Sense organ:vestibular organ

• The purpose of the vestibular system is to keep tabs on the

position and motion of your head in space.
• There are really two components to monitoring motion,
however.
• You must be able to detect rotation, such as what happens
when you shake or nod your head.
• In physics, this is called angular acceleration.
• You must also be able to detect motion along a line - such as
what happens when the elevator drops beneath you, or on a
more subtle note, what happens when your body begins to
lean to one side.
• This is called linear acceleration.
• The vestibular system is divided into two receptor organs to
accomplish these tasks.
• The semicircular canals detect angular acceleration.
• There are 3 canals, corresponding to the three dimensions in
which you move, so that each canal detects motion in a single
plane.
• Each canal is set up as shown below, as a continuous
endolymph-filled hoop.
• The actual hair cells sit in a small swelling at the base called the
ampula. The hair cells are arranged as a single tuft that projects
up into a gelatinous mass, the cupula.
• When you turn your head in the plane of the canal, the inertia
of the endolymph causes it to slosh against the cupula,
deflecting the hair cells. Now, if you were to keep turning in
circles, eventually the fluid would catch up with the canal, and
there would be no more pressure on the cupula.
• If you stopped spinning, the moving fluid would slosh up against
a suddenly still cupula, and you would feel as though you were
turning in the other direction.
• Naturally, you have the same arrangement (mirrored) on both
sides of the head. Each tuft of hair cells is polarized - if you
push it one way, it will be excited, but if you push it the other
way, it will be inhibited.
• This means that the canals on either side of the head will
generally be operating in a push-pull rhythm; when one is
excited, the other is inhibited
• It is important that both sides agree as to what the head is
doing. If there is disagreement, if both sides push at once,
then you will feel debilitating vertigo and nausea.
• This is the reason that infections of the endolymph or damage
to the inner ear can cause vertigo.
• However, if one vestibular nerve is cut, the brain will gradually
get used to only listening to one side - this can actually be a
treatment for intractable vertigo.
• A large role of the semicircular canal system is to keep your eyes
still in space while your head moves around them.
• If you nod and shake and swivel your head, you will find that
you have no trouble staying focused on this page.
• But hold a piece of paper in front of you and shake it around,
and your eyes will not be able to keep up with the quick
movements.
• The reason is that the semicircular canals exert direct control
over the eyes, so they can directly compensate for head
movements.
• Recall that the eye is controlled by three pairs of muscles; the
medial and lateral rectus, the superior and inferior rectus, and
the inferior and superior oblique.
• You may also remember that their directions of motion seemed
to be at crazy diagonals.
• Those same crazy diagonals are matched closely by the three
planes of the semicircular canals, so that a single canal (in
general) interacts with a single muscle pair.
• The entire compensatory reflex is called the vestibulo-ocular
reflex (VOR).
• Although the VOR works on all three muscle pairs, the medial-
lateral rectus pair, coupled to the horizontal canal, is
geometrically the easiest to draw.
• Here is the setup, looking down at a person's head: The lateral
rectus muscle will pull the eye laterally, and the medial rectus
will pull the eye medially, both in the horizontal plane. The
horizontal canal detects rotation in the horizontal plane.
• If you move your head to the left, you will excite the left
horizontal canal, inhibiting the right.
• To keep your eyes fixed on a stationary point, you need to fire
the right lateral rectus and the left medial rectus, to move the
eyes to the right.
• The pathway is as follows:
• the vestibular nerve enters the brainstem and synapses in the vestibular
nucleus.
• Cells that received information from the left horizontal canal project to
the abducens nucleus on the right side, to stimulate the lateral rectus.
• They also project to the oculomotor nucleus on the left side, to stimulate
the medial rectus. Although not shown on the diagram, the same
vestibular cells also inhibit the opposing muscles (in this case, the right
medial rectus, and the left lateral rectus).

• What about the other side?

• The right horizontal canal is wired to the complementary set of muscles.
Since it is inhibited, it will not excite its target muscles (the right medial
rectus and the left lateral rectus), nor will it inhibit the muscles you want
to use (the right lateral rectus and the left medial rectus). Got it? OK, then
draw out what would happen if you turned your head to the right.

• A great deal of the VOR axon traffic travels via a fiber highway called the
MLF (medial longitudinal fasciculus). The integrity of this tract is crucial for
the VOR to work properly. It is occasionally damaged by medial brainstem
strokes.
• The utricle and saccule detect linear acceleration.
• Each organ has a sheet of hair cells (the macula) whose cilia are
embedded in a gelatinous mass, just like the semicircular canals.
• Unlike the canals, however, this gel has a clump of small crystals
embedded in it, called an otolith (yes, all along you've had rocks in your
head).
• The otoliths provide the inertia, so that when you move to one side, the
otolith-gel mass drags on the hair cells. Once you are moving at a constant
speed, such as in a car, the otoliths come to equilibrium and you no longer
perceive the motion.

• The hair cells in the utricle and saccule are polarized, but they are arrayed
in different directions so that a single sheet of hair cells can detect motion
forward and back, side to side. Each macula can therefore cover two
dimensions of movement.
• The utricle lays horizontally in the ear, and can
detect any motion in the horizontal plane.
• The saccule is oriented vertically, so it can detect
motion in the sagittal plane (up and down,
forward and back).
• A major role of the saccule and utricle is to keep
you vertically oriented with respect to gravity. If
your head and body start to tilt, the vestibular
nuclei will automatically compensate with the
correct postural adjustments.
• This is not just something that happens if the
floor tilts - if you watch someone trying to stand
still, you will notice constant small wavers and
rocking back and forth.
• There are four vestibular nuclei within the brain stem (superior, lateral,
medial, and inferior).
• All four can not be seen in the same cross section, since they are present
for a considerable rostrocaudal distance from the rostral medulla to the
middle of the pons.
• You only have to be able to identify the MEDIAL and INFERIOR vestibular
nuclei, both of which are present at level #4 (shown below on the left).
The vestibular nuclei receive their primary input from the vestibular
portion of C.N. VIII (vestibular-auditory).
• The axons in the vestibular nerve are the central processes of neurons that
lie in the vestibular or Scarpa’s ganglion which lies in the internal auditory
meatus.
• These central processes terminate in the vestibular nuclei and the
cerebellum. The peripheral processes of these cells receive information
from the receptors of the vestibular labyrinth, i.e. hair cells located in the
semicircular canals and the saccule and the utricle (otolith organs).
• The three (on each side) membranous semicircular canals lie
within the bony labyrinth and contain endolymph.
• As shown in two of the drawings below, the canals, one
horizontal and two vertical, lie in three planes that are
perpendicular to each other.
• The HORIZONTAL or lateral canals on the two sides lie in the
same plane, while the plane of each anterior canal is parallel
to that of the posterior canal of the opposite side.
• The horizontal semicircular canals communicate at both ends
with the utricle, which is a large dilation of the membranous
labyrinth. The vertical canals (anterior and posterior)
communicate with the utricle at one end, and join together at
the other end (the common canal communicates with the
utricle).
• At one end of each semicircular canal is a dilation called the ampulla (L.,
little jar, is labeled “amp” in the upper right drawing below).
• The ampulla of a horizontal semicircular canal has been enlarged in the
drawing below (upper left). Each ampulla contains a crista (crista
ampullaris; ridge), which is a transversely oriented ridge of tissue.
• The upper surface of the crista contains ciliated sensory hair cells that are
embedded in a gelatinous material called the cupula (L., little tube). These
ciliated sensory hair cells contain vesicles that possess neurotransmitter.
• When the neurotransmitter is released from the hair cell, the peripheral
process of a cell in the vestibular ganglion is turned on.
• Interestingly, the hair cells release transmitter even when they are not
stimulated, so the axons in the vestibular nerve are always firing at a
baseline rate.
Each hair cell of the crista possesses several shorter stereocilia and a
single tall kinocilium at one margin of the cell as shown in the lower
figure. Deflection of the stereocilia TOWARD the kinocilium results in an
INCREASE in the firing rate of the vestibular fiber associated with the hair
cell, while deflection AWAY from the kinocilium results in a DECREASE in
the firing rate of the vestibular fiber.
• While semicircular canals respond to angular acceleration in specific
directions, hair cells in the utricle and saccule respond to linear
accelerations. The utricle and saccule are saclike structures that contain a
patch of sensory hair cells called the macula (L., spot). The hair cells in the
macula, which are similar to those in the cristae, are embedded in the
otolith (ear stone) membrane, a gelatinous structure that contains a large
number of hexagonal prisms of calcium carbonate called otoconia (ear
dust). Since the density of the otoconia is greater than the surrounding
endolymph, the otolith membrane will be displaced by the force of gravity
or other linear accelerations. Such displacement bends the stereocilia and,
depending on the polarity of the cell, either causes an increase or a
decrease in the number of impulses in the associated vestibular fiber.
• Once vestibular input from the semicircular canals and otolith
organs has reached the vestibular nuclei, the information is
used to maintain balance and to stabilize the visual image on
the retina during head movements.
• First we will consider only the projections of the vestibular
nuclei that reach the spinal cord in order to help us maintain
our BALANCE.
• For example, let’s say that you are walking to lecture this
morning and slip on the icy sidewalk. Your feet fly to the
RIGHT and your upper body and head fly to the LEFT (left ear
down). Information coming out of your semicircular canals
will be related to the accelerating head.
• This particular angular acceleration affects several different
canals (beyond what you need to know for this course), but
what I want you to know is that the LEFT vestibular nuclei are
turned on.
• Once your head is not moving the information will come from
the utricles. Again, you do not have to know the specific
pattern from each side, but only that the LEFT vestibular
nuclei are turned on.
• The increased activity in the LEFT vestibular nuclei can affect the body musculature
via the LEFT LATERAL VESTIBULOSPINAL TRACT. This will result in increased activity
in the LEFT arm and leg in order to right ourselves after slipping. The cells of origin
of the lateral vestibulospinal tract lie in the lateral vestibular nucleus (you can not
see this nucleus in your sections). Axons arising from this nucleus descend through
the caudal brain stem (you don’t see these fibers on the cross sections) and upon
reaching the spinal cord course within the ventral funiculus and innervate neurons
for the ENTIRE length of the cord. This projection is UNCROSSED. Through this
tract, the vestibular apparatus—which detects whether the body is on an even
keel—exerts its influence on those muscles that restore and maintain upright
posture. Such muscles are proximal rather than distal.
• The increased activity in the LEFT vestibular nuclei can also affect body
musculature via a second, smaller, descending pathway to the spinal cord.
• This smaller pathway is called the MEDIAL VESTIBULOSPINAL TRACT (or
descending medial longitudinal fasciculus [MLF]).
• Cells within the medial vestibular nucleus possess axons that descend
bilaterally (the ipsilateral projection is denser) in a position just off the
midline near the dorsal surface of the pons and medulla. These
descending axons course caudally and enter the spinal cord, where they lie
within the medial part of the ventral funiculus.
• This pathway makes connections with cervical and upper thoracic motor
neurons that play a role in maintaining the normal position of the head via
innervation of spinal cord neurons that innervate neck musculature.
• Thus when your head flies to the LEFT, it will reflexively be brought to an
upright position via information flowing out of the LEFT medial vestibular
nucleus.
• REMEMBER—LATERAL VESTIBULAR NUC.—
LATERAL VESTIBULOSPINAL TRACT—
UNCROSSED—ENTIRE LENGTH OF CORD—
VENTRAL FUNICULUS—PROXIMAL MUSCLES—
MAINTAINS BALANCE BY ACTING MAINLY ON
LIMBS

• REMEMBER—MEDIAL VESTIBULAR NUCLEUS—

MEDIAL VESTIBULOSPINAL TRACT—BILATERAL—
CERVICAL AND UPPER THORACIC SPINAL CORD
ONLY—MAINTAINS HEAD ERECT.