HISTOLOGY OF THE EAR

The paired ears are

specialized organs
adapted for...

1. The reception and

transduction of vibration
into sounds (audition)

2. Detection of position of
the head (and body)
relative to gravity

3. Detection of motion and

acceleration of the head
adapted across species to
meet environmental needs
Human
– unequalled in the ability
to detect rapidly
articulated speech
- well adapted for the
localization of sounds in
space
(surpassed only by echolocators).
The Ear is traditionally ♣
divided into 3 parts
1. EXTERNAL EAR =

- Auricle (pinnae)
- External Auditory Meatus
- Tympanic Membrane

2. MIDDLE EAR =
- Tympanic Cavity
- Ossicles
- Eustachian Tube

3. INNER EAR =
- Bony Labyrinth
- Membranous Labyrinth
 ♣
EXTERNAL EAR:
Collects sound vibrations –
directs them towards the
sensory transducers.
3 parts

1. AURICLE:

- irregular plate of elastic

cartilage - - covered by thin
skin w/ - hair follicles &
sebaceous glands.

In lower animals striated muscle

is present for directing this
structure toward the source of
sound.

involved in the localization of

sound in space.
2. EXTERNAL AUDITORY MEATUS:

2.5 cm tube connected the

temporal bone

- outer 1/3 -- elastic cartilage

continuous with the
cartilage of the auricle.

- inner 2/3 is formed by the

temporal bone

lined by thin skin

- w/ modified apocrine sweat

glands called CERUMINOUS
GLANDS that secrete a brown
waxy protective secretion =
CERUMEN
3. TYMPANIC MEMBRANE (EARDRUM):

closes the innermost aspects of

the external meatus

consists of 2 layers of collagen

(radial and circular) covered
on:

– the meatus side by a layer of

very thin skin

– the inside by a simple

squamous epithelium

sound pressure wave displace

this membrane and its
vibration is transmitted to the
middle ear bones.
MIDDLE EAR:
♣♣
transduces sound pressure
waves into mechanical
displacement of inner ear
structures. 3 major
components

1. TYMPANIC CAVITY:
box-like air filled cavity deep to
the external meatus.

- lateral wall =tympanic

membrane

- medial wall =the bony

labyrinth of the inner ear

acts as a resonance chamber.

Lined with simple squamous

epithelium with a thin lamina
propria
2. EUSTACHIAN TUBE
4 cm cartilaginous tube
- connects the tympanic cavity
with the nasopharynx.

- equalizes pressure between

the outside air and the
middle ear cavity.

-- cartilage, covered by a
pseudostratified columnar
ciliated epithelium

Clinically Important:
- allows direct access to the
middle ear cavity and is thus
a common route for infection
( Otitis Media).
 ♣♣
3. OSSICLES:
Tympanic Cavity is spanned by a series of
three small bones called the Auditory
Ossicle.

From external inward these are the Malleus,

Incus, and STAPES.

Malleus - attached to the tympanic

membrane at one end and the Incus at the
other.

Incus - suspended between the Maleus and

the Stapes

Stapes - attached to the Incus at one end

and a small oval membrane-covered
opening in the medial wall of the tympanic
cavity called the OVAL WINDOW at the other.

The Malleus and the Incus are suspended

by small ligaments from the roof of the
tympanic cavity
ossicles acts like a lever system to
transmit the movements of the
tympanic membrane to the sensory
apparatus of the inner ear.
- decrease the amplitude
- but increase the force of the
mechanical displacements of the
tympanic membrane ="IMPEDANCE
MATCHING".

The chambers of the inner ear are

fluid filled - not compressible
- an outlet must exist for the forces
transmitted by the ossicles to the
Oval Window.
- This function is served by a
second opening in the medial
wall of the tympanic cavity
called the ROUND WINDOW
covered by fibrous membrane called
the Secondary Tympanic Membrane
 ♣♣
INNER EAR:
houses the vibration and
motion/pressure - sensitive
receptors for hearing and
vestibular sensation.

BONY LABYRINTH
a series of bony channels
consisting of 5 parts:

1. VESTIBULE:

large irregular, central cavity

whose lateral wall contain
two membrane covered
openings = "Oval" and
"Round" windows which lead
into the adjacent cochlear
canal.
EXTENDING OUT FROM THE VESTIBULE
ARE 4 BONY TUBE-LIKE CHAMBERS
– 1 Coclear canal
– 3 Semicircular canals

2. COCHLEAR CANAL:
♣
Cochlear canal - anteriorly extending
tube (35mm ) that spirals like a
snails shell for 2 3/4 turns around a
central bony axis called the
MODIOLUS.

-- Extending laterally from the

modiolus like the threads of a screw
is a thin bony ridge called the SPIRAL
LAMINA

-- spaces within the bony modiolus

house blood vessels and the cell
bodies and processes of the Spiral
Ganglion cells and the Acoustic
Branch of the VIII cranial nerve
 3- SEMICIRCULAR CANALS
anterior - posterior - lateral
(superior)

- extend posteriorly from the

vestibule. (armchair)

Each canal has a dilation at its

connection with the vestibule
called the AMPULLA

The Bony Labyrinth is

• filled with a clear fluid called

PERILYMPH
• which resembles extracellular
fluid (and CSF) in composition
(high Na+, low K+)
 ♣
MEMBRANOUS LABYRINTH
The Channels of the Bony Labyrinth
house membranous ducts. which
contain the specialized sensory
organs for hearing and vestibular
sensation termed the membranous
labyrinth.
delicate connective tissue , lined
with a simple squamous epithelium.
The ducts are suspended within the
perilymph of the bony labyrinth by
thin strands of connective tissue
containing blood vessels.
The shape of the membranous
labyrinth follows that of the Bony
Labyrinth
except the large bony vestibule
region is occupied by two
membranous sacs called the
UTRICLE and the SACCULE
The components of the Membranous ♣
Labyrinth can be divided into those
associated with:
1. VESTIBULAR SENSE:
3 Semicircular Ducts +Utricle and
Saccule .
2. AUDITION: Cochlear (Auditory) Duct
membranous ducts are all
• interconnected and share a
common fluid medium ENDOLYMPH
• similar in composition to
intracellular fluid ( K+, Na+)
Endolymph: drains into a small duct
arising from the Utricle and Saccule
=ENDOLYMPHATIC DUCT
- terminates in just above the dura
of the CNS. Endolymph is filtered
back into the CSF through this
duct.
AUDITORY SENSORY APPARATUS

COCHLEAR DUCT- situated in the

Cochlear Canal of the Bony
Labyrinth

suspended medially within the

Cochlear Canal and follows its coiled
path ending as a blind sac at the
apex of the Cochlear Canal

roughly triangular in transverse ♣

section and divides the Cochlear
Canal into three partitions:
S SCALA VESTIBULI
S SCALA MEDIA (COCHLEA)
-- SCALA TYMPANI
 The Scala Vestibuli and Scala
Tympani

- perilymph filled spaces lying on

either side of the Cochlea.
- lined with a thin connective tissue
continuous with the periosteum of
the Cochlear Canal.

-- The scala vestibuli +scala tympani

communicate across a small
aperture at the apex of the Cochlear
Canal known as the HELICOTREMA.

-- The central partition of the Cochlear

Canal is the COCHLEA whose apex
points toward the Bony Modiolus.

- filled with endolymph.

 The Walls of the Cochlear Duct are
formed as follows:

A. Roof = REISSNER'S (VESTIBULAR)

MEMBRANE

-- 2 layers of simple squamous

epithelium separated by a
basement membrane.
B. lateral wall - a thin layer of
connective tissue applied to the
walls of the Cochlear Canal and
covered by a highly vascular
epithelium called the STRIA
VASCULARIS.
-- These vessels are thought to be
the source of endolymph.
C. Floor =BASILAR MEMBRANE
-- composed of amorphous ground
substance infiltrated with
transversely oriented filaments
whose lower surface is covered by a
columnar epithelium.
BASILAR MEMBRANE is not uniform in
its width
-- 100um wide at its basal region
where it is most responsive to high
frequency vibrations
-- 500um wide at its apical region ♣
where it is most responsive to low
frequency vibrations
Supports the auditory apparatus, the
ORGAN OF CORTI which is anchored to
the Bony Labyrinth by the SPIRAL
LIGAMENT at one end and the
OSSEOUS SPIRAL LAMINA at the other.
 ♣♣
THE ORGAN OF CORTI:
This is a complex organ of
hearing which rest atop the
Basilar Membrane.

The functional elements of the

organ of corti are:
A. TUNNEL OF CORTI:

- A triangular shaped tunnel

located midway across the the
basilar membrane

- running the entire length of

the Cochlea.

The base of the tunnel is formed

by the Basilar Membrane and the
walls are formed by Supporting
cells
B. Two major types of
columnar supporting cells

1. PILLAR CELLS:

-- Cone shaped columnar

cells with basally placed
nuclei.

-- rigid cells due to

numerous cytoplasmic
microtubules

-- These cells form the walls

of the Tunnel of Corti
2. PHALANGEAL CELLS:

-- Tall columnar cells that support

the base of the sensory hair
cells

the apical surface of these cells

project a slender cytoplasmic
process called the Phalanx which
extends to the free surface of the
organ of corti alongside the hair
cells to terminate in a plate-like
expansion ♣
The Phalanx envelopes the apical
surface of the sensory hair cells.

The lateral borders of adjacent

phalanxes have well developed
tight junctions.
C. Two types of Sensory Hair Cells

1. INNER HAIR CELLS:

-- Short goblet shaped cells,

numbering about 3500,

-- arranged in a single row along

♣
the length of the cochlea.

50-70 specialized microvilli

known as "stereocilia" extend
from the apical surface of each
cell.

These cells are sensitive to small

changes in sound "intensity".

recieve a dense innervation by

fibers of the 8th nerve

– each hair cells receiving synapses

from an average of 20 nerve fibers.
 ♣
2. OUTER HAIR CELLS
-- cylindrical columnar cells,
with basally located nuclei,
numbering about 20,000
-- form 3 rows extending the
length of the cochlea.
apical surface has 100 - 300
stereocilia of uneven length
arranged in the shape of a “ W “
The tips of the tallest stereocilia
are embedded in the tectorial
membrane.
Outer Hair Cells:
- respond best to "low
intensity" sounds
- have relatively few neuronal
connections.
 A single 8th nerve fiber
innervates and average of 50
outer hair cells.

THE BASE OF BOTH HAIR CELL TYPES

SERVE AS THEIR SYNAPTIC SURFACE.

EACH MAKES NUMEROUS CONTACTS

WITH THE DENDRITIC PROCESSES OF
THE SPIRAL GANGLION CELLS OF THE
VIIITH CRANIAL NERVE .
 ♣

D. TECTORIAL MEMBRANE:
-- thin gelatinous membrane
composed of a "Keratin-like"
protein.

-- Extends from the spiral laminae to

form the superior surface of the
organ of corti.

-- thought to originate as a secretion

of the cells of the spiral laminae.

THE TIP OF THE HAIR CELL STEREOCILIA ARE

EMBEDDED IN THE TECTORIAL MEMBRANE.

THE SHEERING OF THIS MEMBRANE WITH

RESPECT TO THE CELL BODIES OF THE SENSORY
HAIR CELLS TRIGGERS ELECTRICAL IMPULSES IN
THESE CELLS AND RESULTS IN SOUND
TRANSDUCTION.
 ♣
HOW IT ALL WORKS:
The transformation of vibrations in the air
to sound in the auditory apparatus is
thought to occurs as follows:
1. Pressure waves in the air enter the
External Ear
2. These travel down the External Auditory
Meatus resulting in mechanical
displacement of the Tympanic Membrane
3. Vibrations of the Tympanic Membrane
are reduced in amplitude and increased
in force the mechanical leverage of the
Ossicles a processes referred to as
"Impedance Matching".
4. Vibrations are transferred from the
Ossicles to the Oval Window, movement
of the Oval Window induces a traveling
wave in the perilymph of the Cochlear
Canal that proceeds around the Scala
Vestibuli -- through the Helicotrema --
down the Scala Tympani - pressure wave
dissipates at the Round Window
5. As the pressure wave travels around
the Scala Vestibuli and Scala
Tympani, vibrational forces are
transmitted to the Basilar
Membrane.

Movement of the Basilar Membrane

causes a "shearing motion" of the
hair cell stereocilia with respect to
the Tectorial Membrane

6. Oscillations of the stereocilia of the

sensory hair cells results in their
depolarization, which results in the
initiation of afferent impulse in the
VIII nerve (spiral ) ganglion cells in
the Modiolus, which contact them

7. These peripheral neural impulses are

transmitted via a variety of
pathways to the auditory cortex of
the CNS.
CODING STIMULUS ATTRIBUTES OF
SOUND:

1. Sound Amplitude:
Loud sounds produce a larger
amplitude traveling wave that
stimulates more hair cells on the
basilar membrane

2. Sound Pitch:

2 theories of pitch encoding

A. Volley theory - the frequency

of a sound vibration is encoded
by the frequency of discharge
of the 8th nerve afferents. for
higher frequencies ( >200 Hz)
fibers fire in phase with the
stimulus but different groups
take turns skipping a volley.
 B. Place Theory
Traveling waves caused by
different sound frequencies --
"peak" in their displacement of
the basilar membrane in
different regions along the
length of the cochlea.
-- HIGH FREQUENCIES
(20,000 HZ) = BASE
-- LOW FREQUENCIES (500 HZ)
= APEX
Prolonged exposure to loud
sound of a given frequency
results in degeneration of the
hair cells and nerve fibers in
that region of the cochlea.
 ♣
THE VESTIBULAR SENSORY
APPARATUS
arises from the VESTIBULE portion
of the bony labyrinth adjacent to
the cochlear canal. They Consist of:
1. Two endolymph-filled,
membranous sacs, the UTICULE and
SACCULE housed within the
Vestibule of the Bony Labyrinth
-- the sensory epithelium in these
membranous structures detect
changes in head position and
linear acceleration.
2. Three, endolymph-filled
membranous SEMICIRCULAR DUCTS
(suspended within the bony
semicircular canals) which arise
from an expanded region of the
Utricle called the AMPULLA.
 A semicircular duct is suspended
within each semicircular canal
and the 3 canals lie orthogonal
to one another along the three
planes of movement and are
responsible for the detection of
angular movement of the head.

The ducts are part of the

Membranous Labyrinth and thus
composed of thin connective tissue
walls lined by a simple squamous
epithelium.

SEMICIRCULAR DUCTS & THE

AMPULLA:

semi-circular ducts that connect to

the Utricle at either end.

Each exhibits an expanded region

known as the AMPULLA near one of
its junctions with the Utricle.
The Ampulla of each semicircular duct
contains a raised transverse ridge =
CRISTAE AMPULLARIS which extends
part way across the semicircular
canal.
♣♣
The epithelium of the Cristae is
composed of 3 cell types
A. Tall columnar supporting cells =
SUSTENTACULAR CELLS
B. Two types of SENSORY HAIR CELLS
TYPE I:
-- Flask-shaped cells with apical
stereocilia similar to the inner hair
cells of the basilar membrane.
-- stereocilia are flanked by a single
non-motile kinocilium
TYPE II:
-- cylindrical cells with apical
stereocilia similar to outer hair
cells of the basilar membrane. also
possess kinocilia
 Sensory hair cells receive
contacts from neurons of the
Vestibular (Scarpa's) ganglion
located at the base of the
modiolus.
-- Type I cells - receive
funnel-shaped (chalice)
endings
-- Type II cells receive
bouton-like endings.
THE CENTRAL AXONS OF THESE
GANGLION CELLS TERMINATE IN THE
CNS IN THE VESTIBULAR NUCLEI OF
THE BRAINSTEM. ♣
The apical stereocilia of the
sensory hair cells are embedded
in a gelatinous glycoprotein
material = CUPULA which extends
like a hinged flaps from the
walls of the ampullae .
Rotation or acceleration of the head in
the plane of a given semicircular canal
♣♣
• causes the stereocilia of the sensory
hair cells to bend against the inertia
of the endolymph and the Cupula.

• This results in depolarization of the

sensory hair cells and initiation of
action potentials in the Vestibular
Ganglion cells

THE UTRICLE AND SACCULE:

The Utricle and Saccule

• are housed in the vestibule of the

bony labyrinth

• contain a small 3mm x 3mm patch

of sensory epithelium called the
MACULA.
 ♣
The Macula is responsible the
detection of:

• head position with respect to

gravity

• and linear acceleration of the head

and body.

-- The macula of the saccule is

oriented in the vertical plane

-- the macula of the utricle is

oriented in the horizontal plane.

composed of 2 cell types:

1. columnar supporting cells
2. sensory hair cells similar to
♣
those found in the Cristae
 ♣
The apical surface of the
sensory hair cells are covered by
a gelatinous glycoprotein layer
called the OTOLITHIC MEMBRANE

Suspended in the Otolithic

Membrane are small (3-5 um)
crystalline bodies composed of
calcium carbonate called
OTOLITHS

During tilt of the head with

respect to gravity or linear
acceleration..

– the inertia provided by the

Otoliths in the Otolithic
Membrane cause bending of the
stereocilia of the hair cells.
This generates a depolarization of the
sensory hair cells and initiates action
potentials in the Vestibular ganglion
cells.

The central nervous system

connections of the vestibular system
are quite complex and will be studied
in great detail next quarter in the
neuroscience course.
 ♣♣
THE STRUCTURE OF HAIR CELLS:

The basic structure of hair cells is

similar throughout the
auditory / vestibular apparatus
- regardless hair cell type (type I vs
type II) or the organ in which they
are found (basilar membrane vs
macula vs cristae ampularis.
All share common features and
mechanisms of action.
1. columnar cells capable of
tranducing mechanical stimulation
into electrochemical (nerve)
impulses
2. stereocilia arranged hexagonally
on the apical surface
-- stereocilia are composed of actin
filaments surrounded by
paracrystaline structures which
provided substantial rigidity.
3. Stereocilia are graded in height with
the tallest and shortest at opposite
ends of the bundle.

4. A true cilium called a KINOCILIUM is

adjacent to the tallest row of cilia
(except in the mammalian cochlea
where kinocilium are lost soon after
birth).

5. The tips of the stereocilia are ♣♣

extensively cross-linked by protein
bridges called "TIP LINKS."

6. movement of the steroecilia produce

depolarization of the hair cell
cytoplasm.

-- The orientation of the "TIP-LINKS" is

such that they are at an extreme angle
(nearly verticle) which provides for a
unidirectional responsiveness of the
cilia.
-- Hair cell depolarization occurs when
the cilia are bent in the direction of the
tallest cilium which tightens the TIP -
LINKS.

-- Movement in any other direction

loosens the tip-links and has no
effect on the hair cells.

The mechanism by which hair cell

depolarization occurs following
stereocilia movement must be fast
enough to accomodate the rapid
frequncies of vibration that the
auditory/vestibular system is capable
of responding to( > 1000 Hz).

This rules out any potential second

messenger gated ion channel
apparatus in favor of a direct coupling
of cilia with membrane ion channels.

The location of the channels and the

exact mechanism for their regulation is
still not known.
 ♣
BLOOD SUPPLY OF THE EAR:
external ear - middle ear - bony
labyrinth - served by vessels arising
from branches of the external carotid
artery

The blood supply to the inner ear

arises from the LABARYNTHINE ARTERY a
branch of the basilar artery. The
labarinthine artery divides to form:

1. ANTERIOR VESTIBULAR ARTERY --

supplies most of the semicircular
canals

2. COMMON COCHLEAR ARTERY -- supplies

the Cochlea, Utricle and Saccule
and the remainder of the
Semicircular Canals
 Because the Hair cells of the auditory
and vestibular apparatus have a very
high metabolic rate. Interruption of
blood flow in the labarinthine artery for
even brief periods (minutes) can cause
irreversible damage to these cells.

DISORDERS OF THE EAR:

Clinically disorders of the ear are quite

common particularly with advancing
age.

Loss of hearing most commonly results

from:

1. destruction of the hair cells or 8th

nerve fibers in the cochlea.

-- Hair cells (particularly type I)

sensitive to a variety of
antibiotics, diuretics, and
salicylates
 ♣
-- chronic exposure to loud
sounds of a constant frequency
can result in the excitotoxicity
of hair cells and the 8th nerve
fibers which innervate them.

2. fixation or calcification of
the ossicle

3. rupture or puncture of the

tympanic membrane

4. Tumors of the 8th nerve or

it's ganglion
Vestibular dysfuction:

1. Drug toxicity - particularly certain

antibiotics and diuretics

2. overproduction or blockage of
endolymph circulation can result
in abnormal stimulation of the
vestibular and auditory hair cells .

(MENIER'S DISEASE) and is

characterized by intense
debilitating vertigo, nausea, and
vomiting as well as abnormal
sound perception and sometimes
temporary deafness.

3. Tumors of the 8th nerve or it's

ganglion.