VM8054 Veterinary Histology

Special Senses II: The Ear

Author: Dr. Thomas Caceci
Objectives for This Exercise
Special Senses Introductory Page

The ear is an extraordinarily complex organ

with two functions: sound reception and
maintenance of positional equilibrium.
Although at first it might seem odd that both
of these functions should be combined in one
organ, as we'll see in this exercise, the logic
of doing so becomes obvious upon closer
examination. In both cases, nervous impulses
are generated by the physical displacement of
"hairs" inside the ear, and the mechanics of
the auditory and balance sense are
very similar.
Regions of The Ear
The ear has three distinct parts:
the outer ear (the pinna and
external auditory meatus); the middle ear
(the tympanic cavity, the auditory ossicles,
and the Eustachian tube); and the inner ear

(the sealed, fluid filled chambers responsible

for transduction of sound into nervous
impulses and for providing information about
body position).
In this exercise we'll look mainly at the inner
ear, with a view to understanding the
relationship between its microscopic
structure and its twin functions. We'll not
deal with the middle ear at all, and only
briefly with the external ear. Those parts are
best studied using gross anatomical
techniques.
It's exceedingly difficultheck, no, it's flat
out impossibleto visualize the three
dimensional structure of the inner ear solely
from two dimensional microscope slides. You
are urged to compare what you see on the
slides with the model provided.
The Labyrinths of the Ear
The classic description of the ear includes a
discussion of the two "labyrinths" of which
it's composed.
The bony labyrinth consists of the convoluted
channels in the actual bone of the skull.
Inside these channels in bone sits the
membranous labyrinth, components of which

are the actual functional structures for

balance and hearing.
It's necessary to remember that the one
labyrinth is inside the other. Their
relationship is like that between an
automobile tire and an inner tube; the tire is
the bony labyrinth and the inner tube the
membranous labyrinth.
The bony labyrinth is divisible into several
regions: the vestibule, the semicircular
canals and the cochlea. The first two contain
those parts of the membranous labyrinth that
are involved in the balance sense. The last
part contains the portion of the membranous
labyrinth that is involved in hearing
perception.
There is considerable confusion about the
cochlea, by the way. It's NOT the place in
which transduction of sound occurs. That
happens in the cochlear duct, which is that
part of the membranous labyrinth inside the
bony cochlea.
Similarly, the semicircular canals, contrary to
what you may have been told, aren't the
actual site of balance sensation. That occurs
in semicircular ducts, elements of the
membranous labyrinth which run through

those canals. This may seem to be

hairsplitting, perhaps, but it's not. There are
important functional and anatomic
distinctions between the two labyrinths.
Perilymphatic Spaces
The vestibule, cochlea, and semicircular
ducts all are connected. The vestibule is a
fairly large space, and coming off it are the
bony cochlea and the semicircular canals.
The cochlea is a corkscrew shaped
passageway through the bone of the skull,
and the semicircular ducts are simple loops
that come off and return to the vestibule.
The bony labyrinth contains a small amount
of fluid, the perilymph that fills it and in
which the structures of the membranous
labyrinth are bathed. Perilymph somewhat
resembles extracellular fluid. To return to
the tire/tube analogy, it's as if there were a
small amount of water in the space between
the inside surface of the tire and the outside
surface of the tube. It acts as a shock buffer
and a means of transmitting vibrations to
parts of the membranous labyrinth.
The perilymphatic spaces are functionally
connected to the subarachnoid space in the
meningeal covering of the brain, and this may

be the route by which excess perilymph is

removed. The actual site of perilymph
production is a matter of debate.
Endolymphatic Spaces
The actual perception of balance,
acceleration, and sound transduction all take
place inside the spaces of the membranous
labyrinth.
The membranous labyrinth, too, is filled with
fluid. This fluid is endolymph, and it's very
different in composition from perilymph.
Endolymph is secreted by the cells making up
the lining of the membranous labyrinth, and
(unlike perilymph or tissue fluid) it's high in
potassium and low in sodium and soluble
proteins. The functional significance of the
differences between these two fluids is not
clear.
The Apparatus of Sound Transduction
Let's first examine the apparatus for sound
transduction. Begin with slide 1206, which
shows most of the structures of the inner ear.
At low magnification, you will easily recognize
that most of this section consists of a chunk

of bone, with various shaped cavities in it.

These cavities are the bony labyrinth, and
within them are the membranous labyrinth
structures.
To see this region, click here.
Cochlear Spiral: Scala Vestibuli, Scala
Tympani, Scala Media
Look first at the cochlea and the cochlear
duct which fills it. The cochlea is a spiral
channel through the bone, and in the middle
of it is a large bony projection which looks
something like a wood screw, cut
longitudinally. This is the modiolus. Resting
inside the cochlea is the cochlear duct, in
which the events related to sound
transduction occur. The duct follows the
spiral of the cochlear lumen.
To see this in some more detail, click here.
The cochlear duct is filled with endolymph,
but the cochlea which surrounds is filled with
perilymph. (The former is membranous
labyrinth, the latter is bony labyrinth.) In
cross section, each turn of the spiral shows
three chambers; the first is the scala media,
which is the cochlear duct itself, and in which

are located the parts of the sound

transduction apparatus.
Above the scala media, and separated from it
by the very thin vestibular membrane is the
scala vestibuli. This is continuous with the
vestibule, another perilymphatic space, and
so of course the fluids in the two areas are in
continuity.
Below the scala media, and separated from it
by the thick basilar membrane, is the scala
tympani. The scala tympani is also a
perilymphatic space, is also in
communication with the vestibule, and is also
in communication with the scala vestibuli.
The best analogy I can think of here is the
double spiral ramp of a vertical parking
garage. Think of the scala vestibuli as the
"UP" ramp, corkscrewing its way to the top of
the garage, and of the scala tympani as the
"DOWN" ramp, doing exactly the reverse. If
you were to enter the garage at the bottom
you could drive to the top, where the two
ramps meet, and come back down again into
the lobby of the garage (i.e., the vestibule).
To complete the analogy, think of the scala
media as a water filled pipe lying between the
two ramps. Remember this picture, as we will

come back to it later in considering the

mechanism of sound transduction.
Scala Media & Organ of Corti
Inside the cochlear duct (scala media) itself
is the actual organ of sound transduction.
This is the organ of Corti (Marquis Alphonso
Corti, 1822-1888, an Italian anatomist). The
organ of Corti is a long, flat, spiral structure
which follows the turns of the cochlear duct
as it winds up to the top of the cochlear
spiral. It is inside an endolymphatic space.
When the organ of Corti is viewed in cross
section (as you see it in this slide) you'll see
two groups of cells sitting on the basilar
membrane; these are surmounted by a thin,
transparent, and acellular tectorial
membrane. The tectorial membrane is made
of a keratin-like material, and in H&E
sections rather resembles a piece of
fingernail. It's anchored at one end to the
wall of the cochlear duct, and it's quite stiff.
It, too, is spiraled and it covers the two
groups of cells all the way up to the top of the
spiraling scala media.
To see an example of this, click here.

The cells in question are hair cells and they

are the actual transducers of mechanical
movement into neural impulses. Each is an
epithelioid cell with projecting "hairs" that
contact the bottom of the overlying tectorial
membrane. It's important to realize that
these aren't "hairs" at all, of course; they are
microvilli and modified cilia. It should be
fairly obvious that any movement of the
flexible basilar membrane will deform the
hair cell projections against the tectorial
membrane. The arrangement is what you
would have if you laid a playing card against
the bristles of a toothbrush. Any movement
of the brush will cause its bristles to scrape
along the bottom of the fixed, stiff card.
The hair cells are transducers, specialized
detector elements, and they're surrounded by
sensory fibers of neurons whose cell bodies
are in the spiral ganglion. The spiral
ganglion is sensory and the neurons in it are
of the pseudounipolar type, just as with any
other craniospinal ganglion. Deformation of
the hair cell projections causes changes in
their membrane potentials, which are
detected by the cells of the spiral ganglion.
Thus, mechanical movement of the basilar
membrane relative to the fixed tectorial

membrane results in the generation of a

nervous signal.
Mechanics of Sound Detection
Let's at this point go back to the parking
garage model. Imagine you are driving up
the UP ramp (the scala vestibuli) to the top,
and the floor of the ramp is flexing beneath
the weight of your car. The movement is
mechanically transmitted to that drainpipe
underneath the ramp. Then you reach the
top of the spiral, cross over to the DOWN
ramp, and as your car passes along on the
way back to the lobby, the air it displaces
impinges on the drainpipe from the
underside. This is a crude depiction of the
method by which mechanical movement of
fluids is transduced into sound perception.
Sound waves coming in via the external
auditory meatus cause the eardrum to move
back and forth. In the middle ear, a series of
small bones constituting a lever system
magnifies this initial signal
and mechanically transmits it to an opening
in the side of the vestibule. The pulsations of
this movement are then sent into the
perilymph as actual fluid waves.

The waves induced in the perilymph of the

vestibule travel UP the scala vestibuli to the
top of the cochlea. Since the scala tympani
and the scala vestibuli are in communication
at the top of the spiral (the term for this area
is the helicotrema), the fluid pressure wave in
the perilymph "crosses over" and starts back
DOWN again, via the scala tympani. Hence,
there is perilymphatic fluid moving on both
sides of the scala media.
All this sloshing back and forth in the
perilymph causes the suspended cochlear
duct (which is filled with endolymph) to
vibrate as well. This inevitably moves the
basilar membrane up and down, causing
scraping of the hair cells along the bottom of
the tectorial membrane. The hair cells send
the resultant signal out via the cells of the
spiral ganglion.
The cochlear nerve consists of the axonal
projections of the cell bodies in the spiral
ganglion and, like any other sensory nerve,
runs back into the brain. In this case, it
becomes one of the components of cranial
nerve VIII, the vestibulocochlear nerve.
Balance Sensation

So far, nothing has been said about the

organs involved in detection of balance and
acceleration. The mechanics of the balance
sense, while different in detail, are similar in
many ways to those of sound detection. Both
involve hair cells as transducers and both
require fluid movement. Both occur in
endolymphatic spaces. Both send their
output to the central nervous system via
pseudounipolar neurons of the spiral
ganglion.
The Vestibular Apparatus
We have repeatedly referred to the vestibule,
that portion of the bony labyrinth from which
the cochlea and the semicircular canals
arise. As with other bony labyrinth, the
vestibule and the canals are filled with
perilymph, and in them are suspended parts
of the membranous labyrinth, constituting
the vestibular apparatus of balance sensation.
As is the case with the cochlear duct, the
vestibular apparatus is filled with endolymph
and sealed off from the surrounding
perilymphatic spaces. The vestibular
apparatus consists of three large dilated
areas and three semicircular ducts which run
through channels of the bony labyrinth.

Semicircular Ducts
The semicircular ducts (and the semicircular
canals through which they run) are oriented
in three planes of movement: vertical,
horizontal, and anterior-posterior. Inside the
semicircular ducts are elevated areas of
epithelium, covered with hair cells similar to
those of the organ of Corti. These are the
cristae ampullares, and they can be thought
of as analogous to speed bumps in a roadway,
because they project out into the lumen of
the semicircular duct. You will see one of
these structures on slide 1206.
To see an example of a semicircular duct
inside a semicircular canal, click here.
Now look at the crista itself. It's covered with
a pseudostratified epithelium composed of
hair cells and some supporting cell types.
The hair cells, like those of the sound
detection system, are transducer cells with
deformable elements on the free surface. At
their bases, they are surrounded by the ends
of nerve fibers from the vestibular branch of
the vestibulocochlear nerve. The hairs in this
organ are polarizedthat is, they are of
different lengths, so that there is a

directionality in their arrangement with

respect to the axis of the semicircular duct.
Movement of the fluid in the semicircular
duct will obviously cause deformation of the
hairs, and this in turn will cause
depolarization of the hair cells and a change
in their membrane potential. This change is
transmitted via the sensory fibers and the
vestibulocochlear nerve to the brain. As you
can see, this is the same sort of mechanism at
work in hearing; mechanical movement of
fluid is translated into neural impulses.
Movement of the head in any direction causes
the fluid in the semicircular ducts to flow
relative to the hairs, and sends the sensation
of movement to the brain.
To see another example of a crista ampullaris,
click here.
If you examine slide 1206 carefully, you'll be
able to see the vestibular branch of the nerve
where it emerges from the bone, and joins the
cochlear branch to form the
vestibulocochlear nerve coming from the
spiral ganglion.
Function of the System

This system is designed to detect acceleration

i.e., changes in speed or direction over a
given time. Any movement of the head
relative to the three planes in which the
semicircular ducts are fixed produces a
corresponding movement of the endolymph
inside them. This deforms the hairs and
provides information to the brain about the
direction of movement. However, if the
motion becomes constant, (that is, if
acceleration stops) movement becomes
constant, the fluid "catches up" to the
movement of the head, and the stimulus
ends. You experience this every time you
drive your car away from a stoplight: when
your motion settles down to a steady speed,
you no longer perceive the acceleration
because it has ceased.
This can have unexpected consequences.
Many other clues to position and acceleration
exist, principally visual input. It's necessary
for the brain to sort these inputs correctly
and decide what the body is really doing, and
how it's moving. Pilots frequently have
problems with conflicts between the
information coming from visual and
vestibular inputs, especially if they swivel
their heads in the cockpit during some
aerobatic maneuver. This introduces odd

forces on the vestibular apparatus, and the

pilot gets disoriented. The brain can't sort
out a reasonable solution to the questions,
"Where am I? Which way am I moving?" You
get the same effect by spinning rapidly on a
carousel or ice skates and then stopping; the
visual input becomes stable, but the
vestibular sensation tells you are still
moving. This results from a persistence of
the vestibular signal even after motion has
ceased. Since the integration of the signals is
done in the CNS, the system is (as you might
expect) affected by alcohol and many other
drugs.
Which Way Is Down?
There is one other portion of the vestibular
system not on your slides, but a
demonstration has been provided. This is the
maculae. The maculae are located in the
large bulbous expansions of the vestibular
system (also filled with endolymph), and their
function is to provide a reference to the pull
of gravity.
The expanded portions of the membranous
labyrinth are the saccule and the utricle.
Both these regions contain areas with hair
cells similar to those in the cristae.

Gravitational detection is clearly an

important consideration for any terrestrial
animal (especially arboreal ones).
Unlike the semicircular duct system, the
sensory areas of the maculae don't depend on
fluid movement. The maculae have within
them small calcareous secretions, the
otoliths, or "ear stones," which lie on top of
the hairs of the hair cells. Otoliths are
formed in the embryo as part of normal
development.
To see examples of the maculae and the
otoliths, click here.
When an animal is stable with respect to
gravity, the force the otoliths exert on the
hairs is more or less constant, and the brain
interprets this as "I am not moving
downward, I'm sitting still on the ground."
If, however, the animal falls, the otoliths exert
different forces on the hairs, causing a
different input to the CNS. If you've ever
been in an airplane that's in a stall you will
have experienced the peculiarly unpleasant
feeling of having your otoliths exert noforce
on the hairs at allthe sensation of being in
free fall. This often causes disorientation
problems for pilots, too, and for astronauts,

who are constantly "falling" around the earth

while in orbit.
Otoliths in many non-mammals and
invertebrates often contain iron and are
affected by magnets. If you tape a magnet to
the shell of a crab, for example, his otoliths
are attracted to it, and he'll swim upside
down. Some migratory birds make use of the
magnetic field of the earth for navigation, and
have a built in compass in their otoliths.
Reports indicate that taping magnets to the
heads of pigeons and migratory waterfowl
causes them to lose their way.

Lab Exercise List