Vestibular system

The set of five organsthree semicircular canals and

two otolith organs (utricle and saccule) located in each
inner ear that sense head motion and head orientation
with respect to gravity.

Vestibulo-ocular reflex
(a) As your fingertip moves
faster and faster in front of
your face, the fingertip
begins to blur.
(b) When you shake your
head back and forth, the
fingertip remains clearer.
Vestibular system
reflexively rotates eyeballs
in the sockets to compensate
for head rotationthereby
helping to keep visual
images stable on the retina.

Spatial orientation: where is our head and how is it

moving?
Three interacting sensory modalities.
Our senses of:
Angular motion: For example, rotating
head from side to side as if to say no.
Linear motion: For example,
accelerating or decelerating in a car.
Tilt: For example, nodding head up and
down as if to say yes or left to right
over the shoulders.
Otolith organs (utricle and saccule): The mechanical
structures in the vestibular system that sense both
linear acceleration and gravity.
Semicircular canals: The three toroidal tubes in the
vestibular system that sense angular motion.

Coordinate system for classifying direction

x-axis: Points forward and backward, in the direction the
person is facing.
y-axis: Points laterally, out of the persons ears.
z-axis: Points vertically, out of the top of the head and
feet.
Axes are defined relative to the person, not relative to
gravity.

Three directions for sense of angular (rotational) motion

Linear motion
Movements represented in terms of changes in the
x-, y-, and z-axes.
All linear motions can be represented as a change
along these three axes.

Walking forward

Side-stepping

Going up in an elevator

Tilt: When initially upright with respect to gravity there are two directions of tilt.

Unlike the roll and pitch with angular motion, these two are movements
with respect to gravity. There is no yaw because moving in that direction
would not change the tilt of the head with respect to gravity.

The utricle senses horizontal displacement (moving in a car).

The saccule senses vertical displacement (elevator).

Displacement of the Otolithic Membrane of the Utricle

Otolith Organs take their name from otoconia (ear stones).
Otolithic membrane of the utricle is displaced with
head tilt and linear acceleration
After reaching constant velocity, otoliths catch up
with the head and so there is no sense of constant velocity.

Upright

HOW THE UTRICLE detects both STATIC DISPLACEMENTS of the

head and linear accelerations.
Head tilt backwards--excite half of cells in each utricle, inhibit half.
Orientation of hair cell bundles: half facing each way oriented around the
STRIOLA.
Of course, half of the ones on the other side of the head are also excited,
half inhibited.

Figure 15.7 Hair cell responses

Hair cell responses

In the absence of stimulation, hair cells
release neurotransmitter at a constant rate.
When hair cell bundles bend, change in hair
cell voltage is proportional to the amount of
deflection.
Bending toward tallest stereocilia:
Depolarization
Bending away from tallest stereocilia:
Hyperpolarization
Hair cells increase firing to movement in one
direction and decrease firing to rotation in the
opposite direction.

Coding of direction in the semicircular canals

Three semicircular canals in each ear
Each canal is oriented in a different plane
Each canal is maximally sensitive to rotations
perpendicular to the canal plane:
Anterior: Pitch, Posterior: Roll, Horizontal: Yaw

Fluid Displaces the Cupula During Rotational Movement

stationary

SEMICIRCULAR CANALS detect ANGULAR ACCELERATION.

Inside the ampulla: the hairs extend into an overlying cap-like gelatinous layer called the
CUPULA.
The cupula protrudes into the overlying fluid called the ENDOLYMPH within the ampulla.
The cupula sways in the direction of fluid movement like seaweed in ocean waves.
Movement of the head to the left (B) moves the endolymph to the right which bends the hairs
toward the tallest stereocilium and thus depolarizes the hair cells.

Pushpull symmetry
Hair cells in opposite ears respond in a complementary fashion to each
other.
When hair cells in the left ear depolarize, those in the analogous structure
in the right ear hyperpolarize.

Another look at pushpull symmetry

Example: horizontal canals.
Rotation of head clockwise as viewed from the top.
Fluid movement in both canals is OPPOSITE the
movement of the head, because of inertia of the fluid.
This causes the cupulas in the two canals to both
bend in the direction of the head movement.
But because the hair cells are oriented oppositely,
one of these will be stimulated, the other one
depressed.
WHICH IS WHICH?
The hair cells in the canal that is toward the direction
that the head is moving are depolarized, those on the
opposite side are hyperpolarized.
In this example, the right side is depolarized,
indicating clockwise.
Opposite rotation would give the opposite effect.

Throwing Cold Water on the Vestibular System

A view of the activity of afferent nerves in

response to rotation of the head.

Head moves clockwise.

Fluid moves counter-clockwise.

Afferent fiber activity increased on right side.

Afferent fiber activity decreased on left side.

Coding of amplitude in the semicircular canals

In the absence of any rotation, afferent neurons from the semicircular canals
have a resting firing rate of about 75 spikes/s
High firing rate allows canal neurons to code amplitude by decreasing their
firing rate, as well as increasing it
Neural activity in semicircular canals is sensitive to changes in rotation velocity
Constant rotation leads to decreased responding from the canal neurons after a
few seconds. At constant velocity the endolymph catches up with the head.

Pathways for the VOR

Connections underlying the vestibulo-ocular reflex. Example, head moves counterclockwise

(left) and eyes move right. Left medial rectus and right lateral rectus muscles contracted.
Same head movement inhibits left lateral and right medial rectus muscles.
http://www.tutis.ca/Senses/L10Balance/L10Balance.swf Go to labyrinth, otoliths, canals, VOR

Physiological nystagmus: (normal) movement

of the eyes in response to head movement.
Combination of slow eye movement (VOR)
and rapid eye movement in opposite direction
to reset the eyes--called SACCADES.
Firing rate changes reflect head movement.
Normal--physiological nystagmus
Spontaneous: caused by damage to one side
or other: eye movements even when head is
still.
Low level of baseline firing of undamaged
side causes eye movements.

Learn about the vestibular system:

http://sites.sinauer.com/wolfe3e/chap12/vestibularF.htm