THE VESTIBULAR SYSTEM

Introduction
⚫ The vestibular system is a somatosensory portion of
the nervous system that provides us with the awareness
of the spatial position of our head and body
(proprioception) and self-motion (kinesthesia).
⚫ It is composed of central and peripheral portions.
⚫ The peripheral portion of the vestibular system consists
of the vestibular labyrinth, vestibular ganglion,
and vestibulocochlear nerve (CN VIII).
⚫ The central portion of the vestibular system consists of
the vestibular nuclei.
Introduction
⚫ The vestibular labyrinth is comprised of proprioceptive
components located in the inner ear;
⚫ The semicircular canals, which contain the cells that
detect angular acceleration of the head;
⚫ The utricle and saccule, which contain the cells that
detect the linear acceleration of the head and position
of the head in space (spatial orientation).
Introduction
⚫ The stimuli from these receptors are conveyed to
the vestibular ganglion. From here, they travel through the
vestibular portion of the vestibulocochlear nerve (CN
VIII) into the central portion of the vestibular system; the
vestibular nuclei in the brainstem.
⚫ The vestibular nuclei send projections into
the cerebellum, spinal cord, thalamus, and nuclei of
the oculomotor (III), trochlear (IV) and abducens (VI)
nerves.
⚫ Via these connections, the vestibular system contributes to
the adjustments of the head and neck movements, as well as
the posture and balance of the whole body, vestibulo-ocular
reflex and eye movements.
Labyrinth and semicircular canals
⚫ The vestibular labyrinth is a bony cavity located within
the petrous portion of the temporal bone.
⚫ It consists of the bony framework for the cochlea as
well as the three semicircular canals.
⚫ The bony labyrinth houses the three semicircular
canals and the two otolithic organs (the utricle and
saccule).
⚫ Moreover, it contains the cochlea which is a part of the
hearing apparatus.
Diagram showing the labyrinth
Semicircular canals
⚫ The semicircular canals are the three membranous
channels located within the bony semicircular ducts of
the labyrinth.
⚫ They are located in three planes, with each canal
making an angle of approximately 90 degrees with the
other. Thus, the semicircular canals are the:
⚫ Anterior (superior), located in the sagittal plane
⚫ Lateral (horizontal), located in the transverse plane
⚫ Posterior (inferior), located in the frontal plane
Diagram of the showing the ampulla of
the semi-circular canals
⚫ The terminal part of each canal ends with a dilation called
the ampulla, which opens into the vestibule.
⚫ The ampulla of each semicircular canal contains a cluster of
mechanoreceptor cells called the crista ampullaris.
⚫ Each crista is composed of the special sensory receptor cells,
called the hair cells.
⚫ Given that the semicircular canals are filled with endolymph, the
movements of this fluid stimulate the hair cells.
⚫ In this way, each semicircular canal detects when the head
moves during the rotational acceleration along its corresponding
plane.
⚫ In other words, the semicircular canals detect head movements
such as nodding up and down, shaking side to side, or tilting left
and right.
The otolithic organs
⚫ The otolithic organs are the two membranous cavities that lie in
the bony vestibule of the inner ear. Namely, they are the utricle
and saccule.
⚫ The utricle lies in the posterior part of the vestibule. On one end,
it communicates with the semicircular canals, whilst on the
opposite end forms a utriculosaccular duct with the saccule. This
duct passes through the temporal bone and reaches the posterior
surface of the petrous part of the temporal bone.
⚫ The saccule lies anterior to the utricle and is significantly smaller
than it. Besides joining the utriculosaccular duct, the saccule
communicates with the cochlea by ductus reuniens.
⚫ The utricle and saccule contain the clusters of hair cells as well,
except that in them, the clusters are called the macula of the
utricle and the macula of the saccule.
Diagram showing the macula of the
utricle and saccule
The otolithic organs
⚫ The macula of the utricle is located in the horizontal
plane on the inferior surface of the utricle and plays an
important role in sensing horizontal movement (i.e
forward-backwards, left-right movement)
⚫ The macula of the saccule is located mainly in the
vertical plane and is responsible for sensing movement
in the saggital plane (ie up-down movement).
⚫ These neurosensory areas respond to the stimulation
from the endolymph to detect the linear movements of
the head, as well as its position in space while the head
is not moving but the rest of the body is.
Hair cells
⚫ The hair cells are the motion receptors of the inner ear,
embedded within the walls of the semicircular canals
and otolithic organs.
⚫ They are cylindrical in shape and feature many
stereocilia on their apical ends.
⚫ Each cell also has a single kinocilium, which sits on
the lateral most end of the apical surface.
Hair cells in the semicircular canals
⚫ In the semicircular canals, the hair cells project from the cristae
ampullares towards the cavity of the canals. The apical lining of
the hair cells in the semicircular canals is covered by a gelatinous
mass called the cupula.
⚫ The kinocilia of these cells are all oriented in the same direction
in the cupula, and bending the cupula in that direction causes
depolarization of the hair cells, whereas bending it in the
opposite direction hyperpolarizes the cells.
⚫ Then from the hair cells, appropriate signals are sent by way of
the vestibular nerve to apprise the central nervous system of a
change in rotation of the head and the rate of change in each of
the three planes of space.
Diagram of the cristae ampullaris
showing the cupula and the hair cells
Function
⚫ The rotational movements of the head in one of the planes
causes the endolymph within the semicircular canals to
move in the opposite direction by inertia.
⚫ The endolymph then flows against the cupula, which
distends across the apical surface of the hair cells, moving
the stereocilia toward the kinocilium.
⚫ These ciliary movements open the transduction cell
channels, triggering the excitation of the cells. The
movement of stereocilia in the opposite direction reduces
the cell activity.
⚫ The reason why the linear movements don’t stimulate the
cristae ampullares is that the linear movements happen in
the planes in which the endolymph always hits the cupula
from both sides, eventually not causing any displacement
nor hair cell stimulation.
Detection of rotational movements by
the cristae ampullaris
Hair cells in the utricle and saccule
⚫ In the macula of the utricle and saccule are thousands of hair
cells which project their cilia up in to the gelatinous layer of
the macula.
⚫ The bases and sides of the hair cells synapse with nerve
endings of the vestibular nerve.
⚫ Each macula is covered by a gelatinous layer called the
otolithic membrane in which small calcium carbonate crystals
called statoconia ,otoliths or otoconia are embeded.
⚫ The specific gravity of the calcified statoconia have a specific
gravity two or three times the specific gravity of the
surrounding fluid and tissues. The weight of the statoconia
bends the cilia in the direction of gravitational pull.
⚫ Each hair cell has 50- 70 cilia known as stereocilia plus one large
cilium (the kinocilium). The kinocilium is always located to one side
and the sterecilia become progressively shorter toward the other side of
the cell.
⚫ Minute filamentous attachments connect the tip of each stereocilium to
the next longer sterocilium and finally to the kinocilium. Because of
these attachments, when sterocilia and knocilium bend in the direction
of the kinocilium, the filamentous attachements tug in sequence on the
stereocilia pulling them outward from the cell body, this opens several
hundred fluid channels in the neuronal cell membrane around the bases
of the stereocilia which lead to receptor membrane depolarization.
⚫ Conversely, bending the stereocilia in the opposite direction reduces the
tension on the attachments which closes the ion channels leading to
hyperpolarisation.’
Diagram of the macula showing the hair
cells and otoliths
Functions
⚫ Under normal conditions, the nerve fibres leading from the hair cells
transmit contionous nerve impulses at a rate of 100 per second, when the
stereocilia are bent toward the kinocilium, the impulse tranffic increases
often to several hundred per second; conversely, bending the cilia away
from the kinocilium decreases the impulse traffic often turning it off
completely.
⚫ In each macula, each of the hair cells is oriented in a different direction so
that some of the hair cells are stimulated when the head bends forward,
some are stimulated when it bends backward, others are stimulated when it
bends to one side and so forth.
⚫ Therefore, a different pattern of excitation occurs in the macular nerve
fibres for each orientation of the head in the gravitational field. It is this
pattern that apprises the brain of the head’s orientation in space.
⚫ The hair cells within the vestibular organs are oriented in a way that the
whole vestibular apparatus is sensitive to movements in all directions.
Diagramatic representation of the role of
the utricular macula
⚫ The action potentials from the cristae and maculae are carried
by the corresponding branch of the vestibular nerve;
⚫ Crista ampullaris of the anterior duct transmits signals via
the anterior ampullary nerve.
⚫ Crista ampullaris of the posterior duct transmits signals via
the posterior ampullary nerve.
⚫ Crista ampullaris of the lateral duct transmits signals via
the lateral ampullary nerve.
⚫ Macula of the utricle transmits signals via the utricular nerve.
⚫ Macula the saccule transmits signals via the saccular nerve.
⚫ The utricular, anterior ampullary, and lateral ampullary nerves
join to form the utriculo-ampullary nerve. Ultimately, the
utriculo-ampullary, saccular and posterior ampullary nerves
synapse within the vestibular ganglion.
Vestibular ganglion
⚫ The vestibular ganglion (ganglion of Scarpa) lies
within the fundus of the internal auditory meatus.
⚫ It is a cluster of bipolar sensory neurons, that are the
first-order neurons of the vestibular pathway
⚫ The peripheral processes of vestibular ganglion cells
comprise the nerve fibers that receive the stimuli from
the hair cells of the otolithic organs and semicircular
canals, respectively.
⚫ The central processes of the vestibular ganglion
comprise the fibers of the vestibular portion of
the vestibulocochlear nerve (CN VIII).
Diagram showing the superior and
inferior vestibular ganglion
Vestibular nerve
⚫ The vestibular nerve transmits the equilibrium
impulses from the vestibular apparatus.
⚫ It leaves the inner ear through the internal auditory
meatus and enters the posterior cranial fossa.
⚫ Then, it synapses with the vestibular nuclei in the
brainstem.
⚫ Moreover, some of the fibers synapse directly with the
cerebellum as well.
⚫ This makes the vestibular nerve unique, as it is the
only cranial nerve whose first-order neurons synapse
with the cerebellum directly.
Diagram showing the vestibular nerve
Vestibular nuclei
⚫ The vestibular nuclei are the four nuclei that lie within the rhomboid
fossa of the brainstem. They contain the second-order neurons of the
vestibular pathway that synapse with the vestibular portion of the
vestibulocochlear nerve. Namely, the vestibular nuclei are:
⚫ The superior vestibular nucleus (of Bechterew)
⚫ The lateral vestibular nucleus (of Deiters)
⚫ The inferior vestibular nucleus (of Roller)
⚫ The medial vestibular nucleus (of Schwalbe)
⚫ The superior and medial vestibular nuclei receive most of the inputs from
the cristae ampullares of the semicircular canals.
⚫ The inferior and lateral nuclei receive the remaining fibers from inferior
semicircular canals, as well as from the utricle and saccule.
⚫ The vestibular nuclei integrate inputs from the peripheral vestibular
structures, contralateral vestibular nuclei, cerebellum and the other
sensory systems (the visual and somatosensory systems).
Diagram showing the vestibular nuclei
Connections with the nuclei of the
cranial nerves
⚫ The superior and medial vestibular nuclei send fibers that
join the medial longitudinal fasciculus (MLF).
⚫ Via this pathway, they synapse with the motor nuclei of
the oculomotor (CN III), trochlear (CN IV) and abducens
(CN VI) nerves, including the interstitial nucleus of Cajal
and the nucleus of Darkschewitsch.
⚫ This way, the vestibular system mediates the reflexive
activity of the extraocular muscles.
⚫ More precisely, the vestibular system mediates the
vestibulo-ocular reflex, in which the movements of the
eyes are adjusted to the movements of the head.
Fixing the gaze on a static object with
head rotation
Vestibulo-ocular reflex
⚫ The vestibulo-ocular reflex is an involuntary motor
activity mediated by the vestibular system which serves
for adjusting the eye movements while the head moves in
the horizontal plane.
⚫ It serves for fixing the gaze during head repositioning.
The reflex arc is mediated by the medial longitudinal
fasciculus (MLF) in the following way;
⚫ Once the head is turned sideways (e.g. to the left), the
endolymph in the vestibular organs falls behind to the
right due to inertia and stimulates the hair cells of the left
horizontal semicircular canal.
⚫ The information is conveyed to the vestibular nuclei via
the ipsilateral vestibulocochlear nerve.
Vestibulo-ocular reflex
⚫ The neurons from the superior and medial vestibular nuclei join the MLF
and reach the nucleus of the contralateral abducens nerve. This way, the
contralateral lateral rectus muscle contracts and pulls the eyes in the
opposite direction to that of the head.
⚫ Simultaneously to the abducens stimulation, some MLF fibers cross the
midline and synapse with the oculomotor nucleus. This stimulates
the medial rectus muscle on the opposite eye, resulting with the conjugated
movement of both eyes in the horizontal plane.
⚫ At the same time, the ipsilateral medial rectus muscle is inhibited so that it
doesn’t compete with its contralateral counterpart.
⚫ To reiterate, once a person turns their head while wanting to fix the gaze on
a static object, the vestibulo-ocular reflex enables the eyes to turn in the
opposite direction so that the person can maintain focus on the perceived
object.
Diagram showing eye movements in
vestibulo-ocular reflex
Connections with the spinal cord
⚫ The lateral vestibular nucleus sends axons via the lateral
vestibulospinal tract.
⚫ This tract synapses with interneurons along the entire
length of the spinal cord, adjusting the posture of the
body and tone of extensors according to the vestibular
stimuli (vestibulo-spinal reflex).
⚫ The medial and inferior vestibular nuclei project their
fibers via the medial vestibulospinal tract.
⚫ This tract terminates within the cervical spinal cord,
adjusting the posture of the head and neck
(vestibulo-cervical reflex).
Connections with the cerebellum
⚫ The vestibular inputs connect with the cerebellum in
two ways;
⚫ The second-order neurons from the vestibular nuclei
project to the inferior olivary nucleus via
the vestibulo-olivary tract.
⚫ From here, the vestibular inputs are relayed via the
lateral part of the inferior cerebellar peduncle
(restiform body) into the ipsilateral cerebellar vermis,
flocculus and nodulus.
⚫ This connection enables the joint modulation of
balance by the cerebellum and vestibular system.
Connections with the cerebellum
cont’d
⚫ A portion of the first-order neurons from the vestibular
ganglion of Scarpa pass through the medial part of the
inferior cerebellar peduncle (juxtarestiform body).
⚫ They enter the cerebellum as the mossy fibers and
synapse directly with the ipsilateral
vestibulocerebellum, vermis and fastigial nucleus.
⚫ These connections enable cerebellar awareness of the
vestibular sensations and promote necessary movement
modifications by the cerebellum.
Connections with the cortex
⚫ The superior and lateral vestibular nuclei project to the
ventral posterior nuclei of the thalamus to synapse with
the third-order neurons of the vestibular pathway.
⚫ The thalamus then relays the signals to the primary
vestibular cortex (Brodmann area 3a), located in
the cortex of the parietal lobe adjacent to the
primary motor cortex.
⚫ This area integrates the information from the vestibular
system with other proprioceptive systems and passes that
information directly to the primary motor
cortex (Brodmann area 4).
⚫ From here, the motor response to the proprioceptive
stimuli is generated
Connections with the cortex cont’d
⚫ Other connections between the vestibular system and the
cortex are still being investigated. So far, the two
particularly interesting cortical areas have been
identified;
⚫ One of them lies posterior to the primary somatosensory
cortex;
⚫ The second one lies between the somatosensory cortex and
motor cortex
⚫ These cortical areas process the vestibular and visual
stimuli, and they are particularly active when a person
undergoes rotatory movements with eyes closed.
Pathophysiology
⚫ Motion sickness
⚫ Motion sickness is a condition characterized by nausea
and vomiting due to the travelling, it is most often seen
in some people travelling by vehicle. It is due to
fluctuations in the maculae.
⚫ Vertigo
⚫ Vertigo is a condition in which a person has a false
sensation that either him or surroundings are in motion. It
can cause nausea, dizziness, sweating and vomiting. This
condition is associated with vestibular malfunction.
Vertigo is the common symptom of Vestibular neuritis
(labyrinthitis), Meniere’s disease and VIII nerve damage.
⚫ Meniere’s disease
This is caused by blockage in the cochlear aqueduct
and symptoms are tinnitus, hearing loss and vertigo,
sense of pressure in the ear, sound distortions and
noise sensitivity.
⚫ This disease results in endolymphatic volume increase
with ballooning of the cochlea duct, utricle, and
saccule.
Caloric reflex test
⚫ This is a test of the vestibulo-ocular reflex. It is done by
pouring cold or warm water into the external auditory canal
using a syringe. The temperature difference between the body
and the water creates a convective current in the endolymph
of the horizontal semicircular canal.
⚫ Hot and cold water produce currents in opposite directions
and therefore a quick horizontal eyes movement in the
opposite directions.
⚫ If the water is warm, the endolymph in the ipsilateral
horizontal canal rises, thereby increasing the firing rate of the
afferent vestibular nerve. This will result to the eyes turning
towards the contralateral ear, with a quick horizontal eyes
movement to the ipsilateral ear.
Caloric reflex test
⚫ If the water is cold, the endolymph in the semicircular canal
falls, thereby decreasing the firing rate of the afferent
vestibular nerve. This will result to the eyes turning towards
the ipsilateral ear and a quick horizontal movement of the
eyes to the contralateral ear.
⚫ If the caloric test is done on patients with cerebral damage, the
fast phase of quick horizontal eyes movement will be absent
as this is controlled by the cerebrum.
⚫ Also pouring cold water into the external auditory canal of
brain-damaged patients will result in quick horizontal eyes
movement toward the contralateral ear.