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Anatomy of the Auditory System

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 3 Anatomy of the Auditory System
Peter A. Gochee and Hinrich Staecker
The embryology of the auditory system is complex and
begins early in embryological development. The otic placode,
the precursor to the membranous labyrinth, originates
from an ectodermal thickening around the 3rd week of
gestational age. This invaginates to form the otic vesicle by
the 5th week of development. The ventral portion of the
vesicle forms the saccule and cochlear duct by the 10th
week of development. The dorsal portion forms the utricle,
membranous semicircular canals, and endolymphatic duct.1
The cochleovestibular ganglion develops during this time,
and nerve fibers begin to connect to the macule, cristae,
and organ of Corti. Ossification begins to occur around the
membranous labyrinth to form the bony semicircular canals,
vestibule, and cochlea near the 16th week of gestation.
The middle ear begins to develop in the 4th week of
gestation as an outward growth of the first pharyngeal
pouch endoderm. The lateral portion of this outgrowth
forms the tympanic cavity, and the proximal portion
develops into the eustachian tube. Mesoderm from the
first and second branchial arches form the ossicles.1 The first
branchial arch (Meckel cartilage) derivatives include the
malleus head and neck, anterior malleal ligament, incus
body, and short process. The second branchial arch (Reichert
cartilage) derivatives include the manubrium of the malleus,
long process of incus, and stapes (with the exception of the
vestibular portion of the footplate, which is derived from the
otic placode). The surrounding mesoderm dissolves, leaving
endodermal mucosa overlying the middle ear structures.
The external auditory canal develops from the first
pharyngeal cleft ectoderm that grows medially toward
the endoderm of the tympanic cavity. A meatal plug is
formed which dissolves around the 7th month of gestation.
Congenital aural atresia is failure of the meatal plug to
dissolve, leaving a conductive hearing loss.2 As this occurs
late in gestation, the middle and inner ear structures are well
developed by this time. The auricle begins to form around
the 5th week of gestation from the six hillocks of His, which
are derivatives of the first and second pharyngeal arches. The
first hillock forms the tragus, the second hillock develops
into the helical crus, and the third hillock becomes the helix.
The fourth and fifth hillocks become the antihelix, whereas
the sixth hillock forms the antitragus (Fig. 3.1).
Temporal Bone
Detailed knowledge of the temporal bone and its complexity
is essential for the otolaryngologist. It is composed of four
distinct parts: mastoid, petrous, squamous, and tympanic
portions.3 As a whole, the temporal bone forms a portion
Otology and Neurotology.indb 41
of the outer ear, contains the middle and inner ear, forms
the floor of the middle cranial fossa, and defines the anterior
face of the posterior cranial fossa. Its complexity is also seen
in how it contributes to facial features, abuts the carotid
artery and jugular bulb, and how the facial nerve courses
through it.
The mastoid portion is the least formed part at birth.
There is slight pneumatization of the bone, which increases
in early childhood. The mastoid tip is underdeveloped at
birth and elongates over several years due to downward and
anterior traction of the sternocleidomastoid and digastric
muscles. The petrous portion is a pyramid-shaped bone
that extends anteriorly from the mastoid portion toward
the occipital and sphenoid bones. The arcuate eminence
is an important surgical landmark in middle fossa surgery
and denotes the superior semicircular canal.4 The trigeminal
nerve ganglion lies within a depression in the anterior aspect
of the portion near the petrous apex.
The squamous portion is a large plate-like bony segment
that forms part of the superior and lateral aspects of the
temporal bone. The medial aspect is the lateral extent of
the middle cranial fossa. The temporalis muscle attaches to
the lateral aspect, and a bony extension projects anteriorly
to form the posterior portion of the zygomatic process and
glenoid fossa. The tympanic portion forms the majority of the
bony portion of the external auditory canal and the anterior
and inferior aspect of the middle ear. The tympanomastoid
suture line is an important surgical landmark; it helps
identify the main truck of the facial nerve as it exits the
stylomastoid foramen during parotid surgery (Fig. 3.2).5
External Ear
The auricle is the most lateral portion of the external ear.
Apart from the soft tissue within the lobule, the auricle is
mainly composed of cartilage covered by a thin squamous
epithelium. It is usually at a 30-degree angle from the lateral
portion of the head and is approximately 2.5 cm in length.6
The blood supply is predominantly from the superficial
temporal and postauricular arteries, which are branches of
the external carotid artery. The external auditory canal is
contiguous with the auricle and measures approximately
25 mm in length, 9 mm in height, and 6.5 mm in width. It
is composed of two main components: the lateral one-third
consists of fibroelastic cartilage and the auricle and the
medial two-thirds is bony with a thin squamous epithelial
lining. Within the subcutaneous tissues of the lateral one-
third of the external auditory canal, sebaceous and apocrine
glands are found attached to hair follicle units and contribute
41
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 42 Otology and Neurotology

Figure 3.1 The external ear.

to cerumen production. Fissures of Santorini are naturally

occurring spaces within the cartilage of the lateral external
auditory canal that can serve as pathways for infectious
spread into the superficial parotid. The bony medial two-
thirds of the external auditory canal is covered by a thin layer
of squamous epithelium. It is adherent to the underlying bone
and is devoid of the subcutaneous structures found in the
lateral third. Between the lateral one-third and the medial
two-thirds of the external auditory canal lies the isthmus or
the narrowest portion of the canal. The foramen of Huschke
is patent in certain individuals due to incomplete ossification
of this region. It can serve as a pathway for infectious agents
and is malignantly spread from the external auditory canal
into the deep lobe of the parotid gland.
The tympanic membrane is contiguous with the squamous
epithelium of the external auditory canal and is composed of
three main layers. The lateral layer is a squamous epithelium.
The middle layer has two components: radial and circular
fibrous components. The medial layer is a mucosal layer
that is continuous with the middle ear mucosa. The taut
portion of the tympanic membrane is termed the pars tensa.
The pars flaccida is a small region of the superior posterior
tympanic membrane that does not contain the dense fibrous
layers observed in the pars tensa region. A fibrous annulus
surrounds the pars tensa portion of the tympanic membrane
Figure 3.2 Surface anatomy of the temporal bone. On the lateral
view, the mastoid (M) defines the posterior aspect of the temporal
bone. The spine of Henle (Sp) is an approximate surface landmark for
the mastoid antrum. The line made by the zygoma (Z) approximates
the level of the middle cranial fossa. The oval (Ov) and round (Rw) can
be seen in the middle ear. The styloid process (St) can be seen on all
the three views of the temporal bone. On the medial view, the middle
fossa and posterior fossa (PF) can be seen. The arcuate eminence
(AE) is a bony prominence over the superior semicircular canal. The
internal auditory canal (IAC) contains the cranial nerves VII and VIII and
runs almost in line with the external auditory canal. The stylomastoid
foramen (SMF) can be seen on the inferior view and is bounded by
the styloid process and the digastric ridge (DR). The inferior view also
demonstrates the petrous apex (P), the carotid canal (CC), and the
mandibular fossa (MF).

Otology and Neurotology.indb 42 25-02-2013 17:23:30


and secures it into the bony annulus which is the medial
most portion of the bony external auditory canal. Otoscopy
reveals the tympanic membrane to be of a slightly conical
shape that slopes to form an acute angle with the anterior
wall of the external auditory canal. The surface landmarks
include the lateral and short process of the malleus, the pars
tensa, and the pars flaccida. The tympanic membrane has a
slight dull color in a nondiseased ear. The chorda tympani,
incudostapedial joint, and a dark region corresponding
to the round window niche can be visualized through the
tympanic membrane.
Middle Ear
The main function of the middle ear is to impedance match air
sound pressure waves that arrive at the tympanic membrane
into fluid waves within the inner ear. This is performed by a
lever system that is composed of the three ossicles: malleus,
incus, and stapes (Fig. 3.3).
The malleus is the largest of the three ossicles. It is
a hammer-shaped bone that measures approximately
7.5 to 9 mm in overall length.7 The lateral process of the
malleus is directly medial and adherent to the tympanic
membrane. The tensor tympani muscle, which is innervated
by cranial nerve V, attaches to the neck of the malleus
from the cochleariform process. The incus has three main
subcomponents: body, long, and short processes. The short
process is in continuity with the head of the malleus, and
the long process articulates with the stapes via diarthrodial
joints. There is an approximate 1.3-lever arm leverage
gained through the incus. The distal portion of the incus’
long process has a tenuous blood supply which makes it
susceptible to necrosis from repeated middle ear infections.
The stapes has four main components. The superstructure
is composed of the head, which is attached to the anterior and
posterior crus. The fourth component is the footplate, which
is attached to the oval window of the inner ear by the annular
ligament. The stapedius muscle assists in attenuating loud
sounds and is innervated by cranial nerve VII; it attaches to
the stapes via the pyramidal eminence (Fig. 3.3).
The middle ear is divided into three regions based on
their relationship to the annulus of the tympanic membrane.
The region known as the epitympanum lies above superior
aspect of the annulus. It is bordered by scutum laterally and
the tegmen tympani superiorly. Within the epitympanum
lies the head of the malleus, the short process, and the body
of the incus. The suspensory ligaments and mucosa folds
of the malleus and incus also reside in the epitympanum.
Prussak space is a clinically important region of the
epitympanum. It is defined as the space that is contained
within the lateral process of the malleus inferiorly, lateral
mallear fold superiorly, neck of the malleus medially, and
the pars flaccida laterally. In individuals with prolonged
negative middle ear pressure from eustachian tube
dysfunction, the pars flaccida retract into the middle ear
space, where desquamated epithelium from the tympanic
Otology and Neurotology.indb 43
Anatomy of the Auditory System 43
A B
C
Figure 3.3 The ossicular chain consists of the (A) malleus (B) incus,
and (C) stapes. Ant., anterior; Post., posterior.
membrane can become trapped and form a cholesteatoma.
The mesotympanum lies behind the tympanic membrane
and contains the lateral process of the malleus, long process
of the incus, stapes, oval window, and round window. The
chorda tympani nerve travels within the mesotympanum,
passing medial to the malleus and lateral to the incus. The
hypotympanum lies below the level of the inferior aspect of
the annulus. Occasionally, the jugular bulb may be dehiscent
and extends into this region.
A detailed understanding of the facial nerve, which courses
through the middle ear, and mastoid is mandatory for safe
otologic surgery.8 At the geniculate ganglion, the facial nerve
enters the middle ear from the supratubal recess, forming
the tympanic segment of the facial nerve. Anatomically, this
posterior opening of the supratubal recess can be identified
by the cog superiorly and the cochleariform process inferiorly.
The facial nerve then courses posterior and slightly inferior,
above the oval window proximally within the fallopian canal,
and then inferior to the horizontal semicircular canal more
distally within the mastoid. Near the pyramidal eminence
the facial nerve turns inferiorly, forming the second genu,
and continues vertically toward the stylomastoid foramen.
The tympanic segment of the facial nerve is susceptible to
injury during middle ear surgery, whereas the second genu is
prone to injury during mastoidectomy.9
The eustachian tube is in the anterior portion of the
mesotympanum and courses toward the nasopharynx.10 It
functions to equilibrate middle ear pressures to ambient
atmospheric air pressure. The eustachian tube also protects
the middle ear from ascending infections, clearance of
middle ear mucous, and helps minimize sound fluctuations
during speaking and swallowing. The proximal one-third
of the eustachian tube near the middle ear comprises bone,
and the remaining cartilaginous portion extends into the
25-02-2013 17:23:31
 44 Otology and Neurotology
nasopharynx. It terminates with the torus tubarius.10 The
muscle principally involved with eustachian tube opening
is the tensor veli palatini and is innervated by cranial
nerve V3. Opening usually occurs during contraction of
tensor veli palatini during swallowing and yawning. Three
other muscles are also associated with the eustachian
tube: salpingopharyngeus, tensor tympani, and levator
veli palatini. Dysfunction of the eustachian tube can lead
to large differences between middle ear and ambient air
pressures with decreased tympanic membrane mobility,
development of a middle ear effusion, conductive hearing
loss, and potential for cholesteatoma formation.
Inner Ear
The inner ear is composed of three components: the cochlea,
the semicircular canals, and the vestibule. These three
components are encased within the petrous portion of the
temporal bone. The cochlea is a snail-shell–shaped structure
with two and three-fourth turn. The oval window is located
within the vestibule portion of the inner ear, and the round
window is located at the basal end of the cochlea. These are
the two regions of communication between the inner ear and
the middle ear.11 A third communicating region of the cochlea
is the cochlear aqueduct, which is located in the basal turn
of the cochlea and communicates the perilymph of the inner
ear to the posterior cranial fossa. Under normal condition,
sound is transmitted to the inner ear via the footplate of the
stapes, which is located within the oval window.
A membranous labyrinth lies within the bony snail-shaped
portion of the cochlea. Together the bony and membranous
portions of the inner ear form three chambers within the
cochlea: scala tympani, scala vestibuli, and scala media. All
three chambers coil together within the turns of the cochlea,
with the scala tympani located on the basal side of the
chamber and the scala vestibuli toward the apex. The basilar
membrane separates the scala tympani from the scala media,
and Reissner membrane separates the scala media from the
scala vestibuli. Perilymphatic fluid, which has a high sodium
concentration, is contained in the scala tympani and scala
vestibuli. Endolymphatic fluid is contained within the scala
media, which has a high concentration of potassium. A highly
vascular strip located on the lateral portion of the scala media,
termed the stria vascularis, maintains the high potassium
concentration of the scala media. The endolymphatic fluid
within the inner ear communicates with the endolymphatic
sac via the vestibular aqueduct. It is believed that alterations
in the osmotic concentrations between the endolymph and
perilymph may give rise to hydrops.
The organ of Corti is located on the basement membrane
and comprises the cells and supporting cells of the inner ear
that detect sound.12 When viewed as a cross-section of the
cochlea, the inner and outer hair cells are near the center
of the organ of Corti (Fig. 3.4). To detect sound, vibratory
waves are created within the cochlea and are sensed by the
inner and outer hair cells. On the apical surface of both types
Otology and Neurotology.indb 44
Helicotrema
Organ of
corti Stria
vascularis
Scala vestibuli
Scala
media
Scala tympani
Spiral ganglion
Modiolus
Saccule
Utricle
Figure 3.4 Midmodiolar section through the inner ear demonstrating
the cochlea and vestibule. The scala media is bounded by the spiral
ligament, stria vascularis, Reissner membrane, and the basilar membrane.
As can be seen in this osmium-stained specimen, the primary afferent
auditory neurons extend from the organ of Corti into the modiolus.
of hair cells, stereocilia project upward and are in contact
with tectorial membrane. Additional cross-links also occur
between the smaller stereocilia on each hair cell to the larger
stereocilia. This ensures that all stereocilia are deflected
together as a unit with movement of the tectoral membrane.
There are thee outer hair cells for every inner hair cell, with
the inner hair cell lying toward the modiolus of the cochlea.
The inner hair cells are flask-shaped cells, whereas the outer
hair cells have a cylindrical shape. The inner hair cells receive
more innervation than outer hair cells, and outer hair cells are
thought to increase the sensitivity of sound.13 Several additional
cell types are also present within the organ of Corti and are
thought to have a supportive role. These include Deiters cells,
Hensen cells, Claudius cells, and border cells of Held.12
Both inner and outer hair cells are innervated by
auditory nerve fibers. Type I nerve fibers comprise 95%
of auditory nerve fibers. These fibers are large, bipolar
myelinated nerves that innervate inner hair cells; with each
inner hair cell innervated by up to 20 different type I fibers.14
Type II fibers are smaller unmyelinated auditory nerve fibers
that comprise 5% of the estimated 30,000 total type I and
type II fibers. Type II fibers innervate outer hair cells with
one type II fiber innervating many outer hair cells.15
Central Auditory Pathways
Once sound has been detected by the inner and outer hair cells
within the cochlea, the information is transmitted to the brain
25-02-2013 17:23:37

by the cochlear nerve. Both type I and type II auditory fibers
combine to form the cochlear nerve, a component of the eighth
cranial nerve. Cell bodies for these bipolar nerve fibers are
located within the spiral ganglion of the cochlea and have one
projection toward the cochlear nucleus within the brainstem.15
As the cochlear nerve enters the cochlear nucleus, the
fibers bifurcate. One branch goes toward the anterior ventral
cochlear nucleus and the second branch projects to the
posterior ventral cochlear nucleus and dorsal nucleus of
the cochlear nucleus. Each of these subdivisions within the
cochlear nucleus has its own tonotopic frequency map. As the
cochlear nerve fibers synapse in the cochlear nucleus, new
projections are carried to the ipsilateral and contralateral
superior olivary complex via the dorsal, intermediate, and
ventral acoustic stria.16
It is at the level of the superior olivary complex that
sound from one ear is passed to the contralateral brainstem.
It is thought that this synaptic pathway—in which both
ipsilateral and contralateral innervation occurs—allows
binaural hearing with the ability to localize sound in the
environment. The intensity of the neuronal signals received
from both ears as well in the time delay from the contralateral
ear contribute to the ability to discriminate the origin of the
sound in the environment (Fig. 3.5).
From the superior olivary complex, new neuron projections
pass to the inferior colliculus, located in the midbrain, via
the lateral lemniscus. There are three main divisions within the
inferior colliculus: the central nucleus, the external nucleus,
and the dorsal cortex. Although input to the central nucleus
of the inferior colliculus is generally from the contralateral
ear, ipsilateral input also occurs via lateral lemniscus nuclei.
Directional information and the frequency of sound are
processed in the inferior colliculus, where a reflex center
exists that is associated with extraocular eye movements.
Neuron projections then pass through the medial
geniculate body, which is the thalamic relay of the auditory
DCN DCN
VIII VCN VCN
Superior
Olive LL
Inferior
Colliculus
Medial Geniculate Body
Figure 3.5 Simplified illustration of the central auditory pathways.
Input from the cochlear nerve (VIII) is into the ventral cochlear nucleus
(VCN). The VCN sends afferent input to the dorsal cochlear nucleus
(DCN), the lateral lemniscus (LL), and the superior olive. The DCN sends
input to the contralateral inferior colliculus (IC). The IC additionally
receives input from the contralateral DCN and lateral lemniscus.
Otology and
View Neurotology.indb
publication stats 45
Anatomy of the Auditory System 45
system.16 Three subnuclei exist within the medial geniculate
body: the dorsal, ventral, and medial divisions. From the
medial geniculate body, sound information passes to
the auditory cortex in the sylvian fissure of the temporal
lobe. The primary auditory cortex (Brodmann area) is area
41, with the secondary auditory cortex in area 42.17 Other
cortex regions are also associated with the central processing
of sound including Wernicke area (area 22), angular gyrus
and supramarginal gyrus (areas 39 and 40), and Broca area
(areas 44 and 45).
Conclusion
The auditory system is a complex mechanism that converts
sound to neuronal impulses and ultimately leads to human
perception. The various subsystems allow transmission,
amplification, encoding, and processing. It is through this
highly developed system that we are able to understand the
sounds around us and locate their origin.
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