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Anatomy of the vestibular system: A review

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DOI: 10.3233/NRE-130866 · Source: PubMed

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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

The vestibular system is a complex sensory organi-
zation which involves the communication between the
peripheral vestibular apparatus, the ocular system, pos-
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
vestibular centers in the brain to allow the body to
maintain balance and proper spatial orientation during
movement, as well as the correct processing of visual
images during motion.
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
labyrinth consists of: the cochlea, an oval cavity called
the vestibule, and the semicircular canals. The cochlea
is a snail shaped structure containing the Organ of
Corti, which is the receptor for hearing. The struc-
tures of the bony labyrinth are filled with a fluid that
is continuous with and similar in composition to cere-
bral spinal fluid, known as perilymph [5]. This fluid is
drained by the perilymphatic duct into the adjacent sub-
arachnoid space [9]. The membranous labyrinth houses
the sensory epithelium and structures of the vestibular
apparatus, and is suspended in the perilymph within the
bony labyrinth.
Endolymph flows throughout the structures of the
membranous labyrinth and is similar in composition to
intracellular fluid [5]. This fluid is produced by capillar-
ies in the stria vascularis [9] in the wall of the cochlear
duct and absorbed by the endolymphatic sac [9]. The
vestibular apparatus is composed of 5 organs: the utri-
cle, the saccule, and the lateral, superior, and posterior
semicircular ducts. The utricle and saccule are located
in the vestibule. The semicircular ducts are contained
in the bony semicircular canals (Fig. 1).
3. Hair cells
The vestibular system has two types of sensory neu-
roepthelium, the macula and crista ampullaris. Both
structures contain rod-shaped sensory mechanorecep-
tors called hair cells. These receptor cells are embedded
in a membrane of neuroepithelium. The basic structure
of the hair cell includes a single large kinocilium and
approximately 70–100 stereocilia on its apical end [10].
The stereocilia are organized in rows that range from
the tallest, which are the closest to the kinocilium, and
progressively decrease in size to the shortest stereocilia,
which are the farthest from the kinocilium. The kinocil-
ium resembles a true cilium, but is non-motile and has
a 9 + 2 microtubule doublet arrangement [10] (a central
pair surrounded by 9 pairs). In contrast, the stereocilia
are made up of actin rich parallel filaments coated with
various isoforms of myosin [2].
“Tip links” connect the tips of shorter stereocilia
to the body of their adjacent taller stereocilia [2].
When head motion results in tilting of the stereocilia
toward the kinocilium, shifting of the “tip links” causes
mechanical opening of the transduction channels result-
ing in an influx of K + . This results in a depolarization
of the hair cell and opens Ca++ channels at the base of
the hair cell. Ca++ influx stimulates neurotransmitter
release into the synapses with afferent vestibular nerve
fibers increasing their firing rate. The bending of the
stereocilia away from the kinocilium decreases the “tip
link” tension and results in mechanical closure of the
channel. This causes hyperpolarization of the hair cell
which closes the Ca++ channels and decreases the neu-
rotransmitter release [9], reducing the firing rate of the
vestibular nerve fibers (Fig. 2).
 S. Khan and R. Chang / Anatomy of the vestibular system
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”
causes mechanical opening of the transduction channels resulting
in an influx of K and depolarization of the hair cells. The bend-
ing of the stereocilia away from the kinocilium decreases the “tip
link” tension and results in mechanical closure of the channel causing
hyperpolarization of the hair cell.
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
columnar with bouton synaptic connections to their afferent fibers.
There are two structural types of hair cells. Type I
has a rounded base surrounded by a nerve calyx of the
afferent nerve fiber. The majority of hair cells have a
Type II structure. They are columnar cells with bouton
synaptic connections to their afferent fibers. Type I hair
cells are associated with irregular afferents that have a
high variability of resting discharge. Type II hair cells
usually synapse on regular afferents with a low vari-
ability of resting discharge. Both hair cell types also
have efferent connections from vestibular nuclei that
modulate their sensitivity (Fig. 3) [9].
4. Utricle and saccule
The utricle and saccule are structures of the static
labyrinth that sense the orientation of the head in
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
utricle senses motion in the horizontal plane, while the
macula of the saccule senses motion in the vertical plane
[2].
439
Fig. 4. Utricle and Saccule. The stereocilia in the macula are oriented
in relation to a curvilinear line called the striola. This line is an area
of thinning in the utricle and an area of thickening in the saccule.
The kinocilia and stereocilia of the hair cells are oriented toward the
striola in the utricle and away from the striola in the saccule.
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
to a curvilinear line called the striola [13]. This line is an
area of thinning in the utricle and an area of thickening
in the saccule [10]. The kinocilia and stereocilia of the
hair cells are oriented toward the striola in the utricle and
away from the striola in the saccule. This distribution
of hair cells in different directions means that various
patterns of hair cell stimulation may occur based on the
degree to which head tilt occurs. Motion will stimulate
one group of hair cells while inhibiting another group
and yet, also have no effect on another group of hair
cell receptors [13]. This intricate pattern of response is
critical to relating accurate information regarding head
position to the central nervous system (Fig. 4).
Another important physiologic property of the mac-
ula is that of adaptation. When the head tilt stimulus
remains beyond a few seconds, the bent hair cells and
the depolarized membrane potentials begin to return to
normal. This allows the hair cells to be responsive to
further positional changes [10].
5. Semicircular ducts
The semicircular ducts have the same basic struc-
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]
Right Superior Left Posterior
Left Superior Right Posterior
Left Horizontal Right Horizontal
This arrangement allows for a 3-dimensional vector
representation of rotational acceleration. Each duct is
sensitive to movement in its specific plane.
The semicircular ducts open into the utricle. At the
end of each of the ducts there is a dilation called the
ampulla which contains the sensory neuroepithelium,
the crista ampullaris. It is coated by the cupula, which
is a gelatinous substance through which hair cells are
embedded [9]. The crista ampullaris is histologically
similar to the macula. The cupula is thicker than the
gelatinous membrane of the macula, and also does not
contain otoliths [9]. The kinocilia of the hair cells in
the lateral ducts are oriented toward the utricle, and the
kinocilia of the superior and posterior duct hair cells are
oriented toward the duct.
Rotational acceleration causes endolymph motion
that displaces the cupula, and therefore bends the hair
cells in the opposite direction of the rotation [2]. This
results in opening of the ion channels and depolariza-
tion of the hair cell which increases the firing of its
afferent fibers. When the rotational velocity of the head
becomes constant, the cupula returns to an upright posi-
tion, and the membrane potential of the cell normalizes.
Rotational deceleration of the head results in cupula
displacement in the same direction as the head move-
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).
Endolymph flow that causes excitation in one
semicircular duct will inhibit the hair cells of the con-
tralateral duct it is paired with. There are a few main
advantages of this system. First, it allows for sensory
redundancy in the event there is pathology of one semi-
circular duct. In this instance, vestibular input regarding
movement in its plane will be received from the other
duct in its pair. Another advantage is “common mode
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
neuroepithelium, the crista ampullaris and is coated by the cupula,
which is a gelatinous substance through which hair cells are embed-
ded. Rotational acceleration causes endolymph motion that displaces
the cupula, and bends the hair cells in the opposite direction of the
rotation [10].
rejection.” This means that simultaneous firing of both
semicircular ducts of the pair is ignored by the central
nervous system. This situation may occur with eleva-
tion of body temperature, as in a fever, and is not related
to motion. Finally, this physiologic arrangement also
allows for compensation for sensory overload [5].
6. Vestibular ganglion
The vestibular ganglion, also known as Scarpa’s Gan-
glion, is located in the lateral portion of the internal
auditory meatus [11]. It is composed of close to 20,000
bipolar cell bodies that receive afferent impulses from
the hair cells of the crista ampullaris and the maculae.
The vestibular ganglion is divided into a superior and
an inferior division which are connected by an isthmus
[13]. The peripheral fibers of the superior division of the
vestibular ganglion terminate in the crista ampullaris of
the superior and lateral semicircular ducts, as well as
the macula of the utricle. The macula of the saccule and
the crista ampullaris of the posterior semicircular ducts
receive peripheral vestibular branches from the inferior
section of the vestibular ganglion [13].
7. Vestibular nerve
Axons from the superior and inferior divisions of
the vestibular ganglion merge to form the vestibular
nerve. It combines with the cochlear nerve to become
the vestibulocochlear nerve. This nerve travels with the
facial nerve, nevus intermedius, and the labyrinth artery
through the internal auditory canal, which transverses
through the petrous temporal bone to the posterior fossa
 S. Khan and R. Chang / Anatomy of the vestibular system
[5]. The nerve fibers travel past the cerebellopontine
angle and enter the brainstem at the pontomedullary
junction. At this point, the vestibular nerve separates
from the cochlear nerve. The majority of the affer-
ent vestibular fibers project to the ipsilateral vestibular
nuclear complex in the pons. Some of the nerve fibers
project to the flocculo-nodular lobe of the cerebellum
and the adjacent vermian cortex [11].
8. Vestibular nuclear complex
This complex is the primary processor of vestibular
input and consists of four major nuclei: medial, supe-
rior, lateral, and inferior [13]. These nuclei are also
known as: Schwalbe, Bechterew, Deiter, and descend-
ing, respectively [6, 14]. They are located under the
floor of the fourth ventricle and extend from the rostral
medulla to the caudal pons in two major columns.
The medial vestibular nucleus is the largest and
makes up the medial column. The lateral column con-
sists of the superior, lateral, and inferior vestibular
nuclei [13]. The medial vestibular nucleus receives
afferents from the crista ampullaris of the lateral semi-
circular ducts. Ascending axonal fibers travel via the
medial longitudinal fasciculus to the motor nuclei of
the extraocular muscles to mediate the vestibuloocu-
lar reflex. It also functions in controlling the vesibular
spinal reflex via bilateral descending projections in the
medial vestibulospinal tract to the cervical spinal cord
to allow for the coordination of head and neck motion.
The superior vestibular nucleus also receives vestibu-
lar afferent input from the crista ampullaris of the
superior and posterior semicircular ducts [6]. Like the
medial vestibular nucleus, it sends ascending effer-
ent fibers to the extraocular muscles via the medial
longitudinal fasciculus in order to coordinate the
vestibuloocular reflex [6].
The lateral vestibular nucleus contains the largest cell
bodies of all the vestibular nuclei. It receives affer-
ent input from the crista ampulla, the maculae, and
the vestibulocerebellum. Efferent projections from this
nucleus become the lateral vestibular tract in the ipsi-
lateral spinal cord. This tract functions in the vestibular
spinal reflex by coordinating reflexive tone in the trunk
muscles and proximal extensors of the limbs to maintain
posture and balance [6].
The inferior vestibular nucleus receives afferent
information from the maculae of both the utricule and
the saccule. This nucleus has projections that go to the
other three vestibular nuclei and to the cerebellum [6].
441
9. Vestibulocerebellum
The role of the cerebellum in the vestibular sys-
tem is that it functions as an adaptive processor.
It monitors vestibular performance and will readjust
vestibular input through inhibitory input as necessary
[5]. The “vestibulocerebellum” is composed of the floc-
conodular lobe and the vermian cortex. The ipsilateral
cerebellum can send efferent information to bilateral
vestibular nuclei. It has projection fibers that go directly
to the ipsilateral vestibular nuclei, and to the ipsilat-
eral fastigial nucleus. Axons from the fastigial nucleus
project to the contralateral vestibular nuclei via the
juxarestiform body [11]. This area has an important
role in the generation of postural reflexes and orienting
behaviors.
The cerebellar flocculus adjusts the gain of the
vestibuloocular reflex [5]. The cerebellar nodulus
adjusts the duration of the vestibuloocular reflex and
is also involved in processing afferent activity from
the maculae [5]. The anterior superior vermis plays a
role in regulating the vestibulospinal reflex by encod-
ing vestibular signals and proprioceptive input from the
axial muscles.
10. Higher level cortical areas
Specifics regarding the complex cortical vestibular
connections are not clearly understood and consen-
sus regarding the location of a vestibular cortex still
has yet to be defined. In studies of primates, the main
cortical areas receiving vestibular stimuli include the
parietoinsular vestbular cortex (PIVC), area 2v of the
intraparietal sulcus, and area 3av in the central sulcus
[3]. The PIVC appears to be the main vestibular cortical
region in primates as information from other vestibular
cortical areas is integrated here [3].
Studies of humans have suggested that the main
cortical processing region is most likely in or near
the parietal or insular cortex. Kahane electrically
induced vestibular symptoms in humans and found
a lateral cortical temporoparietal area that he called
the “temporo-peri-sylvian vestibular cortex” and
suggested it corresponds to the PIVC in primates [7].
The use of activation likelihood estimation of cortical
activation on neuroimaging in response to multiple
vestibular stimuli by Lopez found the main areas of
convergence were mainly in the retroinsular cortex, but
also in the parietal operculum, and the posterior insula
[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
The vestibuloocular reflex coordinates eye move-
ment in order to stabilize retinal images during head
rotation. It involves a three neuron reflex arc from the
semicircular ducts to the vestibular nuclei and then to
the extraocular muscles to cause conjugate eye motion
in a direction opposite to head turning [3].
For instance, when the head turns to the right,
endolymph flow in the ampulla of the semicircular
ducts will deflect the cupula to the left. This causes
depolarization of the hair cells on the right, and hyper-
polarization of the left hair cells. This results in an
increase in the firing frequency in the afferent fibers
of the right vestibular nerve and impulses are sent to
the ipsilateral superior and medial vestibular nuclei and
the cerebellum. Excitatory impulses are transmitted in
the medial longitudinal fasciculus to the right oculo-
motor nuclei, and in the ascending tract of Deiters to
the left abducens nuclei [5]. This results in ipsilateral
medial rectus and contralateral lateral rectus contrac-
tion which produces eye movement to the left (opposite
to head turning). If eye velocity and head velocity do
not match, input from the cerebellar flocconodular lobe
is sent to the vestibular nuclei to modify their firing rate
to correct this discrepancy (Fig. 6).
12. Vestibulospinal reflex
This reflex involves many complex connections
which integrate input from the macula, crista
ampullaris, visual system, and axial and limb mus-
Fig. 6. Vestibuloocular Reflex. Head turning to the right causes the
endolymph flow in the ampulla of the semicircular ducts to deflect
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
afferent fibers of the right vestibular nerve. The nerve sends impulses
to the ipsilateral superior and medial vestibular nuclei. Excitatory
impulses are transmitted in the medial longitudinal fasciculus to the
right oculomotor nuclei, and in the ascending tract of Deiters to the
left abducens nuclei. This results in ipsilateral medial rectus and con-
tralateral lateral rectus contraction which produces eye movement to
the left.
cles by the brainstem, and cerebellum in order for the
maintenance of posture and balance. It involves both
the lateral and medial vestibular spinal tracts. The lat-
eral vestibular tract is the main pathway and originates
in the lateral vestibular nucleus. Efferent vestibular sig-
nals in response to input from the macula of the otolitic
organs to the lateral vestibular nucleus are carried in
this tract which projects ipsilaterally in the spinal cord
to neurons of all spinal levels. It produces monosynaptic
activation of ipsilateral trunk and proximal limb exten-
sors and disynaptic inhibition of contralateral proximal
extensors [10].
Angular rotation of the head sensed by the semicircu-
lar ducts is transmitted to the medial vestibular nucleus
where the medial vestibulospinal tract originates. This
tract projects bilaterally to motor neurons in the cervi-
cal spinal cord. It activates cervical axial muscles that
coordinate head and neck motion.
Another reflex that is related to the vestibulospinal
reflex is the vestibulocollic reflex. This activates neck
muscles that function to stabilize the head and keep
it properly oriented in space. The exact pathways that
contribute to this reflex have yet to be known [5].
 S. Khan and R. Chang / Anatomy of the vestibular system
13. Conclusion
The vestibular system coordinates head and eye
movement and activates postural muscles that maintain
balance and provide the proper orientation of the head
and body in space. The vestibular nuclear complex is the
primary processor of afferent vestibular input from the
hair cells of the otolitic organs and semicircular ducts.
The cerebellum functions as an adaptive processor that
readjusts this input as necessary. Details regarding the
vestibular cortical connections and functions have yet
to be fully understood. Efferent projections from the
vestibular neurons, cerebellum, and the cortex acti-
vate responses in the motor and ocular system that are
critical for the body to maintain equilibrium with the
environment.
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