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Physiology of Equilibrium

Balance requires :
vestibular system
Input from visual
Input from

Vestibular Apparatus
Membranous and
bony labyrinth
Vestibular nerves
Vestibular nuclei

5 distinct end organs
3 semicircular canals:
superior, lateral,
2 otolith organs:
utricle and saccule

Semicircular canals sense angular

Otolithic organs (utricle and saccule) sense
linear acceleration

Lies in the spherical
recess on the medial
wall of the vestibule
anteriorly to the
cochlear duct by the
ductus reuniens
posteriorly to the
endolymphatic duct
via the
utriculosaccular duct


lies posterosuperiorly in the

elliptical recess of the
medial wall of the vestibule

connected anteriorly via the

utriculosaccular duct to the
endolymphatic duct

The 3 semicircular canals

open into it by means of 5
openings; the posterior and
the superior semicircular
canals share one opening at
the crus commune

Semicircular canals
lateral or horizontal,
superior or anterior,
and posterior or
They are oriented at
right angles to each

Semi-circular canals
One end of each canal is dilated to form
the ampulla, which contains a saddleshaped ridge termed the crista
ampullaris, on which lies the sensory
The nonampulated ends of the superior
and posterior canal form the crus
commune or common crus.
All canals merge into the utricle.

Location of five vestibular end organs


Sensory epithelium of
utricle and saccule
project into the
overlying statoconial
The statoconial
membrane is
comprised of 3 layers:



The otoconial first layer is comprised of calcareous

particles (otoconia), which are inorganic crystalline
deposits composed of calcium carbonate or calcite.
They are distributed in a characteristic pattern and vary
in size from 0.5-30 mcm, with most about 5-7 mcm.
The specific gravity of the otolithic membrane is much
higher than that of the endolymph, about 2.71-2.94.
The second layer is a gelatinous area of
mucopolysaccharide gel.
The third layer consists of subcopula meshwork.

The macula of the utricle

lies mainly in the
horizontal plane and is
located in the utricular
recess, which is the
dilated anterior portion of
the utricle
Saccular macula is an
elliptical thickened area of
sensory epithelium that
lies on the anterior
vertical wall of the

Orientation of two
maculae. The
utricular macula is in
the horizontal plane,
and the saccular
macula is in the
vertical plane

Morphological polarizations of the stereociliary bundles in the maculae of the (A) utricle
and (B) saccule. The on direction of stereo ciliary deflection is indicated by the arrows.
In the utricle (A) hair cells are excited by stereo ciliary deflection toward the striola
(curving central zone). In the saccule (B) hair cells are excited by stereo ciliary deflection
away from the striola.

Otolithic Organs

consists of a crest of sensory epithelium supported on a
mound of connective tissue, lying at right angles to the
longitudinal axis of the canal.
A bulbous, wedge-shaped, gelatinous mass called the
cupula surmounts the crista.
Cilia of the sensory cells project into the cupula.
The cupula extends from the surface of the cristae to the
roof and lateral walls of the membranous labyrinth,
forming a fluid-tight partition


A, The cupula spans the lumen of the ampulla from the crista to
the membranous labyrinth.
B, Head acceleration exceeds endolymph acceleration. The
relative flow of endolymph in the canal is therefore opposite to
the direction of head acceleration. This flow produces a
pressure across the elastic cupula, which deflects in response.

Lateral semicircular canal kinocilium on

utricular side increased discharge rate
by ampullo-petal movement of the cupula
(towards the utricle) and decreased by
ampullofugal movement.
Opposite in other 2 canals

Vestibular hair cells

type I or type II
Type I hair cells correspond to the
inner hair cells of the organ of Corti
Round bottoms, thin necks, and
wider heads
Each cell is surrounded by a nerve
chalice from one of the terminal
branches of a thick or medium nerve
fiber of the vestibular nerve.

Type II hair cells

correspond to the
outer hair cells of the
organ of Corti
Cylinders, with a flat
upper surface
covered by a cuticle.

Schematic drawing of
the two types of
sensory cells in the
mammalian labyrinth
showing fine
structural organization
of type I and type II
sensory cells and
their innervation

Each hair cell bears 50-100 stereocilia and a

single thick and long kinocilium on the apical
Each hair cell is morphologically polarized with
respect to location of the kinocilium.
The basal portions of the cells synapse with
afferent and efferent nerve fibers.

Afferent vestibular pathways

The primary vestibular neurons are bipolar
neurons whose cell bodies comprise the
Scarpa ganglion in the internal auditory canal.
These bipolar neurons lie in 2 linearly
arranged cell masses extending in a rostralcaudal direction in the internal auditory canal.
Each neuron consists of a superior and
inferior cell group related to superior and
inferior divisions of the vestibular nerve trunk.

The superior
division supplies
the cristae of the
superior and lateral
canals, the macula
of the utricle, and
the anterosuperior
part of the macula
of the saccule.

The inferior division

supplies the crista of the
posterior canal and the
main portion of the
macula of the saccule.
Medial to the vestibular
ganglion, the nerve
fibers of both divisions
merge into a single
trunk, which enters the
brain stem

Vestibular nuclei
(first-order neurone)
Most afferent fibers from the hair cells terminate
in the vestibular nuclei, which lie on the floor of
the fourth ventricle.

Afferent pathways to
the vestibular nuclei

Efferent connections
of the vestibular

In the vestibular nuclei, 4 major groups of cell

bodies (the second-order vestibular neurons)
may be identified:
(1) superior vestibular nucleus (SVN) of
(2) lateral vestibular nucleus (LVN) of Dieter
(3) medial vestibular nucleus (MVN) of
(4) descending vestibular nucleus (DVN).

Extend to the cerebellum, extraocular nuclei, and
spinal cord.
The cells of the superior vestibular nucleus project in
an ascending direction to the nuclei of the extraocular
muscles (III and IV). This projection almost, if not
entirely, reaches the ipsilateral eye nuclei by way of
the medial longitudinal fasciculus.
The lateral vestibular nucleus has been shown to be
the sole source of fibers to the vestibulospinal tract.
These fibers terminate near the anterior horn cells of
all the spinal cord levels and mediate trunk and limb
muscle reflexes.

The descending vestibular nucleus appears to

be the nucleus most clearly related to the
The medial vestibular nucleus appears to be the
least specialized of the complex. It receives
afferents from the semicircular canals and
utricle; its projections are both ascending and
descending in the medial longitudinal fasciculus.
The ascending ones course bilaterally to the
extraocular muscles and the descending to the
cervical segment of the cord.

Cerebral cortex
The vestibular area in the cerebral
cortex has not been defined by using
anatomical methods.
Electrophysiologic studies indicate that
the projection area is in the temporal
lobe near the auditory cortex.

Cerebral cortex
Functional MRI and PET studies implicate the
insula as another possible cortical projection of
the vestibular system.
These pathways appear to integrate vestibular,
proprioceptive, and visual signals to provide a
conscious awareness of body orientation

Midline cerebellum plays a pivotal role in modulating the
final motor activity triggered by the VOR and VSR
Some primary vestibular neurons pass directly to the
cerebellum, in particular the flocculonodular lobe and the
Direct and indirect vestibular input reach the cerebellum,
which in turn produces an inhibitory influence via
Purkinje cell activity to fine tune ocular and postural
movements and assure appropriate interaction among
vestibular , visual, and proprioceptive cues

Semicircular canals
Majority of semicircular canals
input synapses in the medial and
superior vestibular nuclei

Second-order neurons course

ipsilaterally and contralaerally in
the medial longitudinal facilulus to
reach 6th, 4th and 3rd oculomotor
A third motor neuron innervates
the extraocular muscles to create
conjugate eye movement equal to
the opposite head movement

As each semicircular canal is connected to

the eye muscles, stimulation of a canal
nerve results in eye movement
approximately in the plane of the canal

Stimulation of left posterior

canal nerve excites the
ipsilateral superior oblique and
contralateral inferior rectus
muscles while inhibiting the
ipsilateral inferior oblique and
contralateral superior rectus
The end result is an oblique
downward movement in the
plane of the left posterior canal.

Macular afferents

Those from utricle and saccule

synapse mainly in the medial and
lateral vestibular nuclei.

Then send second-order neurons

to the anterior horn cells of the
cervical cord (medial
vestibulospinal tract) and the entire
spinal cord (lateral vestibulospinal
tract), respectively (VCR &VSR)

Important for changes in

orientation to gravity and linear
forces to the antigravity muscles of
the neck, thorax, and lower lumbs

A smaller number of fibers carry

information to extraocular muscles to
produce rotational and vertical ocular
adjustments during head tilt

The principal function of the VOR is the
control of eye position during transient
head movements to maintain a stable
visual image

Acceleration or deceleration movement of

the head, producing stimulation of one or
more of the receptors on one side with the
corresponding inhibition on the other side,
results in asymmetrical neural input from
the vestibular nerves.

This asymmetrical input is interpreted by the

central nervous system as either angular or
linear movement
In addition, the asymmetry resulting from action
of the semicircular canals causes a
compensatory reflexive eye movement in the
plane of the canals being stimulated, known as
the first of Ewalds three laws.
This compensatory movement of the eye is
called the vestibulo-ocular reflex (VOR), and is
opposite to the direction of acceleration.

Canal physiology

Neural connections in the direct pathway for the

VOR from excitation of the left horizontal canal
A counterclockwise head rotation (head)
produces relative endolymph flow in the left
HC that is clockwise and toward the utricle.
The cupular deflection excites the hair cells
in the left HC ampulla, and the firing rate in
the afferents increases (inset). Excitatory
interneurons in the vestibular nuclei (vest.
N.) connect to motor neurons for the medial
rectus muscle in the ipsilateral IIIrd nucleus
(III) and lateral rectus muscle in the
contralateral VIth nucleus (VI). Firing rates
for these motor neurons increase (bar
graphs). The respective muscles contract
and pull the eyes clockwiseopposite the
headduring the slow phases of
nystagmus. Inhibitory interneurons in the
vestibular nuclei connect to motor neurons
for the ipsilateral lateral rectus and
contralateral medial rectus. Their firing rates
decrease (bar graphs), and these antagonist
muscles relax to augment the eye

Complementary neural connections in the direct pathway for

the VOR from inhibition of the right horizontal canal

for the right HC this flow is away from the

utricle. The cupular deflection inhibits the hair
cells in the right HC ampulla, and the firing
rate in the afferents decreases
Inhibitory interneurons in the vestibular nuclei
(vest. N.) reverse this inhibition, sending
further excitatory signals (bar graphs) to the
motor neurons for the medial rectus muscle in
the left third nucleus (III) and lateral rectus
muscle in the right VIth nucleus (VI). The
contraction of these muscles is augmented.
Simultaneously, excitatory interneurons (open
symbols) in the vestibular nuclei (vest. N.)
preserve and convey inhibition (bar graphs) to
motor neurons for the lateral rectus muscle in
the left sixth nucleus (VI) and medial rectus
muscle in the right IIIrd nucleus (III). The
inhibitory signals further relax these antagonist

Major Structures Required For

Intact semicircular canals
and otoliths
Intact vestibular nuclei
Intact brainstem pathways
Motoneurons (eye muscles)
Intact cerebellum

Central VOR Pathways

Two major neuron types are involved in the VOR.

FTNs (Floccular Target Neurons) receive input from the cerebellar

flocculus and the semicircular canals.

PVPs (Position Vestibular Pause neurons) receive input from the

canals but not from the cerebellum.

Smooth pursuit system

Permits tracking of a visual target with a smooth
continuous movement of the eye
Provides for stable image projection to the fovea
of the retina
The frequency of movement should be less than
1.2 Hz
E.g. viewing a single bird flying across a persons
visual field or watching a pendulum swing
Vestibulo-cerebellum (flocculus, nodulus and
posterior vermis) plays a dominant role in smooth

Saccadic system
Provides the fast component during the
production of jerk nystagmus
Reposition a visual target of interest onto
the fovea with a single rapid eye motion
Cortical activity, pontine reticular formation
and vestibulo-cerebellum participate in
modulating saccadic eye movement

Postural and Motor system control

Cerebral cortex influences lower motor
centers through Pyramidal system and
Extra-pyramidal system
Pyramidal system is important for fine
isolated versatile movements
Extra-pyramidal system provide the
mechanisms for large gross movement
patterns that are primarily reflexive and
constitute major postural adjustments

These 2 systems function in a coordinated

fashion, e.g. picking up an object from
directly in front of the subject while

Three tracts originate from, or

contributed to, the vestibular nuclei:
1. Medial vestibulospinal tract
2. Lateral vestibulospinal tract
3. Reticulospinal tract

Medial vestibulospinal tract play a major

role in the cervico-vestibulo-ocular
reflexes, coordinating eye-head
Lateral vestibulospinal and reticulospinal
tracts influence on the coordination of
head and upper body movement with the
lower limbs

Three general movement control

1. Myototic reflex (deep tendon reflex)
2. Automatic muscle response (functional
stretch response)
3. Volitional movement

Myototoic reflex
Is stimulated by an externally generated
pull on a muscle, causing stimulation of
tendon and muscle stretch receptors
Is mediated through spinal cord pathways
Regulate muscle force, thereby
maintaining stability at a joint

Automatic muscle response

(functional stretch response)
Is stimulated by and external event,
primarily the somatosensory input
Is mediated through spinal pathways
Is modulated through brain stem and
subcortical pathways, including basal

Provides for coordinated limb and trunk

movements across the joints (coordinated body
segment movements about a joint, such as hip
or ankle)
Provides for the corrective action to recover from
a destabilizing perturbation of the center of
mass, such as when an individual is lightly
bumped from behind while standing, a balance
correction response

Volitional movement
Mediated at all levels, including cortical
sensory and motor areas
Responsible for learned, purposeful
movements, and balance stabilizing