VESTIBULAR SYSTEM

BNS 2
Long Exam 4

OUTLINE • Perilymph – similar to CSF; high sodium content (150 mM) and
I. Overview D. Other Afferent low potassium content (7 mM); bathes the vestibular portion of
II. Peripheral Vestibular Connections CN VIII
Labyrinth E. Other Efferent • Endolymph – covers the specialized sensory receptors of both
A. Vestibular Recepetor Connections the vestibular and auditory systems; high concentration of
Organs VI. Vestibuloocular Network potassium (150 mM) and a low concentration of sodium (16 mM)
B. Membranous Labyrinth A. Rotational • Important to note differences of the fluids -> disturbance often
C. Meniere Disease Vestibuloocular Reflex lead to vestibular disease
D. Semicircular Canal B. Linear Vestibuloocular
Dehiscence Reflex A. VESTIBULAR RECEPTOR ORGANS
III. Vestibular Sensory C. Nystagmus • Complement each other in function
Receptors VII. Vestibulospinal Network • Semicircular canals (Horizontal, Anterior, Posterior) – transduce
A. Hair Cell Morphology A. Lateral Vestibulospinal rotational head movements/angular accelerations
B. Hair Cell Transduction Tract • Otolith organs (Utricle and Saccule) – translational head
IV. Semicircular Canals and B. Medial Vestibulospinal movements/linear accelerations or to the orientation of the head
Otolith Organs Tract relative to gravity
A. Function of VIII. Vestibulothalamocortical
Semicircular Canals Network
B. Function of Otolith A. Vestibular Thalamus
Organs B. Vestibular Cortex
V. Vesitbular Nuclei C. Navigation
A. Vestibular Afferent IX. Dizziness and Vertigo
Inputs A. Benign Paroxysmal
B. Cerebellar Connections Positional Vertigo
C. Commissural B. Vestibular Neuritis
Connections

I. OVERVIEW
• Vestibular System
o Mediates motor activities (posture, movement) through a
network of receptord and neural elements
o Integrates peripheral sensory information from vestibular,
somatosensory, visceromotor and visual receptors as well as
motor info from cerebellum and cerebral cortex
o Provide an appropriate signal to coordinate relevant movement
reflexes
o Mostly conducted at the subconscious level
• Five Components
o Peripheral Receptor Apparatus – resides in the inner ear;
responsible for transducing head motion and position into
neural information
o Central Vestibular Nuclei – comprises a set of neurons in the
brainstem; responsible for receiving, integrating and
distributing information that controls motor activities (eye and
head movements, posture) and spatial orientation
o Vestibuloocular Network – vestibular nuclei; control of eye
movements
o Vestibulospinal Network – coordinates head movements, axial
musculature, postural reflexes
o Vestibulothalamocortical Network – conscious perception of
movement and spatial orientation
II. PERIPHERAL VESTIBULAR LABYRINTH
• Contains specialized sensory receptors and is located lateral and
posterior to the cochlea
• Consists of five receptor structures, three semicircular canals
and two otolith organs, which are contained in the petrous part
of the temporal bone
o Bony Labyrinth – surrounding shell that contains and protects
the sensitive underlying vestibular sensory structures
o Membranous Labyrinth – closed, fluid-filled system;
connecting tubes and prominences
Figure 1. Orientation of the vestibular receptors. In the lateral view (A), the
horizontal semicircular canal and the utricle lie in a plane that is tilted
relative to the nasooccipital plane. In the axial view (B), the vertical
semicircular canals lie at right angles to each other
• When a person walks -> head tilted downwards -> line of sight
directed downwards -> horizontal canal and utricle parallel with
the earth and perpendicular with gravity
• Anterior and posterior semicircular canals and the saccule are
arranged vertically in the head, orthogonal to the horizontal
semicircular canal and utricle
• 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
• Innervation: primary afferent fibers that join those from cochlea
to form the vestibulocochlear cranial nerve/ CN8
o Cell bodies of these vestibular afferent neurons are in the
vestibular ganglion (Scarpa ganglion), which lies in the internal
acoustic meatus
o Central processes enter the brainstem and terminate in the
ipsilateral vestibular nuclei and cerebellum 

• Blood Supply
o Labyrinthine Artery – supplies the labyrinth (branch of AICA);
enters via the internal auditory meatus
o Stylomastoid Artery – also provides to the labyrinth, mainly
semicircular canals
o Interruption of blood supply to the labyrinth will compromise
vestibular and cochlear function -> labyrinth associated
symptoms, such as vertigo or oscillopsia, and clinical signs
such as nystagmus or unstable gait

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Vestibular System
BNS 2 Long Exam 4

B. MEMBRANOUS LABYRINTH o Stereocilia arranged in ascending height, with the tallest lying
• Supported inside the bony labyrinth by connective tissue next to the kinocilium; arise from a region of dense actin, the
• Receptors in the utricle are oriented longitudinally along its cuticular plate, located at the apical end of the hair cell
base o Type I hair cell
• Receptors in the saccule are oriented vertically along the ¨ Chalice shaped
medial wall ¨ Surrounded by an afferent terminal that forms a nerve calyx
• 3 ducts of semicircular canals connect to the utricle, and each ¨ Concentrated in the regions of the crista (saddle shaped
duct ends with the ampulla

 neuroepithelial ridge)
• Endolymph is drained into the endolymphatic sinus via small o Type II hair cell
ducts

 ¨ Cylindrical
• Endolymphatic sinus communicates with the endolymphatic sac ¨ Innervated by a simple synaptic boutons
through the endolymphatic duct 
 ¨ More numerous in the peripheral of the crista
o Aspartate & Glutamate - NTs at the receptor cell-afferent fiber
• Saccule is connected to the cochlea via ductus reuniens
synapses
o Both hair cells receive synapses from the vestibular efferent
fibers
o Acetylcholine & Calcitonin gene related peptide - NTs in the
efferent cell
o Efferent cell bodies are located in the brainstem just rostral to
the vestibular nuclei and lateral to the abducens nucleus.
Activated by behaviorally arousing stimuli or by trigeminal
stimulation

Figure 2. Membranous labyrinth

C. MENIERE DISEASE
• Disruption of normal endolymph volume, resulting in
endolymphatic hydrops (abnormal distension of the membranous
labyrinth)
• Symptoms
o Vertigo
o Positional Nystagmus
o Nausea
Figure 3. Type I (Left) and Type II (Right) Hair cells
o Unpredictable attacks (audi and vesti) including vomiting,
• Cupula
tinnitus, complete inability to make head movements or even
o attaches to the roof and walls of the ampulla, forming a fluid-
stand passively
tight partition that has the same specific density as that of
• First course of treatment: diuretic (hydrochlorothiazide) and salt-
endolymph
restricted diet
o cupular movements displace the stereocilia (and kinocilium) of
• Second course of treatment: implantation of a small shunt into the the hair cells in the same direction
abnormally swollen endolymphatic sac, delivery of a
• Macula
vestibulotoxic agent such as gentamicin into the perilymph
o Structure analogous to crista
D. SEMICIRCULAR CANAL DEHISCENCE o Contains receptor hair cells
• Portion of the temporal bone overlying the anterior or the • Hair cell stereocilia of otolith organs extend into a gelatinous
posterior semicircular canal thins so much that it dehisces next to coating called the otolith membrane, which is covered by
the dura ® exposes the normally closed bony labyrinth to the calcium carbonate crystals called otoconia
extradural space o Otoconia
• Symptoms ¨ 3x as dense as endolymph
o Vertigo ¨ Not displaced by normal endolymph movements. Instead,
o Oscillopsia changes in head position relative to gravity or linear
o Tullio Phenomenon - precipitation of vertigo and nystagmus by accelerations (forward-backward, upward-downward)
a loud noise. produce displacements
• Eye movements evoked by these stimuli align with the plane of
dehiscent superior canal
• Surgical closure by bone replacement often performed

III. VESTIBULAR SENSORY RECEPTORS

A. HAIR CELL MORPHOLOGY

• Hair cell
o Sensory receptor cells of the vestibular system
o Contains 60-100 hexagonally arranged stereocilia and a
single longer kinocilium

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Vestibular System
BNS 2 Long Exam 4

Figure 5. Physiologic processes explained earlier. Depolarize, Resting,
Hyperpolarize

Morphologic Polarization of Hair Cells

• Horizontal Semicircular Canal
o Hair cells are arranged with their kinocilium on the side closer
to the utricle
o Movement of endolymph toward the ampulla -> stereocilia
deflects toward the kinocilium -> depolarization
• Vertical Semicircular Canals (A&P)
o Hair cells are arranged with the kinocilium father from the
utricle
o Movement of endolymph toward the ampulla -> kinocilium
will be deflected toward the stereocilia to the utricle ->
hyperpolarization
o Ampullipetal – towards the ampulla; Ampullifugal – away from
the ampulla (Know the difference, depo/hypo will happen
depending on which movement and on which canal)

Figure 4. Ampulla and Otolith Organ

B. HAIR CELL TRANSDUCTION

• Depolarization
o Deflection of the stereocilia toward the kinocilium causes
potassium channels in the apical portions of the stereocilia to
open. Potassium flows into the cell from the endolymph,
depolarizing the cell membrane. This depolarization in turn
causes voltage-gated calcium channels at the base of the hair
cells to open, allowing calcium to enter the cell. The influx of Figure 6. Para magets nyo lang yung sinasabi sa taas J
calcium causes synaptic vesicles to release aspartate or • Striola – small curving depression that roughly bisects the
glutamate into the synaptic clefts, and the afferent fibers macula
respond by undergoing depolarization and increasing their rate • Saccule macula
of firing. o Hair cells are arranged in such a way that the kinocilium is are
o Increased rate of firing in the vestibular afferent fibers located at the edges far from striola
• Hyperpolarization o Depolarization if hair cells are deflected away from striola
o If stereocilia are deflected away from the kinocilium • Utricle macula
o Stereocilia and kinocilium return to their resting position, o Hair cells are arranged in such a way that the kinocilium is
allowing most calcium channels to close and voltage-gated near the striola
potassium channels at the base of the cell to open. Potassium o Depolarization if hair cells are deflected towards the striola
efflux returns the hair cell membrane to its resting potential
o Decreased afferent firing rates
• Aminoglycoside (streptomycin, gentamicin) cause ototoxic effects
due to its direct reduction of the transduction of the hair cells

Figure 7. Explanation above. Note the arrows. Away vs Toward the striola.

IV. SEMICIRCULAR CANALS AND OTOLITH ORGANS

• SemicirculaR – Rotational
• otoLith – Linear
o Most prominent linear acceleration is the force of gravity
o Gravitointertial Acceleration - the vector sum of all linear
accelerations and the equivalent acceleration of gravity
• Pitch – forward and backward tilting
• Roll – Side-to-side
A. FUNCTION OF SEMICIRCULAR CANALS

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Vestibular System
BNS 2 Long Exam 4

• Transduce rotational head movements (angular accelerations) ¨ Lies superolaterally in the central pons
• Opposite polarization ¨ Bordered by the restiform body and the fourth ventricle
o Medial vestibular nucleus
¨ Lies in the lateral floor of the fourth ventricle
o Lateral vestibular nucleus
¨ Lies lateral to the medial vestibular nucleus
¨ Contains some large neurons, Deiters cells
o Inferior/ Descending vestibular nucleus
¨ Lies lateral to the medial vestibular nucleus
¨ Extends through much of the medulla

Figure 8. A. At rest B. Left turn, left side depolarize, left excited, right
inhibitedC. Right turn, right side depolarize, right excited, left inhibited
• "Push-pull" concept
o States that directional sensitivity to head movement is
coded by opposing receptor signals
o 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 Figure 9. Vestibular Nuclei (Description is above text)
their afferents show a decreased rate of firing. A rightward
head turn produces the opposite pair of responses A. VESTIBULAR AFFERENT UNITS
o Neurons act as comparator units that interpret head rotation • Enters the brainstem at the pontomedullary junction
on the basis of the relative discharges of left and right canal • Traverse the restiform body and the bifurcate into ascending and
afferents descending branches
o Head trauma, lesion on CN8 (glomus tumor or vestibular • Semicircular canals- project primarily to the superior and medial
schwannoma) may reduce impulses in the ipsilateral afferent vestibular nuclei; lesser input to the lateral and inferior vestibular
fibers or block their impulses transmission nuclei
¨ Akala mo nagrorotate yung ulo, stationary lang pala • Otolith organs- project primarily to the lateral, medial, and inferior
¨ Vestibular nuclei will consistently receive higher impulse vestibular nuclei
frequency from the intact side, which will be interpreted as
turning away from the side of the lesion. B. CEREBELLAR CONNECTIONS
• Vestibular labyrinth - only sensory organ in the body that sends
B. FUNCTION OF OTOLITH ORGANS direct primary afferent projections to the cerebellar cortex and
• Respond to translational head movements (linear acceleration) or nuclei
to orientation of the head relative to gravity • All passes through juxtarestiform body
• When the head is moved with respect to gravity (rolled or • Regulatory mechanism for the control of eye movements, head
pitched), the otoconia crystals are displaced because of their movements, and posture
density with respect to the surrounding endolymph. This
displacement shifts the underlying gelatinous coating on the
maculae and produces stereocilia deflection in the hair cells Primary vestibulocerebellar fibers
• Course through juxtarestiform body
V. VESTIBULAR NUCLEI
• Send collaterals to the dendate nucleus
• Set of neurons in the brainstem
• Terminate as mossy fibers in the nodulus, uvula, flocculus
• Responsible for receiving, integrating, and distributing information
that controls motor activities (such as eye & head movements,
postural reflexes, and gravity dependent autonomic reflexes) and
Secondary vestibulocerebellar projections
spatial orientation • End on the flocculonodular lobe, uvula, immediately adjacent
portions of the paraflocculus, fastigial, and dendate nuclei of the
• Four vestibular nuclei:
cerebellum
o Superior vestibular nucleus

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Vestibular System
BNS 2 Long Exam 4

Cerebellar corticovestibular fibers • Compensatory because they are equal in magnitude and opposite
• Purkinje axons from the nodulus, uvula, flocculus in direction to the head movement perceived by the vestibular
system
• GABAergic, inhibitory
• Occurs for any direction or speed of head movement
• Can be suppressed at will
Fastigiovestibular fibers
• End on the flocculonodular lobe, uvula, immediately adjacent A. ROTATIONAL VESTIBULOOCULAR REFLEX
• Projections from fastigial nucleus • 3 types of rotationally induced eye movements: horizontal, vertical,
torsional
• Glutamate or aspartate, excitatory
• Vertical semicircular canals and saccule - responsible for
controlling vertical eye movements
• Horizontal canals and utricle - control horizontal eye movements
C. COMMISURAL CONNECTIONS • Vertical semicircular canals and utricle - controls torsional eye
• Commisurral vestibulovestibular fibers movements
• Arise from all vestibular nuclei but most prominently from the
superior and medial nuclei
B. LINEAR VESTIBULOOCULAR REFLEX
• Depend on input from the otolith organ receptors
• GABA or glycine, inhibitory
• Side to side head movement result in a horizontal eye movement
• provide the pathways by which information from pairs of opposite to the head movement
corresponding semicircular canals and otolith organs can be
compared C. NYSTAGMUS
• Large head rotations like 360 degree body turn
• Vestibular compensation - process by which reflexes and
• Slow phase
postural control that are impaired as a result of unilateral loss of
o Initially, vestibuloocular reflex directs eyes slowly in the
vestibular receptor function (through trauma or disease) are direction opposite of head motion
restored gradually by means of central adjustment • Fast phase
o When eyes reach limit of how far it can turn, it returns to
D. OTHER AFFERENT CONNECTIONS central position, moving in the same direction as with the head
• Spinovestibular fibers • Nystagmus is combination of this slow and fast phase
o From all levels of spinal cord • Caloric test
o Provide proprioceptive input to the medial and lateral vestibular o Warm (40C) or cold (30C) water is dropped into external
nuclei auditory canal
• Accessory optic system o Normal person: eyes toward ear w/ warm water; eyes away
o Information concerning movement of the head through the ear w/ cold water
visual world o COWS (cold- opposite; warm- same)
• Reticular formation o Unilateral lesion in the vestibular system: nystagmus will be
o Proprioception reduced or absent on the side of the lesion
• Serotogenic cells in the raphe nuclei
o Modulatory or arousal effects
VII. VESTIBULOSPINAL NETWORK
• Coordinates head movements, axial musculature, and postural
• Thalamus
reflexes
E. OTHER EFFERENT CONNECTIONS
A. LATERAL VESTIBULOSPINAL TRACT
• Reticular formation
• LVST
• Posterior pontine nuclei • From neurons in the lateral and inferior vestibular nuclei
• Nucleus of solitary tract (NTS) • Project to all levels of ipsilateral spinal cord
o Cells in the medial and inferior vestibular nuclei project to the • Topographically organized
NTS and ventrolateral medullary reticular nuclei o Anterorostral areas of the lateral nucleus - project to cervical
o Mediate changes in the respiration and circulation that occur cord
with changes in posture o Posterocaudal regions - project to lumbosacral cord
o Stabilize respiration, heart rate, and blood pressure during • Receive input from the orthogonal semicircular canal pairs, the
otolith organs, the vestibulocerebellum, the fastigial nucleus, and
normal body position changes relative to gravity and during
proprioceptive inputs from the spinal cord
locomotion • Fibers course through lateral medulla dorsal to the inferior olivary
• Acute vestibular disorders complex and through the anterior funiculus of the cord
o Dysfunction of the vestibular autonomic response • Terminate directly on the alpha and gamma motor neurons and
o Orthostatic hypotension, postural tachycardia, and falls on the interneurons of lamina VII to IX
o Induction of motion sickness and emesis • Acetylcholine or glutamate; excitatory to the extensor muscle
*Please refer to first figure in Appendix for full understanding J motor neurons
• When you begin to tilt to the right, ipsilateral LVST fibers elicit
VI. VESTIBULOOCULAR NETWORK extension of the left axial and limb musculature. Concurrently,
• Arises from the vestibular nuclei right extensor muscles are inhibited. These actions stabilize the
• Control of eye movements - keep eyes fixed on an object while body's center of gravity and preserve upright posture.
head is moving (nakatitig palagi kay friend na may kaakbay at
B. MEDIAL VESTIBULOSPINAL TRACT
holding hands na iba)
• Primarily from medial vestibular nucleus, lesser projections from
Vestibuloocular reflex the inferior and lateral vestibular nuclei
• Stabilizing eye movements • Receive input from the vestibular receptors, cerebellum, and
spinal cord
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Vestibular System
BNS 2 Long Exam 4

• Descend bilaterally through the medial longitudinal fasciculus THALAMIC REGIONS FUNCTION
• Terminate in lamina VII to IX of cervical spinal cord, neck flexor Ventral posterolateral • Receive somatosensory input
and extensor motor neurons nucleus (VPL) • Project to the following:
• Carry both excitatory and inhibitory signals o primary somatosensory
cortex (area 2v, 3a)
o secondary somatosensory
cortex
o posterior parietal cortex
Ventral (areas 5,7)
posteroinferior o insula of temporal cortex
nucleus (VPI) • Many of these thalamic cells
are multimodal and respond
to both vestibular and
somatosensory stimulation
• Some are purely vestibular
• Located adjacent to the
medial geniculate body
• Receives additional auditory
afferents from the inferior
Posterior nuclear
colliculus and inputs from the
group (PO)
superior colliculus and spinal
cord
• Integration of multiple
sensory signals
• Located posterior to VPL
Anterior pulvinar • Projects to cortical area 3a,
posterior insula, and PIVC
• Lesions: misperceptions of visual vertical and postural instability

Figure 10. Vestibulospinal System
VIII. VESTIBULOTHALAMOCORTICAL NETWORK
A. VESTIBULAR THALAMUS
• The cognitive perceptions of motion and spatial orientation arise
through convergent information from the vestibular, visual, and
somatosensory systems at the thalamocortical level.
• Neurons in all four vestibular nuclei project bilaterally to the
thalamus
• Thalamic regions responding to vestibular stimuli:
B. VESTIBULAR CORTEX
• Multimodal; no primary vestibular cortex
• 4 main areas that respond to vestibular stimulation:
o Areas 2v and 3a
¨ Area 2v
à Located in the intraparietal cortex, at the base of the
intraparietal sulcus just posterior to the hand and mouth
parts of the postcentral gyrus
à Electrical stimulation produces sensations of whole
body motion
¨ Area 3a
à Lies on the base of the central sulcus, adjacent to the
motor cortex
à Receives input fibres from VPL and VPI thalamic
neurons
à Project to Area 4 of the motor cortex
■ One of the main functions of area 3a is to integrate
motor control of head and body
o Area 7 of parietal cortex
¨ Vestibular neurons in posterior parietal cortex
¨ Ventral intraparietal area (VIP) – multimodal neurons
involved in spatial coding
¨ Medial intraparietal area (MIP) – respond to both visual
and vestibular motion signals to integrate cues about body
motion through space
¨ Medial superior temporal area (MST) – same functions as
MIP
*lesions of the parietocortical areas can result in confusion
in spatial awareness
o Parietoinsular vestibular cortex (PIVC)
¨ Many of the cells here are multisensory, responding to body
motion, somatosensory, proprioceptive, and visual motion
stimuli
¨ PIVC lesions: vertigo, unsteadiness, and a loss of
perception for visual vertical
o Frontal eye field (FEF)
¨ Area 6 and the superior frontal gyrus receive vestibular
signals and are related to the FEF

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Vestibular System
BNS 2 Long Exam 4

¨ Control of saccades and smooth pursuit eye movement optimally achieved through which
¨ There are direct corticofugal projections to the vestibular structure?
nuclei, parabrachial nuclei, and prepositus hypoglossi 3. High amplitude reception is
• See Appendix Figure 11 best in the apex of the cochlea
Flexible
due to this characteristic of the
C. NAVIGATION structure
• Cognitive map – internal model of the spatial representation of NONE OF THE ABOVE
our environment (other choices: Opening of
• 3 cell types contributing to spatial orientation: 4. Deflection of the stereocilia
Clacium channels, Opening
away from the kinocilium
of Sodium channels,
CELL TYPE LOCATION RESPONDS TO produces
Opening of potassium
Location in the channels)
Place cells Hippocampus 5. Reflexive in Nature Vestibular System
environment
Entorhinal 6. Associated with language
Grid cells Spatial location Auditory System
cortex centers
Heading direction, 7. The outer portion of the ear has
Anterior dorsal CN 7,9,10
Head cells independent of sensory supply coming from
thalamus 8. Aspartate is the excitatory
location and behavior
o These regions have interconnections that are thought to neurotransmitter to the ganglion Vestibular Hair cells
function together to provide for spatial orientation, spatial cells.
memory, and our ability to navigate to learned locations
• Patients with trauma or disease to the vestibular system, REFERENCES
hippocampus, dorsal thalamus regions may have severe deficits
in their ability to orient in familiar environments Haines, D.E. (2013). Fundamental Neuroscience for Basic and
Clinical Applications. 4th ed. Elsevier Saunders.
IX. DIZZINESS AND VERTIGO
• Dizziness – nonspecific; spatial disorientation that may or may Lu Chiu 2C 2019
not involve feelings of movement
o May be accompanied by nausea or postural instability
o Many of these are not vestibular in origin
• Vertigo – specific perception of body motion (often spinning or
turning) experienced when no real motion is taking place
o Subjective vertigo – the patient experiences the sensation of
spinning
o Objective vertigo – sensation of objects spinning while the
patient is not moving
*examination of eyes during this phase will reveal a
nystagmus that beats in the direction opposite to the
original direction of rotation

A. BENIGH PAROXYSMAL POSITIONAL VERTIGO

• Brief episodes of vertigo that coincide with particular changes in
body position
• Can be triggered by turning over in bed, getting up in the morning,
bending over, rising from a bent position
• Pathophysiologic mechanism not yet understood but abnormality
in posterior canal is implicated
o Cupulolithiasis
¨ Otoconial crystals from the utricle separate from the otolith
membrane and become lodged in the cupula of the
posterior canal à increased density of cupula à abnormal
cupula deflections when head changes position relative to
gravity

B. VESTIBULAR NEURITIS
• Severe vertigo, nausea, vomiting with NO accompanying hearing
loss or other CNS abnormalities
• Edema of the vestibular nerve/ganglion
o Most commonly believed to be due to acute viral infection
such as herpes simplex
• Treatment: antiemetics, vestibular suppressants, corticosteroids,
antiviral agents

SAMPLEX
1. Low amplitude reception in the
cochlea is secondary to this Shorter stereocilia
characteristic of the hair cells:
2. Impedance matching is Oscicles

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APPENDIX

Figure 11. Vestibulothalamocortical

Pathway

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