The vestibular system

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Status Completed

Week 6

Property Chapter 14

The peripheral portion of vestibular system incl. inner ear structures - function as
initial guidance device → made up of small linear accelerometers & angular velocity
sensors
~ report info. about the motions & position of head → to integrative centers in the
brainstem, cerebellum, cerebral cortices.
→ extensive connections w. brainstem & cerebellar structures.

The vestibular sys. - key component in postural reflexes & eye movements ~ in
concert w. the visual & proprioceptive sys.
→ central role in distinguish self-generated ‘active’ movements of head & body from
passive movements - externally applied forces.

The vestibular nuclei - innervate motor meurons controlling extraocular, cervical,

postural muscles

→ mediating stabilization of gaze, head orientation, posture during movement.

The vestibular labyrinth

Labyrinth - main peripheral component of vestibular sys. ~ elaborate set of

interconnected chambers.
→ derived from the otic placode of the embryo - uses the same spec. set of sensory
cells/ hair cells → transduce physical motion into neural impulses.

The vestibular system 1

 deep in temporal bone - consist otolith organs (utricle & saccule) + semicircular
canals.

utricle * saccule - prim. to respond to translational movements of head & static

head position relative to gravitational axis.
~ spec. responding to rotations of head.

The intimate rela. btwn cochlea & labyrinth - beyond their common embryonic
origin.

membranous labyrinth - sacs w/in the bone - filled w. fluid (endolymph)

collectively.

perilymph - btwn bony walls & membranous labyrinth → similar in composition

to cerebrospinal fluid.

ampullae - vestibular hair cells located in the utricle & saccule in 3 jug-like
swellings.

tight junctions - seal the apical surfaces of vestibular hair cells → ensuring
endolymph selectively bathes the hair cell bundle
~ remaining separate from perilymph surrounding the basal portion of hair cell.

Vestibular hair cells

movement of stereocilia toward kinocilium in vestibular end-organs → open

mechanically gated transduction channels located at the tips of stereocilia.
~ depolarise the hair cell → cause neurotransmitter release onto the vestibular
nerve fibers.

movement of stereocilia in direction away from the kinocilium closes the

channels
~ hyperpolarising the hair cell → reduce vestibular nerve ability.

Biphasic nature of the receptor potential means - some transduction channels open
in the absence of stimulation ~ result that hair cells tonically release transmitter

→ generate considerable spontaneous act. in vestibular nerve fibers.

The vestibular system 2

 The firing rates of vestibular fibers incr./ decr. in a manner → mimics the
receptor potentials produced by the hair cells.

Adaptation in vestibular hair cells - Ca enter thru. mechanoelectrical transduction

(MET) & voltage-gated Ca channels
→ imp. to vestibular function - allow hair cells to continue to signal small changes in
head position.

The hair cell bundles in ea. vestibular organs - respond to displacements in all
directions.

Semicircular canal - hair cells in the ampulla all polarised in the same direction

→ 3 semi-canal oriented to detect rotational motion around the 3 cardinal axes.

Striola divides hair cells ~ into 2 populations having opposing polarities.

→ principle of organisation in the vestibular sys. - become apparent in the following
descriptions of indv. vestibular organs.

Otolith organs (utricle & saccule)

detect tilting & translational movements of the head.

Both contains sensory epithelium:

1. macula - consists of hair cells - associated supporting cells.

2. otolith membrane - gelatinous layers of hair cells + bundles → fibrous

structure - embedded crystals of Ca2+ (otoconia).

→ make otolithic membrane heavier than the structures & fluids surrounding it.
(Figure 14.4)

when the head tilts - gravity causes the membrane to shift relative to macula.

⇒ resulting shearing motion btwn the otolithic membrane & macula displaces the
hair bundles - embedded in lower, gelatinous surface of the membrane.

displacement of the hair bundles - generates a receptor potential in the hair

cells.

The vestibular system 3

 shearing motion btwn macula & membrane - occurs when the head undergoes
translational movements.

⇒ greater relative mass of the membrane - causes it to lag behind the macula
temporarily ~ lead to transient displacement of hair bundle.

Otolith afferents - cannot convey info. that distinguishes btwn 2 types of stimuli.
→ expect these diff. stimuli - rendered equivalent when visual feedback is absent.

eg. in the dark when eyes are closed.

Blindfolded sub. - discriminate btwn 2 stimulus categories → depends on the

integration of info. from otolith organs & semi-canals in the central vestibular sys.

The orientation of hair cell bundles - organised relative to striola → demarcates the
overlying layer of otoconia.

a head tilt along striola axis - excite the hair cells on one side ~ inhibiting the
hair cells on the other side.

⇒ tilts of head on 1 side - opp. effects on corresponding hair cells of 2 utricular

maculae.

Saccular macula - oriented vertical → respond to vertical translational

movements of the head - upward/ downward head tilts.

Utricular macula - horizontal → respond to translational movements of the

head in horizontal plane + sideways head tilts.

The vestibular system 4

 How otolith neurons sense head tilts

The mass of otolith membrane - related. to surrounding endolymph + physical

uncoupling from the underlying macula
→ hair bundle displacement - occur transiently in response to translational head
movements/ head tilts.

⇒ hair bundle displacement - reflected in the response of vestibular nerve fibers -

innervate the otolith organs.

nerve fibers steady & relatively high firing rate - head upright.

otolith afferent fiber (axons) - high firing rate → incr./ decr. depending on the
direction of the tilt.

remain high level - tilting force remains constant → encode the static force
being applied to the head.

original position - firing returns to baseline value.

tilt in opp. direction - neurons respond by decr. their firing rate below the resting
level → remain depressed ~ static force continues.

Translational head movements

transient incr./ decr. in firing rate from spon. level → signal the direction of
translational movement.

orientation of hair bundles + otolith organs → enables to transmit info. about linear
forces in every direction head move.

hair cell polarity in otolith organs - produce patterns of vestibular nerve fiber act. →
encode head position ~ forces that infl. it.

Semicircular canals

sense head rotational components of head movements - arising either from self-
induced head rotations/ angular accelerations of head by ex. forces.

The vestibular system 5

 Ea. works in concert w. the partner located on other side of the head → its hair cells
aligned opp.

semircircular canals can be excited by rotations that occur during

the initiation of a head tilt as well as rotations resulting from other
active or passive movements of the head.

ampulla - houses the sensory epithelium (crista) ~ contains the hair cells

cupula - bridges the width of ampulla → form a viscous barrier thru. which
endolymph cannot circulate.

⇒ endolymphic fluid distort the relatively compliant cupula.

the inertia of endolymph - produce a force across cupula ~ distending it away from
the direction of head movement.

→ cause a displacement of hair bundles w/in the crista.

translational movement of the head - produce equal forces on 2 sides of cupula ~

hair bundles w/in the ampulla are not displaced.

cupula moves in appro. direction - entire pp. of hair cells → depolarised & act.
in all of the innervating axons incr.
vv. → hyperpolarised and neuronal act. decr.

The vestibular system 6

 Deflections orthogonal to excitatory - inhibitory direction → produce litt. ~ no
response.

Head rotation - deforms the cupula in opp. directions for 2 partners → results in
opp. changes in their firing rates.

hair cells in the canal toward head turning → depolarised.

other side → hyperpolarised.

net result - sys. provide info. about the rotation of the head in any direction.

How semicircular canal neurons sense head rotations

Vestibular fibers - innervate the semi-canals + otolith organs.

→ exhibit a high level of spon. act.

transmit info. by. either incr/decr their firing rate → allows to encode rotational
head movements.

Bidirectional response of fibers - monkey’s study of vestibular nerve.

3 phases of head movement

1. initial acceleration - max. firing rates observded during this period → cupula
deflected ~ suggest incr. neural act. during this plane.

2. constant velocity - rates return to baseline level → cupula returns to its

undeflected state ~ process infl. by cupular elasticity + viscosity of endolymph.

3. sudden deceleration - min. rates → cupula deflected in opp. direction ~ decr.

neural act.

→ the return to undistorted state of cupula → happen while head is turning ~

constant angular velocity is maintained.

cupula-endolymph sys. - prolonged acceleratory arcs can occur.

The vestibular system 7

 smooths the conversion of head accelerations into neural signals → make the
sensory response more adaptive.

High-freq. rotational movement - head angularly accelerated to const. velocity

rapidly → vestibular units associated w. the affected canal - generate a velocity
signal.
~ reflected in the linear rise in axon firing rate during the acceleration phase.

Low- freq. - head moved const. → firing rate returns to spon. level - represent an
acceleration of 0.

Central pathways for stabilising gaze, head, posture

organisation of central vestibular pathways - reflects multifunctional role ~ pathways

display 2 features → distinguish them from the pathways ~ convey info. imp. to
sight hearing & touch.

central vestibular process - inherently multisensory bc. manny neurons receive

visual input.

many neurons in vestibular nuclei function - premotor neurons → giving rise to

ascending sensory projections.

→ provide very short-latency sensorimotor arc - drive extremely rapid

compensatory eye + head movements ~ response to vestibular stimulation.

The vestibular end-organs communicate - targets in the brainstem + cerebellum →

process necessary info. ~ compute head position and motion.

The vestibular nerves - originate from bipolar neurons.

Cell bodies located in the vestibular nerve Sacrpa’s ganglion.

The peripheral processes of these cells - extend to innervate the semi-canals

+ otolith organs in inner ear.
→ structure crucial for detecting head movements & orientation in space.

The central processes of vestibular neurons - project thru. the cranial nerve VIII
→ reach the vestibular nuclei in the brainstem.

The vestibular system 8

 (direct connections to cerebellum)

Afferent pathways from semi-canals + otolith organs separate in periphery →

sig. amount of convergence btwn canal + otolith inputs

brain processes info. from these 2 types of vestibular receptors in coordinated

manner.

→ encoding head orientation + motion.

💡 otolith organs respond to head tilts and translational movements

semicircular canals are specifically excited by rotations accompanying head

tilts.

The vestibulo-ocular reflex (VOR) - a key reflex mechanism → allows for

compensatory eye movements - counter head movements

~ enabling the gaze to remain fixed on a specific point during head movement.
→ project to the vestibular nuclei - signals are transmitted to eye muscles ~
maintain. gaze stability during head movements.

responsible for responding to linear vertical head oscillations ~ during

locomotion & vertical angular lead accelerations.

The vestibular system 9

 VOR - temporarily diminished during voluntary movements - redirect gaze →
prevent it from interfering w. goal-directed movements.

VOR displayed plasticity - response toaltered visual/ vestibular input → manifest

after unilateral damage to vestibular periphery.

triggered by placing optics over the eyes - magnify, miniaturize, invert the
optical image on the retina.
→ maintain stable image on the retina - enable coordinated movements under
visual control.

Loss of VOR - severe consequences (oscillopsia) → patients struggle to fixate on

visual targets during head movements.

unilateral damage - lead to recovery.

bilateral damage - persistent difficulties in perceiving head movements ~

maintaining stable gaze.

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 VCR (head postural adjustment) & VSR (body) - fast reflexes → requiring small
no. of synapses btwn vestibular organs + motor neurons.
→damage - lead to impaired head & postural stability, gait deviations, balance diff.

💡 The balance system - relies on a combination of vestibular, visual, and

proprioceptive inputs

→ become more critical in low-light conditions or when navigating uneven

surfaces.

VSR regulates head position + posture ~ thru. act. of neck and trunk muscles →
response to stimulation of semi-canals.

helps to stabilise head position in various situations - reflexively act. neck

muscles to pull the head up.

The vestibular system 11

 head movements - initiate VSR can trigger additional reflexes (forelimb
extension & hindlimb flexion) → protect against falls - stabilise the body.

The VSR involves several pathways - including the lateral and medial
vestibulospinal tracts and the reticulospinal tract.

→ receive inputs from the otolith organs and semicircular canals.

inputs projects to lateral vestibular nucleus - send axons in the lateral

vestibulospinal tract → to ipsilateral ventral horn of spinal cord.
⇒ directly excite extensor motor neurons ~ inhibit flexor motor neurons -
lead to activation of extensor muscle (antigravity).

Activation of the medial part of the ventral horn by otolith organs - helps maintain
balance and upright posture

~ by innervating extensor muscles in the trunk and limbs.

Decerebrate rigidity - characterized by rigid limb extension → occurs when the

brainstem is transected above the level of the vestibular nucleus.

Lesioning the vestibular nuclei - relieve this rigidity → highlighting the imp. of the
vestibular system in maintaining muscle tone.

Vestibular-cerebellar pathways

Cerebellum - major target of ascending vestibular pathways

→ provides descending input to the vestibular nuclei ~ result in imp. to modulate

vestibular act.

Vestibular-cerebellar circuit - integrating + modulating vestibular signals

→ enable adaptive changes to VOR ~ distinguish head tilts from translational
movements/ passive movement of the head & body.

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 💡 The flocculus and paraflocculus contain Purkinje cells - transmit inhibitory
signals to the vestibular nucleus
→ these neurons change their firing properties in a manner ~ adaptation of
the VOR.

recording in the nodulus & uvula - reveal indv. P cells → integrate

signals from 2 vestibular sources to unambiguously encode head tilts.
translational movements.

Vestibular pathways to thalamus & cortex

The superior and lateral vestibular nuclei - project axons to the ventral posterior
nuclear complex of the thalamus.

→ serves as a relay station for transmitting vestibular information to various cortical

areas ~ responsible for perceiving and processing vestibular sensations.

The thalamic projections - relay vestibular information to cortical areas, including:

Brodmann's area 2v - located just posterior to the face representation in the

somatosensory cortex.

Two regions in Brodmann's area 3a - located in the fundus of the central sulcus.

The parietoinsular vestibular cortex (PIVC) - associated with the perception of

self-motion and spatial orientation.

PIVC - sense of self-motion and orientation in space → elicit strong vestibular

sensations.

Neurons in cortical regions - respond to variety of sensory inputs incl.

proprioceptive, visual & vestibular stimuli
→ reflect the multisensory nature of central vestibular process.

activated by moving visual stimuli + body rotation → cortical regions respond to

perceiving body orientation in extrapersonal space.

The vestibular system 13

 💡 Lesions in the right parietoinsular vestibular cortex - lead to altered
perception of personal and extrapersonal space
→ affecting our sense of self-motion and orientation in the environment.

Spatial orientation perception and multisensory integration

Vestibular sys. - invol. in auto-reflexes ~ its functioning goes unnoticed unless it

damaged.
→ sig. role in perception of spacial orientation & self-motion.

1. detect direction & mag. of self-generated motion ~ differentiate it from

movement of obj. as well as static tilts.

2. works in conjunction w. visual sys. - provide critical info. for spacial orientation +
navigation.

3. may not match the physical attributes of associated stimulus.

eg. person rotating in a chair at a constant speed - perceive the rotation ~
slowing down and eventually stopping → may not directly correspond to the
actual physical rotation.

Vestibular + visual sys. - work together to est. self-motion.

→ visual flow across retina during translational movement - strongly infl. our
perception of self-motion (vection).

💡 Vestibular damage - lacking gravitational cues from otolith organs

→ exp. heightened vection in response to visual flow info.

The vestibular system 14