SPECIAL SENSES PHYSIOLOGY OF HEARING AND BALANCE
Initially, the auricle/pinna collects sound waves from the
AY 2019 -2020 Dr. Calaquian
Module 8 02/19/2020
SOUND
 Vibrations produced by waves of compression and
decompression transmitted in air
 Being waves, they have the properties of frequency and
amplitude
o Frequency: how quickly the wave occurs
 Unit: Hertz (Hz)
 Associated with pitch, how high or low the
sound
o Amplitude: how strong the waves are
 Related to sound pressure, loudness, how loud
or soft is a certain sound
 Unit: Decibel (dB)
 Travel of vibration through the ear and its corresponding
structures: a) conduction – external and middle ears; b)
transduction – internal ear; c) neural structures – auditory
nerve, brainstem, subcortical & cortical auditory structures
 20-20,000 Hz normal young human ears are sensitive to
this range of frequencies
 65 dB (300-3500 Hz) normal speech
 >100 dB can damage peripheral auditory apparatus
 >120 dB can cause pain and permanent damage
 Presbycusis reduced ability to hear high frequency tones
(common in elderlies)
PATHWAY OF SOUND
The ear converts sound waves in the external environment
into action potentials in the auditory nerves.
1. Pinna
2. Auditory Canal
3. Tympanic Membrane
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environment
Then, sound waves will enter the eardrum via the external
auditory meatus and will strike on the tympanic
membrane causing it to vibrate.
Next, vibration will be transmitted to the inner ear via the
air- filled middle ear cavity by the three auditory ossicles,
namely: malleus (hammer), incus (anvil), and stapes
(stirrups)
After that, the malleus transmits its vibration to the incus.
The incus articulates with the stapes which sits on the
membrane of the foramen ovale (oval window) (vibration
from malleus  incus  stapes)
Note: oval window is located at lateral wall of vestibule
Meanwhile, vibration is transmitted to the perilymph of the
scala vestibule, which travels along the spiral of the
cochlea to a point corresponding to the frequency, crosses
the scala media, then returns along the scala tympani to
the round window.
Afterwards, Hair cells of the Organ of Corti is stimulated
which lie on the vibrating basilar membrane
Eventually, vibration of the Organ of Corti induces a
generator potential which exactly duplicates the sound
vibration (sound  nerve impulse)
Finally, excess pressure waves are dissipated at the area
of the secondary tympanic membrane which covers the
round window.
REVIEW OF FUNCTIONAL ANATOMY OF THE EAR
A. EXTERNAL EAR
 Collects sound, directs it into auditory canal
 Pinna Shadow effect
 Sounds coming from the front reach the ear canal
almost directly, whereas sounds coming from behind
are obstructed by the pinna and therefore attenuated
before they reach the ear canal. ... Sounds from
behind are attenuated by 3-4 dB relative to sounds
coming from the front.
 Important in localization of sound
o Unique shape of pinna helps the brain know where the
sound is coming from
o Sound hits the pinna in a particular way and it changes
the sound depending on the location
 Conduction of sound waves into the tympanic membrane
 Resonance frequency (vibrates maximally) 2-3 kHz-per Doc
Calaquian
3500 Hz- Berne and Levy
 The surface area is about 55 mm2
 Creates harmonic overtones to hear clearly or detect a
particular sound
o E.g. Talking to your friend in a noisy room, even
though it’s noisy, you can still hear your friend clearly
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B. MIDDLE EAR
 Conducts and amplifies sound
 Transmits vibrations of the tympanic membrane across
the cavity of the middle ear to the internal ear
1. Auditory Ossicles
 Malleus (Hammer): attached to the TM, producing the
umbo of the tympanic membrane
 Incus (Anvil)
 Stapes (Stirrup)
 Purpose: Impedance Matching
IMPEDANCE MATCHING
 The middle ear acts as an impedance matching device to
transfer sound energy efficiently from air to a fluid medium
in the cochlea
**Because fluid has far greater inertia than air does. Increase amount
of force are necessary to cause vibration in the fluid and overcome
mismatched impedance. Therefore, the tympanic membrane and the
ossicular system provides impedance matching, between the sound
waves in air and the sound vibration in the fluid of cochlea.
 The system actually increases the force movement by
several mechanisms: Lever mechanism & Attenuation
reflex
LEVER MECHANISM
 The difference in the amplitude of movement of the stapes
faceplate and the handle of malleus leads to an increase of
force movement and reduced the distance.
 Unique shape of the ossicle system (stapes & malleus) acts
as a lever system to amplify the force 1.3 times
o 17 fold size difference x 1.3 fold force due to lever
system = 22 times
2. Muscles
 Ossicles are connected to two muscles:
o Tensor tympani – tenses the tympanic membrane;
innervated by CN V (Trigeminal nerve)
o Stapedius – connected to stapes; when it contracts, it
pulls the stapes away from the cochlea; innervated by
CN VII (Facial nerve)
**the function of these two muscle is to dampen or decrease vibration
from the tympanic membrane
 Perform Tympanic Reflex or Attenuation Reflex
ATTENUATION REFLEX
 Mediated by the opposing forces of the tensor tympani and
stapedius muscle
o Tensor tympani: pulls the handle of malleus inward
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o Stapedius: pulls the stapes outward
 Increases the rigidity of the ossicular system, thereby
decreasing their ability to do impedance matching
 The reflex can reduce intensity of low—frequency sound
transmission by 30—40dB
 Function:
o Protects the cochlea from damaging vibrations
caused by excessively loud sound
o Mask low – frequency sounds in loud environments
 Removes major share of the background
noise
 Allows a person to concentrate on sounds
above 1000 cycle/sec
o Decrease a person’s hearing sensitivity to his or her
own speech
3. Eustachian Tube
 Equalize the pressure between external and middle ear
**99.9% of vibratory energy does not pass from middle ear to
inner ear because of the mismatch between air and fluid thus
vibrations are reflected back to external ear
C. INNER EAR
 Transforms mechanical vibrations into electrical impulses
 Important structures: vestibule, cochlea (responsible for
hearing), semicircular canal (responsible for balance)
Cochlea
 A system of coiled tubes; Fluid-filled
Figure 1. A section through the cochlea
 Reissner’s membrane: separates vestibuli and media
 Basilar membrane: separates tympani and media
 Oval window: found within scala vestibuli; covered by stapes
 Round window: a free membrane
 Endolymph: within scala media; produced by stria vascularis
 Perilymph: within scala vestibuli and scala tympani; similar with
cerebrospinal fluid (CSF)
 Basic concept: stapes moves→ produces fluid wave→
wave travels to cochlea → stimulates Organ of Corti on the
basilar membrane
  Round window is a flexible membrane which bulges out
when the stapes pushes → makes fluid movement possible
due to its flexibility
TRANSDUCTION OF SOUND
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 o Internal hair cells: single row
o External hair cells: 3 to 4 rows
o These hair cells synapse with a network of cochlear
nerve endings

Figure 2. Sound Pathway: pinna auditory canal  tympanic

membrane  ossicles cochlea  auditory nerve

 Sound vibrations enter the scala vestibuli from the

faceplate of the stapes at the oval window. Figure 4. Organ of Corti
 The faceplate covers this window and is connected with the  The spiral ganglion neuronal cells send axons—a total of
window’s edges by a loose annular ligament so that it can about 30,000—into the cochlear nerve and then into the
move inward and outward with the sound vibrations. central nervous system at the level of the upper medulla
 Inward movement causes the fluid to move forward in the  Hair cell apex bathed in endolymph, hair cell base
scala vestibuli and scala media, and outward movement bathed in perilymph
causes the fluid to move backward
**When there is vibration in your organ of corti, the basilar fiber will
move upward and downward while the reticular lamina will move
sideways stimulating your hair cells. Once stimulated your hair cells
will send signal to your spiral ganglion to your cochlear nerve going to
your brain for processes.

EXCITATION OF HAIR CELLS

Figure 3. Movement of fluid in the cochlea after forward thrust of the
stapes
Basilar Membrane
 Fibrous membrane that separates the scala media and
scala tympani
 Contains 20,000-30,000 basilar fibers
→ Stiff, elastic, reedlike structures
**Different frequencies are located in differently in cochlea. Low
frequency is located at the apex (with long and thin fibers). High
frequency at the base (with shorter and thicker fibers). Intermediate
frequency is in between the area of base and apex
 Stereocilia projects upward from the hair cells and either
touch or embed in the surface gel coating of the tectorial
membrane
 Bending of the hair cells towards the kinocilium
depolarizes it
 Once their movement is going towards your kinocilium, tip links
would open up your potassium channel, there would be influx of
potassium inside your hair cells. The increase concentration of
potassium would then trigger the exchange of your potassium to
your calcium because the calcium is the one going to be secreted
by your hair cells going to your synapse to send action potential to
your afferent axon to your spiral ganglion to cochlear nerve to the
brain for processing.
 Bending of the hair cells away from the kinocilium
hyperpolarizes it
 These in turn excites the auditory nerve fibers synapsing
with their bases

**The taller one is the kinocilium beside it is your stereocilia and the
small spiral kinks on top is your tip links which is important because it
is connected to your mechanical-gated potassium channels. The
bending will produce the action potential.

Organ of Corti
 Turns the vibration of the basilar membrane into nerve
impulses by stimulating receptors
 Actual sensory receptors are the hair cells

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BALANCE
 VESTIBULAR SYSTEM
 Detects angular and linear acceleration
o Acceleration = Change in velocity/time
o Angular motion: turning along an angle (e.g. circular,
turning)
o Linear motion: moving along a line (e.g., upward and
downward, forward and backward)
 Concerned primarily with the position of the head in space
 Triggers head and eye movements to stabilize visual
image, maintain balance
 VESTIBULAR APPARATUS
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 The sensory organ for detecting sensations of
equilibrium.
 It is encased in a system of bony tubes and chambers
located in the petrous portion of the temporal bone, called
the bony labyrinth. Within this system are membranous
tubes and chambers called the membranous labyrinth.
 The membranous labyrinth is the functional part of the
vestibular apparatus.
o Membranous – endolymph fluid
o Bony – perilymph fluid
 Functions:
o Controls eye muscles, despite changes in head
position, eyes remain fixed on same point
o Maintain upright position
o Providing consciousness of:
 Position of acceleration area
 Perception of surrounding space
Vestibulo-Ocular Reflex
-operate to maintain the gaze on a selected target.
e.g. Our ability to gaze at the period (.) at the end of a
sentence, while moving the head about.
 Sudden rotation of the head
o signals from the semicircular ducts cause the eyes to
rotate in a direction equal and opposite to the rotation
of the head
o reflexes transmitted through the vestibular nuclei and
the medial longitudinal fasciculus to the oculomotor
nuclei
C. PARTS
Figure 5. Sensory organ for detecting sensation of equilibrium
Three Semicircular Canals
 Joined by ampulla
 Cristae Ampullaris end organs- signals head movement
 Angular acceleration
 Contain endolymph and perilymph
 Superior – detecting movements in the same plane in
conjunction with the posterior semicircular canal
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  Posterior – detecting movements in the same plane in  Reason why it detects upward and downward
conjunction with the superior semicircular canal motions
 Lateral/Horizontal – not really “horizontal”, angled up at 30 o Cilia oriented differently
degrees  Semicircular canals - cilia oriented on the
same direction
Two Otolithic Organs  Utricle/Saccule - cilia face on a bony ridge
 Joined by ductus reuniens → Endolymphatic duct and sac called striola
 Linear acceleration
 Utricle – horizontal acceleration
 Saccule – vertical acceleration

Bony Labyrinth Vestibule Semicircular Canal

Membranous Utricle and Saccule Semicircular Duct
Labyrinth
Specialized Macula Cristae ampullaris
Sensory Structure
Function Linear acceleration Rotational/angular
acceleration
D. DETECTION OF ACCELERATION
Cristae ampullaris
 Detects angular acceleration
 Vestibular hair cells embedded in the cupula
 Cupula
o Gelatinous matrix
o Same specific gravity as endolymph; not moved by
gravity / linear acceleration
o Head rotates → endolymph moves on the opposite
direction as a result of inertia → cupula is deflected
o Bending opens ion channels → local potential → action
potential
o Bends towards the tallest cilium = depolarized Linear Acceleration
o Bends towards the shortest cilium = hyperpolarized  The utricle responds to horizontal acceleration and the
saccule to vertical acceleration
 Otoliths (aka statoconia) are more dense than the
endolymph (2-3x specific gravity)
o acceleration in any direction causes them to be
displaced in the opposite direction, distorting the hair
cell processes and generating activity in the nerve
fibers
 macula also discharge tonically in the absence of head
movement
o Due to pull of gravity on the otoliths
o Because of their greater density, otoliths are displaced
when there is a change in position of the head. This
displacement is transferred to the underlying hair cells
via the gelatinous otolithic membrane.

Angular Acceleration
 Rotational acceleration displaces endolymph in a direction
opposite to the direction of rotation.
 The fluid pushes on the cupula → deforms and bends the
processes of the hair cells
 When constant speed of rotation is reached
o the fluid spins at the same rate as the body and the
cupula swings back into the upright position
 When rotation is stopped,
o deceleration produces displacement of the
Macula Utriculi/Sacculi endolymph in the direction of the rotation
 Detects linear acceleration
 Pouch / cul de sac in utricle / saccule VESTIBULAR PATHWAY
 Also has vestibular hair cells, with stereocilia embedded in
gelatinous mass  there are 4 vestibular nuclei on each side of the
 Differences: brainstem: superior, medial, lateral and inferior (also called
o Gelatinous mass has otoliths on top (CaCO3 crystals) descending)
= otolithic membrane  they all receive input from both the semicircular
 Affected by gravity canals and the otolith organs
 they all project to all pathways

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4 important vestibular pathways to consider:
1. the primary sensory pathway from the vestibular
nuclei (particularly the superior and lateral) to the VP
nucleus of the thalamus and then to cortex
there is debate over exactly where the vestibular cortex is:
some people put it close to auditory cortex in the superior
temporal gyrus, others at the end of somatosensory cortex
where it joins with motor cortex

2.vestibulospinal reflexes

3. the vestibulo-ocular reflex (VOR)

4. vestibulo-cerebellar connections

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