Inner Ear

The inner ear is divided into 3 parts:

a) Pars superior: scc + utricle
b) Pars inferior: saccule + cochlea
c) Endolymphatic sac + duct
Osseous labyrinth:

Located in the petrous portion of the temporal bone

Surrounds the membranous labyrinth
Contains perilymph
Connects the middle ear via fenestra vestibule & fenestra
cochleae
Divided into 3 parts:
1. Vestibule
2. Cochlea
3. Semicircular canal

Membranous labyrinth:

Located within the bony labyrinth

Contains endolymph
Divided into :
1. Cochlear duct= sacula media
for hearing
2. vestibular apparatus
for balance and motion sense which is made
of
a) Semicircular ducts
b) Saccula
c) Utricle

Note : The oval window and round window open into the vestibule, at the base of the cochlea

Cochlea

The cochlea is coiled around the modiolus

The Modiolus:

The spongy bone axis around which the cochlea is colied

(central axis of the cochlea)

Contains:

1. cochlear nerve
2. Spiral ganglia: comprised of bipolar neurones that:
a) peripherally innervate the hair cells
b) centrally form the cochlear nerve
3. Blood vessels

The cochlear coil extends up from its base (2 3/4 X)

The chambers spiral and exhibit 5 cross sectional profiles.

Different pitches (frequencies) of sound are detected at
different positions along this long coiled structure. (high
frequency at the base, low frequency at the tip)

Cross section through 1 turn of the cochlea shows that the

cochlea is dividied by Reissners membrane and the basilar
membrane into 3 scalae(chambers):
1. Scala vestibule
2. Scala media (cochlear duct)
3. Scala tympani

Note:
o The perilymph of The scala vestibuli and scala tympani are continuous with
one another at the tip of the cochlea (helicotrema)

o
o
o

The bony wall of the cochlea has 2 defects, each covered by a thin membrane
PA( Round & oval window)
Scala vestibule is connected to the oval window
Scala tympani connected to the round window

Cochlea: 3 scala: scala media

Scala media is more or less triangular:

formed by :
a. Reissners membrane ( above)
b. basilar membrane (below)
c. stria vascularis:
(laterally/outer part) base of the triangle lying against the bony wall of the
cochlea
derived from the proper cochlear artery
made of basal & marginal cells
marginal cells are the one responsible 4 formation + absorption of endolymph (
hence the source of action potentials in the inner ear=endocochlear potential)
It contains endolymph ( produced by sacula vascularis)

a) The Reissner membrane:

3 layered structure
Consisting of 2 cell layers separated by a basal lamina
Attached
medially to the modiolar edge of the spiral imbus
laterally : spiral ligament at the apical edge of stria vascularis

b) The basilar membrane contains the organ of corti

The side of the duct where the nerve fibers exit is the inner /modiolar side of the duct.
The opposite side is theouter side
The spiral ligament: the prominent cells in the matrix is fibroblast type I

c) Stria Vascularis :
It extends from the attachment of Reissner's membrane to the spiral prominence.
it is an unusual epithelium in that it lacks a basement membrane.
It is a stratified epithelium containing primarily three cell types:
1. Marginal
2. Intermediate
3. basal cells
intraepithelial capillaries

1. marginal cells:
the primary functional units of the stria vascularis
produce the positive DC endocochlear potential
They are the only cells of the stria vascularis that directly face the endolymph

their basolateral surface is convoluted into numerous mitochondria-packed folia that

are rich in Na+ and K+-ATPase and Na-K-Cl cotransporter
2. Intermediate cells:
stellate-shaped
contain melanin
have phagocytic activity
contain carbonic anhydrase enzyme activity
3. Basal cells:
The basal surface of the stria vascularis is sealed from paracellular transport of materials
by several layers of tightly interdigitated basal cells.
the basal cells exhibit numerous tight junctions with adjoining cells of the stria vascularis
and the spiral ligament.
It contains lipid inclusions and has apical processes that interdigitate with processes of
the marginal and intermediate cells.
4. melanocytes
Intermediate cell + intraepithelial capillaries comprise the middle layer of the stria vascularis:
The capillaries are formed by nonfenestrated endothelial cells that are surrounded by a dense
homogeneous basement membrane .

This basement membrane may be formed by the fusion of epithelial and endothelial basement
membranes during development.
The capillaries are also surrounded by pericyte cells.

Organ of Corti/ spiral organ

Specialized epithelium (sensory receptor for sound) Located within the inner part of the basilar
membrane

Components of the organ of Corti :

a) cells :
1. hair cells: inner + outer
2. support cells (pharyngeal cells):
Deiters, Hensen, Claudius

provide structural and metabolic support for the organ of Corti.

The phalangeal processes of the Deiters cells form tight cell junctions of the
reticular lamina.
b) tectorial membrane
c) reticular lamina

Hair cells of corti organ

1. Hair cells:
Receptor cells ( the ones that transduce sound)
It possesses stereocilia.
Name of pillar cells
Inner hair cells

Outer Hair cells

Site
Number of rows

Modiolar side
1 row

Outer side of the tunnel of corti

3 rows

Total number

3.000

12.000

Shape of the cell

Bulbous(flask)

Columnar (cylineric)

Shape of cilia

gentle curve (From inside toward outside

Each stereocilia is taller than the next

W shape (with a very shallow

central notch)

Number of cilia

Few

Many

Attachment of cilia to
tectorial membrane
Function

Non attached

Longest stereocilia is firmly attached

Relation to the
support cells
Afferent innervations

principal transducer of motion from the

Cochlear amplifier: amplify motion
basilar membrane to a nerve impulse (Send inf from the basilar membrane at the
about hearing to the brain)
point of maximum response (which
increase sensivity & frequency
selectivity)
controlling and fine tuning
mechanisms
Contains actin and myosin to adjust
the size of the cell
Surrounded completely by the supporting
Only the base
cells
Type I
Type II

Afferent innervations
%
Efferent source

90%

10%

Lateral superior olivary complex

Medial superior olivary complex

Post synaptic target

Denterics of the afferent fibers

Base of the hair cells

The tips of the stereocilli are mechanically linked by tip filaments, which are directly connected
to mechanically gated cation channels in the plasma membrane covering the stereocilia.

When the bundle of stereocilia is deflected in the direction towards the longer
stereocilia, the tension on the tip filaments is increased and the mechanically gated
ions channels are opened, allowing potassium and calcium to enter the cell, which
becomes depolarized.

When the bundle of stereocilia is deflected in the direction towards the shorter stereocilia,
tension is reduced on the tip filaments (also referred to as gating springs) and the ion channels
close and the cell becomes hyperpolarized.
The state of polarization of the hair cell controls the release of neurotransmitter at the cell's basal
end, where the cell synapses with a nerve ending.

The tops of the hair cells form the reticular lamina, which isolates the hair cells stereocilia from
Their cell bodies

The probable site of noise induced hearing loss is stereociliary rootlets of the hair cells.
Tectorial membrane (non-cellular/extracellular matrix membrane) is in intimate contact with the
stereocilia on the apical surface of the outer hair cells.

2. Support cells:
Support hair cells
Such as the Deiter,Claudius and Hensen.
Notice that :
o The Deiters cells support the Outer hair cells at their base so the outer hair cell walls
are surrounded by fluid.
o The inner hair cell is compleately surrounded by support cells

ACOUSTIC NERVE

auditory nerve has approximately 30,000 fibers

it is made of:
1. afferent Fibers
2. efferent Fibers

1. Afferent fibers

Carries information:
from the hair cells in the inner ear to the Ipsilateral cochlear nucleus in the brainstem

Bipolar neurons
Their cell body reside in the spiral ganglion in the
modioludus

Types of afferent neurons:

Type I neurons

Type II neurons

Innervate

Inner hair cells

Outer hair cell

% of afferent

90%

10%

Mylination of the central

& peripheral process
Other name

Mylinated

unmylinated

Radial fibers

Outer spiral fibers

Differences in afferent nerve (typeI, typeII)innervations of (IHC,OHC) pattern:

A. convergent manner of Type I innervating IHC:

Each IHC has its own private set of fibers ( each inner hair
cell is connected to 20 afferent fibers that does not share with
surrounding cells)
Type I afferent innervates the IHC via a peripheral process
that terminates on the hair-cell base as a bulb.
B. Divergent manner of Type II innervating OHC:
peripheral processes enter the organ of Corti through the
same route as that of the type I
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
branching to innervate up to ten outer hair cells each

Note:
The peripheral process of type 1 neurons 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

Most, if not all, recordings from auditory nerve fibers are from the larger type I fibers in
contact with inner hair cells.

low frequency fibers occupy the centre of the auditory nerve while the high frequency
occupy the peripheral part

this tonotopic organization is continous throughout the central auditory complex

the 1st synapse is at dorsal & ventral cochlear nuclei (so the
1st obligatory relay center for CN VIII afferent fibers is the
cochlear nucleus)

then fibers goes to the ipsilateral/contralateral superior olive

majority ascend in the contralateral lateral leminiscus to

inferior colliculus ; medial genigulate ganglion: auditory
cortex ( note it relays=synapse in all of them)

The primary auditory centers are located in pons:

1. Inferior colliculus ( inferior corpora quadrigemina)
2. Medial geniculate ganglion

the following mnemonics outline the neural pathway:

E COLI:
(Eighth nerve, Cochlear nucleus, superior Olivary nuclei, Lateral lemniscus, Inferior colliculus)
S-L-I-M:
(sup olivary n,lateral lemniscus,inferior colliculus,medial geniculate)

The primary auditory cortex is:

Brodmann's area 41 in the superficial area of temporal lobe
The blood supply of the auditory cortex is the middle cerebral artery

Type 1 fibers: sparsely granulated

Patterns of efferent Fibers (olivochochlear fibers):

carries inf from auditory cortex into SOC then into hair cells
highly granulated

monopolar

Note: both IHCs and OHCs receives bilateral innervations

The majority of innervations into the outer hair cells
Efferent fibers to OHC are mylinated while IHC unmylinated
Medial efferent system

Lateral efferent system

Origin

medial superior olivary complex

lateral

Crossed vs uncrossed

Majority cross midline

Do not cross

Type of innervated cells

outer hair cells directly

Inner hair cells indirectly

Mylination

Mylinated

unmylinated

Function

suppress outer hair cell motility to make

the cells less sensitive, providing protection
from very loud sounds

1-inducing either slow increases or decreases in

the magnitude of the response of auditory
nerve fibers.
2-Since these fibers 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

Implicated in the production of the

otoacoustic emission + tinnitus

Note that efferent fibers terminate in the basal turn of the cochlea
Related to otoacoustic emission & tinnitus

The action of the outer hair cell in more details

The motile response of outer hair cells:
consist of a length change in the outer hair cell body.
part of the mechanism by which the outer hair cells amplify the travelling wave to increase the
cochlea's sensitivity and frequency selectivity
this response is nonlinear
The motile mechanism resides in the membrane, and is a property of the protein prestin which is
situated in the membrane
prestin changing its configuration as the electric potential across the membrane varies:
Cl- ions occupy a position in the intracellular domain of the membrane-bound prestin protein
Cl move along an inward electrical gradient away from the extracellular domain during
depolarization.
This charge movement produces, in turn, a conformational change in the protein that reduces
its surface area and shortens the cell;
the opposite action occurs during hyperpolarization, when Cl- ions move toward the
extracellular space within the domain of the protein (although the anion is never translocated to
the extracellular space), causing an increase in surface area and lengthening of the cell
depolarization: contraction
hyperpolarization: increase the length

so the process is dependent on prestin changing its configuration voltage generated by intracellular
electrolyte movement

Fluid filled Spaces of the inner ear

1. The perilymphatic space:

in which the membranous labyrinth is suspended
located within located scala tympani + scala vestibuli
contains the perilymph
a) similar to extracellular fluid+ CSF (rich in Na= 150/ k=7)
b) formed by infiltrate of blood
c) similar to the blood but different from CSF
d) Connected to the CSF via cochlear aqueduct
2. The endolymphatic spaces:
Located within the membranous labyrinth (scala media)
Separated from the perilymphatic space by reissner membrane & basilar membrane
Contains endolymph:
a. the only extracellular fluid that resemble intracellular fluid component (rich in
K=150/Na=16)
b. produced by perilymph infiltrate ( produced by sacula vascularis)
endocochlear potential in the sacla media is generated by sacula vascularis =+ve 80 mEqv
Note:
there is a negative intracellular potential, which is:
-45 m V for inner hair cells and -70 m V for outer hair cells
3. The cortilymphatic space:
lying within the organ of Corti
separated from the endolymphatic space by the reticular membrane
contains corticolyph:
a) similar to perilyph
b) derived from scala tympani via foramina in the osseous spiral lamina

Location
Source

Electrolyte
component

PH

Perilymph

Endolymph

corticolymph

Sacla tympani & vestibule

(Surrounding the
membranous labrynthin)
Blood infilterate

Sacal media

Corti organ

Cochlea:
marginal cells of the
stria vascularis
Vestibular system
except saccule:
Dark cells
Rich in K (similar to
intracellular)

derived from scala

tympani via foramina in
the osseous spiral
lamina

1-Rich in Na
2-Low in amino acids &
glycerin in comparasion to
blood & high in comparasion
to the CSF
3-Does not contain CL
7.2

Rich in Na

Rich in chlorid
7.5

Note:

Perilymph:
The site of perilymph production is still controversial.
It is unclear whether perilymph is derived as an ultrafiltrate of blood, from cerebrospinal
fluid (CSF) or from both.
CSF can reach the vestibule by means of:
1. vestibular aqueduct
2. perivascular channels
3. perineural channels.
There is more convincing evidence in favor of perilymph originating as an ultrafiltrate from blood.
1.
In terms of chemical composition, the amino acid content of perilymph, particularly glycine
and alanine, is low compared with blood but much higher than CSF.
2.
changes in blood composition are reflected much more rapidly in perilymph than in CSF.
Kellerhals has proposed a dual origin of perilymph. He found that the majority was derived from
blood and the remainder from CSF

Perilymph leaves the ear by drainage through:

1. venules
2. through the middle ear mucosa

Endolymph:
dark cells (of the cristae and maculae) & marginal cells of the stria vascularis have:
1. Ultrastructural morphologic properties of secretory cells:
deeply invaginated nuclei (indicating a cell actively engaged in production and
secretion);
many microvilli on their apical surface
large number of free ribosomes and many vesicles (again indicating active production
and packaging of secretory products);
deep infoldings on their basal surface containing many mitochondria(Presumably, the
mitochondria provide the fuel for the energy-consuming process that pumps the
endolymph out of the cell)
long, thin cytoplasmic extensions.
2. biochemical properties of secretory cells,they have a high concentration of enzymes associated
with active ion pumps and fluid transport:
Na+=K+=ATPase
adenylate cyclase
carbonic anhydrase
the "dark cells" of the cristae and maculae,which are separated from the neuroepithelium by a
transitional zone
absorption of endolymph :
site: endolymphatic sac
Morphologically:
the columnar cells located here are specialized for absorption. Like intestinal cells, they
have long microvilli on the luminal surface and contain many pinocytotic vesicles and
vacuoles

Energy transduction:

Transform the mechanical energy (sound wave pressure) into electrochemical Energy ( nerve
impulses)
The process of transduction occurs in the structures within scala media, sitting on the basilar
membrane (organ of Corti).

I.

Summary of the Pathway for sound

A.
Sound waves in the air enter the outer ear canal.

B.
C.
D.
E.
F.
G.
H.
I.
J.

Vibration of the tympanic membrane is transmitted through the chain of three (3) ossicles
(which amplify the sound 20 X) to the oval window.
Vibration of the oval window induces pressure waves in the perilymph of the scala
vestibuli.
Vibration of the vestibular (Reissner's) membrane transfers
pressure waves from perilymph to the endolymph of the scala
media (cochlear duct).
Pressure waves of the endolymph induce the basilar membrane
to vibrate.
Vibration of the basilar membrane (which is the critical step in
the transduction process) induces a shearing of the apical ends
of the hair cells against the tectorial membrane.
This will bends the bundle of stereocilia on each hair cell
resulting in the bending of the bundle of stereocilia and the
opening of mechanically gated ion channels in the membrane of the stereocilia
Influx of K+ ions, and depolarization of the hair cell depolarizing the hair cell, which
releases neurotransmitters.
This results in an action potential being propagated along nerve fibers of sensory
neurons in the spiral ganglion.

Vibration in the perilymph are transmitted to the base of the cochlea where it
displace the round window

INTENSITY RESOLUTION
Low Intensity sounds

When the organ of Corti moves in response to sounds of low intensity the cilia of the inner hair
cells do not contact the tectorial membrane.
The brain receives impulses from the afferent fibres of the outer hair cells
therefore, the brain will be aware that there is a sound but without perception of sound as
there are no impulses from the inner hair cells.
Signals are sent along the efferent nerve fibres to the outer hair cells, causing them to shorten
in length.
This pulls the tectorial membrane closer to the basilar membrane, allowing the cilia of the inner
hair cells to impact the tectorial membrane.
Deflection of the cilia leads to depolarisation and action potentials from the inner hair cells and
perception of sound.

High-intensity sounds

High intensity sound lead to more movement of the organ of Corti.

In order to protect the cilia, efferent impulses cause the outer hair cells to lengthen and push
the tectorial membrane away from the basilar membrane

The vestibular system:

main functions of the vestibular system:
1. It is the primary organ of equilibrium and thus plays a major role in the subjective sensation
of motion and spatial orientation.
2. Vestibular input to areas of the nervous system involved in motor control elicits adjustments
of muscle activity and body position to allow for upright posture.
3. Vestibular input to regions of the nervous system controlling eye movements helps stabilize
the eyes in space during head movements. This reduces the movement of the image of a fixed
object on the retina.

Reflexes used to Counter head motion with to maitain stable vision & posture are:
a) vestibulo-occular reflex: stabilize gaze despite head movement
b) vestibulocollic reflex: sabilize head despite body movement
c) vestibulo-spinal reflex: stabilize posture and fascilate gait

Basic elements needed for conversion of acceleration into useful signals for the nervous system are:
1. inertial mass
2. one or more sensory hair cells
3. the nerve fibers connected to the hair cells through synaptic junctions
Inertial mass:
For saccule + utricle: otolith
For semicircular canals: endolymph cupula

Components of the vestibular system

This comprises the 3 semicircular canals and the saccule and utricle= membranous labrynithin
except the cochlear duct.
The semicircular canals are concerned with the rotation and the saccule and utricle with
gravitation
Semicircular Canals (SSC):
senses rotational (angular) acceleration
Note; constant-velocity rotation will not stimulate SCC
Orientation of the semicircular canals
horizontal canals and the utricle:
lie parallel to the line between the EUC and the outer canthus of the eye, which is inclined 30
degrees above the horizontal axial plane (anterodorsally relative to the naso-occipital plane )
so in the anatomical position the plane of the lateral semicircular canal is at angle of 30 to the
horizontal tilting downward Posteriorly
This orientation causes the plane of the
horizontal canal and utricle to be parallel
with the earth horizontal and
perpendicular to gravity.

Caloric tests:
Test the function of the lateral SCC
Fitzgerald-Hallpike test: the pt lies supine with head tilted 30 forward so the
lateral SSC assumes a vertical position ( the most sensitive to a thermal gradient)
Kobrak test: the pt sits with head tilted backward 60 degree

The anterior and posterior semicircular canals and the saccule are arranged vertically in the head,
orthogonal to the horizontal semicircular canal and utricle

Vertical/Superior canals:

When looking down at the top of the head, the anterior canal is oriented at approximately 45
degrees off midsagittal and 45 degrees anterior to the intraaural line.
Located at right angles to:
a) the horizontal canals
b) to each other.
Posterior canal:
aligned roughly 45 degrees behind the intraaural line

The two vertical canals in each ear are positioned orthogonal to each
other
whereas the plane of the anterior canal on one side of the head is
coplanar with the plane of the contralateral posterior canal

This orthogonal pattern creates functional pairs in that increased output from one canal results in
decreased output from its paired canal thus creating a push-pull mechanism that enhances the
sensitivity of the system

 Each end of the canals opens into the utricle but there are only 5 openings as the anterior and
posterior canals unite Posteriorly.
Ampulla: pear-shaped expansion of the membranous labyrinth located at one end of each SSC
near the vestibular opening (where the semicircular canal meets the utricle

The location of the ampullated end of the semicircular canals

1. lateral semicircular canal : at the anterior end of the canal

2. superior semicircular canal is also anterior
3. posterior semicircular canal is lateral.

Location of nonampullated end of semicircular canals:

1. lateral semicircular canal: enters the vestibule posterolaterally.
2. The nonampullated ends of the posterior and superior semicircular canals join to form
the common crus and enter the vestibule posteromedially

Within each ampulla there is the sensory organ : the crista ampullaris.

Crista ampullaris is a saddle shaped neuroepithelium end organ

Crista ampullaris has 2 types of hair cells :

1. Type I hair cells:
Flask shaped (rounded base and narrow neck)
The body is entirely engulfed by one afferent
terminal.
Efferent innervation is indirect, as the efferent
nerve has its synapse on the afferent nerve ending
2. Type II hair cells:
Cylindrical in shape
have one or more afferent nerve endings on the
body of the cell.
Type II hair cells can also be directly or indirectly
innervated by vestibular efferent terminals

Both types of hair cells apex contain:

1- stereocilia
2- 2- kinocilium

Kinocilium : A true (but non-motile) cilium, taller and larger in X-S than the stereocilia, having an
array of 9 + 2 microtubules in its core
Stereocilia are arranged by height relative to kinocilium in a step-like manner (Tallest clostest,
shortest farthest, from kinocilium)
there are both afferent and efferent nerve fibres to each hair cell.

the hair cells and their sup porting cells lie embedded in a saddle-shaped neuroepithelial ridge, the
crista, which extends across the base of the ampulla
Type I hair cells are concentrated in central regions of the crista, Type II hair cells are concentrated
in peripheral areas.
Arising from the crista and completely enveloping the stereocilia of the hair cells is a gelatinous
structure, the cupula.
The cupula attaches to the roof and walls of the ampulla, forming a fluid-tight partition that has the
same specific density as that of endolymph.

Afferent fibers of the vestibular system:

The efferent fibers to the vestibular end organs arises from:
a small group of about 200 neurons
lateral to the abducens nucleus and the genu of the facial nerve, the so-called group e
These neurons project both ipsilaterally and contralaterally
The contralateral pathway crosses the midline at the level of the facial genu and joins the
ipsilateral pathway
both pass ventral to the vestibular nucleus.
At this point, they are joined by cochlear efferents originating from the superior olivary complex
(the olivocochlear bundle).
All of the efferents then enter the vestibular nerve, coursing through the middle of the nerve in
a small distinct bundle.
At the end organs, these relatively few fibers branch profusely to innervate the entire sensory
epithelium.
the ipsilaterally projecting efferents supply the central regions of the crista, while the
contralaterally projecting efferents supply the peripheral zone.
The efferent fibers terminate as highly vesiculated boutons, making synaptic contacts with hair
cells and afferent fibers

Note that the vestibular nucleus

in the brain is made of:
1. Superior vesibular
nucleus
2. Descending V.N
3. LATERAL
4. MEDIAL
There is similarity between hair cell innervations in the auditory system and the vestibular system
Auditory system: inner & outer hair cell
Vestibular system: Type I & Type II hair cells
Inner hair cell/type 1

Outer hair cell/type II

Shape

Flask shape

slender

Afferent

Completely surround the hair cell

Efferent

Contact the hair cell indirectly through the synapse with

afferent/its extension
Note the # of afferent vestibular fibers:18000
# of efferent vestibular fibers: 200

Contact the hair cell directly

There is postganglionic sympathetic efferent innervations of the vestibular end organ of unknown
function (originates from the superior cervical ganglion)

Physiology:

Any change in rotation speed of the head results in motion in the endolymph.
This causes the cupola to move and brings about a shearing movement of the cilia.
Direction of bending of the stereocilia, relative to kinocilium, is significant:
bending away from kinocilia hyperpolarization of receptor cell (inhibition)
bending toward kinocilia depolarization of receptor cell & thus generation of an
action potential (impulse) in the vestibular branch.
Because of the orientation of the semicircular canals rotation will produce an increase in firing
rate in the vestibular nerve on one side of the head and a reduction on the other side.

Horizontal (Lateral) SSC:

30 from horizontal
Kinocilia is located on the utricle side
So displacement of the steriocilia to ward the
kinocilium by ampullopetal (toward vestibule) flow
of endolymph increases vestibular neuron firing
rate
Vertical (Superior and Posterior) SSC:
both canals share a nonampullated common crus,
kinocilia is located on the semicircular canal side
ampullofugal (away from vestibule) flow of
endolymph increase vestibular neuron firing rate

When the head is stationary (no angular acceleration), the endolymph and the cupula remain still,
and the afferents from the two horizontal semicircular canals fire at the same (resting) rate 90.

When the head turns to the right or left, the horizontal semicircular ducts turn with it, but the
endolymph lags owing to inertial forces and the viscous drag between the fluid and the duct wall.
The lagging endolymph deflects the cupula, which in turn deflects the stereocilia of the hair cells.
leftward turn of the head causes the stereocilia in the left horizontal canal ampulla to be deflected
toward their kinocilia, resulting in an increase in the discharge rate of the eighth nerve afferents on
the left side.
Simultaneously, the hair cells in the right horizontal canal ampulla are hyperpolarized, so their
afferents show a decreased rate of firing.
Scarpas Ganglion: bipolar vestibular nerve cell bodies located in internal acoustic meatus

Plane of semicircular canal stimulation:

1. Horizontal scc: head turning in horizontal plane 45 degree off the midsagittal plane
2. Posterior SCC: nose is pitched upward & turning 45 degree off the midsagittal plane
3. Superior SCC: nose is pitched down & 45 degree off the midsagittal plane

Clinical application:

each canal has a resting basal discharge rate,and by modulating the nonezero baseline firing of
vestibular afferent fibers,the semicircular canal encode rotation of the head so a lesion of the
eighth nerve, such as that produced by a glomus tumor or acoustic neuroma may reduce the
frequency of impulses in the ipsilateral afferent fibers or block their impulse transmission entirely.
The comparator units of the vestibular nuclei will then consistently receive a higher impulse
frequency from the intact side, which will be interpreted as a head turn away from the side of the
lesion
The semicircular ducts are particularly concerned with reflex control of visual movements through
the vestibulo-ocular reflex at the brainstem level to allow the individual to maintain optic fixation
in the presence of movement(the head moves in one direction the eyes are moved in the opposite
direction so they can remain focused on the same point)

The jerky eye movements caused by rotation of the head (rotatory nystagmus) are dependent on
the defections of the cupulae. The slow component of the nystagmus is always in the direction of
the cupula deflection
Nystagmus:
The nystagmus is named after the fast component of the nystagmus
Nystagmus components:
1. Slow component: vestibular in origin toward the defective lesion
2. Fast component: neural in origin
In caloric test:
COWS:
1. Cold: toward the opposite ear
2. warm: toward the same ear, it causes ampullopetal/uriticopetal flow
cold water makes the labyrinthine hypoactive=labrynthiectomy
vestibular neuritis has the same effect of warm water on labyrinthin

THE UTRICLE AND SACCULE:

The utricle and saccule both contain endolymph.

Saccule is smaller than the utricle,& lies in a depression below & infront of the utricle
Within each structure is a gravitational sense organ: the macula (otolith organ).
The hair cell stereocilia of otolith organs extend into a gelatinous coating called the otolith
membrane, which is covered by calcium carbonate crystals called otoconia (ear stones).
Otoconia:
3x times as dense as the surrounding endolymph
not displaced by normal endolymph movements.
Instead, changes in head position relative to gravity, or linear accelerations (forward-backward,
upward-downward) produce displacements of the otoconia, resulting in bending of the
underlying hair cell stereocilia
Any linear motion displaces this membrane in the opposite direction.
This again produces deflection of the cilia and depolarization of the hair cells.

The macula is situated in:

Utricle: on the floor: respond to horizontal, side-to-side motion-axial plane-inter aural
saccule : on to the medial wall: responds to movement in the vertical plane.-sup inferior
The hair cells within these structures are orientated with reference to a central plane ( the striola).
In the utricle the cilia are arranged so that the kinocillium is towards the striola whereas in the
saccule the kinocilia are arranged away from the striola.see p 100 fig4-4
There is again both an afferent and an efferent nerve supply to the hair cells of the maculae.

Otolith organs alone are unable to distinguish linear acceleration from vertical acceleration

So the brain uses other clues such:

1. Rotational information from SCC
2. Visual input

Summary of the Sensory Receptors of the Membranous Labyrinth:

6 regions of sensory receptors project from the wall of the membranous labyrinth:
a) 3 cristae ampullaris within the ampullae of the semicircular ducts
sensitive to angular acceleration (turning of the head)
b) 2 maculae (macula utriculi & macula sacculus)
sensitive to position relative to vertical (gravity); linear acceleration
c) the spiral organ (of Corti) of the cochlear duct
sensitive to sound receptor
All of these sensory receptors share similar structural specializations and characteristics, including hair
cells that:

are non-neuronal, mechanoereceptor, epithelial cells

possess numerous stereocilia (modified microvilli) called sensory hairs
are associated with both afferent and efferent nerve endings
are all transducers, i.e., they convert mechanical energy (bending of stereocilia) into electrical
energy transmitted by the vestibulocochlear nerve to the brain
the means by which bending or flexing occurs varies for different receptors

stretching of the plasma membrane caused by bending generates transmembrane potential

changes in the receptor cell that are conveyed to the afferent nerve endings associated with the
cell

Summary of the plane of stimulation

The max sensitivity to motion for the vestibular end organs occur along an axis oriented
1. Perpendicular to plane of the SCC
2. Parallel to the plate of otolith organ
Movements that stimulate semicircular canal:
Angular rotation
1. Horizontal scc: head turning in horizontal plane 45 degree off the midsagittal plane
2. Posterior SCC: nose is pitched upward & turning 45 degree off the midsagittal plane
3. Superior SCC: nose is pitched down & 45 degree off the midsagittal plane
Movements that stimulate the otolith organ:
Linear acceleration:
1. Utricle: horizontal ( ant-pos, Rt-Lt)
2. Saccule: vertical ( change in head position in relation to gravity) tilting the head

Vestibular nuclui

The vestibular nuclei is divided into 4 subdivisions:

1.
2.
3.
4.

Superior vestibular nuclei

Medial vestibular nuclei
Lateral vestibular nuclei:subdivided into dorsal & ventral
inferior vestibular nuclei

The superior vestibular nucleus :

located dorsally and rostrally in the vestibular complex.

involved in VOR pathways.
Neurons in this nucleus fire in relation to eye movements as well as head movements
a prominent efferent projection of this nucleus is to the oculomotor nucleus via the medial
longitudinal fasciculus.

The lateral vestibular nucleus:

subdivided into two subnuclei:
a) dorsal lateral vestibular nucleus (or Deiters' nucleus)
b) ventral lateral vestibular nucleus.
dorsal lateral vestibular give rise to:
lateral vestibulospinal tract
do not give efferent fibers to the cerebellum & brainstem
ventral lateral vestibular nucleus give rise to:
vestibuloocular pathways
medial vestibulospinal tract
vestibulothalamic pathways.
The medial vestibular nucleus :
extends rostrocaudally in the brainstem from the level of the abducens nucleus to the hypoglossal
nucleus. It is bounded medially by a functionally related nucleus and the nucleus prepositus hypoglossi,
and laterally by the inferior vestibular nucleus. Recent studies indicate that the medial vestibular
nucleus can be subdivided into a rostral magnocellular region (MVmc) and a parvocellular region (MVpc)
and a caudal region (MVc) with small- to medium-sized neurons.[23][33] The rostral medial vestibular
nucleus, like the superior nucleus, contains many neurons that project to the oculomotor nuclei and

whose firing behavior is related to eye movements.[60][68][69] Little is known about the functions of the
MVc, although many of the cells in this region appear to project to the cerebellum. [24] The medial
vestibular nucleus also gives rise to the medial vestibulospinal tract, which descends in the medial
longitudinal fasciculus to terminate on interneurons in the cervical spinal cord and ascends to terminate
in the eye motor nuclei. This tract is particularly important for cervicovestibuloocular reflexes.
The inferior (or descending) vestibular nucleus

one of the primary recipients of vestibular afferents that innervate the otolithic organs.
major projections from this area are to the cerebellum and reticular formation.
Efferent fibers from the vestibular nucleus

 Reflexes are:
1. vestibulo-occular reflex:
Rotates the eye in the same speed but in the opposite direction of the head rotation so

it stabilize the image on the fovea despite head movement

vestibular nerve: Ipsilateral medial vestibular nuclui (medulla) : stimulatory impulse into
contra-lateral abducent nerve(pons):
1. lateral rectus
2. indirectly via medial longtidudinal fasciculus into the Ipsilateral oculomotor
nucleus in mid brain
loss of vestibuloccular reflex in comotoze patient post head trauma indicates brain
stem injury

2. vestibulo-spinal reflex:
stabilize posture and fascilate gait

Head position
control
Semicircular canals

Body position
control
Otolith organs

Vestibular
nuculeus

Medial

Dorso-Lateral

Tract

Medial longtidunal
fascicular tract
Medial
vestibulospinal tract
Medial part of the
ventral horn

Lateral
vestibulospinal tract

Neck & axial muscles

Ipsilateral proximal
limb muscles

Stimulation

Horn

Muscles

Ant horn

Classes of eye movement:

The aim of eye movement is to bring the image into the fovea:
If the head is steady & object steady:
visual fixation :
stationary object
vergene angle:
nearer stationary object
If the head is steady & the object is moving
smooth pursuit:
saccule

slowly moving object

rapidly moving object

If the object is steady & the head is moving:

vestibular : rapid,brief,high frequency head rotation
optokinatic: slow,low frequency head rotation

Note: that optokinetic reflex depends on the visual integrity

other reflexes do not depend on visual input & can work in complete darkness