EAR EMBRYOLOGY,

ANATOMY AND
PHYSIOLOGY
NOTES

BY : Amani Tawfiek Allam

 EMBRYOLOGY OF THE EAR

1. THE OUTER AND MIDDLE EARS(derivatives of the first and second branchial arches)

A. Auricle

 The external auricle or pinna develops from a series of small

cartilaginous tubercles (tubercles of His) that surround the first
pharyngeal groove. These enlarge and coalesce, although it seems
that the majority of the auricle is derived from the second arch
cartilages and that the tragus is the only contribution from the first arch.
(some authors say 3 tubercles from 1st & 3 tubercles from 2nd)
 The rudimentary pinna has formed by 60 days although it apparently
continues to grow throughout life.

B. External auditory canal

 The external ear canal develops from the first pharyngeal groove as
invagination into afunnel shapped pit (primary external auditory canal).
 The meatus deepens by proliferation of its ectoderm and that an
anteriorly placed bud of epithelial cells expands vertically to form the
skin, which will cover the future tympanic membrane.
 This clump of cells then opens up as a slit to form the canal lumen and
produce the pars tensa and deep external canal epithelium (defective
external auditory canal) (8th:10th weak IUL).
 These two types of skin both have migratory properties so that the ear
canal becomes self-cleansing.
 Anomalies : Complete atresia, Shallow depression stenosis, Changes
in the curvature of the canal.
C. Tympanic membrane
 The endoderm of the slit-like sac that is the precursor of the middle ear
lies against the ectoderm of the first pharyngeal groove by the fourth
week.
 Mesenchyme grows in between these two layers to form the middle
layer of the future tympanic membrane.

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D. Eustachian tube, epitympanum and mastoid antrum
tubotympanic recess(Middle ear cleft)
 Outpouching of the 1st pharyngeal pouch (expansion) due to the rapid
growth of the surrounding mesenchyme and, after dragging in some of
the second pouch endoderm, results in the formation of the Eustachian
tube (proximal narrow part), middle ear and mastoid antrum (Disital
dilated part).
 The future Eustachian tube lumen and middle ear spaces are formed
by eight months gestation.
 Epitympanum and mastoid antrum are developed by birth (mastoid
antrum(at birth 3 mm thick & it increase 1 mm every year to reach 15
mm adult size.
 A few mastoid ‘air cells’ may be present at birth, albeit filled with
amniotic fluid. However, development of the mastoid air cell system
does not occur until after birth, with about 90 percent of air cell
formation being completed by the age of six with the remaining 10
percent taking place up to the age of 18.

E. Ossicles
 The ossicles develop from the outer ends of the first arch (Meckel’s)
and second arch (Reichert’s) cartilages that lie above and below the
first pharyngeal pouch. The process begins at four weeks and adult
shape, size and ossification is present by 25 weeks.
 Meckel’s catilage forms the head of the malleus and the body of the
incus.
 Reichert’s cartilage forms the manubrium (handle) of the malleus and
the long process of the incus and the crurae of the stabes.
 The foot plate of stapes develops from three sources namely:
1. The outer periosteal layer of the otic capsule.
2. Middle endochondral layer of the otic capsule.
3. Inner endosteal layer is same as the endosteum of the bony
labyrinth and develops from the periotic endosteum.

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F. Muscles
 The muscles attached to the ossicles arise from the arches that give
rise to that part of the ossicle to which the muscle attaches.
 Thus the tensor tympani is attached to the upper part of the handle of
the malleus, which is derived from the first arch and is, therefore,
supplied by a branch of the Vth (mandibular) nerve.
 By the same argument the stapedius muscle is supplied by the VIIth
(facial nerve).

G. Chorda tympani
 The chorda tympani, which is the pretrematic nerve of the second arch
that supplies endodermal structures of the first arch, i.e. taste to the
anterior two-thirds of the tongue and submandibular gland
secretomotor fibres, has to run through mesoderm since it cannot run
through air.
 This mesoderm subsequently becomes the middle layer of the
tympanic membrane and is the physical connection between the first
and second arches.

H. Mucosal folds
 When the sac of endoderm expands and as it reaches the developing
ossicles and labyrinth, the epithelium is draped over these structures
and their associated muscles, tendons and ligaments, so that a
complex series of mucosal folds is formed.
 Compartments and folds of the tympanic cavity:
1. Compartments of mesotympanium: Upper part of mesotympanium:
Is areduplication of middle ear mucosa and divided into:
a. Ant. Malleolar fold: Neck of malleu to ant. margin of tympanic
sulcus.
b. Post. Malleolar fold: Neck of malleu to post. Margin of tympanic
sulcus.
2. Attic compartments:
a. Anerior compartment
b. Posterior compartment: Superior incudal space & medial
incudal space.

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FAILURE of formation of the first and second branchial arch structures
gives rise to problems e.g.

- Failure of adequate skin migration with ‘ear canal cholesteatoma’ (which

are very rare).
- Bat ears: Defective devlopment of the 4th tubercle = absence of antihelix
leading to bat ear deformity.
- Accessory auricles
- Preauricular sinuses (Defective fusion of the tubercles)
- Complete failure of formation of the external ear canal and middle ear.
- There are also the rare first arch fistulae (collaural fistulae), which
comprise a tract between the external ear canal and the skin of the
cheek, with the fistula path passing through the parotid gland and
frequently between branches of the facial nerve.
- Because of the dual origin (Sound conductive apparatus from branchial
apparatus. Sound perceptive apparatus from ectodermal otocyst)
conginital anomalies rarely co-exist.

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2.THE INTERNAL EAR (LABYRINTH)

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 The internal ear initially develops independently of the middle and external
ears, although the two become interconnected by the stapes super-
structure becoming attached to the stapes footplate thereby giving
continuity to the auditory pathway. The development of the labyrinth can be
thought of as the initial development of the generalized structure of the
membranous labyrinth, followed by a period of encasement by the bony
labyrinth and the production of a further series of spaces within this bony
shell that in turn become the perilymphatic spaces of the complete
structure. These different activities are happening at different times in
different parts of the labyrinth so that damage or derangements at specific
times give rise to many peculiar and varied abnormalities.
 MEMBRANOUS LABYRINTH:
o Within the first few days of embryonic life that is at about day 22–23
(preembryonic life is the first 21 day of gestation, embryonic life 35 day,
in pre embryonic and embryonic life definitive systems and fetus shape
develops, until end of gestational period fetus enlarge in size)
ectodermal thickening forms on the side of the head end of the embryo
close to that part of the developing neural tube and neural crest cells,
which will later become the brain and brainstem and the cranial nerves,
respectively. The ectodermal thickening is the otic placode, which
deepens and then sinks below the surface as the sides close in to form
an otic pit which eventually loses connection with the surface and
forms the otocyst.
o Associated with the otocyst is the cluster of neural crest cells that later
become the separate facial (geniculate), auditory (spiral) and vestibular
(Scarpa’s) ganglion cell bodies.

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o The otocyst then undergoes a series of spectacular changes, which
result in the full-sized outline of the adult membranous labyrinth by 25
weeks gestation.
 The semicircular canals start to develop at around 35 days as three
flattened pouches that grow out at right angles from each other from
the utricle. At the centre of each semicircular ridge, the opposing
epithelial surfaces meet, fuse and then coalesce to be replaced by
mesoderm. The superior canal is the first to be fully formed by six
weeks.
 The cochlea starts to be formed. The saccule, which has separated
from the utricle, starts to put out a single pouch-like process that
grows and then begins to coil from base to apex to reach its full 2.5
coils by 25 weeks.
 Within the membranous labyrinth the sensory cells of the three
cristae, two maculae and the organ of Corti are beginning to
develop from areas of ectodermal specialization. This is either
encouraged by the ingrowth of nerve endings from the
cochleovestibular ganglion, which was originally outside the otocyst,
or the development of the sensory cells encourages neural
ingrowth.
 The organ of Corti starts developing as a single block of heaped up
ectodermal cells at about 11 weeks. Within this mass develop inner
and outer hair cells and then specialized supporting cells. The
tunnel of Corti appears between inner and outer pillar cells as
clusters of stereocilia and a single kinocilium are developing on
each hair cell. The cochlear kinocilium regresses leaving the adult
configuration of stereocilia and the spaces between the outer hair
cells open up as the supporting cells (the Deiter’s cells) change
shape. Differentiation progresses from base to apex so that at any
one time various stages of development can be seen in
appropriately prepared material.
 Epithelium close to the sensory regions develops into the
specialized cell groups that maintain the ionic and electrical stability
of the endolymph. These regions are the stria vascularis of the
cochlear duct and the ‘dark cell’ regions of the vestibular sensory
epithelium.

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 THE BONY LABYRINTH
o The mesenchyme enclosing the otocyst becomes chondrified to form
the otic capsule.
o As the membranous labyrinth expands, the otic capsule remodels and
in places undergoes dedifferentiation to form fluid-filled spaces that
eventually become the perilymphatic spaces. This dedifferentiation
does not occur where nerves enter the sensory cell regions.
Elsewhere, the perilymphatic spaces become continuous and a
communication with the cerebrospinal fluid is formed by the
development of the cochlear aqueduct, which runs to the posterior
cranial fossa from the scala tympani in the base of the cochlea.
o Ossification of the cartilaginous otic capsule begins in or around week
16 from a variable number of centres(14) that finally fuse without
leaving telltale suture lines. This dense bony mass is the petrous bone
and is frequently the last part of a whale to decompose and is
sometimes the only remains of those eaten by sharks.
o There are certain channels that remain within the otic capsule with one
of the most important being the oval window where part of the otic
capsule becomes the stapes footplate and the annular ligament,
thereby allowing sound from the middle ear to enter the labyrinthine
fluids.

3.The temporal bone

o Four separate elements eventually fuse to form the temporal bone, which
is one of the most complex and interesting parts of the bony anatomy of
the body.
o These four parts are the petromastoid complex, the squamous portion, the
tympanic bone and the styloid process.
o The petromastoid is derived from petrous bone described above and the
continuing growth of its outer layers which form the roof of the middle ear,
the lateral wall of the Eustachian tube, the floor of the middle ear, the
canal of the facial nerve and the petrous apex.
o The squamous bone develops in mesenchyme and starts to ossify from a
single centre close to the roof of the zygoma as early as eight weeks. The
posteroinferior portion grows down behind the tympanic ring to form the
lateral wall of the foetal mastoid antrum.

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o The tympanic bone also forms in mesenchyme, but from several centres
around the external meatus. These form the major part of a ring with a
groove on the inner aspect that becomes the sulcus for the tympanic
membrane. Even by late foetal life the bony ear canal is unformed and
only after birth do anterior and posterior protuberances grow to form the
floor of the canal. These crescentic swellings eventually fuse laterally
leaving a gap in the middle of the floor. This is the foramen of Huscke –
which is usually closed by adolescence.
o The tympanic bone fuses with the mastoid process of the petromastoid
and with parts of the squamous and petrous bones as the
tympanomastoid, tympanosquamous and petrotympanic sutures. The
petrotympanic suture has a canal for the chorda tympani nerve.
o The styloid develops from two centres at the cranial end of the second
arch (Reichert’s) cartilage. The part close to the base of the skull starts to
ossify before birth, but the styloid process itself does not start to ossify
until after, and fusion of the two parts may not occur until after puberty.
o Much of the growth of temporal bone occurs after birth with enlargement
of the whole structure and major growth of the mastoid process so that the
stylomastoid foramen, which was initially close to the surface, becomes
buried by the development of the mastoid tip. At the same time there is
generalized downwards rotation of the petrous bone so that the tympanic
membrane moves from being almost horizontal in the neonate to an angle
of about 550 with the horizontal in the adult. With these changes, the floor
of the middle cranial fossa flattens and the relations of the geniculate
ganglion, the semicircular canals and the internal auditory meatus change
so that middle cranial fossa surgery in the baby is anatomically a different
experience from that in the adolescent.
o Temporal bone of newly borne:
 No mastoid process.
 The antrum is superficial 1 mm from outer cortex.
 Superfacial stylomastoid foramen.
 Facial nerve lies under skin.
 Tympanic bone is thin and formed by a ring

1-5 years : Partial formation of mastoid process.

5years: Mastoid process covers stylomastoid foramen and tympanic
bone is thicker.

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 ANATOMY OF THE EAR
1-EXTERNAL EAR

A.AURICLE
 The lateral surface of the auricle has characteristic prominences and
depressions, which are different in every individual even among identical
twins. This unique pattern is comparable to fingerprints and can allow the
identification of persons on the physiognomy of their auricles.
 The body of the auricle is formed from elastic fibrocartilage and is a
continuous plate except for a narrow gap between the tragus and the
anterior crus of the helix, where it is replaced by a dense fibrous tissue
band (Incisura terminals). This gap is the site for an endaural incision which,
properly performed, should not damage cartilage or its perichondrium and
which by splitting the soft-tissue ring surrounding the bony ear canal allows
wide exposure of the deeper parts.
 The cartilage extends about 8mm down the ear canal to form its lateral
third.
 The cartilage of the auricle is covered with perichondrium from which it
derives its supply of nutrients, as cartilage itself is avascular.
 Stripping the perichondrium from the cartilage, as occurs following injuries
that cause haematoma, can lead to cartilage necrosis with crumpled up
‘boxer’s ears’.
 The skin of the pinna is thin and closely attached to the perichondrium on
the lateral surface with minimal or no subcutineous tissue , so it is more
liable to frost bite. On the medial (cranial) surface, skin loosly adherent
there is a definite subdermal adipose layer that allows dissection during
pinnaplasty surgery, cysts as sebacoes cyst can be found because sub
cutineous tissue is more in this surface. The skin of the auricle is covered
with fine hairs and, most noticeably in the concha and the scaphoid fossa,
there are sebaceous glands opening into the root canals of these hairs. On
the tragus and intertragic notch coarse, thick hairs may develop in the
middle-aged and older male.

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 The cartilage of the auricle is connected to the temporal bone by:

1- Two extrinsic ligaments.

o The anterior ligament runs from the tragus and from a
cartilaginous spine on the anterior rim of the crus of the helix to
the root of the zygomatic arch.
o A separate posterior ligament runs from the medial surface of the
concha to the lateral surface of the mastoid prominence.
2- Intrinsic ligaments connect various parts of the cartilaginous auricle;
o that between helix and tragus
o another runs from the antihelix to the posteroinferior portion of the
helix.
3- Extrinsic muscles There are three extrinsic muscles:
o Auricularis anterior,
o Auricularis superior
o Auricularis posterior: supplied by the posterior auricular branch of
the facial nerve.

All three radiate out from the auricle to insert into the epicranial
aponeurosis. Temporal and posterior auricular branches of the
facial nerve supply the extrinsic muscles and, while being
functionally unimportant, they do give rise to the post-auricular
myogenic response following appropriate auditory stimulation.

1- Intrinsic muscles
The intrinsic muscles – six in number – are small, inconsistent and
without useful function.

Extrinsic and Intrinsic muscles are attached to the perichondrium of

the cartilage.

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 LATERAL SURFACE OF AURICLE

a) Helix: Rolled edge of the periphery, has asmall prominence in its

posterosuperior aspect called Darwins tubercle.
b) Antihelix : "Y – shaped " , another prominence Anterior to and parallel
with the helix is. Superiorly, it divides into two crura, between which is the
triangular fossa; the scaphoid fossa lies above the superior of the two crura
c) Concha : bowel – shaped depression . In front of the antihelix, and partly
encircled by it. This is divided into two portions by the descending limb of
the anterior superior portion of the helix,( known as the crus of the helix),
which rests just above the external auditory meatus. The smaller superior
portion is the cymba conchae and is the direct lateral relation to the
suprameatal triangle of the temporal bone. The larger inferior portion is
known as the cavum conchae.
d) Meatus: Entrance of the external auditory canal ( S shaped) .
e) Tragus: Below the crus of the helix and overlapping the external auditory
meatus, which is a small blunt triangular cartilage prominence pointing
posteriorly..
f) Antitragus: Opposite the tragus, at the inferior limit of the antihelix
g) The intertragic notch: separates the tragus from the antitragus.
h) Lobule: Fleshy inferior portion devoid of cartilage. The lobule lies below
the antitragus and is soft, being composed of fibrous and adipose tissue.

 THE MEDIAL (CRANIAL) SURFACE OF THE AURICLE

Has elevations corresponding to the depressions on the lateral surface,
and possesses corresponding names, for example the eminentia
conchae..

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BLOOD SUPPLY OF AURICLE
Arterial branches of the external carotid supply the auricle.
1- The posterior auricular appears to be the dominant artery and supplies
the medial surface (except the lobule), the concha, the middle and lower
portions of the helix and the lower part of the antihelix.
2- The anterior auricular branches of the superficial temporal supply the
upper portions of the helix, antihelix, triangular fossa, tragus and lobule.
3- The superior auricular artery has a constant course and connects the
superior temporal artery and the posterior auricular artery network. This
branch can provide a reliable vascular pedicle for retroauricular flaps.
4- A small auricular branch from the occipital artery may assist the
posterior auricular in supplying the medial surface.

NERVES SUPPLY THE AURICLE

Both cranial branchial nerves and somatic cervical nerves supply the auricle.
Their distribution is heterogeneous and the overlap may be extensive;
however, the greater auricular nerve prevails on the lateral and medial
surfaces.
1- Greater auricular Cervical plexus C2, 3 lower third of both surfaces.
2- Lesser occipital Cervical plexus C2, 3 Superior portion (upper two thirds)
of medial surface
3- Auricular Vagus X Concha and antihelix
4- Auriculotemporal Vc mandibular Tragus, crus of helix and adjacent
helix, upper two thirds of lateral surface
5- Facial VII Probably supplies small region in the root of concha (Evident
in herpes zoster inf). Some authors say twig from the facial nerve is
incorporated with the tympanic branch of vagus and distributed along with
it.

LYMPHATIC DRAINAGE OF THE AURICLE

1- Posterior surface is to the lymph nodes at the mastoid tip,
2- From the tragus and from the upper part of the anterior surface
to the preauricular nodes.
3- From the rest of the auricle to the upper deep cervical nodes.

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B. External auditory canal (E.A.C.):

 The external auditory canal extends from the concha of the auricle to the
tympanic membrane and is approximately 2.4 cm long,s shaped, oval in section.
 The diameter of the canal varies greatly between individuals and between
different races.
 The supporting framework of the canal wall is cartilage in the lateral one-third and
bone in the medial two-thirds.
 Cartilaginous portion:
o In adults, the cartilaginous portion runs inwards slightly downwards and
forwards. The canal is straightened, therefore, by gently moving the
auricle upwards and backwards to counteract the direction of the
cartilaginous portion.
o In the neonate, there is virtually no bony external meatus as the tympanic
bone is not yet developed, and the tympanic membrane is more
horizontally placed so that the auricle must be gently drawn downwards
and backwards for the best view of the tympanic membrane.
o In the adult, the lateral cartilaginous portion is about 8 mm long and is
continuous with the auricular cartilage. The medial border of the meatal
cartilage is attached firmly to the rim of the bony canal by fibrous bands
(imp).
o Frunculosis, more common in outer part of the canal as cartilagenous
part has epidermis and dermis but bony part has only epidermal layer.
o Fissure of santorini: from where infection from mastoid and parotid can
pass from each other.
o Cartilagenous part lined by pseudostratified columnar ciliated epithilium ,
but toward the tympanic memb. It lined by ciliated cuboidal epithelium
o The narrowest part of the estachian tube is the junction between bony
and cartilagenous part.

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 The bony canal wall:
o About 1.6mm long, is narrower than the cartilaginous portion and itself
becomes smaller closer to the tympanic membrane.
o The medial end of the bony canal is marked by a groove, the tympanic
sulcus, which is absent superiorly.
o Anterior recess: when anterior wall goes sharply to tympanic membrane
form it (common site of f.b)
o Foramen of Huschke: deficiency in antroinferior part of bonny canal : till
the age of 4 it remains.It located in anterior tympanic plate along non
ossified portion of plate.
o The anterior portion ,floor,part of posterior portion of the bonny canal
are formed by the tympanic part of the temporal bone & the rest of
posterior and roof formed by squamosa.

 Two suture lines in the canal wall:

o Tympanosquamous anteriorly.
o Tympanomastoid suture (posteriorly): is a complex suture line between
the anterior wall of the mastoid process, a portion of the squamous
bone and the tympanic bone.
These suture lines may be more or less developed; they project into
the canal with overlying closely adherent skin, which can make raising
an intact tympanomeatal flap a challenge.

 Two constrictions:
o One at the junction of the cartilaginous and bony portions
o The other, the isthmus, 5mm from the tympanic membrane where a
prominence of the anterior canal wall reduces the diameter. Deep to
the isthmus, the anteroinferior portion of the canal dips forward forming
a wedge-shaped anterior recess between the tympanic membrane and
the canal. This recess can be a difficult spot for access either in the
clinic or at surgery.

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 Skin:
o The external canal is lined with skin.
o There is outward, oblique growth of the epidermis of the canal skin and
pars flaccida so that the surface layers effectively migrate towards the
external opening of the canal. The normal rate of migration is about 0.1
mm/day, although this range is hugely variable and in some conditions
there is complete failure of migration with a consequent build-up of shed
keratin in the ear canal.
o The skin of the pars tensa has a different derivation from that of the
deep canal and cell divisions occur randomly within the layer of basal
cells. The effect of this, in a circular sheet with the handle of the malleus
forming a central boundary extending halfway down the membrane, is to
create outward mass migration of the skin of the pars tensa. Ink dots
applied to the surface have an outward pattern of movement. However,
if a hole is made in the tympanic membrane and a graft laid underneath
the membrane (an underlay graft) then migration of the skin from the
outer edge of the perforation is directed centrally to cover the graft. This
occurs because the boundary conditions have altered and fortunately
provides the basis for the healing of grafts and for the reepithelialization
of mastoid cavities. Even a small piece of pars tensa skin has this ability
and so is a precious material and needs to be preserved during ear
surgery if a bare area needs covering. However, the property of canal
skin to migrate can also bring problems with the formation of
cholesteatoma if the skin becomes displaced into the middle ear cleft.

 Ceruminous glands:
o At the outer limits of the ear canal are some short hairs that project
towards the opening of the canal. In this region are clusters of
ceruminous and sebaceous glands.
o The ceruminous glands are modified apocrine sweat glands that open
into the root canal of the hair follicles and produce a watery, white
secretion that slowly darkens, turning semi-solid and sticky as it dries.
o Since these glands are apocrine sweat glands they respond to many
stimuli such as adrenergic drugs, fever and emotion which, along with
direct mechanical stimulation, can all produce an increase or altered
secretion.

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 Sebaceous glands
o The sebaceous glands produce an oily material (sebum) from the
breakdown of their fat-containing cells which is usually excreted into
the root canals of the hair follicles.

 Wax
o The mixture of desquamated cells, cerumen and sebum forms wax.
Human earwax is a Mendelian trait consisting of wet and dry forms.
o Dry wax, lacking cerumen, is yellowish or grey and brittle, while wet
wax is brownish and sticky.
o The wet phenotype is dominant over the dry type, and is frequently
seen in populations of European and African origins.
o East Asians show the dry phenotype and there are intermediate
frequencies among the Native American and Inuit of Asian ancestry.
o A singlenucleotide polymorphism in the ABCC11 gene is responsible
for the determination of earwax type, with the AA genotype
corresponding to dry wax and GA and GG to wet wax.
o There is dispute with regard to the specific antibacterial activity of
cerumen in in vitro studies. However, the areas of skin that take part in
cerumen production have all the components of an active local immune
system and probably protect the canal by an antibody-mediated local
immune response.
o Wax is not usually found in the deep ear canal and a lump of ‘wax’
overlying the upper portion of the tympanic membrane (pars flaccida or
attic region) is rarely true wax, but is nearly always associated with an
underlying cholesteatoma as it is, in fact, dried-up, oxidized keratin.
The sense of the old adage ‘beware the attic wax’ is still just as true
today as it was in the past.

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THE ARTERIAL SUPPLY OF THE EXTERNAL MEATUS

Is derived from branches of the external carotid:

1- The auricular branches of the superficial temporal artery:

Supply the roof and anterior portion of the canal.
2- The deep auricular branch of the first part of the maxillary artery arises
in the parotid gland behind the temporomandibular joint:
Pierces the cartilage or bone of the external meatus and supplies the
anterior meatal wall skin and the epithelium of the outer surface of the
tympanic membrane.
3- Auricular branches of the posterior auricular artery:
Pierce the cartilage of the auricle and supply the posterior portions of the
canal.

VENOUS DRAINAGE

The veins drain into the external jugular vein, the maxillary veins and the
pterygoid plexus.

LYMPHATIC DRAINAGE
The lymphatic drainage follows that of the auricle.

BOUNARIES (RELATIONS) OF EUC.

1- Anterior: mandibular fossa& parotid

2- Inf : parotid
3- Posterior:mastoid
4- Superior :epitympanic recess(medially) , cranial cavity (laterally)

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 2-MIDDLE EAR CLEFT
The middle ear cleft consists of the tympanic cavity, the Eustachian tube and
the mastoid air cell system. The tympanic cavity is an irregular, air-filled
space within the temporal bone between the tympanic membrane laterally
and the osseous labyrinth medially. It contains the auditory ossicles and their
tendons that attach them to the middle ear muscles. Other structures,
including the tympanic segment of the facial nerve, run along its walls to
pass through the cavity.

A- THE TYMPANIC MEMBRANE (T.M.):

 The tympanic membrane lies at the medial end of the external auditory
meatus and forms the majority of the lateral wall of the tympanic cavity.
 It is slightly oval in shape, being broader above than below, forming an
angle of about 550 with the floor of the meatus so that the anteroinferior
portion is deeper , and the light directed onto the T.M. appears coned
anteroinferiorly .
 It lies obliquely because roof and posterior wall of external auditory canal
shorter than the floor and anterior wall.

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 Its longest diameter from posterosuperior to anteroinferior is 9–10mm,
while perpendicular to this the shortest diameter is 8–9mm.
 Semitranslucent, pearly white in color
 Most of the circumference is thickened to form a fibrocartilaginous ring,
the tympanic annulus, which sits in a groove in the tympanic bone, the
tympanic sulcus. The sulcus does not extend into the notch of Rivinus at
the roof of the canal, which is formed by part of the squama of the
temporal bone.
 From the superior limits of the sulcus, the annulus becomes a fibrous
band which runs centrally as anterior and posterior malleolar folds to the
lateral process of the malleus, the handle of which is clearly visible within
the tympanic membrane. This leaves a small, triangular region of tympanic
membrane above the malleolar folds within the notch of Rivinus, called the
pars flaccida (charpnells membrane), which does not have a tympanic
annulus at its margins. So TM formed of pars tensa and pars flacida
separated by ant. & post. Malleolar folds.
 The pars tensa forms the rest of the tympanic membrane and is concave
towards the ear canal but each segment is slightly convex between the
lateral attachment of the annulus and the centre of the membrane where
the tip of the malleus handle is attached at the umbo.
 Both the pars tensa and pars flaccida comprise three layers.
1- Outer epithelial layer, the epidermis, which is continuous with the skin
of the external meatus
2- Inner mucosal layer continuous with the lining of the tympanic cavity.
3- Middle, mainly fibrous layer, the lamina propria. The lamina propria of
the pars tensa has radially oriented fibres in the outer layers and
circular, parabolic and transverse fibres in the deeper layer. This
arrangement probably accounts for the complex pattern of tympanic
membrane displacement during sound stimulation. In the pars flaccida,
the lamina propria is less marked and the orientation of the collagen
fibres seems random.
 Average total area of tympanic membrane 70:80 mm2
Average vibrating surface of tympanic membrane 55mm2
 The T.M. is 0.1 mm thick in adults. It is thicker in infants and young
children.
 Umbo:(the tip of handle of malleous on the inner surface): It is the point of
gratest curvature:

20
The arterial supply of the tympanic membrane
Arises from branches supplying both the external auditory meatus and the
middle ear. These two sources interconnect through extensive anastomoses
within the connective tissue layer of the lamina propria.
a- The epidermal vessels originate from the deep auricular branch of the
maxillary artery coming from the external auditory meatus
b- Mucosal vessels arise from
o The anterior tympanic branches of the maxillary artery
o The stylomastoid branch of the posterior auricular artery
o Probably from the middle meningeal artery.

Venous drainage of tympanic membrane

a- Outer surface: external jugular-
b- Inner surface: transverse sinus and venous plexus located around the
eustachian tube.

Nerve supply of the tympanic membrane

These also run in the lamina propria and, while variations and overlap are
considerable, both the vascular supply and innervation are relatively sparse
in the middle part of the posterior half of the tympanic membrane.

c- Branches of the auriculotemporal nerve (Vc)

d- Auricular branch of the vagus
e- Tympanic branch of the glossopharyngeal nerve supply the tympanic
membrane.

21
B-The middle ear cavity ( Tympanic cavity )
It is an air-filled space similar to a rectangular box ( i.e. six-walled ). It is
wedge shaped with the posterior wall wider than the anterior wall . It is
rather narrower from side to side (15mm ) than anteroposteriorly (2-6mm) .

The cavity is divided into three parts :

1- Epitympanum(Attic ):
o This is the superior portion located above the level of the T.M.
It is connected with the mastoid antrum through the aditus ad
antrum . So, its outer wall is bony ( part of squamous temporal
bone) and not the pars flaccid as sometimes mis understood.
o Boundaris of epitympanium:
Ant: zygomatic arch
Posterior: aditus
Medial: lateral semicircular canals& F.N.
Lateral: squamosa(scutum)
Superior: tagmen
Inferiorly: fossa incudis.
2- Mesotympanum : This lies opposite the T.M.
3- Hypotympanum: This is the most inferior portion below the level of
the T.M.

22
  Walls of the middle ear

1 . ROOF :
 The roof of the epitympanum is the tegmen tympani, a thin bony plate that
separates the middle ear space from the middle cranial fossa. It is
continuous with roof of the antrum .
 Both the petrous and squamous portions of the temporal bone form it; and
the petrosquamous suture line, which does not close until adult life, can
provide a route of access for infection into the extradural space in children.
Veins from the tympanic cavity running to the superior petrosal sinus pass
through this suture line.

2 . FLOOR :
 The floor of the tympanic cavity may consist of compact or pneumatized
bone and separates the hypotympanum from the dome of the jugular bulb.
 Its thickness can vary according to the height of the jugular fossa.
Occasionally, the floor is deficient and the jugular bulb is then covered
only by fibrous tissue and a mucous membrane.
 At the junction of the floor and the medial wall of the cavity there is a small
opening that allows the entry of the tympanic branch of the
glossopharyngeal nerve into the middle ear from its origin below the base
of the skull.

23
 3 . ANTERIOR WALL :
 The anterior wall of the tympanic cavity is rather narrow as the medial and
lateral walls converge.
 The lower-third of the anterior wall consists of a thin plate of bone
covering the carotid artery as it enters the skull and before it turns
anteriorly. This plate is perforated by the superior and inferior
caroticotympanic nerves carrying sympathetic fibres to the tympanic
plexus, and by tympanic branches of the internal carotid artery.
 The middle-third comprises the tympanic orifice of the Eustachian tube,
which is oval and 5*2mm in size. Just above this is a canal containing the
tensor tympani muscle that subsequently runs along the medial wall of the
tympanic cavity enclosed in a thin bony sheath.
 The upper-third is usually pneumatized and may house the anterior
epitympanic sinus, a small niche anterior to the ossicular heads, which
can hide residual cholesteatoma in canal wall up surgery

24
5. LATERAL WALL:
 Formed by
a- Bony lateral wall of the epitympanum superiorly, The lateral
epitympanic wall is wedge-shaped in section and its sharp inferior
portion is also called the outer attic wall or scutum (Latin: ‘shield’). It is
thin and easily eroded by cholesteatoma, leaving a telltale sign on a
high resolution coronal computed tomography (CT) scan.
b- The tympanic membrane centrally
c- Bony lateral wall of the hypotympanum inferiorly.
 THE PETROTYMPANIC FISSURE = GLASERIAN FISSURE
o Is a fissure in the temporal bone that runs from the temporomandibular
joint to the tympanic cavity.
o The mandibular fossa is bounded, in front, by the articular tubercle;
behind, by the tympanic part of the bone, which separates it from the
external acoustic meatus; it is divided into two parts by a narrow slit, the
petrotympanic fissure.
o It opens just above and in front of the ring of bone into which the
tympanic membrane is inserted; in this situation it is a mere slit about 2
mm. in length.
o It lodges the anterior process and anterior ligament of the malleus, and
gives passage to the anterior tympanic branch of the internal maxillary
artery.
o The contents of the fissure include:
- Communications of cranial nerve VII to the infratemporal fossa.
- Chorda tympani, runs through the fissure to join with the lingual nerve, it
enters the medial surface of the fissure through a separate anterior
canaliculus (canal of Huguier) which is sometimes confluent with the
fissure. It then runs posteriorly between the fibrous and mucosal layers
of the tympanic membrane, across the upper part of the handle of the
malleus and then continues within the membrane, but below the level of
the posterior malleolar fold. The nerve reaches the posterior bony canal
wall just medial to the tympanic sulcus.

25
-

26
MEDIAL WALL :
 The medial wall separates the tympanic cavity from the internal ear.
 THE PROMONTORY
o Is a rounded elevation occupying much of the central portion of the
medial wall.
o It covers part of the basal coil of the cochlea
o Usually has small grooves on its surface containing the nerves which
form the tympanic plexus. Sometimes the groove containing the
tympanic branch of the glossopharyngeal nerve may be covered by
bone, thereby forming a small canal.
o The promontory gently inclines forwards to merge with the anterior wall
of the tympanic cavity, but is more steeply sloped posteriorly.

 OVAL WINDOW (fenestera vistubuli)

o Behind and above the promontory is the oval window; a nearly kidney-
shaped opening that connects the tympanic cavity with the vestibule, but
which in life is closed by the footplate of the stapes and its surrounding
annular ligament.
o Its size naturally varies with the size of the footplate, but on average it is
3.25mm long and 1.75mm wide.
o The oval window lies at the bottom of a depression or niche that can be
of varying width depending on the position of the facial nerve superiorly,
and the prominence of the promontory inferiorly.

27
 ROUND WINDOW NICHE (fenestera chochlae)
o The round window niche lies below and a little behind the oval window
niche from which it is separated by a posterior extension of the
promontory called the subiculum.
o Occasionally, another ridge of bone – the ponticulus – leaves the
promontory above the subiculum and runs to the pyramid on the
posterior wall of the cavity.
o The round window niche is most commonly triangular in shape, with
anterior, posterosuperior and posteroinferior walls. The latter two meet
posteriorly and lead to the sinus tympani.
o The round window membrane is usually out of sight, obscured by the
overhanging edge of the promontory forming the niche and mucosal
folds within it. The membrane is roughly oval in shape, about 2.3*1.9 mm
in dimension and lies in a plane at right angles to the plane of the stapes
footplate. It tends to curve towards the scala tympani of the basal coil of
the cochlea, so that it is concave when viewed from the middle ear, and
appears to be divided into an anterior and posterior portion by a
transverse thickening.

 THE FACIAL NERVE CANAL (OR FALLOPIAN CANAL)

o It runs above the promontory and oval window in an anteroposterior
direction.
o It has a smooth rounded lateral surface that often has microdehiscences
and when the bone is thin or the nerve exposed by disease, there are
two or three straight blood vessels clearly visible along this line of nerve.
These are the only straight blood vessels in the middle ear and indicate
quite clearly that the facial nerve is very close by.
o The facial nerve canal is marked anteriorly by the processus
cochleariformis (a Pully like projection (curved projection) of bone,
concave anteriorly, which houses the tendon of the tensor tympani
muscle as it turns laterally (900) to the handle of the malleus).
o Behind the oval window, the facial canal starts to turn inferiorly as it
begins its descent in the posterior wall of the tympanic cavity.
o The region above the level of the facial nerve canal forms the medial
wall of the epitympanum.

28
 DOME OF THE LATERAL SEMICIRCULAR CANAL
o The dome of the lateral semicircular canal is the major feature of the
posterior portion of the epitympanum, lying posterior and extending a
little lateral to the facial canal.
o During a cortical mastoidectomy, the triangular relationship of the lateral
semicircular canal, the short process of the incus and the seventh nerve
is often quite helpful.
o In wellaerated mastoid bones, the labyrinthine bone over the superior
semicircular canal may be prominent, running at right angles to the
lateral canal and joining it anteriorly at a swelling which houses the
ampullae of the two canals.
 GENICULATE GANGLION
o In front and a little below this, above the processus cochleariformis, may
be a slight swelling corresponding to the geniculate ganglion, with the
bony canal of the greater superficial petrosal nerve running for a short
distance anteriorly.

A specimen cut axially at the level of the labyrinthine and intratympanic segments of the facial nerve
(VII), showing a small cholesteatoma (C) lying between the head of the malleus and anterior
epitympanic space. The ossicular heads resemble a scoop of ice cream in a cone, which can be
readily seen on a high-resolution axial CT scan. I, incus; M, malleus.

29
The medial wall of the mesotympanum seen after removal of the tympanic membrane. The
tympanic branch of the glossopharyngeal nerve (IX) can be seen crossing the
promontory. Note the notch of Rivinus in the roof of the canal.

Endoscopic view of the posterior medial wall of

the tympanic cavity showing the relationship between the oval
window niche (OWN), round window niche (RWN), promontory
(P), facial nerve (VII) and facial recess (FR). The sinus tympani
(ST) lies medial to the facial nerve between the ponticulus
(PONT) and subiculum (SUB).

30
A specimen cut to show the medial wall of the tympanic cavity. The processus cochleariformis (PC)
marks the anterior position of the intratympanic portion of the facial
nerve, which passes between the lateral semicircular canal (LSCC) and oval window niche (OWN).
Note the jugular bulb (JB) in the floor of the tympanic cavity.

31
POSTERIOR WALL :
 The posterior wall is wider above than below.
 THE ADITUS AD ANTRUM
o In upper part of posterior wall
o It is a large irregular opening that leads back from the posterior
epitympanum into the mastoid antrum.
 THE FOSSA INCUDIS
o A small depression below the aditus, which houses the short
process of the incus and its suspensory ligament.
 PYRAMID
o Below the fossa incudis and medial to the opening of the chorda
tympani nerve.
o It is a small hollow conical projection with its apex pointing
anteriorly.
o This houses the stapedius muscle and tendon, which inserts into the
posterior aspect of the head of stapes.
o The canal within the pyramid curves downwards and backwards to
join the descending portion of the facial nerve canal.
 THE FACIAL RECESS (SUPRATYMPANIC RECESS)
o Is a groove which lies lateral to F.N.
o bounded by
- Fossa incudis superiorly and
- Chorda tympani nerve inferiorly.
- Posterosuperior meatal wall laterally,
- Pyramid medially.
o It can be directly accessed via posterior tympanotomy approach(The
chorda always runs medial to the tympanic membrane, which means
that the angle between the facial nerve and the chorda allows a
posterior tympanotomy to be cut, thereby accessing the middle ear
from the mastoid without disruption to the tympanic membrane. This
angle can be small or large depending on the site of origin of the
chorda from the facial nerve).
o This is shallower lower down where the facial nerve canal forms only
a slight prominence on the posterior wall.

32
 THE SINUS TYMPANI (INFRAPYRAMIDAL RECESS)
o It is a posterior extension of the mesotympanum and lies deep to both
the promontory and the facial nerve.
o This extension of air cells into the posterior wall can be extensive, and is
probably the most inaccessible site in the middle ear and mastoid.
o The sinus can extend as far as 9mm into the mastoid bone when
measured from the tip of the pyramid.
o The medial wall of the sinus tympani becomes continuous with the
posterior portion of the medial wall of the tympanic cavity where it is
related to the oval and round window niches and the subiculum of the
promontory.
o On rare occasions it can communicate with the mastoid air cells.
o Its importance is that cholesteatoma which has extended here from the
mesotympanum is extremely difficult to eradicate.
o The worst region for access is above the pyramid, posterior to an intact
stapes and medial to the facial nerve.
o A retrofacial approach to this region via the mastoid is not possible
because the posterior semicircular canal blocks access.

A specimen cut coronally at the level of the

long process of incus. The aditus to the mastoid antrum lies
posterosuperiorly to the incus, (I). LSCC, lateral semicircular
canal; M, malleus; S, stapes; VII, facial nerve.

33
 (a) The facial recess (FR) and sinus tympani (ST) at different levels in the middle ear. Section AA is at the level of the pyramid
where the facial recess is relatively deep. In section BB, at the level of the round window, the facial recess is quite shallow. The
extent of the sinus tympani, deep and posterior to the facial nerve is variable. (b) A specimen in section AA at the level of the
pyramid (P). S, stapes; VII, facial nerve.

Posterior tympanotomy in a left ear, showing the relationship of the facial nerve (VII), corda tympani
(CT), annulus (A) and lateral semicircular canal (LSCC). The promontory is readily accessible via this
approach in cochlear implant surgery.

34
  CONTENTS OF TYMPANIC CAVITY
1. AIR .
2. THREE OSSICLES :

A- Malleus:
 The malleus is the largest of the three ossicles, measuring up to 9mm in
length.
 It comprises a head, neck and handle or manubrium.
 The head lies in the epitympanum and is suspended by the superior
ligament, which runs upward to the tegmen tympani. The head of the
malleus has a saddle-shaped facet on its posteromedial surface to
articulate with the body of the incus by way of a synovial joint.
 The neck of the malleus connects the handle with the head and amputation
of the head by cutting through the neck leaves both chorda tympani and
tensor tympani intact.
 Below the neck of the malleus, the bone broadens and gives rise to the
lateral process, the anterior process and the handle.
 The lateral process is a prominent landmark on the tympanic membrane and
receives the anterior and posterior malleolar folds from the tympanic
annulus.
 A slender anterior ligament arises from the anterior process to insert into
the petrotympanic fissure.

35
 The handle

o Runs downwards, medially and slightly backwards between the mucosal

and fibrous layers of the tympanic membrane.
o While it is very closely attached to the membrane at its lower end, there
is a fine web of mucosa separating the membrane from the handle in the
upper portion before it becomes adherent again at the lateral process.
This can be opened surgically to create a slit without perforating the
membrane to allow a prosthesis to be crimped around the malleus
handle in certain types of ossicular reconstruction.
o On the deep, medial surface of the handle, near its upper end, is a small
projection into which the tendon of the tensor tympani muscle inserts.
o The chorda tympani crosses the upper part of the malleus handle on its
medial surface above the insertion of the tendon of tensor tympani, but
below the neck of the malleus itself.

B- THE INCUS
 The incus articulates with the malleus and has a body and two processes.
 The body lies in the epitympanum and has a cartilage-covered facet
corresponding to that on the malleus. The body of the incus is suspended
by the superior incudal ligament that is attached to the tegmen tympani.
 The short process projects backwards from the body to lie in the fossa
incudis to which it is attached by a short suspensory ligament.
 The long process descends into the mesotympanum behind and medial to
the handle of the malleus, and at its tip is a small medially directed
lenticular process. This has sometimes been called the fourth ossicle
because of its incomplete fusion with the tip of the long process, thereby
giving the appearance of a separate bone or at least a sesamoid bone.
 The lenticular process articulates with the head of the stapes.

36
C- THE STAPES
 The stapes is shaped like a stirrup and consists of a head, neck, the
anterior and posterior crura and a footplate.
 The head points laterally and has a small cartilage-covered depression for
a synovial articulation with the lenticular process of the incus.
 The stapedius tendon inserts into the posterior part of the neck and upper
portion of the posterior crus.
 The two crura arise from the broader lower part of the neck and the
anterior crus is thinner and less curved than the posterior one. Both are
hollowed out on their concave surfaces, which gives an optimum
combination of strength and lightness. There is great variation in the shape
of the two crura.
 The two crura join the footplate, which usually has a convex superior
margin, an almost straight inferior margin and curved anterior and posterior
ends. The average dimensions of the footplate are 3mm long and 1.4mm
wide, and it lies in the oval window where it is attached to the bony margins
by the annular ligament. The long axis of the footplate is almost horizontal,
with the posterior end being slightly lower than the anterior.

3. Two muscles :

37
A- The stapedius muscle
 Origin: from the walls of the conical cavity within the pyramid and from
the downward curved continuation of this canal in front of the
descending portion of the facial nerve.
 Insertion: A slender tendon emerges from the apex of the pyramid and
inserts into the stapes.
 Neve supply: The muscle is supplied by a small branch of the facial
nerve.
 Action: dams down excessive movements of stapes.

B- The tensor tympani muscle

 Origin: This is a long slender muscle arising from the walls of the
bony canal lying above the Eustachian tube. Parts of the muscle also
arise from the cartilaginous portion of the Eustachian tube and the
greater wing of the sphenoid.
 Insertion:From its origins, the muscle passes backwards into the
tympanic cavity where it lies on the medial wall, a little below the level
of the facial nerve. The bony covering of the canal is often deficient in
its tympanic segment where the muscle is replaced by a slender
tendon. This enters the processus cochleariformis (Figure 225.9)
where it is held down by a transverse tendon as it turns through a
right angle to pass laterally and insert into the medial aspect of the
upper end of the malleus handle.
 Neve supply: The muscle is supplied from the mandibular nerve by
way of a branch from the medial pterygoid nerve.
 Action: pulls the tympanic membrane inwards protecting it from
excessive movement.

N.B.
Thus the 2 muscles produce an action of damping down the intensity of
high pitched sounds (Acoustic reflex) but loudness basilar memb.

38
4. Nerves :

A- The chorda tympani nerve

 This branch of the facial nerve enters the tympanic cavity from the
posterior canaliculus at the junction of the lateral and posterior walls.
 It runs across the medial surface of the tympanic membrane between
the mucosal and fibrous layers (below the level of the posterior
malleolar fold) and passes medial to the upper portion of the handle
of the malleus above the tendon of tensor tympani to continue
forwards it enters the medial surface of the petrotympanic fissure
through a separate anterior canaliculus (canal of Huguier) which is
sometimes confluent with the fissure. It runs through the fissure to
join with the lingual nerve deep to the medial ptrygoid muscle..

B- The tympanic plexus

 The tympanic plexus is formed by the tympanic branch of the
glossopharyngeal nerve (Jacobson’s nerve) and by caroticotympanic
nerves, which arise from the sympathetic plexus around the internal
carotid artery.
 The nerves form a plexus on the promontory and provide the
branches to the mucous membrane lining the tympanic cavity,
Eustachian tube and mastoid antrum and air cells.
 The plexus also provides branches to join the greater superficial
petrosal nerve and the lesser superficial petrosal nerve that contains
all the parasympathetic fibres of the glossopharyngeal nerve.

5. Small blood vessels :

 Plexus of vessels of stylomastoid artery and from corticotympanic
artery.

39
 THE MUCOSA OF THE TYMPANIC CAVITY
 The middle ear mucosa is essentially mucus-secreting respiratory
mucosa bearing cilia on its surface.
 The extent of the mucociliary epithelium varies in normal middle ears,
being more widespread in the young.
 Three distinct mucocilary pathways can be identified – epitympanic,
promontorial and hypotympanic, the latter being the largest. Each of
these pathways coalesces at the tympanic orifice of the Eustachian tube.
 The mucous membrane lines the bony walls of the tympanic cavity, and
it extends to cover the ossicles and their supporting ligaments in much
the same way as the peritoneum covers the viscera in the abdomen.
 The mucosal folds also cover the tendons of the two middle ear muscles
and carry the blood supply to and from the contents of the tympanic
cavity. These folds separate the middle ear space into compartments.
 As a result, the only route for ventilation of the epitympanic space from
the mesotympanum is via two small openings between the various
mucosal folds – the anterior and posterior isthmus tympani.
 Likewise, Prussak’s space (potential space) is found between the pars
flaccida and the neck of the malleus
Boundaries:
- Laterally: pars flaccida= sharpnell's membrane
- Medially:by the neck of malleus
- Superiorly : lateral malleolar fold
- Inferiorly: lateral process of malleus
This space can play an important role in the retention of
keratin and subsequent development of cholesteatoma(the
first to involve during extension of Cholesteatoma).

40
 THE BLOOD SUPPLY OF THE TYMPANIC CAVITY

Arteries supplying the walls and contents of the tympanic cavity arise from
both the internal and external carotid system. The overlap between branches
is extensive and there is great variability in the supply between individuals.
anterior tympanic and stylomastoid arteries are the biggest.

1- Anterior tympanic artery: branch of maxllary artery.supply

TM,malleus,incus and anterior part of tympanic cavity
2- Posterior tympanic artery(stylomastoid a.): branch of posterior
auricular artery.supply posterior part of tympanic cavity and stapedius
muscle.
3- Mastoid a.: branch of stylomastoid supply mastoid air cells.
4- Petrosal a.: branch from middle meningeal a. supply roof of mastoid and
roof of epitympanium.
5- Superior tympanic artery: branch of middle meningeal arery.supply
malleus,incus and tensor tympani.
6- Inferior tympanic artery: branch of ascending pharyngeal artery.supply
mesotympanium
7- Branch from artery of ptrygoid canal: supply meso and hypotympanium
8- Tympanic barches from internal carotid: supply meso and
hypotympanium

 Lymph drainage of middle ear:

 The lymphatics of the middle ear and mastoid antrum drain into parotid
lymph nodes and upper deep cervical lymph nodes= preauricular and
retropharyngeal lymph nodes.
 Ventilatory anatomy of middle ear:
 Normally middle ear cleft is well ventilated
o Air comes via E.T. to anterior mesotympaniumgoes up to anterior
epitympanium via ismus tympanic anticus goes to posterior
epitympanium part goes across aditus to ventillate mastoid air
cells and part goes to ventillate posterior mesotympanium across
ismus tympanic posticus (in awell ventillated mastoid).

41
C. THE EUSTACHIAN TUBE ( E.T. )

 The Eustachian tube is a dynamic channel that links the middle ear with
the nasopharynx.
 In adults, it is about 36 mm in length, a size that is normally reached by
the age of seven.
 Where it is flattend its diameter 2mm.
 It runs downwards from the middle ear at 450 and is turned forwards and
medially.
 The tube is lined with respiratory mucosa containing goblet cells and
mucous glands, having a carpet of ciliated epithelium on its floor. At its
nasopharyngeal end, the mucosa is truly respiratory; but in passing along
the tube towards the middle ear, the number of goblet cells and glands
decreases, and the ciliary carpet becomes less profuse.
 The tube can be considered to consist of two unequal cones, connected at
their apices.
o The lateral third is bony and arises from the anterior wall of the tympanic
cavity.
o A medial cartilaginous part, which makes up two-thirds of the tubal
length
o Both parts join just after cartilagenous narrowest portion, called the
isthmus.

 The bony portion

o Is about 12mm long.
o Widest at its oval-shaped orifice in the anterior wall of the tympanic
cavity.
o It runs through the squamous and petrous portions of the temporal
bone, gradually tapering to the isthmus, where the diameter is only
0.5mm or less.
o A thin plate of bone forms the roof, separating the tube from the tensor
tympani muscle above.
o The carotid canal lies medially and can impinge on the bony Eustachian
tube.
o In cross section, the tube is triangular or rectangular with the horizontal
diameter being the greater.

42
 The cartilaginous part
o Is about 24mm long.
o Consists of a fibrocartilaginous skeleton to which is attached the
peritubal muscles.
o At its upper border, the cartilage is bent over to resemble an inverted ‘J’,
thereby forming a longer medial cartilaginous lamina and shorter lateral
cartilaginous lamina.
o The cartilage is fixed to the base of the skull in a groove between the
petrous part of the temporal bone and the greater wing of the sphenoid,
which terminates near the root of the medial pterygoid plate. Thus, the
back (posteromedial) wall is composed of cartilage and the front
(anterolateral) wall comprises cartilage and fibrous tissue.
o The apex of the cartilage is attached to the isthmus of the bony portion,
while the wider medial end protrudes into the nasopharynx, lying directly
under the mucosa to form the torus tubarius.
 In the nasopharynx, the tube opens 1–1.25 cm behind and a little below
the posterior end of the interior turbinate.The opening is almost triangular
in shape and is surrounded above and behind by the torus.
 The salpingopharyngeal fold stretches from the lower part of the torus
downwards to the wall of the pharynx.
 The levator palati, as it enters the soft palate, results in a small swelling
immediately below the opening of the tube.
 Behind the torus is the pharyngeal recess or fossa of Rosenmu¨ller(Fossa
of rosenmuller lies behind the nasopharyngeal orifice of E.T. is normally
packed with small but well organised lymph nodes. it is the most common
site of nasopharyngeal malignancy).
 Lymphoid tissue is present around the tubal orifice and in the fossa of
Rosenmu¨ller, and may be prominent in childhood.
 The tube is normally closed at rest and opens during yawning and
swallowing by the action of levator palate and to a lesser extent the
tensor palate
 The main functions of the E.T. is to
o Equalize the air pressure on both sides of the tympanic
membrane (protective )
o Allow drainage of secretions of middle ear .
o Prevent reflux of nasopharyngeal secretions

43
 Ostmann's pad of fat: fibrofatty tissue related to membranous part of
cartilagenous tube especially in the region of nasopharynx by keeping the
E.T.closed poecting it from nasopharyngeal reflux.
 In infants and young children , the tube is wider, shorter and more
horizontal allowing easier spread of infection to the middle ear .

MUSCLES ATTACHED TO THE EUSTACHIAN TUBE

 The tensor palati muscle:
o Arises from the bony wall of the scaphoid and from along the whole
length of the lateral cartilaginous lamina that forms the upper portion
of the front wall of the cartilaginous tube.
o From these broad origins the muscle descends, converges to a short
tendon that turns medially around the pterygoid hamulus and then
spreads out within the soft palate to meet fibres from the other side in
a midline raphe.
o The tensor palate separates the tube from the otic ganglion, the
mandibular nerve and its branches, the chorda tympani nerve and the
middle meningeal artery.
o It is supplied by the mandibular nerve.

 The salpingopharyngeus
o Is a slender muscle attached to the inferior part of the cartilage of the
tube near its pharyngeal opening, and it descends to blend with the
palatopharyngeus.
 The levator palati
o Contains a few fibres that arise from the lower surface of the
cartilaginous tube and originates from the lower surface of the petrous
bone, just in front of the opening for the entrance of the carotid, and
from fascia forming the upper part of the carotid sheath.
o It first lies inferior to the tube, then crosses to the medial side and
spreads out into the soft palate.
o Both the salpingopharyngeus and the levator palati are supplied from
the pharyngeal plexus.

44
Arterial supply of E.T.:
 The ascending pharyngeal
 Middle meningeal arteries supply the Eustachian tube.
 Artery of pterygoid canal

Venous drainage of E.T.

 The veins drain into the pharyngeal plexus and the

Lymph drainage of E.T.

 lymphatics pass to the retropharyngeal nodes.

Nerve supply of E.T

 The nerve supply arises from
o the pharyngeal branch of the sphenopalatine ganglion (Vb) for the
ostium
o the nervus spinosus (Vc) for the cartilaginous portion
o from the tympanic plexus (IX) for the bony part.

45
 D. THE MASTOID AIR CELL SYSTEM:

 It is a three sided pyramidal bone within the petrous part of the temporal
bone with its apex directed caudally.
 It contains air-filled cavities ( Mastoid air cells )
 In normal ears, the lining of the mastoid is a flattened, nonciliated
epithelium without goblet cells or mucus glands.
 According to the degree of cellularity , the mastoid process can be
divided into three types :
1. Pneumatic: Cellular 85%.
2. Diploic: less cellularity: Children,or may be due to closure of E.T.
3. Sclerotic: acellular .15%
 The mastoid air cells are arranged in groups :

1. THE MASTOID ANTRUM

o It communicates with the middle ear by way of the aditus.
o Aditus ad antrum: Is ashort canal connecting epitympanium with mastoid
antrum. Its floor is short process of incus. Medial wall is lateral
semicircular canal $ the bone lateral to aditus like abridge during
surgery.
o Is well developed at birth and by adult life has a volume of about 2mL.
o MEDIAL WALL relates to the posterior semicircular canal. It shows two
elevations in its medial wall : the bulge of the semicircular canal,
and the 2nd genu of the facial nerve anteroinferior to it (always
present and important land mark during surgery).

46
o THE ROOF of the mastoid antrum and mastoid air cell space form the
floor of the middle cranial fossa. More deeply and inferiorly is the dura of
the posterior cranial fossa and the endolymphatic sac which emerges
through the operculum on the posterior surface of the petrous bone and
derives from the endolymphatic duct, which has passed through the
vestibular aqueduct.
o POSTERIOR to the endolymphatic system is the sigmoid sinus, which
curves downwards only to turn sharply upwards to pass medial to the
facial nerve and then becomes the dome of the jugular bulb in the middle
ear space.
o BASE: The posterior belly of the digastric muscle forms a groove in the
base of the mastoid bone. The corresponding ridge inside the mastoid
lies lateral not only to the sigmoid sinus but also to the facial nerve and is
a useful landmark for finding the nerve itself. The periosteum of the
digastric groove on the undersurface of the mastoid bone continues
anteriorly and part of it becomes the endosteum of the stylomastoid
foramen and subsequently of the facial nerve canal.
o LATERAL WALL: The outer wall of the mastoid lies just below the skin
and is easily palpable behind the pinna.
MacEwen’s triangle (suprameatal triangle) is a direct lateral relation to
the mastoid antrum and is formed by
- Temporal line of supramastoid crest (a posterior prolongation of the
line of the zygomatic arch)
- Posterosuperior border of the external auditory meatus (spine of
henle).( Spine of henle is the land mark of MacEwen’s triangle)
- Tangential line between them

During surgery: to reach antrum , korner's septum must e removed.

korner's septum in mastoid separates squamous cells from the deeper
petrosal cells
korner's septum = persistent petrosquamous suture.

o The mastoid antrum may be the only airfilled space in the mastoid
process when the name acellular or sclerotic is applied. This condition
occurs in perhaps 20 percent of adult temporal bones and is seen in
individuals with chronic ear disease.

47
2. SINODURAL AIR CELLS
o between the dura of the roof and sigmoid sinus.
3. Tip cells :
o At the tip of the mastoid .
4. Retrofacial cells ;
o Behind the vertical portion of the facial nerve .
5. Perisinus cells :
o Over the sigmoid sinus .

6. Petrous apex
o Only 1/3 population has apnematized petrous portion of temporal bone.
o Surgically, the apex of the petrous bone is the most inaccessible
portion of the temporal bone. It is shaped like a foreshortened pyramid
and points anteriorly and medially.
o The posteromedial surface of the petrous apex is part of the posterior
cranial fossa, while the superior aspect of the bone forms the floor of
the middle cranial fossa.
o The internal carotid artery and the internal auditory meatus run through
the bony petrous apex.
o At the apex of the petrous bone (superiorly) is the trigeminal nerve
(gasserion ganglion) running into Meckel’s cave with the abducent
nerve passing close to its roof ( In Dorello's canal : is between the
petrous tip and the sphenoid bone)
o Gradenigo’s syndrome is the result of infection at the petrous apex and
comprises a lateral rectus palsy (diplopia), facial pain and discharging
ear.

7. Accessory group:
o Squamous
o Zygomatic (arch of zygoma)
o Styloid
o Occipital
o Peritubal

N.B. antrum is central group all others is prepheral

48
RELATIONS OF MIDDLE EAR CLEFT:

1- Temporal lobe of brain and meninges are above the antrum, aditus and
epitympanium separated by tagmen plate.
2- Cerebellum is posteromedial to mastoid air cells
3- Inner ear is medial to the antrum , aditus, tympanium, external ear is
lateral.
4- F.N :-its horizontal part runs downwords in the medial wall of the
tympanium and horizontal cemicircular canal runs posterosuperior to it.
Vertical part behind terminal.
5- 5th &6th C.N. lie close to the apex.
6- Jugular bulb at the floor.
7- ICA anterior to tympanium.
8- lateral sinus: posterior to mastiod cells .

49
THE INTERNAL AUDITORY MEATUS

 This is a short canal, nearly 1 cm in length and lined with dura, which
passes into the petrous bone in a lateral direction from the
cerebellopontine angle.
 It transmits the facial, cochlear and vestibular nerves and the internal
auditory artery and vein.
 The meatus is closed at its outer lateral end, or fundus, by a plate of bone
that is perforated for the passage of nerves and blood vessels to and from
the cranial cavity. The bony plate separating the fundus from the middle
and internal ears has a transverse crest – the crista falciformis – on its
inner medial surface, which separates a small upper region from a larger
lower area.
 Above the crest and anteriorly is the opening of the facial canal carrying
the facial nerve (VII). This is separated, by a small vertical ridge known as
Bill’s bar, from the posterior region that transmits the superior vestibular
nerve through several small foramina to the superior and lateral
semicircular canals, to the utricle and a part of the saccule.
 Below the transverse crest, the cochlear nerve lies anteriorly and leaves
the meatus through the cochlear area, which comprises a spiral of small
foramina and a central canal. The inferior vestibular nerve passes through
one or two foramina behind the cochlear opening to supply the saccule.
 Just behind and below the inferior vestibular foramen is the foramen
singlare, which contains the singular nerve. This runs obliquely through the
petrous bone close to the round window to supply the sensory epithelium
in the ampulla of the posterior semicircular canal.

50
N.B.
There are 4 openings in temporal bone
1- Internal auditory meatus= Porus acosticus: is the mouth of
internal auditory canal.
2- Vestibular aqueduct
3- Cochlear aqueduct: bonny channel connecting the scala tympani of the
basal turn with the sub arachenoid space of the poserior cranial cavity.
The average adult cochlear aqueduct is 6.2 mm long. Cochlear and
vestibular aqueduct: through which pus from supp. labyrinth go to brain.
4- Subarcuate fossa.

51
GROSS MORPHOLOGY

 The human cochlea is situated in the inner ear and consists of a spiral
tubular duct embedded in the petrous bone, lined by membranes.
 The membraneous compartments are filled with fluid and open into the
vestibule of the inner ear at the cochlear base.
 There are two membrane-covered openings in the bone at the base of the
spiral, the oval and round windows. The oval window membrane is attached
to the stapes bone of the middle ear, through which sound pressure waves
enter the cochlear fluids. The round window functions to release the
pressure induced by sound stimulation in the incompressible internal fluids
preventing rupture of the internal membranes.
 Pressure waves caused by sound travel through the cochlea and are
analysed by a complex and delicate sensory epithelium situated within the
cochlear duct, the organ of Corti. The basic function of the organ of Corti is
to transduce and process sound stimuli, converting them into electrical
signals in the auditory nerve for transmission to the higher auditory
pathway. During this process, the stimulus is analysed for frequency
content and amplitude, both properties being encoded in the individual and
ensemble discharge patterns of auditory nerve fibres.

52
ULTRASTRUCTURE OF THE COCHLEA
1. COCHLEAR DUCT
The cochlear duct is subdivided by two longitudinally running membranes
that separate three chambers, the scala tympani, scala media and scala
vestibule.

(a) Schematic cross section of the cochlea in situ. The cochlear duct, sectioned in five places, contains three
longitudinal chambers, scala tympani (st), scala vestibuli (sv) and scala media (sm) bounded by Reissner’s
membrane (Rm), the basilar membrane (bm) and the stria vascularis (stv). The central bony axis of the spiral, the
modiolus (mod) contains the spiral ganglion (sg) comprised of bipolar neurones that peripherally innervate the hair
cells and centrally form the cochlear nerve (co nv). Afferent fibres representative of low (blue), middle (green) and
high (red) frequency illustrate the tonotopic arrangement within the nerve. The sensory epithelium is indicated by the
blue box in one section. (b) Scanning electron micrograph of a dissected guinea pig organ of Corti (oC) showing its
location spiralling around the modiolus (mod). As indicated, high frequency sounds are detected near the base of the
spiral and low frequencies near its apex. Scale bar = 1 mm. (c) Light micrograph of the region represented by the
blue box in (a). The basilar membrane (bm) stretches from the osseus spiral lamina (osl) to the spiral prominence.
Nerve fibres (nf) enter through the osl, underneath the spiral limbus (sl) and inner sulcus (is). The organ of Corti lies
on the basilar membrane and contains one row of inner hair cells (IHC) separated by the tunnel of Corti (tc) from
three rows of outer hair cells (OHC). Stereocilia (s) project from the hair cells towards the tectorial membrane (tm)
which has retracted from its contact with outer hair cell stereocilia during the processing for histology. The outer hair
cells are supported by Deiters’ cells (DC) and outside of both are Hensen’s cells and Claudius’ cells (CC). Scale
bar = 100 mm.

53
A. THE SCALA MEDIA
Is triangular in section, boundaries represented by Reissner’s
membrane, which runs obliquely with respect to the basilar
membrane from a ridge of tissue, the spiral limbus near the
modiolus, to the lateral wall that runs along the inside of the bony
wall.

The basilar membrane

o Acellular layer stretches across the cochlear duct from a bony shelf
spiralling around the central bony modiolus, the osseous spiral lamina,
to a bony promontory, the spiral prominence, on the inside of the outer
wall of the cochlea.
o Although the cochlear duct widens substantially towards the basal end
of the spiral, the width of the basilar membrane decreases, the
difference being accounted for by the more rapidly increasing width of
the osseous spiral lamina.
o Underneath the basilar membrane is a layer of spindleshaped cells, the
tympanic cells, whose long axes are orientated in an apical-basal
direction along the cochlea.
o A branching spiral vessel lies under the basilar membrane.

Spiral limbus
o The longitudinal ridge of the spiral limbus is composed of a layer of
epithelial cells, the interdental cells, forming its upper surface and a
main body containing blood vessels and connective tissue cells
embedded in extracellular matrix.
o The side of the limbus facing the organ of Corti is concave.
o The concavity is lined by cells and forms a longitudinal groove, called
the inner sulcus, which borders the region of the organ of Corti
containing the sensory cells and the supporting cells.

Tectorial membrane
o An acellular flap.
o Forms a thin layer over the weakly convex top of the spiral limbus and
projects over the inner sulcus and across the organ of Corti.
o It widens substantially in crosssectional area as it does so, to a
maximum, then tapers again to a thin edge that lies over the outer side
of the organ of Corti.

54
Reissner’s membrane
o Consists of two layers of cells separated by a basement membrane.
o The layer facing into the scala tympani is the mesothelial cell layer and
consists of cells with an extremely thin cytoplasm and prominent central
nucleus.
o The layer facing the scala media is the endothelial cell layer, consisting of a
greater density of thicker cells covered by a dense mat of microvilli. The cells
within each layer are joined by tight junctions which act as an impermeable
barrier to ions and small molecules.
The lateral wall
o Consists of the stria vascularis, composed of three layers of cells on the
external side of which is a layer of fibrocytes and connective tissue called the
spiral ligament. Both regions are supplied with blood vessels.
o The three layers of the stria vascularis are composed of marginal cells facing
the scala media, the intermediate cells and the basal cells.
o Intermediate cells tend to have highly convoluted membranes.
o The marginal cells also have tight junctions connecting them together.
o The cells of the lateral wall contain a variety of ion pumps, enzymes and
transport proteins associated with homeostatic mechanisms for maintaining
the ionic composition of the fluids of the cochlea.
Endolymph
o In fact, the composition of the fluid within the scala media, endolymph, is
unusual for an extracellular fluid, containing high potassium but low sodium
levels at an unusually high positive electrical potential (180mV) called the
endolymphatic potential (EP).
o Maintenance of the EP is crucial to hearing. As with both the stria vascularis
and Reissner’s membrane, the cells of the organ of Corti facing the scala
media are joined by tight junctions. Thus the whole of the scala media is
chemically and electrically isolated from the other scalae, the only
communication being through ion channels in the sensory cells of the organ
of Corti. The EP is involved in driving currents through transduction channels
that are fundamental to hair-cell function and is thus a vital component
required for producing the high sensitivity to the cochlea. One of the initial
pathological changes found in animal models of age-related hearing loss is
damage to and loss of the fibrocytes in the lateral wall, which in turn leads to
reduction in EP. Subsequent age-related damage to other structures may
then, at least in some cases, result from failure of lateral wall homeostatic
mechanisms.

55
Cellular architecture and function of The organ of Corti
o Organ of corti Runs in a spiral along the floor of the scala media, situated
on basilar membrane.
o The organ of Corti extends across the upper surface of the basilar
membrane from the spiral limbus situated over the osseous spiral lamina to
the Claudius’ cells that lie between the edge of the sensory region and the
outer anchorage of the basilar membrane.
o The organ of Corti extends with a repetitively patterned structure along the
spiral for approximately 35mm in humans. However, there are gradual
variations longitudinally along the cochlear spiral that systematically
change the mechanical characteristics of the organ of Corti–basilar
membrane complex.
o The sensory region consists of two types of sensory hair cell that are
characterized by an apical bundle of hairs called stereocilia. In both cases,
the stereociliary bundle projects into the overlying endolymph from the
apical flattened surface.
o Whilst the apical regions of both supporting cells and hair cells reach the
upper surface of the organ of Corti, only in the supporting cells do the
basal ends rest on the basilar membrane, those of the hair cells held
above the basilar membrane within the supporting cell framework.

o Inner hair cells

 Usually form a single longitudinal row running along the inner side of
the sensory region with respect to the centre of the spiral, there may
also be additional inner hair cells outside the normal single row.

o Outer hair cells

 Outer hair cells form three rows running along the outer side of the
epithelium.
 The number of outer hair cell rows may increase to four or five over
short, apparently random, lengths of the organ of Corti, especially in
basal regions of the cochlea.
 As well as the gradual narrowing of the basilar membrane, the length of
outer hair cells and their supporting cells gradually decreases towards
the basal end. They also tend to be longer in the outermost row and
shorter in the innermost row, differences that are more pronounced
near the apex of the spiral and diminish towards the base.
 Other systematic changes include a gradual decrease in the length and
increase in the number of the stereocilia on both hair cell types towards
the base and other ultrastructural features

56
o Pillar cells
 Outer and inner hair cell are separated by two rows of pillar cells
forming a triangular arch-like structure in cross section and enclosing
the tunnel of Corti running lengthwise.
 The inner pillar row borders the inner hair cell row, and the outer pillar
row borders the innermost of the three outer hair cell rows.

o Deiters’ cells and phalangeal cells

 The outer hair cells are enclosed only at their bases and their apices,
The hair cells have specific types of supporting cell associated with
each of them.
 The base is held in the cup-shaped body of a Deiters’ cell which
extends a thin phalangeal process up to contact the apical part of
adjacent outer hair cells.
 The origin of the phalangeal process is from the top of the cell body
and it projects at an angle towards the reticular lamina.
 In the inner two rows of outer hair cells, the phalangeal process passes
alongside the neighbouring outer hair cell to join the reticular lamina
and form tight junctions between the more apical outer hair cell in the
row behind, and its neighbour.
 Thus, the Deiters’ cell belonging to the base of one hair cell supports
the apices of the next two along the cochlea, but in the row behind.
 In the case of the third row of outer hair cells, the Deiters’ cells have
the same basic pattern, but the process terminates along the outer
edge of the outermost row of outer hair cells thus forming a border to
the hair-cell region.
 The arrangement of the Deiters’ cells and outer hair cells produces a
repetitive triangular structure underneath the reticular lamina which is
presumed to provide strong mechanical support for the outer hair cells.
 Inner phalangeal cells lying next to the pillar cells and border cells
lying next to the first inner sulcal cells completely enclose the inner hair
cells.

57
o Reticular lamina

Tight junctions firmly connect the outer hair cells, at the level of their cuticular
plates, to pillar cells and to phalangeal process of Deiters' cells. This structure
presents a remarkable stiffness for bending in the radial direction. The
cytoskeletal rarefaction in the phalangeal processes of Deiters' cells is
responsible for a relative compliance of the reticular lamina in the longitudinal
direction.
 The upper surfaces of hair cells and their supporting cells, and the pillar
cells, are held together by tight junctions.
 This system of junctions when viewed from above the organ of Corti
gives the appearance of a network in which the hair cells and supporting
cells are embedded, which has been named the reticular lamina.
 The reticular lamina, which is oblique with respect to the basilar
membrane near the apex, becomes more parallel to it near the base as
the length of the outer hair cells decreases.

o Hensen’s cells
 Beyond the third row of outer hair cells lie Hensen’s cells which, at least
in guinea pigs, can contain large lipid droplets.
 Hensen’s cells become smaller and the tectorial membrane changes in
cross-sectional area, becoming thicker near the basal end of the
cochlea.

o Cuboidal Claudius’ cells

 The outer edge of the organ of Corti curves down from the reticular
lamina, lined by the Hensen’s cells, to a single layer of cuboidal
Claudius’ cells which cover the remainder of the basilar membrane
out to the spiral prominence.

58
FUNCTION OF THE ORGAN OF CORTI
Detect sounds and decompose them into their component frequencies
o In the process converting the physical vibrations into an electrical
response (mechanoelectrical transduction) and then causing neural
signals to be transmitted along the auditory nerve and higher auditory
pathway for central processing.
o Sound pressure waves in cochlear fluids set up waves of motion along
the basilar membrane (called travelling waves) which peak in different
parts of the spiral according to frequency, resulting in mapping of high to
low frequency components along its length.
o The morphological gradients of organ of corti contribute to this mapping
by gradually varying the mechanical resonance properties of the basilar
membrane-organ of Corti complex.
o The hair cells detect the basilar membrane motion, being stimulated
more strongly at the point coinciding with the peak of the travelling wave
than elsewhere.
o The innervation pattern of the organ of Corti strongly suggests that inner
hair cells contribute most to the neural signalling representing sensory
transduction and processing by the cochlea and it is assumed that they
perform a more or less passive sensory role.
o However, the cochlea contains an active amplifier that enhances the
ability to detect and separate frequencies in sound. The outer hair cells
are thought to represent this amplifier, mechanically boosting sound-
induced vibrations of the basilar membrane to produce a sharply tuned,
highly sensitive displacement pattern along the basilar membrane that is
reported by inner hair cells.

59
HAIR-CELL ULTRASTRUCTURE AND FUNCTION
A.Outer hair cells

1. Nucleus, 2. Stereocilia, 3. Cuticular plate, 4. Radial afferent ending

(dendrite of type I neuron), 5. Lateral efferent ending, 6. Medial
efferent ending, 7. Spiral afferent ending (dendrite of type II neuron)

"W" pattern of OHC stereocilia

60
 a)Stereocilia (around a hundred) are generally arranged in three rows of graded lengths. In addition to
thin tip links (shown here in red) which are involved in the mechano-transduction process, stereocilia
are attached by transverse (lateral) links, both in the same row and from row to row.Tip (red arrow) and
lateral (blue arrow) links between two stereocilia.b) With transmission electron microscopy (TEM), the tip
link (red arrow) and a lateral link (blue arrow) between medium and tall stereocilia are clearly visible. At both
ends of the tip link, a membrane condensation is seen.

Electro-mechanical transduction

61
- The apical part of the outer hair cell is the sensory end, bearing the
stereociliary bundle, whilst the basal end consists of the rounded synaptic
pole where the cell connects with afferent and efferent nerve fibres of the
auditory nerve.
- The apical surface is flattened and, when viewed from above, triangular or
heart-shaped whilst the cell bodies are cylindrical
- The STEREOCILIA are arranged in a very pronounced W- or V-shape.
- The rows increase in height across the bundle, like a staircase, along a
radial axis from the modiolus to the lateral wall. In the case of the outer
hair cells, the top of the bundle is in contact with the underside of the
tectorial membrane whilst the inner hair stereocilia are probably free
standing.
- The stereocilia are cylindrical, narrowing sharply to an ankle region where
they join the cell and, at least in the intermediate rows, with a bevelled tip.
They are angled towards each other so that they converge at their tips.
The shorter stereocilia are slightly narrower than the intermediate and
taller stereocilia. Stereocilia contain a core of parallel actin filaments
cross-linked by the actin associated proteins plastin and espin. In the
lower third of the stereocilium, dense material becomes associated with
the centre of the core. Along with a group of actin filaments, this extends
down as a rootlet which contains tropomyosin into an actin-rich
filamentous mat, the cuticular plate, located beneath the apical
membrane. The stereocilia also contain several unconventional
(nonmuscle) myosins and are enriched in calmodulin and other calcium
handling proteins.The stereocilia are connected together by extracellular
filaments. A single filament, the tip link runs from the tip of each
stereocilium of the shorter rows to the side of the adjacent stereocilium
immediately behind. At the upper attachment of the tip link, there is a
distinctive electron dense plaque lying between the membrane and the
actin core. At the lower attachment there is dense material over the actin
core, separated by a gap from the membrane of the tip. Below the lower
attachment is a zone called the contact region where the membranes of
the two converging stereocilia approach very closely. Lateral links connect
the shafts of adjacent stereocilia, both within and between rows. Where
the lateral links connect to the stereocilia, the membrane and the adjacent
actin filaments inside show an increased density. The stereociliary

62
 membrane contains proteins associated with calcium control (plasma-
membrane calcium ATPases) and mechanosensitivity (mechanoelectrical
transduction channels). There may also be other ion channels (for
example, ATP gated receptors, acid sensing channels) in the stereociliary
membrane, but their contribution to hair cell function is unclear. Although
the precise location of the transduction channels has still to be confirmed,
it seems likely that they are associated with the regions near the tips of the
shorter stereocilia. During transduction, they are gated (opened)
mechanically by movements of the hair bundle, which modulate their rate
of opening and closing. Deflections of the bundle are driven by the motion
of the basilar membrane–organ of Corti complex that causes shearing of
the tectorial membrane parallel to the surface of the organ of Corti. This in
turn directly drives the outer hair cell stereociliary bundle backwards and
forwards. Deflections in the direction of increasing stereociliary height
depolarize the outer hair cells by causing the channels to open and allow
an influx of cations, whilst opposing deflections hyperpolarize the hair cells
by closing the channels and orthogonal deflections have little effect. The
magnitude of the receptor potentials produced by depolarizing deflections
is increased by the high positive EP (180mV) which, together with the
resting membrane potential of the hair cell (around #70mV), results in a
much greater potential difference than normally present even in neurones
of 150mV between endolymph and the inside of the hair cell. This large
driving force increases the sensitivity of the sensory cells substantially.
The tip link has been postulated to represent a ‘gating spring’ for the
transduction channels, a physical mechanism to open them. It is located in
such a position that it would be stretched in the case of depolarizing
deflections and could pull channels open, and relaxed in the case of
hyperpolarizing deflections, which would allow them to close.
- The hair bundle may also have an active role in mechanical amplification.
In a range of vertebrate hair cells the bundles show evidence of force
production that produces active motion which adjusts the position of the
bundle (adaptation). Recent studies suggest that rapid force production
driven by a calcium-dependent process could enhance the
mechanoelectrical transduction response to amplify the very smallest
stimuli.

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- Again, the high EP would make this amplification more effective.
- A slower adaptation has also been noted in a range of hair cells.
- The unconventional myosins could interact with actin in the stereocilia
providing one source of such force production, as acto–myosin
interactions are the basis of muscle contraction and other motile
processes throughout the body. However, the acto–myosin interaction is
probably too slow to account for the rapid force production of the bundle.
- The cuticular plate contains actin filaments that are more randomly
organized into a meshwork than the parallel bundle found in the
stereocilium. Nevertheless, there are regional differences and structural
features in the cuticular plate that indicate a high level of organization in
both the actin filaments and other actin-associated proteins that are
present. For example, a ring of circumferential actin filaments has been
described in some species and there are nodes of dense material
arranged in layers. The cuticular plate is interrupted in the region just
behind the stereociliary bundle and adjacent to its centre. Here, a channel
of cytoplasm runs through the plate to the apical membrane. This channel
contains a basal body.
- In hair cells from non-mammalian species and mammalian vestibular hair
cells, the basal body underlies a kinocilium that projects alongside the hair
bundle. Other, usually narrower, cytoplasmic channels run around the
apical perimeter of the plate and penetrate through it. The cuticular plate
acts as an anchoring structure for the stereocilia but could also be a site
where the mechanical properties of the hair bundle are modified via the
rootlets.
- In the apical part of the cochlear spiral in guinea pigs, a long projection
from the underside of the cuticular plate extends down into the cytoplasm.

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- In the cell body, there is a concentration of Golgi apparatus, mitochondria
and endoplasmic reticulum just beneath the cuticular plate and many outer
hair cells contain a large body (Hensen’s body) composed of multiple
concentric layers of membrane, especially in cells near the cochlear apex.
These layers appear to be contiguous with a stack of submembraneous
cisternae lying just beneath the plasma membrane and extending down
the side from near the cuticular plate to the level of the nucleus. The latter
is spherical and situated in the basal region of the cell. Between the
outermost of these and the plasma membrane is a cortical lattice complex
of actin and spectrin filaments and rows of short pillars.
- The plasma membrane contains a protein called prestin that changes
shape when subjected to a voltage change.
- Isolated outer hair cells can be made to undergo very fast contraction–
elongation cycles by electrical stimulation, a property called electromotility.
The cortical lattice is believed to act as a cytoskeletal spring involved in
converting the voltage-evoked conformational changes in prestin into the
movement of the whole hair cell. This outer hair cell motility is generally
regarded as the main source of active cochlear amplification although, as
noted above, an alternative or additional source of amplification may be
the hair bundle.
- Mitochondria are also found lining the subsurface cisternae and a few are
scattered in the cytoplasm. They tend to be concentrated beneath the
nucleus of the cell, interspersed with a range of membraneous coated and
uncoated tubules and vesicles, and microtubules.
- Synaptic specializations associated with afferent and efferent terminals
are present in the basal pole (see under Innervation of the organ of Corti
and Descending pathways).

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- As well as the gross morphological gradients noted under Cellular
architecture and function of the organ of Corti above, the outer hair cells
show other gradients in ultrastructural morphology along the length of the
cochlear spiral and also across the three rows. In the guinea pig, they
display more layers of subsurface cisternae near the cochlear apex and
the subcuticular projection of the cuticular plate decreases in extent
towards the cochlear base, and from the outermost to the innermost row in
parallel with the differences in hair-cell length. These gradients are
presumed to reflect differences in cell physiology and micromechanics that
contribute to the systematic changes in properties of the organ of Corti–
basilar membrane complex.

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B.INNER HAIR CELLS

1. Nucleus, 2. Stereocilia, 3. Cuticular plate, 4. Radial afferent ending

(dendrite of type I neuron), 5. Lateral efferent ending, 6. Medial efferent
ending, 7. Spiral afferent ending (dendrite of type II neuron)

Linear pattern of IHC stereocilia

- The inner hair cells have a flattened or slightly concave apical surface that
is ovoid when viewed from the scala media and a flask-shaped cell body
with a wide centre tapering basally and apically.
- The inner hair cell stereociliary bundle consists of three to four relatively
linear rows of stereocilia, their long axes running parallel with the hair-cell
row. However, there is often evidence of a shallow notch approximately
halfway along where the rows indent slightly, and the ends of the bundle
tend to curve round to some degree towards the modiolus. The effect of
these features is to produce a shallow version of the more pronounced W-
shape of outer hair cells, although the stereociliary arrangement is less
precise.

67
- Each row contains stereocilia of generally similar height, though tending to
shorten near the ends of the row. As with outer hair cells, the height of the
rows increases in a step-wise manner across the bundle from the modiolar
to the strial side. The stereocilia themselves are cylindrical actin-
containing structures as in outer hair cells, with bevelled tips in the second
tallest row and rounded tips in the others, and narrow ankles. The
stereocilia have dense rootlets that penetrate into an apical cuticular plate
and thus appear to be anchored within, as in outer hair cells. Externally,
the stereocilia are again connected together by filamentous lateral links,
which bind them both sideways and across the rows, and a contact region
and tip links as seen in outer hair cells.
- The process of transduction by hair cells is similar to that described for
outer hair cells. However, unlike the tallest stereocilia of the outer hair
cells, the inner hair cell stereocilia do not appear to insert into the tectorial
membrane. Thus, sound-induced motion of the basilar membrane is
thought to stimulate the inner hair cell bundle via fluid motion of the
endolymph between the tectorial membrane and the hair cell apex.
- The cell body contains a region rich in Golgi bodies, endoplasmic
reticulum, mitochondria and other evidence of synthetic activity just below
the cuticular plate and the neck of the hair cell.
- The centre of the cell is occupied by a large spherical nucleus, with
clusters of mitochondria and endoplasmic reticulum around it.
- Beneath the lateral plasma membrane in the neck region and extending
down to the level of the nucleus is a single layer of cisternal membranes.
The space between these membranes and the plasma membrane
contains a cortical lattice of actin and spectrin filaments and rows of pillars
connecting to the membrane similar to that of outer hair cells. However,
there is no evidence of prestin in the membrane of the inner hair cells, and
they do not appear to display electromotility like that of outer hair cells.
- The basal end of the cell is the synaptic pole where the terminals of the
afferent auditory nerve fibres make synaptic contact. This region is filled
with vesicles and coated and uncoated membraneous tubules and has
synaptic specializations called synaptic ribbons with associated synaptic
vesicles.

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- Depolarization of the hair cell is believed to result in calcium-dependent
vesicular release of the amino acid glutamate onto the postsynaptic
afferent terminal, which contains glutamate receptors. This depolarizes the
nerve ending, resulting in the generation of action potentials. Efferent
contacts directly onto inner hair cells are rare.

ULTRASTRUCTURE OF SUPPORTING CELLS

Deiters and phalangeal cells
The supporting cells that are directly associated with the two hair-cell types
were introduced earlier. Deiters’ cells are associated with outer hair cells,
and are also called outer phalangeal cells. The latter name arises from the
presence of the phalangeal process that extends up from the body of the
Deiters’ cell. The process is supported by one or more microtubular bundles
that appear to originate in basal filamentous material and end in the apical
junctional complex and associated material. The Deiters’ cell cup contains
dense material directly beneath the outer hair cell base which is rich in actin
and appears to be physically strongly attached to the hair-cell base. The
inner phalangeal and border cell lateral membranes are closely associated
with that of the inner hair cell on the strial and modiolar sides respectively.
Both cells are longer than the hair cell and jointly surround the basal pole
before extending down to attach to the basilar membrane next to each other
where their nuclei are located. At their tops, they are connected to the hair
cell and other supporting cells by junctional complexes. Both cells display
finger-like projections that interdigitate with afferent auditory nerve fibres as
they approach the base of the inner hair cell. These cells are presumed to
have a protective role because they strongly express transporters
associated with glia-like uptake of glutamate after its release from the hair
cells. This uptake is believed necessary to prevent glutamate-induced nerve
cell damage that can occur with various auditory trauma. These supporting
cells are also interconnected by gap junctions, which could provide a route
of potassium ion removal from around the hair cells.

69
Pillar cells
Between the two hair cell types are the two rows of inner and outer pillar
cells, or rods of Corti. These cells are supported by a thick microtubular
bundle emanating from apical and basal filamentous zones composed of
actin and other cytoskeletal proteins. The microtubular bundles also contain
actin filaments interspersed with the microtubules. The two pillar cells have
radial feet resting on the basilar membrane above which the pillars are
angled towards each other, forming the arch over the tunnel of Corti, the
rounded head of the outer pillar cell contacting the concave underside of the
head of the inner pillar cell in a strong joint held together by junctional
complexes. Thus, the inner pillar cell head curves over the top of the outer
pillar cell head and forms a rectangular profile obscuring the top of the outer
pillar cell when viewed from above the reticular lamina, and lying between
the inner hair cells and first row of outer hair cells. The outer pillar, however,
produces a process from the side of the head region, which contains a
microtubular bundle, and extends up to the reticular lamina in the direction
of the stria vascularis. It surfaces between two adjacent outer hair cells of
the first row. Like all other supporting cells in the organ of Corti, the outer
pillar cell has an apical surface exposed to the endolymph. The cytoskeletal
organization of pillar cells strongly suggests that they have an important role
in providing physical support to the organ of Corti.

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INNERVATION OF THE ORGAN OF CORTI

The radial afferents (blue) and the lateral efferents (pink) innervate the inner
hair cells; the spiral afferents (green) and the medial efferents (red) innervate
the outer hair cells.

The IHC is synaptically connected to all type I spiral ganglion neurons (refs. a1, c5)
forming the radial afferent system (blue) going to the cochlear nuclei (CN). The lateral
efferent system (pink) arising from small neurons in the ipsilateral lateral superior olivary
complex (LSO) brings a feedback control to the IHC/type I afferent synapse.

The OHC synapses with a few (at least in basal and mid-portions of the cochlea) small
endings from type II spiral ganglion neurons (ref. c1), forming the spiral afferent system
(green). In turn, large neurons of the medial efferent system (red), from both sides of the
medial superior olivary complex (MSO), form axo-somatic synapses with the OHC.

71
 Type I (blue) spiral ganglion neurons (95% of the ganglion
neurons) have a single ending radially connected to IHCs.
Type II (green) small, unmyelinated neurons spiral basally
after entering the organ of Corti and branch to connect about
ten OHCs, generally in the same row.

o Acoustic information from the hair cells is transferred by the auditory

portion of the VIIIth cranial nerve (the vestibulocochlear nerve) to the
ipsilateral cochlear nuclear complex in the brain stem.
o The auditory nerve is composed of afferent fibres projecting from spiral
ganglion neurones, the cell bodies of which reside in the modiolus, just
central to the osseus spiral lamina.
o The spiral ganglion thus follows the course of the organ of Corti inside the
modiolus.
o The neurons are of two types, type I that innervate the inner hair cells and
type II that innervate the outer hair cells.
o The afferent innervation of the two hair cells types differs considerably in
both number and distribution of fibres.

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o The majority of spiral ganglion neurones (up to 95 percent) are TYPE I and
innervate the inner hair cells in a convergent manner.
 Up to 20 type I neurones innervate each inner hair cell via a peripheral
process that terminates on the hair-cell base at a ribbon synapse.
 These synapses consist of post-synaptic density on the afferent terminal
and a presynaptic density on the hair cell membrane, attached to which
is a dense presynaptic ribbon or bar. A single layer of synaptic vesicles
usually clusters around the bar.
 The cell body of the type I spiral ganglion neurone, its peripheral
process and the central axon, which projects to the cochlear nucleus,
are myelinated.
 The peripheral process becomes unmyelinated in the osseus spiral
lamina just before it enters the organ of Corti through a hole (the
foramen nervosum) in the upper border of the spiral lamina, the
habenula perforata, to approach the inner hair cell.
 At the hair-cell base, each process widens into a bulb where the
synapse forms.
 The central process enters the modiolus.
 At the apex of the cochlear spiral, the central processes enter the
middle of the modiolus and, progressively down the spiral, more are
added successively to the periphery of the nerve.
 Thus, as the nerve grows to its maximum diameter where it exits
through the internal auditory meatus at the base of the spiral, low
frequency fibres occupy the centre of the auditory nerve, with fibres of
increasingly higher frequency found towards the periphery.
 The responses of the type I auditory nerve fibres reflect the input from
inner hair cells.
 When the inner hair cell is depolarized it releases a neurotransmitter,
generally believed to be glutamate, from the presynaptic vesicles onto
post-synaptic glutamate receptors on the nerve ending which itself then
becomes depolarized.
 Provided the amount of depolarization is sufficient, this triggers action
potentials in the nerve fibre.

73
 As noted above, a fibre originating near the base of the cochlea will
have its best response to a high frequency, and one originating near the
apex to a low frequency.
 For any tone, the peak of the travelling wave will occur at the point of
maximum resonance on the spiral for that frequency.
 If the tone is loud enough after cochlear amplification, there will be
sufficient displacement of the inner hair cell stereocilia to depolarize the
hair cell and evoke action potentials in the attached cochlear nerve fibre.
 The number of action potentials per second increases with increasing
sound intensity and for frequencies below 5 kHz, the action potentials
can become phase locked (i.e. occur at a particular point in the cycle)
and so provide information on the frequency content of the stimulus as
well as the intensity.
 Each individual nerve fibre is thus defined by its characteristic
frequency, which is the frequency for which it has its lowest threshold.
 For each fibre, the threshold increases (i.e. there is a weaker response)
for tones that are higher or lower than the characteristic frequency,
reflecting the change in position of the peak of the travelling wave and
the narrowness of the peak. The latter depends on the cochlear
amplifier.
 Thus, the sharpness of tuning of the basilar membrane organ of Corti
complex is represented by the sharpness of tuning of the inner hair cell
and the nerve fibre.
 This, in turn, ultimately determines the ability of the auditory system, to
distinguish between sounds of different frequency.
 When the amplifier is impaired, for example through loss of outer hair
cells, fibres will have higher thresholds and broader tuning, making
separation of frequencies (frequency selectivity) poorer and requiring
louder sounds to evoke a response.

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o The TYPE II NEURONES innervate the outer hair cells in a divergent
manner and tend to be smaller than the type I neurones.
 Their peripheral processes and central axons are unmyelinated.
 The peripheral processes enter the organ of Corti through the same
route as that of the type I processes, and traverse the inner hair cell and
inner pillar cell rows, crossing the floor of the tunnel of Corti as basilar
fibres.
 They then travel towards the cochlear base for varying distances as
outer spiral fibres, before branching to innervate up to ten outer hair
cells each.
 The form of the synapse with the outer hair cell is not dissimilar to that
of the inner hair cell.
 There are presynaptic dense bodies and evidence of some associated
vesicles.
 The relatively small number of fibres to the outer hair cells, which are
three times more numerous than inner hair cells, suggests they are not
the primary signalling pathway from the cochlea, and indeed type II cells
are difficult to record from using microelectrodes, unlike the type I cells.
What they signal to the brain is uncertain, although there is evidence
that they respond to very loud sounds.
o As well as afferent innervation, the auditory nerve contains myelinated
and unmyelinated efferents that represent descending projections from
the brainstem. The majority of the myelinated fibres end on outer hair
cells directly, whereas the majority of unmyelinated efferent fibres end on
the peripheral processes of afferent auditory nerves just below the inner
hair cell.
o Primates, including humans, are noted in particular for having nerve
terminals that form reciprocal synapses with outer hair cells. These have
features of both afferent and efferent endings. Moreover, very recent work
in rodents suggests that there may be multiple synaptic contacts between
inner hair cells, the type II afferents to outer hair cells and lateral efferents
suggesting the possibility of local circuitry within the organ of Corti
between the inner and outer hair cell innervation. The functions of this
putative local circuitry have yet to be determined.

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SUMMARY OF FUNCTIONAL PROPERTIES OF THE ORGAN OF CORTI–
BASILAR MEMBRANE COMPLEX
The cochlea performs frequency analysis, splitting complex sounds up into
component tones and signalling that information to the brain. Thus the
cochlea responds to sound stimuli by:
o Producing travelling waves along the basilar membrane that peak more
apically for decreasing sound frequency and whose amplitude reflects the
intensity of the sound;
o Detecting the motion of the basilar membrane through deflection of the
stereocilia which produce receptor potentials in the hair cells graded in size
with the amplitude of the motion;
o Enhancing the motion of the basilar membrane with a biomechanical
amplifier residing in the stereociliary bundles and/or the outer hair cell lateral
wall (somatic motility); enhancement is nonlinear and maximally amplifies
motion at the point of maximum response, hence producing high frequency
selective peaks in the travelling waves;
o Detecting the resultant basilar membrane motion through deflection of inner
hair cell stereocilia which results in neurotransmitter release and the
production of action potentials in the auditory nerve fibres. The frequency is
encoded by (1) the place of origin of the nerve fibres (i.e. those connected to
the hair cells which are being stimulated most) and (2) the timing of action
potentials (for frequencies below 5 kHz), whilst intensity is coded by the rate
of action potential firing. The main evidence for the duplex activity of the hair
cells in the cochlea is:
o The differential distribution of innervation with afferent fibres mainly to inner
hair cells and efferent fibres mainly to outer hair cells, indicating inner hair
cells are the main signalling pathway
o Selective loss of outer hair cells produced by aminoglycoside antibiotics
which causes a reduction in sensitivity and frequency selectivity (broadening
of tuning), indicating their primary role in determining these properties
compared with inner hair cells.
o Otoacoustic emissions (sound generated within the cochlea) which is
evidence of active amplification.
o Evoked motility of the outer hair cells in vitro15 and in vivo, the latter also
capable of generating otoacoustic emissions16 which suggests they are the
site of an active process.
o Active force production by the outer hair cell stereocilia that has been
observed in the adult mammalian cochlea in vitro.

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 AUDITORY PATHWAY
A. THE ASCENDING AUDITORY PATHWAY

pathway. Schematic diagram of the auditory pathway represented in slices of

the cerebrum (upper), midbrain and brainstem. The main nuclei between the
cochlea and cerebral cortex are shown (orange). The major pathways from
each cochlea are coloured black and green respectively. More minor pathways
are represented with finer black or blue arrows.

1ST ORDER NEURON:

 located in spiral ganglion and are bipolar
 Prepheral process: receive from organ of corti
 Central process: terminate in dorsal and ventral cochlear nuclei

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2ND ORDER NEURONE:
 Lie in dorsal and ventral cochlear nuclei
o The cochlear nuclear complex is subdivided into dorsal cochlear nucleus
(DCN) and ventral cochlear nuclei.
o Ventral cochlear nuclei composed of anteroventral cochlear nuclei (AVCN)
and posteroventral cochlear nuclei (PVCN).
o These three regions are distinguishable on the basis of their location and
cytoarchitecture (the range of cells of different morphology).
o The central processes of type I spiral ganglion neurones enter the cochlear
nuclear complex and immediately bifurcate, sending branches to the DCN
or PVCN and the AVCN.
o Low frequency fibres divide ventrally, and high frequency fibres dorsally so
that the cochleotopic map of frequency, represented anatomically by the
distribution of fibres in the auditory nerve, is maintained across the
cochlear nuclei as a tonotopic map of neurones responding to
progressively higher frequency from one side to the other.
o The auditory nerve afferents in the AVCN terminate on the principal
projection neurones of the cochlear nuclear complex, the spherical/bushy
cells, so called because they have round cell bodies and bushy dendritic
fields.
o The end of most auditory nerve fibres expands into a single very large
terminal, the end bulb of Held, which cups around the soma of the
spherical cell (very low frequency neurones, o1 kHz, may branch to form
two endbulbs).
o One or two such terminals are found on each spherical cell. This large
excitatory terminal contains large numbers of round neurotransmitter
vesicles typical of glutamatergic terminals and ensures rapid transmission
of the signal from the auditory nerve fibre that faithfully preserves the
original frequency selectivity and sensitivity of the cochlear response.
o Accordingly, these cells have electrophysiological responses to sound that
are called primary-like because they reflect the primary input from auditory
nerve fibres.

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o However, in the DCN or PVCN, several auditory nerve fibres may contact a
single multipolar (or stellate) cell, which have more complex responses.
This multiple input means these cells may analyse the pattern of sound, or
determine the intensity.
o Also in the PVCN are octopus cells, which have an extended dendritic field
lying across a number of auditory nerve fibres, so that they receive input
representing a range of frequencies. These cells respond rapidly and may
be responsible for determining the precise time of arrival of sounds. They
also send signals to motor nuclei in the brain stem and midbrain so they
may be involved in acoustic startle responses, where loud or unexpected
sounds evoke movement.
o As well as these projection neurones, the cochlear nuclei contain
interneurones and receive inputs from higher up the auditory pathway that
produces inhibition and generates more complex responses in some
neurones.
o In particular, it has been suggested that these complex responses in the
DCN are important in determining what sounds are.

 Most of axons cross over trapezoid body and terminate in superior olivary
nucleus.
 Many end in trapezoid body and lateral leminiscus
 Some remain uncrossed.

3RD ORDER NEURONE :

 Lie in superior olivary nucleus
o The auditory pathway splits as it leaves the cochlear nuclear complex. The
dorsal pathway projects directly to the inferior colliculus, the ventral
pathway divides further and projects to both the ipsilateral and contralateral
superior olivary complex. This makes the superior olivary complex the first
part of the ascending auditory pathway where major binaural comparisons
can be made.

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o The superior olives receive binaural information from spherical/bushy cells.
This arises from collaterals from the output fibres of the cochlear nuclei on
the same side that then cross over to the opposite superior olivary
complex. This enables the superior olives to function in sound localization.
o Each superior olivary complex contains an S shaped lateral olivary
nucleus, a disc-shaped medial olivary nucleus and the medial nucleus of
the trapezoid body together with smaller periolivary nuclei.
o Within the medial olivary nucleus, there are neurones that use the binaural
inputs to compare the time of arrival of sounds to each ear. For example,
for a sound coming from the left of a person’s head, the left ear would
receive the sound first, the right ear second because of the difference in
distance to the two ears.
o The further to the left, the greater the difference in time of arrival. From
experimental work in owls, computation of the interaural time delay would
allow the medial olivary nucleus to localize a sound. In fact, a spatial map
seems to be present, represented by gradual changes in the responses of
the neurones across the anterior–posterior axis of the medial olive to
specific interaural time differences.
o This method of localization would work for discontinuous sounds and for
relatively low frequency continuous tones, but breaks down at higher
frequencies for continuous tones because individual auditory nerve fibre
responses cannot encode accurate timing information above 5 kHz due to
limitations on the rate of firing of action potentials. Sound localization at
higher frequencies may be carried out by comparing sound intensities.
o If a sound source is on the left of a person’s head, as before, it is closer to
the left ear than to the right and so sounds louder.
o Neurones that detect differences in sound intensity are located in the
lateral superior olives. Most of these neurones receive an excitatory input
from the ipsilateral cochlear nucleus and an inhibitory one from the
contralateral cochlear nucleus. Thus if a sound is of equal intensity in both
ears because it originates on the midline, the inhibition and excitation of
the binaural neurones between the two nuclei balances out. If, however, a
sound is louder in the left ear, the excitation will be stronger for the
neurones in the ipsilateral olive and the inhibition weaker enhancing the
ipsilateral response, whilst the converse will hold true for the neurones in
the contralateral olive, which will have a much weaker response

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 Axons pass through lateral lemniscus and reach inferior colliculus.

The ascending projections from the SOC pass along the lateral lemniscus (C) and arrive at
the external nucleus of the inferior colliculus (A and B). The fibers that send low
frequency information (in red and marked with number 3) project along the lateral side
of the lemniscus (ipsilaterally) and reach the external nucleus of the IC. The fibers that
send high frequency information (in blue, and number 1) cross over in the superior
olivary complex and project up inside the contralateral lemniscus to the medial region of
the IC. The colour scale at the bottom indicates sound frequency.

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 Nuclei of lateral lemniscus

o These nuclei are composed of different neuronal groups organised

between the ascending fibers of the lateral lemniscus. They are organised
into two regions:
o The ventral nucleus, composed of monaural neurons that receive
information from the ipsilateral ventral cochlear nucleus and are not
arranged tonotopically. This nucleus has a role in processing the duration
of complex sounds, which is very important for understanding language.
o The dorsal nucleus, composed of binaural neurons that recevei
information from both ears, thanks to the information exchange that
happens at the commissure between the lemnisci and the olivary complex.

4TH ORDER NEURONE

 Lie in the inferior colliculus

o function of inferior colliculus:-

 It is the center of various auditory reflexes.
 The inter connection between the inferior colliculi of both sides allows
bilateral cortical representation of the auditory sensation.
 bilateral representation of the auditory sensation.
 The auditory impulses from each ear are bilaterally represented in both
temporal lobes due to:
1- Inferior colliculus
2- Pass to 2 lateral leminiscus
So,unilateral cortical lesion lowers hearing power in both sides.

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o There are four bumps on the surface of the midbrain, which together form
the corpora quadrigemina. These are composed of the two superior and two
inferior colliculi.
o The inferior colliculi:
 Receive direct input from the brainstem auditory nuclei via a tract called
the lateral lemniscus.
 Each of the inferior colliculi consists of a central nucleus which receives
the major auditory input, and an outer region composed of a dorsal cortex
and an external lateral cortex.
 The external portions of the inferior colliculus receive connections from
cerebral cortex and from mutimodal sources respectively.
 In the inferior colliculi, the two pathways that emerge from the cochlear
nuclear complex join together again for further analysis.
 More complex responses are found in inferior collicular neurones, and
further features are extracted and mapped towards understanding ‘what a
sound is’.
 The central nucleus is layered into isofrequency bands. Along each band,
the cells have flattened dendritic fields and respond best to approximately
the same frequency.
 The higher frequency bands are found towards the midline of the brain,
low frequency bands more towards the outside, producing a tonotopic
map.
 Superimposed on each band is another map that relates to intensity. This
is best visualized by thinking of each isofrequency band as a disc.
 The cells in the centre of the disc have low thresholds which means they
respond to quiet sounds, whilst moving out to the periphery of the disk
there are concentric areas in which the threshold of the neurones
increases, hence requiring louder and louder sounds to stimulate them.
 There are also neurones that respond to timevarying stimuli, such as
changes in frequency (frequency modulated – for example, a sound
moving towards you or away, such as an aeroplane) or in intensity
(amplitude modulated – for example, an air-raid siren). These responses
also seem to be mapped approximately from front to back across each
inferior colliculus.
 These intersecting maps in the inferior colliculi are thus able to extract
complex features of sounds.

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  At the next nucleus in the auditory pathway, cells have been found that
respond to particular complex sounds, for example, the mew of a kitten.
 These maps in the inferior colliculus thus provide a basis for recognizing
patterns in sound.
 There are also neurones in the inferior colliculus that are involved in
sound localization; as many as threequarters of the neurones may have
binaural responses. These neurones may provide input to the superior
colliculus in which there are visual and auditory space maps that can
therefore be compared so that sounds can be assigned to specific
objects.
 The inferior colliculus is also involved in auditorymotor responses, for
example, controlling middle ear muscles, which can be used to attenuate
loud sounds and protect the ear. In addition, there are projections to
motor nuclei that contribute to turning the head or moving the eyes in
response to sound.

 Axons pass through inferior brachium to reach medial geniculate body.

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5TH ORDER NEURONE:
 Lie in the medial geniculate body.

 Their axons form the auditory radiation which passes through the part of
the internal capsule to reach the auditory area in the temporal lobe.

(a) Diagram of the brain showing the region of primary auditory cortex (area A1). (b) Diagram of
A1 showing the arrangement of isofrequency laminae, the numbers representing best frequency
(in kHz). (c) Block of auditory cortex showing the isofrequency planes orthogonal to the normal
cortical layering (I–VI) and across the planes, alternating strips of cells that show either
excitatory–excitatory responses (Ex–Ex) or excitatory–inhibitory responses (Ex–In). Redrawn
from Roberts D. Signals and Perception: The Fundamentals of Human Sensation, 2002, with
permission of Palgrave Macmillan.

o The thalamus contains three regions where auditory influence is known to

occur, the medial geniculate body, the posterior nucleus and part of the
reticular nucleus of the thalamus.
o The most important for auditory function are the geniculate nuclei.
o Geniculate nuclei which are bilateral rounded regions lying on the surface
of the thalamus.
o These nuclei have three major divisions each receiving a separate, parallel
pathway from the inferior colliculus.
o The ventral division is organized tonotopically into isofrequency layers
within which there are ordered maps of neurons responding to different
auditory cues and is the tonotopic pathway, receiving its input from the
central nucleus of the inferior colliculus.

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o Secondly, the diffuse pathway enters the dorsal division and is not
tonotopically organized, arising from the dorsal cortex of the inferior
colliculus. The cells here respond to many different frequencies, many only
to complex sounds.
o The medial division receives multimodal inputs from external lateral cortex
of the inferior colliculus involving several other sensory systems including
the auditory system. Neurones within this division thus respond to one or
more modality and can be modified by learning.
o The main projection to the primary auditory cortex arises from the ventral
division of the medial geniculate nucleus and terminates in area A1,
corresponding to Brodmann’s area 41 in the human brain, within the lateral
fissure of the temporal lobe.
o The dorsal division of the medial geniculate nucleus projects to the non-
primary auditory areas around A1.
o The medial division projects diffusely to the whole region and to
surrounding cortical fields.
o A1, like the preceding auditory nuclei, is organized into isofrequency layers
arranged tonotopically from low frequency in the rostral end to high
frequency in the caudal end. Most cells within A1 respond to binaural
stimulation. There are two main types of response:
 Neurones that summate excitatory responses from both ears
 Neurones that receive excitatory stimulation from one ear and inhibitory
stimulation from the other.
o Bands of cells displaying excitation–excitation and excitation–inhibition
responses run alternately across the isofrequency layers.
o The main function of these cells, and of primary auditory cortex in general,
appears to be sound localization.
o Complex responses can be found in neurones in the areas surrounding A1.
These include responses to features such as specific delays between
significant parts of a complex sound, and the simultaneous occurrence of
harmonically related frequencies. These types of feature extraction are
likely to be important in the analysis of time-varying acoustical signals such
as human speech.

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Auditory cortex

1. Auditory sensory area(area41,42)

 Located in the upper part of the temporal lobe"mostly hidden
in the floor of the lateral fissure".
 Its anterolateral part receives impulses from the apex of the
cochlea(i.e. low pitched sounds)
 Its posteromedial part recieves impulses from the base of the
cochlea(i.e. high pitched sounds)

2. Auditory psychic area (area 22=association or inter

prretative)
 Lies adjacent o the auditory sensory area on the lateral
surface in the superior temporal gyrus.

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Functions of auditory cortex
a) Function of auditory sensory area:-
 Perception of the pitch,intensity and quality of sounds"without
understanding their meanings"
 Perception of the source of sound: localization of source depend on:

1-Difference in time of arrival of sound to both ears

2- Difference in loudness of sound in both ears.
3- Intact audiory sensory area.

b) Function of auditory psychic area:-

Explains and understands the meaning of various sounds and words
,so alesion affecting this area on the left side(in right handed
individualy leads to auditory aphasia, or word deafness.

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B. DESCENDING PATHWAYS
 There are descending projections from each of the stations of the
ascending auditory pathway, down as far as the cochlear nuclei and from
the superior olivary complex to the cochlea.
 Some of these descending projections may participate in attention level and
anticipation of signals.
 The olivocochlear feedback loop is a major descending projection.
o Following afferent input to the superior olives, fibres completing this loop
originate from neurones in or around the superior olivary complex and
project back along the auditory nerve into the cochlea.
o Those originating adjacent to the contralateral medial superior olives,
which cross the midline and are myelinated, form the majority and
constitute the crossed olivocochlear bundle. They contribute largely to the
efferent projection to the outer hair cells, also known as the medial
efferent system.
o The fibres end in relatively large primarily cholinergic nerve terminals next
to a subsynaptic cistern in the hair cells, and contain large numbers of
vesicles.
o It is believed they function to suppress outer hair cell motility to make the
cells less sensitive, providing protection from very loud sounds.
o A smaller number of unmyelinated efferents originate from the lateral
superior olive ipsilaterally and contribute mainly to the efferent projection
synapsing with the peripheral processes of type I spiral ganglion neurons
beneath the inner hair cells, although a few terminate directly on the hair
cells. This has been called the lateral efferent system and may comprise
two functional subdivisions, capable of inducing either slow increases or
decreases in the magnitude of the response of auditory nerve fibres.
Since these fibres originate in the lateral superior olive, which is involved
in sound localization, they may be useful in maintaining accurate binaural
comparisons during slow changes in interaural sensitivity.

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ANATOMY OF VESTIBULAR ORGAN

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INTRODUCTION
Vestibular receptors for sensing position, motion and gravity evolved early in
nature together with effector systems to maintain equilibrium.
In humans, the inner ear has five different organs which are functionally
related and establish neural connections in the brain with the oculomotor and
postural control systems. The vestibular system plays a lesser role for the
control of human posture and balance, but is of great importance clinically
due to its common and dramatic involvement during pathological conditions
affecting the inner ear.
The vestibular organ consists of a delicate system of membranous ducts
containing sensory epithelia or mechanoreceptors important for the sense of
gravity and balance.
The sensory epithelium is located in the three ampullae of the semicircular
canals and in the maculae of the saccule and utricle.
The organ contains endolymph and is surrounded by perilymphatic fluid.
The membranous system, or ‘labyrinth’ due to its complexity, is connected to
the
cochlea through the thin reunion duct.
In addition, the various portions of the vestibular organ are interconnected by
small canals: the utricular duct and the saccular duct which merge and form
the utriculosaccular duct. This canal runs posteriorly and enters the internal
aperture of the bony vestibular aqueduct. Here, it forms the endolymphatic
duct which widens triangularly within the bone of the posterior surface of the
petrous bone into the endolymphatic sac.
The endolymphatic sac ends blindly within a dural duplicature in the posterior
cranial fossa in close association with the sigmoid sinus and venous drainage
of the brain. The endolymphatic sac consists of both an intraosseous and an
extraosseous portion. The surface of the intraosseous portion of the sac is
irregular and tubular, while the surface of the intradural portion is usually more
smooth and flat.
The endolymphatic duct and sac, which are closely related to the brain, are
thought be involved in the reabsorption of endolymph and regulation of
endolymph pressure.

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VESTIBULAR SENSORY SYSTEM
The vestibular sensory organ is a so-called ‘open’ system that rests on a
balance of function between the left and right ear. There is a continuous influx
of nerve impulses (over one million per second) from each vestibular nerve
to the central nervous system (CNS). This resting discharge is the source of
vestibular tonus. It is therefore not surprising that a viral, fracture or vascular
disturbance (enhancement or impairment) in one of the two organs will result
in an imbalance between the sides and give rise to clinical symptoms not
seldom perceived as violent by the patient. Typical is a rotatory type of vertigo.
Appearing alone, without any concomitant neurological or cardiovascular
signs, these symptoms are generally of a benign nature.

THE VESTIBULAR NERVE

 The human vestibular nerve contains approximately 18,000 afferent nerve
fibres whose size varies between 2 and 15 mm.
 The number of ganglion cells in humans has been estimated to be
approximately 18,000 in young adults, decreasing to around 12,000 at the
age of 80 years.
 Numbers of both vestibular hair cells and nerve cells in Scarpa’s ganglion
are found to be reduced in the ears of older people.
 Changes in the vestibular sensory epithelia appear to precede those seen
in Scarpa’s ganglion.
 The vestibular ganglion or Scarpa’s ganglion consists of bipolar neurons
located in the lateral part of the internal auditory canal. It consists of a
superior and an inferior group of cells associated with the superior and
inferior vestibular nerve.
 The superior branch innervates the cristae of the superior and lateral
canals, the macula of the utricle and the anterosuperior part of the macula
of the saccule.
 The inferior branch supplies the crista of the posterior canal and the main
portion of the macula of the saccule.
 The nerves contain both large and small nerve fibres.
o Large nerve fibres predominantly involved in the innervation of type I
hair cells located in the central region of the maculae (striola) or the
summit of the crista.
o The small fibres are more abundant at the slope and the peripheral
region of the maculae.

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 Like the cell bodies of the human spiral ganglion, the perikarya of the
vestibular ganglion cells are unmyelinated and surrounded by a thin
sheath of Schwann cell. Some cells lie in groups impinging upon each
other, suggesting that electrical activity could be transmitted from one cell
to another.
 In the internal auditory meatus, the vestibular and cochlear nerves merge.
There is usually a small groove between the major division which can be
developed to a greater or lesser extent. During their course to the
brainstem, the facial nerve becomes located further up the brain (VIIth and
VIIIth cranial nerve). A small arterial branch from the anterior inferior
cerebellar artery (AICA) runs between the VIIth and VIII nerve on the
brainstem. It can be seen during vestibular schwannoma surgery and be
used as a landmark.
 The human vestibular nerve contains efferent nerve fibres terminating in
both the cochlear and vestibular sensory organs.
 Inner ear efferents of central origin may arise from the general visceral
efferent cell column of the facial nerve and are parts of the nervus
intermedius having autonomic functions. The origin of cochlear and
vestibular efferents from the facial branchiomeric cell column.
 The human vestibular labyrinth is supplied by approximately 200–300
efferent nerve fibres located ventromedially to the ventral portion of the
lateral vestibular nucleus.
 There is both an ipsi- and contralateral origin of vestibular efferents in the
brainstem.
 In mammals, the auditory and vestibular efferents are completely
separate. Cochlear efferents can be divided into at least two distinct
pathways, descending medially and laterally.
 During development, inner ear efferents appear to be a specific motor
neuron phenotype, but unlike motor neurons have contralateral
projections, innervate sensory targets, and, at least in mammals, also
express noncholinergic neurotransmitters.
 Efferent cochlear and vestibular fibres converge as they enter the
vestibular root in the brainstem.
 Efferent running to the vestibular organs split off and follow the course of
the afferents to the neuroepithelium of the maculae and cristae.

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 Efferents to the labyrinth are small and generally branch heavily before
reaching the peripheral receptors. Efferents typically demonstrate
acetylcholinesterase activity which can be used for demonstrating their
location.
 Interestingly, as in the auditory system, there is a bimodaltype of
innervation where the efferents only make direct contact with the
phylogenetically older type II cells (similar to outer hair cells), while on the
type I cells (similar to inner hair cells) the efferent terminals always contact
the afferent nerve fibre or terminal.
 Thus a ‘higher’ or central control of afferent sensory activity could act both
at a presynaptic level on the afferent fibres, but also as a direct modulation
on the peripheral receptor as can be observed in other sensory systems.
 The effect of the efferent system on the afferent system is rather weak and
inhibitory in nature. It may allow the primary afferent receptors to both
increase and decrease their activity as a response of stimulation.
However, extralabyrinthine stimulation especially those involved in
initiating body or eye movements cause efferent activity preceding
movements, suggesting that modulation may be caused by higher motor
centra.
 The efferent nerve fibres to the cochlea (mostly unmyelinated) leave the
vestibular trunk just beyond the saccular ganglion.
 The vestibulocochlear anastomosis in the adult human was found to
contain both myelinated and unmyelinated axons. The number of
myelinated axons ranged from 223 to 695, with a mean of 360, while the
unmyelinated axons varied from 638 to 1453, with a mean of 1005.

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Vestibular sensory cells, type I and type II

 The sensory epithelia contain two types of sensory cells, the cells are
classified as type I and type II sensory cells.
 THE TYPE I CELLS:
o Are flask-shaped and surrounded by a nerve chalice formed by the
terminal end of the afferent nerve fibre of the vestibular nerve. Many of
the calyces have collateral extensions that end on type II hair cells.
o Type I cells, correspond to the inner hair cells of the organ of Corti.
Some of the sensory cells are low threshold mechanoreceptors, showing
a high degree of adaptation, and are believed to be presynaptic to the
large-diameter myelinated neurons in the vestibular nerve.
o These highly adaptive cells are thus considered to be associated with
neurons showing an irregular discharge pattern.

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 TYPE II HAIR CELLS
o Synapse with collateral extensions of calyces and also directly with the
outer membrane of calyces surrounding type I hair cells.
o Efferent nerve endings also terminate on the afferent nerve calyce and
on type II hair cells.
o Type II cells, which are thought to be phylogenetically older, resemble
outer hair cells of the organ of Corti. They are cylindrical in shape, but
have the same arrangement of stereo- and kinocilia as the type I cells.
 THE UPPER SURFACE OF THE HAIR CELL
o The upper surface of the cell contains a thicker region, called the
‘cuticular plate’, into which the stereocilia extend their rootlets.
o Stereocilia:
 Upper surface of hair cell contains approximately 70 stereocilia and
one kinocilium arranged with the longest stereocilia positioned
adjacent to the kinocilium.
 The stereocilia in the macula are a few microns long, while in the
crista they measure up to over 35 mm.
 The stereocilia contain actin filaments.
 All the cilia within a bundle move together as if joined to one another.
 The membrane surrounding the cilia has a surface coat of negatively
charged molecules.
 BASE OF HAIRS CELL
o Around their base Hairs cell appear quite stiff and pivot. When
subjected to force, they break as if they are brittle.
 There are now data suggesting that the sensory vestibular epithelia may
regenerate in mammals including man.This may explain the return of
symptoms after gentamicin treatment in patients with Meniere’s disease.

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SACCULE AND UTRICLE

The utricle is oblong, irregular and slopes anteriorly upwards at an angle of

approximately 300. It lies superior to the saccule. The macula utriculi, which is
the largest, lies mostly in the horizontal plane located in the dilated superior
portion of the utricle. The mean area in adults and children is 4.30mm2.
the macula utriculi contains approximately 33,000 hair cells. This was thought
to be an overestimation since the total number of vestibular
neurons is around 18,000. The right and left macula lie in the same plane.

The human saccule lies in a spherical recess in the medial wall of the
vestibule, is hook-shaped and lies virtually in a vertical position. The mean
area in adults is 2.4mm2. The saccular macula contains approximately 18,000
hair cells.
The pars interna is slightly but significantly smaller than the pars externa.

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Overlying the neuroepithelium is a calcareous material consisting of a
multitude of small cylindrical and hexagonally shaped bodies with pointed
ends, usually named otoconia or better ‘statoconia’. These otoconia are
anchored and partially embedded in a gelatinous substance forming the
otoconial
membrane. The hair processes of the sensory cells
project into the otoconial membrane and are displaced
mechanically by the otoconial mass relative to the sensory
epithelium.

Each macula may be divided into two areas by a narrow curved line in its
centre termed the ‘striola’. The striola area contains a higher concentration of
type I
cells. The sensory cells change orientation along this line so that cells are
often ‘mirror-shaped’ with opposite polarity as to the location of the kinocilium.
In the
utricle, the kinocilia face the striola. However, in the saccule the kinocilium is
polarized away from the striola.

Each sensory cell has a polarization vector with maximal sensitivity to

mechanical deviation of its sensory hairs in a certain direction. Due to the
polarity and the curvilinear shape of the striola, this would seem to result in a
wide range of angles in all three dimensions.
Thus, sensory cells may be more or less ‘tuned’ to certain geometric vectors
and the concerted action of macula cell activity may offer the CNS a wide
degree of neural information or head ‘spatial images’ for optimal perception
and compensatory correction movements during tilt, translational head
movements and positioning. In this regard neck muscular, joint and ligament
receptors may also play a significant role combined with visual stimuli.

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Otoconial layer

o The otoconial membrane consists of a gelatinous layer, a subgelatinous

space and otoconia.
o The otoconia measure 3–19 mm in length.
o They consist of an organic protein matrix together with inorganic calcium
carbonate crystallized in the form of calcite, which is a stable and
common mineral with a specific gravity of about 2.71. Secretion of organic
material occurs from the apical cytoplasm of adjacent supporting cells and
may form the core matrix on which the inorganic material is seeded.
o The enzyme carbonic anhydrase is located in the epithelium which
catalyzes the chemical reaction H2O+CO2H2CO3.The carbonic acid
combines with calcium to form crystals of CaCO3 which is trapped in the
matrix leading to the formation of otoconia.
o Human otoconia seem to undergo degenerative and chemical alterations
with ageing and disease. The otoconial degeneration which occurs with
age is sometimes related to degenerative changes of the neuroepithelium
similar to presbyacusis.
o The otoconia are believed to undergo turnover, whereby the dark
epithelial cells in the utricle may play an important role. A certain amount
of degradation may also occur in the endolymphatic duct and sac, where
degenerated otoconia may also sometimes be seen.

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STRUCTURE AND FUNCTION OF THE AMPULLA
AND CUPULA

o The three semicircular canals (superior, posterior and lateral) provide a

complete coordinate system.
o The horizontal canal generally lies in a horizontal plane when the head is
oriented in an active position. The macular organ is responsible for this.
o The superior opening of the horizontal and superior canal, and the inferior
opening of the posterior canal widen into an ampulla where the crista
ampullaris forms a perpendicularly running crest in relation to its
longitudinal axis. The crest is covered by sensory cells whose kinocilia are
directed in the same direction.
o In the crista of the lateral semicircular canal, the polarization is towards the
utricle, while it is polarized away from the utricle in the superior and
posterior canal cristae.
o The afferents in the horizontal canal are stimulated when endolymph flows
against the utriculus, while the two vertical canals are stimulated when
endolymph moves against the ampulla.

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o The sensory epithelium on the crista is covered by a gelatinous mass
called the cupula. The cupula in the ampulla of the semicircular canal helps
in transferring endolymph fluid movement stimuli to the hair cells. This
gives rise to kinetic reflexes. It contains proteoglycans arranged in a
filamentous network and it is thought to be secreted by the supporting
cells. Firm cupula attachment to the ampulla wall is a physiological
necessity. The cupula is assumed to adhere firmly to the ampulla wall, with
a diaphragm like displacement in the central section and at the base.
Whether the endolymph of the semicircular canal passes the cupula
through the subcupular space. Displacement of the midpoint of the cupula
was as small as 0.76 mm. In addition, the most responsive afferents had a
threshold sensitivity of 10–3 Pa.
o The structural relationship between perilymph and peripheral neurons may
be essential for understanding the pathophysiology of the Menie`re’s
attack. According to Dohlman, the nerves and not the hair cells are
engaged in the production of those symptoms dependent on potassium
increase. The clinical signs of a similar reaction of the cochlear hair cells in
Menie`re’s cases seem to support the assumption of asimilar cause.
Experimentally the objective symptoms of a Menie`re’s attack may comply
with the experimental findings of an endolymphatic potassium
contamination of the perilymph spaces around the afferent nerve branches.
o A remarkable feature of the vestibular neurons is their high frequency of
resting discharge up to 200 impulses per second with an average of 90 per
second (both in the semicircular canals and in otoconia controlled units).
The units are mostly regular but sometimes irregular, especially in the
canals. The irregular units showing high adaptation are believed to
represent sensory cells innervated by large-diameter axons.

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SUPPORTING AND ‘DARK’ CELLS
SUPPORTING CELLS

o Surround sensory cells.

o Function: May be partly secretory in nature. They may provide support
and insulation for the sensory cells and may also form precursor cells for
sensory hair cells.

‘DARK’ CELLS

o Found in all sensory epithelia of membranous labyrinth except the saccule

which is believed to lack production of endolymph.
o The cells have an irregular surface and resemble the marginal cells of the
stria vascularis.
o In addition, pigmented cells or melanocytes are often associated with the
dark cells, which is similar to the situation in the stria vascularis in the
cochlea.
o These cells are important for the development and maintenance of the
unique chemical composition of the endolymph adjacent to the vestibular
mechanoreceptors thereby playing a role for the proper function of the
electric activity of the sensory cells and initiating conductive neural
responses of afferent nerves.
o Degrading otoconia can often be seen on the surface of the dark cells,
suggesting that these cells are involved in the degradation and resorption
of dislodged otoconia.

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VASCULAR SUPPLY OF THE VESTIBULAR ORGAN

Vascular disturbances of the labyrinth probably play a much more important

role in vestibular disease and balance disorders than generally believed. The
inner ear blood supply derives from Labyrinthine artery:
Labyrinthine artery:
o Branch of the AICA and Vertebrobasilar artery system.
o It is an end artery. This may have relevance since vascular obstruction
may not be compensated by collateral blood supply.
o The labyrinthine artery divides into:
 Superior vestibular artery: Supplies the vestibular nerve, utricle and
parts of the semicircular canals.
 common cochlear artery: Which divides into:
- The cochlear artery
- Vestibulocochlear artery: divides into vestibular branches at the
basal turn of the cochlea and supplies the saccule and the
semicircular canals.
o The vestibular labyrinth is drained both through a labyrinthine vein running
through the meatus and through the vein of the vestibular aqueduct
running parallel to the endolymphatic duct and sac in a separate bony
channel named the accessory canal of the vestibular aqueduct.

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 Physiology of hearing
i. SOUND AND ITS ANALYSIS
Sound is produced by setting off its source (e.g a tuning fork) into avibrational
motion. Such vibrations should be transmitted through a medium (as air or
water) to be perceived and heard by the ear.

In air sound is produced as a result of vibration of the air molecules and is

transmitted in the form of longitudinal waves i.e. as alternate phases of
compression ad decompression (or rarefaction) of the air molecules .
These waves travel in air at a speed of about 344 meters/ second (at a
temperature of 20C in the sea level), and it increases with both temperature
and altitude in fresh water, sound is transmitted at a speed of about 1450
meters/ second.
Sound is the sensation produced when sound waves strike the tympanic
membrane. It is asequence of pressure waves that propagates through
compressible media such as air or water
Hearing is the ability to perceive and interpret the sound waves.
The ear functions as a transducer, because it transforms the pressure energy
of the sound waves into nerve impulses, and the auditory cortex converts
these impulses in the sensation of soun
A sound wave
has two basic properties: intensity, which has the subjective correlate of
loudness; and frequency, which has the subjective correlate of pitch.

104
1- The pressure and intensity of sound waves(Amplitude):

The amplitude of sound wave is the average amount of sound energy

passing through a unit area per unit of time. It determines the intensity or
volume of the sound (i.e. its loudness or audibility). The greater the
amplitude, the louder will be the sound and vice versa. However, the
loudness of a certain sound is not only determined by the wave amplitude
but also by its frequency e.g. a high pitched sound is heard louder than
another sound having the same amplitude but is low-pitched.

105
In practice, sound intensity is measured in bels (B) or decibels (dB) one B= 10
dB (i.e. one dB= 1/10 B).
The intensity of a certain sound in bels equals the logarithm of the ratio of the
intensity of the sound to that of a standard sound i.eg.
int ensity of the given sound
The sound intensity in bels= log
int ensity of the s tan dard sound

The standard sound is a sound having a frequency of 1000 Hz and an

intensity that is just audible in a young adult (thus representingthe hearing
threshold). Accordingly, a sound would have an intensity of zero bels
(because log 1= zero). Therefore, a value of zero bels (or dB) does not
mean absence of sound but a sound having an intensity equal to that of the
standard on the other hand, in case of a whispering sound, the ratio of the
intensity of the whisper to that of the standard sound is 100, therefore The
intensity of the whisper = log 100 (102) = 2 bels = 20Db.
During ordinary conversations, the intensity of the sound is about 50 dB (5
bels) i.e. 105 (=100000) times greater in intensity than the standard sound.
Sound intensities of about 120 dB produce discomfort while those of 140 dB
are painful and those of 160 dB (as those produced by a jet plane) have a
very high energy (they increase the body temperature and may cause
damage to the delicate tissues of the ear).
Higher sound intensities (specially at high frequencies) lead to tissue
destruction e.g. the ultrasonic waves(which have a frequency of 1000000
Hz). These waves are inaudible and are clinically used to destroy localized
pathological masses (e.g. a localized brain tumour) without serious damage
to the intervening tissues.

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DECIBELS
 Is alogarithmic unit that indicates the ratio of intensity as aphysical quantity
relative to aspecified or implied reference level.
 If logarithms to the base 10 are used, the resulting units are called bels,
after Alexander Graham Bell, the inventor of the telephone, since the scale
was first used in telephony.
 The bel turns out to be rather too large to be convenient: therefore, the
numbers are multiplied by 10 to obtain units in terms of a smaller unit, one-
tenth the size, known as the decibel.
 The formula for calculating decibels (dB) is therefore as follows:
 Number of dB= 10 log10 (sound intensity / reference intensity)
 Because it is usual to measure pressures rather than intensities, and
because the intensity varies as the square of the pressure, decibels can be
expressed in terms of pressure ratios, multiplying the logarithm by 20
instead of by 10.
 Finally, a reference pressure has to be chosen. Any appropriate reference
can be used.
 One scale in common use, the decibel sound pressure level (dB SPL)
scale, uses a reference pressure of 2*10-5 N/m2 RMS (20 mPa or 2*10-4
dyn/cm2).
 At standard temperature and pressure, air impedance is such that this
corresponds to a power flow of approximately 10%12 W/m2. In this scale,
therefore:
 Intensity )dB SPL( = 20 log10 (RMS sound pressure/2 * 10-5 N=m2)
 Any other convenient reference pressure may be used. If the reference
pressure is the subject’s own absolute threshold at the frequency in
question, the measure is known as decibels sensation level. Here, the
subject’s threshold is by definition 0 dB sensation level. The International
Standards Organization (ISO) scale uses as a reference the ISO standard
human absolute threshold for the frequency being considered. Scales
constructed in these ways are convenient for describing auditory
performance. For instance, negative values of decibels sound pressure
level (dB SPL; that is pressures below the reference) rarely have to be
considered, since the reference is near the lowest absolute threshold. The
range of numbers that has to be used is small, since 130 dB SPL is the
human being’s pain threshold. Similarly, step sizes less than 1 dB rarely
have to be taken into consideration, since 1 dB is approximately the
minimum intensity step detectable. Moreover, changes in intensity of equal
numbers of decibels correspond to approximately equal steps in loudness.

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2- THE FREQUENCY OF SOUND WAVES
frequency, wave length and velocity

This is the rate at which the air waves oscillate per unit time. It is
expressed in Hertz (Hz) i.e. the number of cycles (waves) per second, and it
determines the pitch of sound (high frequencies produce high- pitched
sounds and vice versa).
The human ear can discriminate frequencies (i.e. pitches) ranging
from 20 to 2000 Hz (best in the range of 1000-3000Hz). Some animals (e.g.
Dogs and bats) can discriminate much higher frequencies.
During an ordinary conversation, the pitch of a male voice is about
120 Hz while that of a female voice is about 250 Hz. A normal average
individual can distinguish as much as 2000 different pitches but trained
musicians can discriminate a much grater number of pithes.

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 3- THE PROPAGATION OF SOUND WAVES (wave form)

Different sound wave forms are perceived as a difference in the quality

(timbre) of the sound.
Simple sound waves produce sounds with pure tones. Sound waves that
have repeating patterns even though the individual waves are complex are
perceived as musical sounds. most musical sounds are made up of a wave
with a primary frequency (which determines its pitch) plus a number of
harmonic vibrations (=overtones) that give the sound its characteristic
timbre.
On the other hand, a periodic non repeating vibrations (i.e. wave forms that
have no regular repeated pattern) cause a sensation of noise
ATTENUATION BY DISTANCE

The way that sound waves progress through a medium depends on:
1- The nature of the medium, on the irregularities and in homogeneities it
contains and on the boundaries of the medium.
o The sound waves spread out evenly in all directions, so that the wave
fronts make a series of expanding spheres centred on the source.
o In the idealized situation, the wave front will decreases with distance,
sound intensity falls. Thus sound gets quieter with distance.
o Obstructions will hinder the passage of the sound waves. Objects may be
present at the side, and often these serve to reflect the sound and prevent
it spreading. Under these circumstances, the intensity will fall less rapidly.

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2- Compression of air :when the air is compressed, at the peak of the pressure
wave, its temperature rises, and some of the energy of the sound wave is
stored as heat. The reverse process takes place, and the energy is passed
back into the wave, when the pressure is reduced in the through of the wave.
However, if heat flows from the warmer to the cooler regions of the wave,
some of the energy in the wave is irredeemably lost. This factor is particularly
important for high frequency waves, where the short wavelength allows flow
of heat between the peaks and the troughs of the wave.
TRANSMISSION BETWEEN DIFFERENT MEDIA

 If a medium light and compressible(air is an example), only small sound

pressures will be needed to give a certain velocity of vibration, and hence
displacement, of the air molecules. The pressures will be inadequate to give
similar velocities of vibration in a denser, less compressible medium
(medium with a higher impedance).
 If, therefore, a sound wave in air meets a medium of higher impedance, the
sound pressures developed on the air side of the boundary will be
inadequate to give the same amplitude of vibration of the medium on the
other side of the boundary. The result is that much of the sound is reflected,
with only a small proportion being transmitted. The pressure at the
boundary stays high, but because the reflected wave travels in the reverse
direction, it produces movements of the molecules in the reverse direction.
Near the boundary, the movements arising from the incident and reflected
waves substantially cancel one another, and the net velocity of the air
molecules will be small.
 This point is important physiologically when considering the transmission of
sound from the air into the cochlea. The impedance of the cochlea is much
higher than that of air. If the sound waves meet the oval window directly,
then, as can be calculated from the measured input impedance of the
cochlea, only approximately 1 percent of the incident energy would enter
the cochlea, with the rest being reflected. However, the middle ear
apparatus, by its action as an impedance transformer, improves this
proportion considerably.

110
Fourier analysis

Def : Analysis of a complex sound into sine waves is known as Fourier analysis.
 There are several reasons why it is useful to analyse waveforms into sine
waves (sinusoids) rather than into any other waveforms.
o A primary reason is mathematical, as any realistic waveform can be
made out of sums of sinusoids.
o A second reason is that sinusoidal sound waves behave in a relatively
simple way in many complex environments, such as a reflecting
environment (e.g. the external auditory meatus), or one that has complex
mechanical properties, such as the middle ear.
o A third reason, which is very important in understanding how the ear and
the brain analyse sounds, is that the cochlea itself seems to perform a
Fourier analysis. Therefore, if a complex sound has been analysed into
sinusoids, it is often possible to understand how the auditory system
itself would analyse it.
PERIODIC AND NON PERIODIC SOUNDS
 Periodic signals means signals that have a waveform shape that repeats.
The time taken for the waveform to repeat is its period (T), and the number
of periods in one second is called the repetition frequency (or the
fundamental frequency) (F=1/T).
 Non periodic signals do not have a repeating waveform shape: they are
either signals that occur once (impulsive signals) or signals that change
randomly (noise signals).
LINEAR SYSTEMS
If a system, e.g. the middle ear, is linear it means that it transmits sounds without
distortion (amplitude distortion, is meant). Linearity means that the presence of
one signal does not change the responsiveness of the system to other signals. It
means that the only frequencies that are produced by the system are the ones
that are put into it.

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ii. MECHANICAL CONDUCTION OF SOUND = ACOUSTIC
TRANSFORMER:

A. THE EXTERNAL EAR

Pinna:-
Collect sound waves from air and reflect them in the external auditory
meatus.
Conchae:-
Concentrate sound at intrance of the auditary canal. This action increase
sound pressure as much as 6 dB" 2 times" " mega phone".
EAC:
Increase sound pressure at tympanic membrane by 15:22 dB at 4000 Hz.

B. THE GAIN IN SOUND PRESSURE AT THE TYMPANIC MEMBRANE

(CATENARY LEVER)
 Gain 2 times of sound pressure (x2)
 The annular bone is immobile so the malleus is the recipient of
magnified energy directing it into the ausicular chain for transmission.
 The main function of the tympanic membrane is to convert sound
waves into mechanical vibrations of the same frequency which are
transmitted to handle of malles, incus & stapes to its force to enable
the vibration.

112
C. THE MIDDLE EAR
The middle ear couples sound energy to the cochlea. As well as
providing physical protection for the cochlea, the middle ear serves as an
acoustic transformer to match the impedance of the air to the much
higher impedance of the cochlear fluids. The middle ear apparatus also
serves to couple sound preferentially to only one window of the cochlea,
thus producing a differential pressure between the windows, required for
the movement of the cochlear fluids.
 The mode of vibration of the middle ear structures

khanna

o Bekesy Suggested that tympanic membrane moves like a stiff plate up to 2

kHz, hingeing around an axis of rotation at one edge. He found that the
inferior edge of the membrane was flaccid and it was here that the
movements were greatest.
o Khanna did not confirm this pattern of movement at any frequency; rather,
there were two maxima of vibration, one on either side of the manubrium (it
buckled in the regions between the manubrium of the malleus )and the
anterior and posterior edges. At frequencies above 6 kHz, the pattern
becomes much more complex.
o The axis of rotation of the ossicles and the axis of suspension by their
ligaments nearly coincide with their centre of rotational inertia. Therefore,
the bones are able to vibrate with very little loss through the suspending
ligaments. The relatively massive head of the malleus and incus aid
determination of the appropriate centre of inertia. However, this factor is
dominant only at mid frequencies. At low frequencies, where the mass
effects are small, the ligaments play an important role in maintaining the
position of the ossicles. Bekesy showed this by cutting the ligaments.
Changing the suspension in this way affected transmission below, but not
above 200 Hz. At high frequencies, above 4 kHz, the axis of rotation
changes in a complex way so that there is no simple axis of rotation.

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 The impedance transformer action of the middle ear

o Impedance or acoustic resistence : Resistance of sound passage through

amedium.
Impedance matching:- The middle ear transfers the incoming vibration
from tympanic membrane from large, low impedance, to the much
smaller, higher impedance, at reaching oval window.
o In matching the impedance of the tympanic membrane to the much
higher impedance of the cochlea, the middle ear uses two principles.
1- Hydraulic lever (ratio between TM& stapes footplate) The area of the
tympanic membrane (60mm2) is larger than that of the stapes
footplate in the cochlea (3.2mm2). The pressure on the stapes
footplate is therefore increased by 60/3.2 = 18.75 times. This is the
most important factor in achieving the impedance transformation.

2- Ossicular lever (ratio between length of malleus and incus) : The

geometrical length of the malleus is approximately 2.1 times that of
the incus,26 and so the lever action multiplies the force 2.1 times.
However, the velocity is decreased 2.1 times. The lever action
therefore increases the impedance ratio (being the pressure/velocity
ratio) 2.12 = 4.4 times.

3- Ratio of specific impedances :The final transformer ratio, calculated

here as a ratio of specific impedances, can be obtained by multiplying
these two factors together. The transformer ratio determined in this
way is assessed as 18.75 * 4.4 = 82.5.

 Influence of the middle ear muscles

 Tensor tympani:-Adjust the degree of stretch of the T.M. according to
the different sound notes.
 Stapedius muscle: Pulls the stapes away from the foramen ovale in
violent movements of ossicles otherwise the membrane may be torn out
and one would be deaf.
 The drum can move inwards more readily than outwards and there fore,
acts as arectifier.

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 Transmission through damaged middle ears

1. In the case of a total removal of the middle ear apparatus, the impedance
transformer action (lead to an increase of some 15 dB in auditory
thresholds in the mid-range of frequencies) .also the differential
application of pressure to the round and oval windows are lost (pressures
delivered to the round and oval windows were nearly equal). (in the normal
circumstances pressure go from round to oval window because of 2 factors
the first is the scala vestibuli was more yielding than the scala tympani,
because its blood vessels could be displaced out of the osseous labyrinth,
the second factor, the small compliance of the annular ligament of the
stapes, in comparison with the much larger compliance of the round
window membrane, will tend to enhance the differential movement. At the
present time, it is not known whether the 40–60 dB loss observed clinically
can be explained by known mechanisms. In the case of these severe
losses, hearing by bone conduction may become significant.

2. If the middle ear apparatus is lost, but the round window is protected in
some way from the incoming sound waves, then a differential pressure can
be set up across the round and oval windows.

3. If the tympanic membrane is intact and connected directly to the oval

window, either by direct contact between the drum head and the stapes or
by means of a prosthesis, the major component of the impedance
transformer, the area ratio, remains, although the lever action of the
ossicles and possibly the buckling action of the tympanic membrane will be
lost. As these two latter factors have a comparatively small influence on
the impedance transformation, the transformation produced by the middle
ear will be only slightly affected, and good hearing is theoretically possible.

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4. A hole in the tympanic membrane will reduce the effective area of the
membrane in contact with the sound wave. Holes will also reduce the
pressure differential across the tympanic membrane and, depending on
their position, reduce the mechanical coupling between the remaining
intact portions of the membrane and the malleus. Holes in the tympanic
membrane led to air flow through the hole, leading primarily to losses
which were largest at low frequencies and which increased as the
perforation size increased. In the human cadavers, holes of 0.5mm across
produced transmission losses only below 600 Hz, while holes 3.3mm
across produced losses below and up to 1.5 kHz. The largest holes,
5.0mm across, gave losses below and up to 2 kHz and also gave a second
band of loss above 2.5 kHz. While there were some losses due to changes
in coupling between the tympanic membrane and the manubrium of the
malleus, these were generally less than 5 dB.
o Small and moderate lesions (10–40 percent of tympanic membrane) had
far more severe effects when placed on the posterior and superior margin
of the membrane than when placed on the anterior and inferior margin,
presumably due to changes in coupling.
o Static pressure changes across the tympanic membrane, whether positive
or negative, will increase the tension in the tympanic membrane and so
increase its stiffness in response to applied sound. The increased stiffness
will attenuate the transmission of sound in the frequency region in which it
is stiffness limited, i.e. below approximately 1 kHz. The lowest stiffness is
recorded when the static pressure differential across the membrane is zero
and, in the normal case, transmission decreases symmetrically for positive
and negative changes in the pressure differential.

116
 Mechanisms of bone conduction
Bone conduction is the normal route for hearing some of the components of one’s own
voice, it is useful with severe conductive loss and is used as a tool of considerable
diagnostic power. The mechanisms of bone conduction have been a matter of
controversy over the years, it was determined by a multiplicity of factors, arising in the
internal ear, the middle ear and the external ear.

a- INTERNAL EAR FACTORS

o Distortional vibration factors: intrinsic detection of distortional vibrations of
the cochlear bone. Differential distortion of the bony structures of the cochlea,
and of the walls of the other labyrinthine spaces closely connected to it,
would produce a movement of the cochlear fluids. This is proved when the
skull was stimulated with a bone vibrator, and microphonics were measured
in a cochlea from which the middle ear apparatus had been entirely removed
and the cochlear aqueduct blocked. If both the oval and round windows were
sealed with rigid cement, bone conduction was substantial, although several
decibels worse than in the normal animal. If, however, only the oval window
was occluded, the microphonic responses of the cochlea were better than
normal by some 10 dB over the frequency range below 1 kHz. The fact that
substantial responses to bone conduction remained with the cochlear outlets
sealed would suggest that differential flow of cochlear fluids between the
windows, or to another opening such as the aqueduct, is not entirely
necessary for the detection of bone vibrations.
Further evidence: direct transmission of vibration from the skull via the brain
and cerebrospinal fluid to the cochlear fluids.
o Differential compliance : leaving the round window open would release
pressure and produce movements in phase with those resulting directly from
the distortional vibration. On the other hand, increasing the mobility of the
oval window would produce a shunt for the pressure changes, allowing them
to bypass the cochlear partition and so reduce the response. This gives an
explanation of the Weber test: an increased stiffness of the ossicular chain
will reduce the shunting and so increase the detection of bone vibration. The
good response with one window sealed was seen, however, only when the
vibrations were introduced into the bone.
o Cochlear third window: If the cochlea was stimulated by air vibrations led
directly into the middle ear, it was necessary to have both windows open to
obtain a response. Release of pressure through the cochlear aqueduct, and
hence also through the blood vessels, would influence the fluid flow resulting
from bone vibration. Sealing the aqueduct reduced the response to vibration.
This supports the notion of a ‘third window’ in the cochlea.
o Inertia of the cochlear fluids: Evidence was also presented for its role in the
production of movements resulting from the Inertia of the cochlear fluids, with
a differential flow between the third window and the oval window.

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b- MIDDLE EAR FACTORS:

o The centre of inertia of the middle ear bones does not coincide exactly with
their points of attachment.
- Transational vibrations of the skull will therefore produce arotational
vibration of the bones, which can be coupled to the internal ear.
- Obviously, this factor will be affected by any pathology in the mobility of
the middle ear apparatus. It will also be affected by any natural
resonances in the middle ear system.
o The middle ear acts as a broadly tuned band pass filter with peak
transmission around 1 kHz at its natural resonant frequency.
- If the degree of resonance of the system is reduced by damping the
stapes, the main losses will be produced around the natural resonant
frequency. This is the explanation given for the appearance of Carhart’s
notch with mild stapes fixation, although the notch usually appears at
rather higher frequencies, namely 2 kHz.

c- EXTERNAL EAR FACTORS

Vibration of the bone is coupled to the walls of the ear canal and so to the air
within it. These vibrations escape externally when the ear canal is open.
When, however, the ear is occluded, the sound pressure changes in the canal
are increased and are transmitted to the internal ear by the normal middle ear
route.
Occlusion therefore increases bone conduction, but only if a significant area
of the canal, or other part of the external ear which can radiate, is included
behind the occlusion. Because the external radiation of sound is normally best
for low frequencies, the change with occlusion is greatest for those
frequencies.

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D. TRANSDUCTION FUNCTION OF THE COCHLEA
 Cochlea: Is aspiral coiled tube containing the membranous cochlea, about
2.5 turns around acentral bony axis called "modulus" which contains the
fibers of cochlear nerve which leaves its base to enter the internal auditory
meatus
 Abony shelf called osseus spiral lamina is attached to the modulus in the
middle of lumen of long canal
 The spiral lamina contains acanal continous with the cavity of the modulus.
This canal contains the spiral ganglion.
 Spiral ganglion formed of bipolar nerve cells the dendrites of the cells
extend to the organ of corti and their axon outer the modulous to form the
fibers of the cochlear nerve.
 Amembrane: basilar membrane attaches the bonny shelf "spiral lamina"to
the external wall of the canal.
 Bonny cochlea diveded into 3 compartments:
1- Scala media= cochlear duct
2- Scala vestibuli… above
3- Scala tympani…. Below
 Scala media =cochlear duct.
 Separaed from scala vestibuli by resiner membrane.
 Separated from scala tympani by basilar membrane and osseus spiral
lamina.
 The 2 scala contain perilymph and are continous with each other at the
apex of cochlea "helicotrema"
 PerilymphCSF
 Endolymphby stria vascularies.
 Basilar membrane:
- formed of fibers; their number is 24.000 in the human cochlea. The
fibers increase gradullay in length from the base to the apex….length
at base 65:160 micron….legnth at apex 350:500 micron.
- the base of basilar membrane vibrates in response to high pitch, and
its apex responds to low pitch
- organ of corti lies on it
- thick at prephery= zona pictinata…. Thinner inner part = zona
arcuata.

119
 Organ of corti:
1- Supporting cells: pillar,dieter,hensens cells
2- Hair cells :
Short cylindrical cells which don't rest on the basilar membrane but are
supported by supporting cells.
Arranged in 2 rows of cells
Single inner row consising of one cell
Outer rows consisting of 3 cells
The tips of the hairs lie in contact with astrucurless membrane.
From the base of hair cells nerve fibers pass towards the modulus to form
the cochlear division of 8th nerve.

3- Tectorial membrane: the membrane is formed of gelatinous connective

tissue acting as aroof on the hair cells.

Perilymph Endolymph=the only extracelluar

fluid in the body.
Origin Perilymphatic capillaries ..may
Secreted by stria vascularis
be from C.S.F.
Na+ 150 1
K+ 3 150
cl- 125 130
Found in Scala tympani,scala vestibuli
Scala media

N.B. Endolymph is sole source of oxygen supply for the organ of corti
which it self has no blood supply.

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 Transduction function of cochlea:
 Transduction: convertion of mechanical energy of movement of sound to
electrical energyelectrical event in cochlear nerve.
 Stapes passed into oval windowpressure on perilymph of scala
vestibulitransferred to scala mediadownword movement of basilar
membranepressure on scala tympanitransmitted to round window
 Stapes move outupword movement of basilar membrane.
 Up and down movement"elastic tension"wave.
Each wave is weak at onset but becomes stronger as it reaches its natural
resonant frequency

 Transduction by hair cells:-

Auditory nerve ending stimulated both electrically and chemically.it
transduce sound by following steps:
1-basilar membrane and organ of corti move up and
downchearing action between tectorial membrane and
reicular laminabending of sterocilia sideway.
2-bending of hair bundelsopen channelsk+ enter hair
cellsdepolarization.
3- depolarization spreads to lower part of cellopen
Ca++channels.
4-Ca+ causes transmitter vesicles to fuse with the basal part of
cell membranerelease transmitter substance
5- Transmitter substance"aminoacid glutamate"diffuse across
synaptic cleftintiate action potential in auditory nerve
fibers.

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 ELECTRIC POTENTIAL OF COCHLEA
Several electric potential have been demonstrated:

1-Cochlear microphonic potential:

o It is genarated by deformation of the process of the hair cells and it is

linearly proportionate to the magnitude of the displacement of the
basement membrane.
o It was known that Cochlear microphonic potential was the same action
potential, but recently it has been proved that it is not because it can be
recorded from arecently killed animal in which neural activity has ceaed

2- Summating potential:

o Arising from inner hair cells with small contribution of outer hair cells.
o The negative and positive Summating potentials have been recorded in
aresponse to acoustic stimulation.
o The positive has been thought to originate from the outer hair cells,
whereas the negative from the inner hair cells.

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3-Endolymphatic potential:

o It is apositive electric potential recorded from the endolymph in the

cochlear duct.
o At rest , it is +80 mV potential in the scala media with respect to the tissues
of the head.
o It is dependent upon ionic composition, and also upon oxygen supply
because it decreases in asphexia.

4-Cortilymphatic potential:

o It is anegative potential(-20 to -80 mV) recorded in organ of corti with

respect to perilymph.

5-Action potential from the auditory lymph:

o At threshold intenisty, the nerve endings give response only to anarrow

range of frequencies. With the increase of intensities, the nerve endings
respond to an increasingly wide range of sound frequencies which lie
below their maximal response.
o The frequency of the action potential in single auditory nerve fiber is
proportionate to the loudness of the sound stimuli.
o Stimulusneural discharge in the auditory nerve amplitude increases
while latency decreases with intensity over 40:50 dB range.

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 THEORIES OF PITCH ANALYSIS(THEORIES OF HEARING)

1- Theories of cochlear analysis

At present,two types of theories try to solve the proplem of the site and
mechanism by which different sound pitehes reaching the ear are
differentiated (anaiysed).
Some theories assume that pitch anaiysis occurs in the cochiea,e.g.
Heimholt 's theory. Others suggest that analysis oc-curs in the auditory
cortex,e.g.Rutherford's theory. Theories of cocliear analysis are
generally accepted at present.

A- Helmholtz's"resonance "theory:

This theory is based on the phenomenon of "resonance"which is explaned

as follows. If avibrating tuning fork (say of 200 C.P.S.) is brought near the
strings of apiano,certain strings are immediately set into vibration.
The fibers of the basilar membrane are similarly stretched between the
spiral lamina and ligament. The fibers near the apex of
cochlea(helicotrema) are longer and less stretched than those at the base.
All fibers act as resonators, each group of which can be set into vibration
only by aparticular pitch. So, the organs of corti lying on this group of fibers
alone are stimulated while all other organs remain silent.
Accordingly,the longest fibers near the apex of cochlea respond by
vibration to sounds of the lowest pitch(30 to 40 C.P.S.).
The shortest basal fibers respond to sounds with the highest pitch(30 000
C.P.S.).The fibers between the tip and the base of the cochlea respond to
intermediate pitches (between 30-40 and 30 000 C.P.S.).
Stimulated nerve fibers coming from these organs end in aparticular spot in
the auditory cortex./ it is evident, therefore,that Helmholt's theory cosiders
that the analysis of different sound pitches occurs in the cochla and not in
the brain.

124
EVIDENT IN FAVOUR OF HELMHOLT'S THEORY:
o The direct microscopic observation of the movements of the basilar
membrane.it shows that the basal segment of the membrane maximally
moves in response to high pitch sounds while the apical segment shows
the opposite(Bekesy).
o Workmen exposed to high pitch sounds, e.g. shipbuilders and poiler
makers how deafness to high sounds. They can, However, hear normally
low and moderate pitch sounds. Postmortemexamination of those subjects
reveals degeneration in the basal fibers of the basilar membrane as aresult
of prolonged exposure to this particular higher note.
o Damage in the basal fibers is demonestrated in guinea pigs exposed to
high-pitch sounds, and in apical fibers when exposed to low-pitch sounds.
o Electrical potential changes are maximally detected at the apical and when
low- pitch sounds are applied to the ear such changes are detected at the
basal end on applying high notes.
o damage of aparticular segment of the basilar membrane results in
deafness to sounds of aparticular pitch.

EVIDENT AGAINST HELMHOLT'S THEORY:

This is modified by place theory or Helmholt's place theory
o The fibers of the basilar membrane are embedded in ahomogenous
gelatinous material which cannot allow separate fibers to vibrate Fibers are
not stretched because fibers don't shrink on cutting and so they vibrate
according to their load or weight at acertain place or locus this was
overcome by Helmholt'z "Place theory".
o Very important: The fibers corresponding to aparticular tone vibrates at
amaximum rate and so its hair cells are deflected on crushed against the
tectorial membrane.
o The vibration of the fibers is not determined by heir legnth only but also by
their weight and stretch.
o It is expected that the vibration will continue for some time after stoppage of
the sound. This doed not occur.This objection is overcome because the
fluid surrounding the basilar membrane is apowerful damper of sound
vibrations.
o The single fiber of the auditory nerve (like other sensory nerve) is unable to
transmit more than 1000 impulses per second because its refractory period
is about one millisec. According to some authors, However, his is possible
because at high frequencies the nerve fibers responds to the
second,third,fourth or even tenth sound wave. Therefore, considering all
nerve fibers together, the rate of conduction will be very high, but each
nerve fiber conducts at arate below 1000/sec.

125
B- PLACE THEORY
The most widely accepted theory of pitch discrimination.
This theory is baed on :
o Mechanism of basilar membrane:
The basilar membrane and the organ of corti respond o different
frequencies of sound waves. Aparticular region of basilar membrane
resonates to aparticular frequency of sound waves, more easily than
other frequencies.
o Electric potentials of cochlea
The electrical poentials recorded from the auditory nerve reproduced the
frequency and amplitude of the sound waves reaching the ears.
According to this theory the hair cells will respond more easily to one
frequecy than another, those at the apex responding better to low tones,
while those at the base of the cochlea responding better to high tones.

2- THEORIES OF CORTICAL ANALYSIS:

A- Rutherford's telephone theory
This theory suggests that the cochlea is like atelephone and the basilar
membrane resembles the diaphragm of the receiver. Accordingly, the
membrane vibrates as awhole and not in segments. The analysis of
these sounds occurs in the hearing centers in the brain but not in the
cochlea because its basilar membrane vibrates as awhole by all
audible frequencies.
OBJECTIONS TO RUTHERFORD'S THEORY:
o The theory cann't explain partial deafness to particular notes while
other notes are normal as in boilar makers disease.
o It suggests that every nerve cell in the auditory cortex can analyse
(discriminate) all audible sounds.However, it was shown that this is
not true because every cell in these centers can perceive only
aparticular sound pitch.
o It states that an acoustic fiber can transmit 20,000 impulses/sec and
this is impossible.
B- WEVER'S RESONANCE-VOLLEY THEORY
This theory proposes that the whole cochlea is responding to the low
frequencies, but the cochlear analysis occurs for the high frequencies.

126
 PHYSIOLOGY OF EQUILIBRIUM

ROLE OF THE VESTIBULAR SYSTEM:

i. INTRODUCTION
 The main function of the mammalian vestibular system is to generate
information for the central nervous system with a four-fold purpose:
1- provide general orientation of the body with respect to gravity;
2- enable balanced locomotion and body position;
3- readjust autonomic functions after body reorientation; and
4- ensure gaze stabilization(important).

 Several input systems provide information to the brain, such as the

visual, vestibular and proprioceptive systems, as well as other systems
such as hearing are used, albeit to a lesser extent.
 Central processing of all this information in the brain leads to specific
outcomes:
1- the vestibuloocular reflex (VOR) to ensure gaze stabilization;
2- the vestibulospinal reflex (VSR) and the vestibulocollic reflex (VCR)
to ensure maintenance of an upright position of the body and trunk
and head stabilization in space;
3- orientation and, in higher species, navigation and the perception of
self-position with respect to the surroundings and gravity, mediated
by the vestibular cortex;
4- autonomic function adjustments after alterations of body orientation.

N.B.
This list of outcomes is certainly not restricted to these four tasks
since, for example, the vestibular system has also recently been
shown to influence the circadian rhythm, and there are also
connections with cognitive function.

127
  Although somatosensory, visual and vestibular inputs are constantly
processed by the brain, this information is heavily weighted by other
factors, such as learning, memory, drugs, ageing, as well as
environmental conditions.
 Additionally, the weight of the different inputs is constantly adjusted
depending on the circumstances. It is obvious that, for example, walking
in darkness, balance can not rely so heavily on vision and so reliance on
the somatosensory and vestibular senses is increased.
 For optimal functioning in daily life, all systems are needed.
 To test gaze stabilization and the vestibuloocular reflex, shake your
index finger at arm length left and right, with increasing speed. As soon
as you shake it at a frequency above 1 Hz, the eyes are not capable of
following the movement and you see a blurred image of your finger.
Next, shake your head while you fixate your finger, held stable at arm
length. You will notice that the image remains sharp even at higher
accelerations. Up to frequencies of 5–6 Hz of head movements, many
people are able to maintain a clear image of the target. This is the result
of effective function of the VOR.

ii. MOTION DECOMPOSITION AND ORIENTATION IN THE HEAD

 Every motion in space can be broken down into three rotational degrees
of freedom (yaw, pitch and roll) and three translational degrees of
freedom (left–right, up–down, for–aft).
 No event in one degree of freedom can be described by the others,
hence every movement is uniquely and appropriately described by a
combination of all six degrees of freedom.
 The anatomical design of the motion sensors in the peripheral vestibular
system in the inner ear reflects these six degrees of freedom.
 The semicircular canals measure predominantly rotations, whereas the
macules of the utricle and saccule detect mainly translations.

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129
 The orientation of the vestibular system in the head is depicted in this Figure:

Top view of labyrinths as oriented in the human head. The labyrinths are of a gerbil.

 In a first approximation, one can model the left and right canals as parallel
systems, i.e. the left posterior canal is parallel with the right anterior canal
which both lies in a plane denoted as the right anterior–left posterior (RALP)
plane. With the right posterior canal, the left anterior constitutes the left
anterior–right posterior (LARP) plane.
 Both horizontal canals are also parallel with each other in the lateral plane.
The horizontal canal makes an angle of approximately 300 with respect to
the horizontal axis of the upright head.
 The angles of the vertical canal are approximately 450 with the saggital
plane of the head.
 When studied in more detail, there exists however a great variability of
orientation within and between subjects. Additionally, the canals themselves
are often quite curved.

Side view of a right human labyrinth (labyrinth filled with resin and bone cut away). The nose is on the right side. The lateral
canal makes, in general, an angle of 30 degrees with the horizontal axis of the head.

 The distance from vestibule to midline is quite symmetric and very constant
between subjects, i.e. the deviation is less than 5 percent.
 The orientation in the head is a constant factor within every subject, so each
central nervous system (CNS) is totally adapted to the individual geometry
of the peripheral vestibular system for movements which are in the normal
range.
 Anatomical differences however, between left and right, predominantly at
the level of the maculae of the saccule and utricle are hypothesized as a
factor contributing to motion sickness.

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iii. Movement detection
 During daily life, body and head are continuously moving (during each
heartbeat, the head and body are shaking a little) and these movements
are always related to forces, i.e. accelerations. These movements are
sensed in the vestibular organ by a rigid coupling of sensory epithelium to
the bony structure. Thus, a fluid-filled system is attached to the skull and
the inertial forces (upon head acceleration, as well as gravity) drive the
fluid.
 Motion of the fluid lags behind any motion of the head and this relative
displacement is the trigger for movement detection.
 To limit the movement of the hair cells triggered by accelerations that drive
head movements, the canal system is designed such that the deflection of
the hair cells is proportional to the head velocity.

a- SACCULE AND UTRICLE:

Two sacs inside the bony vestibule. They contain a sensory organ called the
macula or otolith organ.
The macula is fored of ridge of columnar epithelium with hair cells having cilia
(hairs) which penetrate into a gelationous material, embedded on its top a
numerous small paticles of calcium carbonate called otoliths or otoconia.
With the head in the erect position, the macula of the utericle is in the
horizontal plane while that of the saccule is vertical.

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 Mechanism of action of macula:
Bending the cilia of hair cells to one side causes stimulation of
the hair cells and increases their discharge, while bending the
cilia to the opposite side causes their inhibition.
As the position of the head in space changes, the weight of the
otoconia bends the cilia and stimulates the hair cells.
In each macula there are thousands of hair cells whih are
oriented in different directions, so that some of them are
stimulated when the head bends forwards, some when it bends
backwards and others when it beds to one side.
Impulses from the sensory hair cells of the macula of the saccule
and utricle are carried along afferent fibres in the vestibular
division of the 8th cranial nerve.

Functions of the Otolith Organs (Maculae):

1. Orientation of the Head in space:

Impulses from both the right and left utericles are in balance
with the head in the normal ercect position. Tilting the head to
one side  displacement of the calcium carbonate particles
(otoconia) by the force of gravity  bending the cilia of hair
cells  change in the rate of discharge of impulses of hair cells.
Therefore, when the head is tilted to the right side, the rate of
discharge of impulses from the right utricle is increased while
that from the left utricle is decreased.

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 2. Receptors for Linear Acceleration:

The otoconia are displaced during linear acceleration in an opposite direction

leading to bending of the cilia.
When the body is suddenly accelearated, e.g. forwad acceleration, the
otoconia which have greater inertia than the surrounding fluids displaced
backwards  bending the cilia  stimulation of hair cells. The subject will
have a feeling of falling backwards which automatically causes him to lean his
body forwards.
3- Receptors for different posture reflex.
4- Receptors for Labyrinthine Righting Reflexes correcting the
Position of the Head.
5- Receptors for orientation during swimming under Deep water:
Swimming should be avoided in vestibular disorders since the visual and
proprioceptive impulses are absent in this condition.

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b- SEMICIRCULAR CANALS:
The semicircular canals (S.C.Cs) are 3 on each side and lie in 3 planes in
space at right angles to each other. They are the horizontal (external),
anterior (vertieal) superior) and posterior vertieal the horizontal canals are
sloping backwards by 30o be low the horizontal line.
Each S.C.C. has a dilated end called the ampulla which contains a
sensory organ called the crista ampullaris. The 3 S.C.Cs open into the
corresponding utricle by 5 openings only because the anterior and
posterior canals have a common opening at their non-ampullary ends.

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Mode of Action of Semicricular Canals:
The crista consists of supporting cells and sensory hair cells.
Their hair fillaments are embedded in a gelatinous material called the cupula.
 During rest; there is equal discharge of impulses from the S.C.C.s of both
sides.
 S.C.Cs maintain posture during angular acceleration or rotation in space.
They are stimulated at the beginning of rotation, end of rotation and by
increasing (acceleration) or decreasing (deceleration) the velocity of
rotation. S.C.Cs are not stimulated by a constant rate of rotation, e.g.
rotation of earth.
 Deflection of the stereocilia in the cupula towards the kinocilium (ampullo-
or utriculopetal) results in hair cell depolarization and consequently the
activity of the primary afferent neurons increases. Deflection of the
stereocilia away from the kinocilium (ampullo- or utriculofugal) results in
hair cell hyperpolarization and decreased primary afferent neuron activity.
The epposite occurs in the anterior and posterior vertical canals. Bending
of the cupula is produced by movement of the endolymph impulses from
the hair cells of the crista ae transmitted along afferent fibres in the
vestibular division of the 8th cranial nerve.
 The left and right semicircular canals are oriented in the head such that
any movement always induces an antagonistic response in both canals.
 The activity of the lateral SCC primary afferent neurons is modulated by
horizontal head rotation. At the leading ear (the ear towards which the
movement is directed), the firing rate increases and at the following ear,
the firing rate decreases. This is called the push–pull principle of the VOR.
 Each semicircular canal is maximally stimulated by movements taking
place in its plane.

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 Angular Acceleration :

If the head is rotated in the horizontal plane from left to

right the following will occur:
a) At the beginning of Rotation: the endolymph will lag behind the moving
canals due to its inertia (resistance to movement). This leads to flow of
the endolymph from right to left, and results in bending of the cupula of
the right canal towards the utricle, i.e. stimulated. At the same time the
cupula of the left canal is bent away from the utricle, i.e. inhibited the
unequal discharge of impulses from the cristae ampullaris of the two
horiznal S.C.Cs gives the person the sensation of rotation to right.
b) After Half Minute: as the rotation is controlled, the endolymph takes up
the same velocity of the bony and membranous S.C.Cs. both cristae
return to their normal central position. This leads to equal discharge
from both cristae. The person does not feel that he is rotated.
c) At the stoppage of Rotation: the movement of the bony and
membranous S.C.Cs stops but the endolymph continues to flow from left
to right in the same direction of rotation due to its momentum force
keeping it moving. This results in bending of the cupula of the left canal
towards the utericle, i.e. stimulated, and that of the right away from the
utericle, i.e. inhibited. The person feels a false sensation of rotation to
the left. This is called vertigo.
d) After Half Minute : the movement of endolymph stops  cupulae return
to the resting position due to their elasticity  sense of rotation stops.

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Methods of stimulation of the semicircular canals
1. Rotational Method:
 The subject is rotated with high speed on a otatory chair, with the head
fixed at certain position to stimulate particular S.C.C. placed in the
horizontal plane.
a. Horizontal canal: If the head is bent forwards by 30o.
b. Anterior vertical canal: If the head is dorsiflexed by 60o.Or
ventroflexed by 120o, i.e. by rotation in the frontal plane.
c. Posterior vertical canal: if the head is bent 90o over one shoulder,
i.e. by rotation in the sagittal plane.
 Typical stimulation patterns exist of
o Sinusoidal rotations, Characteristic for sinusoidal rotations is that the
angular velocity and acceleration are varied with sine and cosine
functions, respectively.
o The Velocity step test (ramp test) is where a sudden acceleration
rotates the subject at a constant velocity and then after a plateau of
constant speed, the chair is suddenly stopped. A variation to this is the
test where gradually the speed is increased by applying a constant yet
slow acceleration (e.g. 5 degrees/second2) for 40 seconds to reach a
speed of 200 degrees/second, followed by a plateau during which the
nystagmus fades away, and then suddenly the chair is stopped.

2. Caloric Methods:
 Stimulates only one canal which must be vertical. The head is bent
backwards 60o to bring the horizontal canals into a vertical plane.
 Douching the ear with cold water at 20oC or warm water at 44oC
causes a greater change in the temperature of the endolymph in the
part of the canal lying near the external auditory meatus than in the
part more deeply situated.
 Convection currents occur and the endolymph moves towards the
ampulla, i.e. stimulation of S.C.C.

3. Electrical Methods :
Galvanic current directly stimulates the cristae of the S.C.Cs of one side.
The stimulating electrode is placed on the mastoid process.

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Effects of stimulation of the semicircular canals

1) NYSTAGMUS:

 The eye response to a head rotation consists of a combination of a slow

phase or drift until the eye reaches the edge of the outer canthus, and a
fast phase to reset the eye in its initial position. This saw-tooth pattern is
called nystagmus.
 The direction of the nystagmus is defined by the fast reset phase, since
this is the most easy for the clinician to identify.
 The slow phase, however, represents the actual vestibular output and is
quantified.
 An upward excursion, by convention on electronystagmography or video
nystagmography, represents eye deviation to the right. If the slope of the
sawtooth is upward to the right, i.e. a positive slope, this corresponds to a
slow drift of the eye to the right, followed by a quick left ward reset
saccade, as represented by a steep downward trace. This is defined as a
left nystagmus. It is quantified by measurement of the slope of the upward
trace, which indicates the speed of the eye movement (degrees/second).
 The semicircular canals and the otoliths provide the inputs for the
vestibuloocular reflex.
 Horizontal VOR compensates for both horizontal rotation(due to the canal
system) and horizontal translation(due to the utricular system). It is
therefore more convenient to use the angular VOR (aVOR) and linear VOR
(lVOR).
 A third type of VOR, the ocular counter-rolling, is provided by the otoliths
as a response to, for example, tilting of the head with respect to gravity or
a stimulus that results in a shift of the gravitoinertial acceleration (the sum
of an acceleration in any direction and gravity).

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Velocity storage mechanism(VSM)
 It is a neurophysiological process, taking place mainly in the nucleus
prepositus hypoglossus and the adjacent medial vestibular nucleus. The
VSM serves to maintain the VOR at low frequencies (below about 0.02
Hz).
 Rotation at relatively high but constant speeds is not sensed by the SCC
because of this mechanism because cupula repositions itself into its center
in approximately five to seven seconds, which implies that after 12
seconds the cupula is almost entirely restored to its central position. The
VSM uses principally the peripheral labyrinthine signal and by a process of
integration, in the mathematical sense, increases the frequency response
of the VOR, by prolonging the time constant (originally seven seconds) of
the decay of the vestibular nystagmus to approximately 20 seconds. This
circuitry is therefore a mechanism that stores neural activity related to head
and eye velocity and discharges it over its own time course.
 The velocity storage mechanism provide transition between VOR& OKN.
VOR reflects high-frequency behaviour, OKN reflects low frequency
characteristics
 ( OKN provides visual information of the surrounding rotating world during
sustained rotations . when sustained rotations reached the optokinetic
reflex comes into play) .

VOR generation
 The right medial vestibular nucleus in the brainstem receives an increased
input from the right lateral SCC primary neurons (no crossing), which
excites the activity of type I position vestibular pause (PVP) secondary
vestibular neurons.
 These excitatory neurons drive the left ward compensatory eye
movements of the VOR, to ensure gaze stabilization. However,
commissural disinhibition from the left lateral SCC primary neurons also
contributes to the excitation of the PVP neurons. Therefore, both the
excitation of the right SCC and the disinhibition of the left SCC are needed
for an optimal VOR.

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 In first approximation, this VOR process is a three arc neuron reflex (first
order vestibular neurons, second order vestibular neurons and oculomotor
plant neurons).
 The simplified principle of VOR generation (yaw plane rotation and
horizontal SCC) is as follows:

1- During head rest, hair cells in both SCC have a resting discharge rate of
90 spikes per second.
2- Head rotation is to the right.
3- Endolymph fluid lags behind, i.e. moves relative to the left within each
SCC due to inertia.
4- The cupula bends to the left in each canal.
5- In the (leading) right SCC, the stereocilia bend towards the kinocilium.
6- In the (following) left SCC, the stereocilia bend away from the kinocilium.
7- The discharge rate increases in the leading right ear (e.g. from 90 to 300
spikes per second).
8- The discharge rate decreases in the following left ear (e.g. from 90 to 20
spikes per second).
9- The vestibular nuclei interpret the difference in discharge rates between
left and right SCC as movement to the right, and therefore trigger the
oculomotor nuclei to drive the eyes to the left to maintain gaze
stabilization.

 Similar to the horizontal canal, a push–pull principle also governs the

vertical canal excitatory and inhibitory functions. For example, the left
anterior canal is excited, while the right posterior canal is inhibited for the
same movement.
 Also, the vertical canals are direction sensitive, but at this stage
ampullopetal movements result in a decreased firing rate. Thus, the
ampullar deflection in the vertical canals, corresponding to excitation, is in
the opposite direction to the horizontal SCC.
 For the horizontal canal, excitation is elicited upon ampullopetal endolymph
movement (towards the vestibulum–utricle), whereas for both the vertical
posterior and anterior SCC ampullopetal flow is inhibitory.
 The horizontal canal is involved in pitch movements to only a small extent.

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 The inclination of the vertical canals is more than 901 to the horizontal, so
that horizontal movements are always detected by the vertical canals.
 However, given the antagonistic response of the anterior and posterior
SCC on each side, horizontal movements produce primarily horizontal
movements.
 There are three types of rotationally induced eye movements: horizontal,
vertical and torsional. Each of the six pairs of eye muscles must be
controlled to produce the desired response.
 The vertical semicircular canals and the saccule are responsible for
controlling vertical eye movements, whereas the horizontal canals and the
utricle control horizontal eye movements. Torsional eye movements are
controlled by the vertical semicircular canals and the utricle.
 Stimulation of a single canal results in eye movements that lie in the plane
of the canals.
 To understand the generation of the different eye movements, it is
necessary to analyse the stimulation of individual canals and their effect on
the eye muscles. The following figures represent the effects of single or
multiple canal stimulation on the eye muscles and consequently the type of
compensatory eye movement is generated. Figures 1 and 2 illustrate the
stimulation of the anterior and posterior canals. Every stimulation in
whichever direction always evokes the push–pull principle. When the head
is tilted sidewards (Figure 3), a torsional nystagmus is generated, as both
posterior and anterior canals are stimulated. When the head is pitched
forward (Figure 4), both left and right anterior canals are stimulated,
whereas both posterior canals are inhibited. This produces an upward eye
movement. Each anterior canal also produces an ocular counter-rolling
movement, but as both are in different directions, the counter-rolling is
cancelled out. The same applies when the head is tilted backwards (Figure
5), but now both posterior canals are stimulated producing a downward
compensatory eye movement.

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 Stimulation of the right anterior semicircular canal, while the left posterior canal is inhibited, occurs during a head movement
in the right anterior – left posterior (RALP) semicircular canal (SCC) plane, i.e. turn the head 451 to the right and move the
head downward in that plane. This generates a counter-clockwise movement of the eyes (seen from the patient’s view),
together with an elevation. The right superior rectus (SR) muscle and the left inferior oblique (IO) muscle are contracted. The
right inferior rectus (IR) muscle and the left superior oblique (SO) muscle are inhibited. 1, excitation; %, inhibition; LR, lateral
rectus muscle; MR, medial rectus muscle. The black arrows show the direction of movement of the head.

Stimulation of the right posterior semicircular canal, while the left anterior canal is inhibited, occurs during a head
movement in the left anterior – right posterior (LARP) semicircular canal (SCC) plane, i.e. turn the head 451 to the
left and move the head upward in that plane. This generates a counterclockwise movement of the eyes (seen from
the patient’s view), together with a depression of the
eye. The right superior oblique (SO) muscle and left inferior rectus (IR) muscle are contracting,
while the right inferior oblique (IO) muscle and the left superior rectus (SR) muscle are relaxing.
, excitation; %, inhibition LR, lateral rectus muscle; MR, medial rectus muscle. The black arrows show the direction of
movement of the head.

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 Stimulation of the right posterior and right anterior semicircular canals (SCC), while the left anterior and left posterior
canals are inhibited, occurs during a rightward sideways tilting of the head towards the shoulder. This generates a
counter-clockwise movement of the eyes (seen from the patient’s view). The right superior rectus (SR) muscle and
superior oblique (SO) muscle, as well as the left inferior oblique (IO) muscle and inferior rectus (IR) muscle, are
contracting while the right inferior rectus (IR) muscle and inferior oblique (IO) muscle and the left superior rectus (SR)
muscle and superior oblique (SO) muscle are relaxing. The elevation and depression cancel out and a pure counter-
clockwise torsional nystagmus is generated. This suggests why during a unilateral lesion, such as a vestibular neuritis that
affects the whole peripheral vestibular system, there is never a vertical nystagmus, but only a torsional and a horizontal
(not depicted here). Spontaneous vertical nystagmus is therefore almost always due to a central neurological pathology.
The black arrows show the direction of movement of the head. LR, lateral rectus muscle; MR, medial rectus muscle.

Stimulation of the right anterior and left anterior semicircular canals (SCC), while the right posterior and left posterior canals are
inhibited, occurs during a forward bending of the head. This generates an elevation of the eyes or a downbeat nystagmus, depending
on the amplitude and speed of bending the head. The right superior rectus (SR) muscle and inferior oblique (IO) muscle, as well as the
left inferior oblique (IO) and superior rectus (SR), are contracting, while the right inferior rectus (IR) muscle and superior oblique (SO)
muscle and the left superior oblique (SO) muscle and inferior rectus (IR) muscle are relaxing. Given the activity of these muscles, any
torsional components are cancelled out.
The black arrows show the direction of movement of the head. LR, lateral rectus muscle; MR, medial rectus muscle.

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 Stimulation of the right posterior and left posterior semicircular canals (SCC), while the right anterior and left anterior canals are
inhibited occurs during a backward bending of the head. This generates a depression of the eyes or an upbeat nystagmus,
depending on the amplitude and speed of bending the head backwards. The right inferior rectus (IR) muscle and superior
oblique (SO) muscle, as well as the left superior oblique (SO) muscle and inferior rectus (IR) muscle, are contracting, while the
right superior rectus (SR) muscle and inferior oblique (IO) muscle and the left superior rectus (SR) muscle and inferior oblique
(IO) muscle are relaxing. Given the activity of these muscles, any torsional components are cancelled out and a pure vertical
nystagmus remains. The black arrows show the direction of movement of the head. LR, lateral rectus muscle; MR, medial rectus
muscle.

Nystagmus induced by acute unilateral deafferentiation acute

peripheral lesions
 The push–pull principle is very convenient to explain the origin of the
nystagmus observed during acute peripheral lesions. In the case of an
acute unilateral vestibular differentiation (uVD) on, for example, the right
side, all three right canals and the otoliths cease spontaneous activity (from
90 spikes per second to zero), while the spontaneous activity of the left
SCC remains at 90 spikes per second. Although the head is not moving, the
brain perceives an apparent imbalance (R, 0 spikes/second versus L, 90
spikes/second) The brain erroneously interprets the abruptly decreased or
absent firing rate of the ipsilateral affected peripheral system as a relative
increase of the contralateral system, resulting in a nystagmus that beats
away from the acute lesion.

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 Additionally, the combination of muscle contraction and relaxation
generates not only a pure horizontal nystagmus, but also a torsional
nystagmus.
 Indeed, when on the right side all the canals are lesioned, this is interpreted
by the brain as a sudden excitation of the contralateral vestibular system,
which generates a contraction of the left eye medial rectus, superior rectus
and superior oblique muscles.
 Eye movements may be considered as additive and so although both
muscles are on the upper surface of the eye, they have an opposite effect
on the eye movement, cancelling any vertical movement.
 Considering the right eye, activation of the superior rectus results in an
elevation, adduction and intorsion, whereas the superior oblique generates
a depression, abduction and again an intorsion.
 When all muscles contract simultaneously, as is the case upon stimulation
of the anterior and posterior canals of the same side, the elevation and
depression cancel out, as does the abduction and adduction. The torsional
movement remains, as well as a horizontal nystagmus due to the
contraction of the medial rectus of the left eye (and the lateral rectus of the
right eye).
 The torsional nystagmus is clockwise (CW) (from the subject’s perspective)
for a left uVD and a horizontal nystagmus beating to the right and a counter-
clockwise (CCW) and leftward nystagmus for a right-sided uVD.
 A torsional and horizontal nystagmus is the clinical sign indicating an acute
whole labyrinthine deficiency.
 Conversely, a pure vertical nystagmus is very unlikely to be produced by an
acute labyrinthine lesion and the clinician should in that case firstly consider
a central neurological lesion rather than a peripheral vestibular lesion.
 The sudden onset of this nystagmus is associated with vertigo and
disorientation, since the absence of real movement constitutes a conflict
between vision, proprioception and the vestibular system.

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2) VERTIGO
ORIGINS OF VERTIGO AND MOTION SICKNESS

 Any deficiency in the system, either in the peripheral sensors or in the central
integration, leads inevitably to a problem with balance or vertigo.
 When one sense is misled by disturbing information, the other senses
compensate in general to maintain balance, otherwise one would not be able
to stand up while closing one’s eyes.
 When two senses are deprived, the task to maintain optimal function is
challenged and may readily lead to vertigo, imbalance or disorientation. Such
a condition exists even in healthy subjects when, for example, one is skiing
on a mountain in severe fog. Absence of visual input as well as
somatosensory input (absence of ankle information due to the immobility in
the skiboots attached to the skis) and slow sliding on the snow can be very
provocative of disorientation, even for perfectly healthy people.
 The interplay of attributing differing reliance upon the different senses is
hampered however in the elderly affected by a vestibular lesion. These
people often suffer from additional decreased somatosensory input (for
example, due to diabetes) which increases the risk for falling. It is therefore of
great importance for the elderly to live in optimal conditions favouring the
other senses by sufficient lighting in the house, avoiding loose carpets, proper
spectacles and a cane for additional tactile information of the surroundings.
 However, internal conflicts also lead to vertigo or disorientation. Motion or sea
sickness is the most common example. When you find yourself as a
passenger on a cruise ship but below deck, the visual environment suggests
a stable world. The vestibular system however senses continuously the
movements of the ship on the ocean, especially under storm conditions. Sea
sickness is then often unavoidable, accompanied in the worst case with
vomiting. This is purely due to the sensory input conflict that is provoked by
contradictory information coming from the eyes and the vestibular sensors,
without any deficit imposed on the systems themselves. When both the visual
and vestibular information are matched, by framing the ship with respect to
the horizon for example, by looking outside and viewing the contours of the
deck or other features of the ship, as well as the motion of the ship against
the sea and the horizon, then it is possible to build up a consistent image of
the real movement of yourself and the ship, which then alleviates the
symptoms of motion sickness. Walking on the ship, however, will still be
challenging since this consists of a combination of several movements in 3D
space, i.e. the self-motion and the ship movement.

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 Vestibular pathway

-Vestibular receptors are the macula of the saccule and

utricle and the cristae of the ampullae.
-They are innervated by the prepheral processes of
bipolar neurons of the vestibular ganglion which is
situated in the internal acoustic meatus.
- The central processes from the vestibular nerve which
ends in the vestibular nuclei.

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1- The sensory receptors: three cristae and two maculae.
2- The First neurone: Vestibular nerve.
3- The second neurone: Lie in the Vestibular nuclei. These
send fibers to the following:
 The occulomotor nuclei
 The anterior horn cells of motor part of spinal cord through
the vestibulo-spinal tract. The impulses arising in the labyrinth
can influnce the movement of eyes,head,neck and trunk
through the vestibular pathway.
 The cerebellar cortex and nuclei: Archicerebellum through
inferior cerebellar peduncle(Vestibulo-cerebellar tract).
 Motor nuclei of brain stem(shiefly 3,4 and 6 ) through the
medial bundle.
 The temporal lobe cortex.
 The autonomic nervous system.

4- Third neurone= Efferent neurone= Motor neurones.

5- The effector organ : e.g. The occular and postural
muscles.

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 NERVOUS CONNECTIONS OF THE

VESTIBULAR APPARATUS
The cell bodies of the vestibular nerve are bipolar cells present in the
vestibula ganglion (Scarpa's ganglion). Their peripheral branches end at the
labyrinthine receptors while their central branches from the vestibular nerve
which enters the brain stem at the level of the medulla and ends in the
vestibular nucleus from the vestibular nucleus impulses pass to:
1. Vestibulo-cerebellar fibres: from the vestibular nucleus and directly from the
vestibular apparatus  inferior cerebellar peduncle  flocculondoular lob 
dentate nucleus  superior cerebellar peduncle  that amus of the opposite
side  cerebral cortex, i.e. vestibulo-carebell-dentato-thalamo-cortical
connection. Which ends in the cortical areas:
a. Motor areas: to control the voluntary movement.
b. Equilibrium centre: in the superior temporal gyrus of the temporal lobe.
This pathway is concenrned with the sensation of vertigo. The
equilibrium centre receives also impulses from: vestibular nucleus 
reticular formation  thalamus of the opposite side  temporal lobe.
2. Medial Longitudinal Bundles: of both sides which connect the motor nuclei of
the cranial nerves 3, 4, 6 that supply the extraocular muscles. These bundles
are the pathways for:
c. Correction movement of the eyes which help to fix objects seen in the
visual field when the head is rotated.
d. Nyatagmus
3. Vestibule-reticular fibres: to the reticular formation of the brain stem and
then to the spinal cord along the reticulespinal tract.
4. Vastibulospiinal tract: to the spinal cord.

Impulses from the vestibular apparatus maintain equilibrium by controlling

the tone of the extensor muscles. These impulses reach the spinal cord
along the reticulospinal and vestibulospinal tracts.
So :-
Impulses arising in the labyrinth can influnce the movement of eyes, head,
neck, trunck through vestibular pathways.

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Reflex responses distributes as follows= Functions of vestibular
nuclei:
1- Sup. V. Nucleus Receive from Cristae of SCC& cerebellum Contral
semicircular Canals & occular reflex .
2- Inf.v.n. receive from macula of utricle&saccluleMainly to cerebellums and
Reticular formation
3- lat v.n. Receive from utricle & cerebellumVestibul spinal activity
4- Medial Vn(have type 1 exitatory neurone, type 1 inhibitory neurone and
type 2 inhibitory neurone) receive from cristea of scc and cerebellumCo-
ordinate eye,head,neck through medial longitidudinal bundle.

Subject awarness by vestibule – cortical projections:

vetibular nuclei connect 5 main systems:
1-Occulo motor nuclei 3, 4, 6
2-Motor pant of spired cord
3-Cerebellum
4-Cenebral cortex
5-Autonmic nervous system

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 Vestibulo-ocular reflex

 Utriculopetal stimulus in one labyrinth is matched by an equal but opposite,

utriculofugal displacement in the functionally paired canal of the other ear.
 Accordingly the laeral canals form one pair while the posterior canal pair
with the opposie side superior canal.
 Taking as example-:- Utriculopetal deviation of the cupula of the right
horizontal canal occurs with an utriculofugal movement of the cupula of the
left horizontal canal.
 This results in an increase in the firing rate of the right afferent nerve and
an equal but opposite response in the left afferent nerve. The right afferent
information exerts an exitatory influence on the agonist muscles(axons
decussate on the contralateral abducent nucleus which innervate the
lateral rectus of the contralateral eye through the V1th nucleus. additionally,
the interneurons of the contralateral abducens nucleus project through the
longitudinal medial fasciculus to the ipsilateral medial rectus subnucleus in
the occulomotor nucleus that activates the medial rectus muscle of the
ipsilateral eye ) and an inhibitory influence on the antagonist muscles.

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 The left afferent information reduces the exitatory influence on the
antagonist muscles and disinhibits the agonist muscles.
 This results in contraction of the left lateral rectus and the right medial
rectus muscles producing deviation of the eyes to the left.
 If the stimulus is large, such that the compansatory movement cannot be
obtained within the confines of the orbit, fast movement in the opposite
direction occurs.
 This fast movement is central intiated and mediated by the reticular
formation, cutting off the incoming flow from the vestibular nucleus and
reticular activating neurons directing the occular muscle nuclei to return the
eyeballs to the point of gaze at which the slow component began.
 The combination of this alternating slow and fast movements in opposite
directions is called nystagmus.
 Three controlled oculomotor systems are of clinical importance with
reference to the vestibular ocular reflex.

The Saccadic system:

Generates rapid eye movement to correct errors in the direction of gaze
and bring the desired object of fixation to the fovea in the shortest possible
time.it is produced volantarily or reflexly.

Smooth pursuit system:

Low velocity accurate tracking eye movement enabling amoving target to
be stabilized on the fovea. It is most efficient in low velocity targets and
cannot occur in the absence of atarget.

Optokinetic nystagmus:
It is the reflex oscillation of the eyes included by movements of large areas
of target, e.g. jerking movements of the eyes of a passenger travelling in
atrain as he views the passing landscape. There are two types of
optokinetic reflex:

a) Persuit : normal and requires foveal vision

b) Non persuit: debends on the peripheral retina and occurs abnormally
with central scotoma.

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 Vestibulospinal reflexes

 The vestibular system influences posture and orientation through neck,

axial and limb motor neurons.
 The basing resting activity of the vestibular labyrinth which discharges
impulses at asteady rate is responsible for maintaining normal tone.
 Stimulation of the semicircular canal causes deviation of the head which
tends to restore it to its former position.
 Backward rotation of the head stimulates the posterior vertical canal and
inhibits the anterior ones. This activates muscles which tend to tilt the head
forwards.
 Similarly, limb and trunk muscles respond to head movements with
contractions appropriate to prevent falling and to maintain balance and
restore normal posture.
 The control of posture is by way of the myotactic reflex.
 The myotatic reflex which is also often called The stretch reflex; knee-jerk
reflex, or deep tendon reflex, is a pre-programmed response by the body to
a stretch stimulus in the muscle. When a muscle spindle is stretched an
impulse is immediately sent to the spinal cord and a response to contract
the muscle is received. Since the impulse only has to go to the spinal cord
and back, not all the way to the brain, it is a very quick impulse. It generally
occurs in 1-2 milliseconds.
 The lateral vestibulo-spinal, medial vestibulo-spinal and reticulo-spinal
tracts are the three major pathways by which the vestibular information
influences spinal anterior horn cells.
 The immediate responses to stimulation are generated by the semicircular
canals but after correction of the initial movement, macular stimuli maintain
the necessary muscle tone.

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 POSTURAL REFLEXES AND EQUILIBRIUM

Postural reflexes are a group of reflexes which maintain the body

position and equilibrium during rest (static) and movement (kinetic)
by adjusting the distribution of muscle tone.
Receptors of postural reflexes:
1. Proprioceptors: in the muscles, tendons and joints.
2. Retinal receptors: which discharge to the occipital cortex.
3. The receptors of the non-auditory memberanous labyrinth.
in general

Vinetic reflexes bring the body into normal stance while it is maintained by
activation of various static reflexes.
Ewaid's law

3 observations in vestibular physiology

1- Head and eye movement always occur in the plane of the canal being
stimulated & in direction of endolymph flow.
2- In the horizontal cala: ampullo petal enolymph flow greater response.
3- Vertical canal: Umpullo Fugal flow cause greater response than
ampullo petal flow.

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Acute peripheral vestibular lesion and vestibular compensation
 After uVD, specific changes occur immediately at the level of type 1 and
type 2 neurons in the MVN on the ipsilesioned site.
 Given the absence of peripheral activity by the SCC afferent neurons, the
firing rate of the type1 neurons decreases. Due to the decreased inhibitory
effect of the ipsilateral type 1 neurons, the contralateral type 2 neurons are
less stimulated so that their inhibitory effect on the contralateral healthy
type 1 neurons is decreased, and thus the healthy type 1 neurons increase
their firing rate.
 This increased type 1 activity on the healthy side in turn activates the
inhibitory type 2 neurons on the lesioned side, so that they additionally
inhibit the neighbouring type 1 neurons on the lesioned side. This
imbalance generates the typical clinical signs of acute labyrinthine lesions,
such as spontaneous nystagmus, i.e. a nystagmus that is present even
under static conditions.
 The direction of the slow phase of the nystagmus is towards the lesioned
side, i.e. the fast phase of the nystagmus beats towards the healthy side.
 The generated nystagmus reflects the situation as if the subject rotates
towards the intact side. Indeed, both the absence of the ipsilesioned tonic
input from the affected labyrinth and the increased contralesioned activity
in the MVN mimic a rotation towards the intact side.
 Stance and gait disturbances, as well as vertigo, are clearly observed in
most patients. The postural disturbance often includes head and trunk
flexion towards the damaged labyrinth with the head tilted so that the
ipsilesioned ear is directed down. The appearance is of the healthy side
being pushed towards the damaged side, which lacks the power to
counteract the push. These disturbances also occur under static, immobile
conditions.
 The dynamic alterations yield a dysfunctional VOR, so that gaze
stabilization during, for example, locomotion is hampered. Whereas the
static symptoms usually improve within a week, the dynamic disturbances
can last much longer, i.e. from weeks to years.

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 The static symptoms decrease over time, as a restoration of the resting
discharge rate is affected at the level of the vestibular nuclei. This results
in the disappearance of the spontaneous nystagmus: a process called
‘vestibular compensation’. In humans, the gain of the VOR is permanently
limited to a variable degree, depending on the acceleration of the head
movements.
 Many VN neurons show convergent input from both canal and otolith
systems. The spontaneous activity of otolith afferents arriving in the
vestibular nuclei is important for the generation of the oculomotor
compensatory responses to SCC stimulation. The consequence of this is
that an otolith loss not only affects the lVOR, but also the aVOR during
angular acceleration. The otoliths are therefore fundamental for the optimal
operation of the entire vestibular system.
 The imbalance in resting discharge rate between the lesioned vestibular
neurons and the healthy vestibular neurons after uVD is the key factor that
generates the acute clinical signs of nystagmus and unsteadiness.
 The short term restoration of the imbalance of resting activity at the level of
the VN results in a recovery of the static clinical symptoms, i.e. acute
spontaneous nystagmus disappears within a week.
 Very soon after uVD (within 52 hours), type 1 neurons in the MVN
generate a spontaneous firing rate to balance the situation, although this
firing rate is not influenced by afferent input from the lesioned side during
rotation towards the lesioned side. It is, however, modulated through the
ipsilesioned type 2 neurons that receive their input from the intact side type
1 neurons. Therefore, the ipsilesioned type 1 neurons are inhibited when
the type 1 neurons on the healthy side are excited during rotation towards
the healthy side and are disinhibited when the contralateral type 1 neurons
are inhibited during rotation towards the lesioned side. The responses of
the ipsilesioned type 1 neurons are only half of their initial responsiveness,
but qualitatively they act in a similar manner to normal. This recovery takes
place over a period of weeks to months and parallels the clinical recovery
of the patient. The type 1 neurons on the intact side regain a firing rate
equal to that before the acute lesion.

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Principle of the head impulse (thrust) test
 The head-impulse or head-thrust test detects severe unilateral loss of
semicircular canal function.
 Clinically, is more sensitive and specific than the traditional Romberg and
similar tests, and is particularly important in the emergency unit where this
test can distinguish between vestibular neuritis and cerebellar infarction
that can both generate similar symptoms of an initial attack of severe acute
vertigo. The result of the head-thrust test is definitely normal in a patient
with a cerebellar infarction, but may be abnormal in a patient with
vestibular neuritis.
 The head-thrust test is based on the fact that inhibition of primary and
secondary vestibular neurons cannot produce fewer than 0 spikes per
second. Excitation can drive the discharge rate from 90 to 300 or more
spikes per second. So when the healthy side is excited for a high
acceleration head movement, the healthy side will generate the larger part
of the VOR, since the disinhibition of the ipsilateral type 1 neurons by the
contralateral SCC contributes relatively little to the VOR.
 Passive head impulses or thrusts are typically rapid, but have a small
amplitude. They have to be unpredictable since the patient very quickly
learns to anticipate and this largely decreases the sensitivity of the test.
Therefore, the examiner should at random thrust the head of the patient
firmly from left to right and a little later, i.e. not immediately, from right to
left. The starting position should be such that the patient’s head is turned
slightly over the midline to thrust it just over the midline to the opposite
side. As such, little amplitude but great acceleration can be imposed. This
execution demands some training, particularly to position the hands on the
side of the head and hold the head firmly.
 The instruction to the patient is to fixate a point in the distance behind the
examiner. When the subject’s head is turned to the lesioned side, the VOR
is deficient and the eyes move together with the head, so that they do not
fixate the point any more. Therefore, the patient has to make one or more
refixation saccades just after the thrust. When the head impulse is
imposed in the direction of the healthy side, the VOR is able to maintain
the target on the fovea and no refixation saccade is needed.

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 The head-thrust test is positive for the side that causes the refixation
saccade upon thrust. Not only can the lateral SCC be examined, which is
in a sense a clinical approximation of the caloric test, but also the other
SCC can be investigated.
 The patient’s head has then to be moved in the RALP or LARP planes.

Head impulse test for horizontal canals. Hold the subject’s head slightly turned to one side ((a) and (c)),
and ask them to maintain fixation on a target behind you. Then, turn the head briskly to the opposite
side. If the gaze is still focused on the target, (as in (b)), then the semicircular canal on the side towards
which the head was turned (in this picture to the right) is functioning properly, generating compensatory
eye movements upon excitation. However, when the eyes rotate together with the head and are no
longer focused on the target, the semicircular canal on the side towards which the head was turned is
not properly functioning (d). The off-target eye movement is quickly followed by a re-fixation saccade, so
that the target is in sight again. Since this happens quickly, careful observation and handling by the
investigator is required to notice abnormalities. When posterior and anterior canals need to be
investigated, brisk head movements need to be made in the right anterior – left posterior (RALP) and left
anterior – right posterior (LARP) planes while observing the gaze stabilization.

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