Anatomy of the Vestibular

System

Djoko Prakosa
Dept. of Anatomy, Embryology &
Anthropology

Vestibular system
Detects motion of the head-in-space and in turn generates reflexes that are
crucial for daily activities, such as stabilizing the visual axis (gaze) +
maintaining head and body posture.
Provides subjective sense of movement and orientation in space.
Located in the petrous part of the temporal bone near the cochlea.

Comprised of two types of sensors:

- macula sacculi and utriculi which sense linear acceleration (i.e., gravity and
translational movements)
static equilibrium
- crista ampularis of semicircular canals, which sense angular acceleration in
three planes.
dynamic equilibrium

They send signals through the vestibular nerve fibers to the

neural structures that control eye movements, posture, and
balance.

Vestibular labyrinth and receptors.

Vestibular signals from the labyrinth are transferred to the vestibular

nuclei (VN). VN is connected to other brain areas: 1) to contralateral
VN, 2) to the abducens nucleus (Abd) to produce the vestibulo-ocular
reflex, 3) to higher brain centers to provide information about spatial
orientation, or 4) to the spinal cord motor neurons that produce
reflexes to stabilize posture.

Afferent Vestibular Pathways

Primary vestibular neurons:
- cell body (vestibular ganglion) in the MAI/IAC
- axons form vestibular nerve
vestibulocochlear
(N. VIII)
enters the brain in the pontocerebellar
angle.
- most axons terminate in the vestibular nuclei
- some pass directly to cerebellum, esp.
flocculonodular lobe and vermis.
- No primary neurons cross midline

Central Vestibular Connections

Vestibular Nuclei (Second order neurons)
- medial (MVN) (Schwalbe)
- superior (SVN) (Bechterew)
- lateral (LVN) (Dieter)
- inferior (or descending) (IVN/DVN)
 Some nuclei receive only primary vestibular afferents
 Most nuclei receive afferents also from cerebellum, reticular
formation, spinal cord, and contralateral vestibular nuclei

SECOND ORDER VESTIBULAR NEURONS

MVN and SVN receive inputs from horizontal and vertical
semicircular canals, respectively. The utricular afferents
terminate in the IVN, and also send projections to the
LVN,MVN, and SVN. Saccule fibers innervate the LVN and
IVN.
Inputs also come from cortical, cerebellar, and other
brainstem structures. They relay somatosensory and visual
inputs and signals related to eye movements and premotor
head movement commands to the vestibular nuclei. These
extra-vestibular inputs modify the processing of vestibular
information at an early stage of sensory processing.

Vestibular nuclei (VN) receive inputs from

multiple brain areas.
A.
Vestibular inputs to the four major
vestibular nuclei.
B.

In addition to inputs from the

vestibular receptors, vestibular
nuclei receive inputs from other
cortical areas, brainstem,
cerebellum, and the spinal cord.

Projections from vestibular nuclei

SVN to nuclei of extraocular muscles (III, IV, VI)
by way of medial longitudinal fasciculus

reaches ipsilateral eye nuclei
LVN
forms vestibulospinal tract
mediate
trunk and limb muscle reflexes
IVN/DVN
to cerebellum
MVN
ascend and descend
ascend bilateral -- to eye muscles
descend to cervical segment of spinal cord

Controlling the axis of gaze

The vestibulo-ocular reflex (VOR) effectively stabilizes

gaze during everyday activities such as walking and
running by moving the eye in the opposite direction
to the on-going head motion. The three neuron arc
consists of projections from vestibular afferents to
neurons in the vestibular nuclei, which in turn
project to extraocular motoneurons.


A. Schematic diagram
of the direct
vestibulo-ocular
reflex (VOR) pathway.
Rotation of the head
to the left excites
neurons in the left
vestibular nuclei.
Position-vestibularpause (PVP) neurons
send excitatory
projections to
contralateral
abducens (ABD)
motoneurons and
inhibitory projections
to ipsilateral
abducens neurons to
generate eye
movements to the
right. Inset, the
latency of the eye
movements evoked
by the VOR is ~5 ms.

Controlling the vestibulo-spinal and

vestibulo-collic reflexes

The vestibular system also coordinates postural reflexes.
- Vestibulo-collic reflex (VCR) maintaining head and body
posture. VCR functions to stabilizes the head relative to
inertial space by generating a command to move the head in
the opposite direction to that of the current head-in-space
velocity.

These neurons also most likely control vestibulo-spinal
reflexes since neurons that project to the cervical spinal cord
can have multiple axon collaterals which also project widely
to other segments in the spinal cord (i.e. thoracic and
lumbar).

HIGHER ORDER VESTIBULAR PROCESSING

Vestibulo-cerebellum
There are five main regions of the cerebellum that receive either primary
(i.e. from afferents) or secondary (i.e. from vestibular nuclei) vestibular
input including:
i) the nodulus and ventral uvula,
ii) the flocculus and ventral paraflocculus,
iii) the oculomotor vermis of posterior lobe,
iv) lobules I-V of the anterior lobe, and
v) the deep cerebellar nuclei.

Vestibulo - Cerebellum
i) The vestibular nuclei are reciprocally interconnected with the
nodulus/uvula. These areas make significant contributions to the
computation of inertial motion
ii) The flocculus and paraflocculus are involved in the generation and
the plasticity of compensatory eye movements, including visual
ocular following reflexes (i.e. smooth pursuit and the optokinetic
reflex) and the VOR.
iii) Neurons in lobules VI and VII (oculomotor vermis) contribute to
visual-vestibular processing. In addition, it receives eye movement
signals from the nucleus preopositus as well as pursuit-related inputs
from the dorsolateral pontine nuclei -

Vestibulo cerebellum
Lobules I-V encodes both vestibular and neck proprioceptive-related
signals and is thought to control of vestibulo-spinal reflexes. The
integration of vestibular and proprioceptive information ensures that
the motor responses produced by these reflexes are appropriate to
maintain body stability.
v) The signal processing done in the fastigial nucleus of the deep
cerebellar nuclei is tightly linked to the vestibular system. It receives
both primary and secondary vestibular inputs as well as input from
the cerebellar vermis. The fastigial nucleus plays an important role in
the generation of postural reflexes and orienting behaviors, and
accordingly projects to brainstem structures that control these
behaviors including the vestibular nuclei and medial reticular
formation. Many neurons in this area integrate vestibular and
propiciocpetive inputs, and in turn, encode vestibular signals in a
body-centered reference frame.
iv)

Vestibular thalamus and cortical projections

Neurons in several areas of cortex respond to vestibular stimulation:

- area 2v of the intraparietal sulcus,

- area 3a in the sulcus centralis

- ventral intraparietal area (VIP)
- medial superior temporal area (MST)
- parietoinsular vestibular cortex (PIVC)

Areas 3a, T3, and PIVC receive vestibular projections through thalamus

PIVC is considered as the main vestibular cortex:
- information from other vestibular cortical areas converge in PIVC
- PIVC receive vestibular input
- stimulation of this area produces vestibular sensation
- lesions of this area impairs perception of subjective vertical
- vestibular aurae in epileptic patients is related to activation of this area
- cerebral blood flow of PIVC area increases during vestibular stimulation

Vestibular thalamus and cortical projections (cont.)

Cortical processing of vestibular input is important for generating
appropriate motor response and provides subjective sense of movement
and orientation in space. Neurons in area VIP of area 7 show differential
activity in response to active and passive head movements including
changes in the strength, timing, and direction selectivity of their responses
under the two conditions. This differential encoding of vestibular
information is important for shaping appropriate motor responses to
guide voluntary movements. Most recently, studies have focused on how
the responses of neurons in the dorsal medial superior temporal cortex
(MSTd) contribute to our perception of self-motion. This cortical area has
long been known to process optic flow information. Experiments
demonstrating a functional link between area MSTd and heading
perception based on vestibular signals further support the proposal that
that MSTd plays a role in self-motion perception.
As a final point, corticofugal projections project back to vestibular nuclei,
thereby also modulating vestibular processing at an earlier stage. How
these feedback projections influence sensory processing at the level of the
VN remains to be determined.

Figure 9: Schematic representation of vestibular

cortical areas. A. Areas of cortex that receive
inputs from vestibular nuclei. B. Cortical areas
that project back to the vestibular nuclei. FEF is
the frontal eye field, MST is medial superior
temporal, VIP is the ventral intraparietal, PIVC is
the parietoinsular vestibular cortex. Striped areas
are deep cortical areas.

Vertigo
Vertigo is a sense of derangement in position and motion of the body or
its mental phenomenon, and it is physically manifested as disequilibrium
and symptoms of dysautonomia.
When unusual stimuli are given to the system of equilibrium maintenance
or the system is damaged, otitic/ocular/cervical vertigoes, psychogenic
vertigo, etc. are generated.
Various vertigoes were generated experimentally by giving a rotary motion
stimulus, a linear motion stimulus, etc., to observe characteristics of
vertigoes through balance tests.
Stimuli to organ of vision, vestibule/semicircular ducts, etc. induce
attitudinal reflex in eyeball/limb/trunk, work to maintain the body balance
and are projected to multiple vestibular cortical centers at the same time,
to construct images of position and motion. When such functions are
damaged, disequilibrium is generated physically, and confusion in
information (vertigo) is made in vestibular cortical centers.

Left. Vestibular afferent input during normal horizontal head rotation to the right. Increased firing rate
from right peripheral vestibular apparatus. Ocular deviation shows slow phase deviation to the left. VN
= vestibular nuclei.Right: Acute left peripheral vestibulopathy with resultant acute vertiginous sensation
simulating head rotation to the right. Slow phase ocular deviation to the left (small arrow) and fast
phase of nystagmus to the right (bold arrow) and away from the side of the peripheral vestibular injury.

Normal adaptation for prior left peripheral vestibulopathy. Despite a reduced firing rate
from the left side, the central nervous system (CNS) has compensated for the disparity and
there is no nystagmus or vertigo. (Right) Abnormal compensation for prior left peripheral
vestibulopathy. The patient continues to experience vertiginous sensations and may have
nystagmus with a fast phase to the right (solid arrow).