [5]. The cerebellar nodulus
medulla to the caudal pons in two major columns. adjusts the duration of the vestibuloocular reflex and
The medial vestibular nucleus is the largest and is also involved in processing afferent activity from
makes up the medial column. The lateral column con- the maculae [5]. The anterior superior vermis plays a
sists of the superior, lateral, and inferior vestibular role in regulating the vestibulospinal reflex by encod-
nuclei [13]. The medial vestibular nucleus receives ing vestibular signals and proprioceptive input from the
afferents from the crista ampullaris of the lateral semi- axial muscles.
circular ducts. Ascending axonal fibers travel via the
medial longitudinal fasciculus to the motor nuclei of
the extraocular muscles to mediate the vestibuloocu- 10. Higher level cortical areas
lar reflex. It also functions in controlling the vesibular
spinal reflex via bilateral descending projections in the Specifics regarding the complex cortical vestibular
medial vestibulospinal tract to the cervical spinal cord connections are not clearly understood and consen-
to allow for the coordination of head and neck motion. sus regarding the location of a vestibular cortex still
The superior vestibular nucleus also receives vestibu- has yet to be defined. In studies of primates, the main
lar afferent input from the crista ampullaris of the cortical areas receiving vestibular stimuli include the
superior and posterior semicircular ducts [6]. Like the parietoinsular vestbular cortex (PIVC), area 2v of the
medial vestibular nucleus, it sends ascending effer- intraparietal sulcus, and area 3av in the central sulcus
ent fibers to the extraocular muscles via the medial [3]. The PIVC appears to be the main vestibular cortical
longitudinal fasciculus in order to coordinate the region in primates as information from other vestibular
vestibuloocular reflex [6]. cortical areas is integrated here [3].
The lateral vestibular nucleus contains the largest cell Studies of humans have suggested that the main
bodies of all the vestibular nuclei. It receives affer- cortical processing region is most likely in or near
ent input from the crista ampulla, the maculae, and the parietal or insular cortex. Kahane electrically
the vestibulocerebellum. Efferent projections from this induced vestibular symptoms in humans and found
nucleus become the lateral vestibular tract in the ipsi- a lateral cortical temporoparietal area that he called
lateral spinal cord. This tract functions in the vestibular the “temporo-peri-sylvian vestibular cortex” and
spinal reflex by coordinating reflexive tone in the trunk suggested it corresponds to the PIVC in primates [7].
muscles and proximal extensors of the limbs to maintain The use of activation likelihood estimation of cortical
posture and balance [6]. activation on neuroimaging in response to multiple
The inferior vestibular nucleus receives afferent vestibular stimuli by Lopez found the main areas of
information from the maculae of both the utricule and convergence were mainly in the retroinsular cortex, but
the saccule. This nucleus has projections that go to the also in the parietal operculum, and the posterior insula
other three vestibular nuclei and to the cerebellum [6]. [7]. In another study by Eulenburg, PET and fMRI
442 S. Khan and R. Chang / Anatomy of the vestibular system

activation to vestibular stimuli demonstrated that the

right hemispheric parietal opercular area was the most
consistent area of activation [4].
Vestibular connections in the thalamus and hip-
pocampus are also thought to exist. Animal studies have
shown vestibular neurons in multiple thalamic regions.
In humans, it is believed that some ascending vestibular
fibers make connections in the ventral posterior nucleus
of the thalamus prior to reaching the cortex [8]. The
hippocampus is believed to play a critical role in the
processing of spatial orientation and spatial memory.
Vestibular input regarding the movement of the head
and body is thought to be necessary for this function
[8]. Further studies are necessary to gain a better under-
standing of the higher level cortical connections and
functions of the vestibular system.

11. Vestibuloocular reflex Fig. 6. Vestibuloocular Reflex. Head turning to the right causes the
endolymph flow in the ampulla of the semicircular ducts to deflect
The vestibuloocular reflex coordinates eye move- the cupula to the left. This causes depolarization of the hair cells
in the right ampulla and an increase in the firing frequency in the
ment in order to stabilize retinal images during head
afferent fibers of the right vestibular nerve. The nerve sends impulses
rotation. It involves a three neuron reflex arc from the to the ipsilateral superior and medial vestibular nuclei. Excitatory
semicircular ducts to the vestibular nuclei and then to impulses are transmitted in the medial longitudinal fasciculus to the
the extraocular muscles to cause conjugate eye motion right oculomotor nuclei, and in the ascending tract of Deiters to the
left abducens nuclei. This results in ipsilateral medial rectus and con-
in a direction opposite to head turning [3]. tralateral lateral rectus contraction which produces eye movement to
For instance, when the head turns to the right, the left.
endolymph flow in the ampulla of the semicircular
ducts will deflect the cupula to the left. This causes cles by the brainstem, and cerebellum in order for the
depolarization of the hair cells on the right, and hyper- maintenance of posture and balance. It involves both
polarization of the left hair cells. This results in an the lateral and medial vestibular spinal tracts. The lat-
increase in the firing frequency in the afferent fibers eral vestibular tract is the main pathway and originates
of the right vestibular nerve and impulses are sent to in the lateral vestibular nucleus. Efferent vestibular sig-
the ipsilateral superior and medial vestibular nuclei and nals in response to input from the macula of the otolitic
the cerebellum. Excitatory impulses are transmitted in organs to the lateral vestibular nucleus are carried in
the medial longitudinal fasciculus to the right oculo- this tract which projects ipsilaterally in the spinal cord
motor nuclei, and in the ascending tract of Deiters to to neurons of all spinal levels. It produces monosynaptic
the left abducens nuclei [5]. This results in ipsilateral activation of ipsilateral trunk and proximal limb exten-
medial rectus and contralateral lateral rectus contrac- sors and disynaptic inhibition of contralateral proximal
tion which produces eye movement to the left (opposite extensors [10].
to head turning). If eye velocity and head velocity do Angular rotation of the head sensed by the semicircu-
not match, input from the cerebellar flocconodular lobe lar ducts is transmitted to the medial vestibular nucleus
is sent to the vestibular nuclei to modify their firing rate where the medial vestibulospinal tract originates. This
to correct this discrepancy (Fig. 6). tract projects bilaterally to motor neurons in the cervi-
cal spinal cord. It activates cervical axial muscles that
coordinate head and neck motion.
12. Vestibulospinal reflex Another reflex that is related to the vestibulospinal
reflex is the vestibulocollic reflex. This activates neck
This reflex involves many complex connections muscles that function to stabilize the head and keep
which integrate input from the macula, crista it properly oriented in space. The exact pathways that
ampullaris, visual system, and axial and limb mus- contribute to this reflex have yet to be known [5].
 S. Khan and R. Chang / Anatomy of the vestibular system 443

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