THE ANATOMY OF THE EAR
Part II: Histology
Dr. Art Dalley February 22, 2002
(10:10 am)
Review Anatomy of the Ear, Part I: Gross Anatomy notes from Dec. 5, 2000 [or read
Ross et al]
Added information regarding the external and middle ear:
I. External Ear:
Basic tissues: elastic cartilage skeleton covered with skin
A. External auditory meatus has, in addition to typical features of skin (hair
follicles and sebaceous glands), ceruminous glands:
Modified sweat glands (cuboidal epithelium) in skin of lateral part of canal
cerumin (ear wax) is a combination of the apocrine secretion from these
glands, plus that of sebaceous glands and desquamated skin cells.
B. Tympanic membrane (trilaminar construction):
1 cm in diameter, 0.1 mm thick.
1. Outer layer = a continuation of the skin of the ext. auditory meatus.
thus, it is innervated by nerve V superiorly and X inferiorly.
2. Middle connective tissue core (a very thin lamina propria):
elastic fibers of pars tensa arranged in a double layer: a concentric
parabolic circular arrangement (stratum circulare) crossed by radial
fibers (stratum radiatum) from a central hub, much in the form of a
orb spider's web, but with some additional transverse fibers
inferiorly; peripherally, theres a fibrocartilagenous ring (yet another
anulus tendineus), commonly incomplete superiorly (in region of
pars flaccida)
3. Inner layer = a continuation of the mucous membrane of the middle ear.
thus, it is innervated by nerve IX.
II. Middle Ear:
Lined with mucous membrane, including ossicles
Low cuboidal epithelium
Note: ossicles, first fully-formed bones (via endochondral ossification),
lack periosteum
Pharyngotympanic (auditory) tube (Eustachius)
In X-S, hyaline cartilage (becomes elastic near nasopharnx) has an inverted
J-shape, with long & short crura
Mucous membrane changes from that of middle ear to respiratory
epithelium (ciliated pseudostratified columnar)
III. Internal ear:
comprised of two major components: (A) the osseous labyrinth and (B) the
membranous labyrinth -- the latter housed inside the former.
A. The Bony (Osseous) Labyrinth:
a complex cave in solid bone
the bone which forms the immediate walls of the cave (the otic
capsule) is the hardest bone in the body. Hence, the experienced
dissector can isolate the walls of the bony labyrinth by removing the
softer bone which surrounds it. This may be done with a dental drill.
1. Components of the bony labyrinth:
a. A vestibule, or large 'central room' or compartment into which the
other components open.
b. Three semicircular canals (anterior (or superior), posterior and
lateral).
c. The cochlea ('snail's shell) or cochlear canal.
2. Openings (communications) into the vestibule:
a. The oval window (fenestra ovale or vestibulum).
b. The round window (fenestra rotundum or tympanicum).
c. The six ends of the three semicircular canals by means of five
openings (sup. & post. canals have a 'common crus', merging to
enter together at the non-ampullary end.
d. The cochlear canal.
e. The vestibular aqueduct giving passage to the endolymphatic
duct.
f. The cochlear canaliculus giving passage to the perilymphatic
duct
By means of this duct, the osseous labyrinth is in open
communication with the subarachnoid space.
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since the bony labyrinth is actually a space inside of the bone, it is
perhaps more realistically demonstrated by means of a cast of that
space.
the bony labyrinth is filled with a fluid, the perilymph, in which the
membranous labyrinth is suspended by filaments similar to, but more
delicate than, those spanning the subarachnoid space. Perilymph
resembles extracellular fluid in composition (sodium salts are the
predominate positive electrolyte);
B. The Membranous Labyrinth:
extends into all parts of the osseous labyrinth, but occupies only a
fraction of the volume of the osseous labyrinth (approx. 1/16th).
walls formed by a delicate membrane (ectodermally-derived
neuroepithelium)
filled with endolymph. Endolymph resembles intracellular fluid
(potassium is the main positively-charged ion).
The osseous labyrinth with its contained membranous labyrinth may be
subdivided into an auditory system, serving the sense of hearing, and a
vestibular system or apparatus, for balance and sense of orientation. Each
component is innervated by its own, separate division of the vestibulocochlear
nerve (CN VIII).
C. Auditory System:
The cochlear canal (bony labyrinth) is partially divided by a bilaminar
bony shelf (the spiral lamina) which winds around a hollow central bony
core (the modiolus). The cochlear duct (membranous labyrinth) is
attached to the 'free' edge of the spiral lamina and extends to the opposite
wall of the canal, creating separate tube-like compartments above (the
scala vestibuli) and below (the scala tympani). These two tubes, and the
cochlear duct between them (also referred to as the scala media), spiral
about the modiolus for 2 3/4 turns. Scalae vestibuli and tympani
communicate at the apex by means of the helicotrema. The cochlear duct,
being part of the membranous labyrinth, is filled with endolymph while
scalae vestibuli and tympani, on either side, are filled with perilymph.
The vibrations of the footplate of the stapes are converted to variations
in fluid pressure at the oval window of the vestibule of the bony labyrinth.
Waves of the varying pressures are conducted up the scala vestibuli to the
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helicotrema and then down the scala tympani to the secondary tympanic
membrane of the round window where they are 'lost' to the airspace of the
middle ear. The ascending and descending waves of fluid pressure distort
the intervening cochlear duct, which contains the sensory spiral organ (of
Corti) that converts the mechanical forces to nervous action potentials.
D. Vestibular Apparatus:
consists of the membranous labyrinth minus the cochlear duct, i.e.:
1. Saccule
2. Utricle
a tube connects them together (utriculosaccular duct).
3. Three semicircular ducts -- anterior, posterior, and lateral
occupy ~1/16 of the corresponding semicircular canal
like the semicircular canals, the ipsilateral semicircular ducts are
arranged at right angles to each other, and duct (like each canal) is
perpendicular to its contralateral partner; however, they do NOT lie
within the major axes or anatomical planes of the body:
the anterior and posterior ducts diverge 45 from the medial and
frontal planes
the lateral canal is tilted 30 posterolaterally from the horizontal
plane (i.e., to make the right lateral canal horizontal, the head
must be tipped forward and to the left 30).
also like the semicircular canals, each has an independent dilated
end or ampulla opening into the utricle; two share a non-ampullary
end (common crus), so that there are only five openings of the ducts
into the utricle.
E. Sensory Receptors of the Membranous Labyrinth:
6 regions of sensory receptors project from the wall of the membranous
labyrinth:
3 cristae ampullaris within the ampullae of the semicircular ducts
sensitive to angular acceleration (turning of the head)
2 maculae:
(1) macula utriculi
(2) macula sacculus
sensitive to position relative to vertical (gravity); linear acceleration
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the spiral organ (of Corti) of the cochlear duct
sound receptor
All of these sensory receptors share similar structural specializations and
characteristics, including hair cells that:
are non-neuronal, mechanoereceptor, epithelial cells
possess numerous stereocilia (modified microvilli) called sensory
hairs
vestibular apparatus cells & inner hair cells of organ: ~50-
100/cell
outer hair cells of the spiral organ ~100-300/cell
are associated with both afferent and efferent nerve endings
are all transducers, i.e., they convert mechanical energy (bending
of stereocilia) into electrical energy transmitted by the
vestibulocochlear nerve to the brain
the means by which bending or flexing occurs varies for different
receptors
stretching of the plasma membrane caused by bending
generates transmembrane potential changes in the receptor cell
that are conveyed to the afferent nerve endings associated with
the cell
1. Sensory receptors of the vestibular apparatus:
Two varieties of hair cells occur:
(1) Type I hair cells:
Piriform in shape with rounded base and narrow neck
Surrounded by an afferent nerve chalice & a few efferent
nerve endings
(2) Type II hair cells:
Cylindrical in shape
Afferent and efferent bouton nerve endings synapse basally
Both types possess, in addition to the stereocilia, a kinocilium
A true (but non-motile) cilium, taller and larger in X-S than the
stereocilia, having an array of 9 + 2 microtubules in its core
Stereocilia are arranged by height relative to kinocilium in a step-
like manner
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Tallest clostest, shortest farthest, from kinocilium
Direction of bending of the stereocilia, relative to kinocilium, is
significant:
bending away from kinocilia hyperpolarization of receptor cell
(inhibition)
bending toward kinocilia depolarization of receptor cell & thus
generation of an action potential (impulse).
Both types are surrounded by columnar supporting cells w/ basally-
located nuclei, irregular microvilli and secretory granules
(a) Cristae ampularis
thickened, linear, epithelial ridge in ampulla of semicircular ducts
with both Type I & II hair cells, aligned in same direction (i.e., all
kinocilia are on same side).
ridge is oriented perpendicular to long axis of duct
a gelatinous cupula with canals that receive stereocilia extends from
ridge to opposite side of ampulla
deflected by angular acceleration which produces a movement
differential between the crista (fixed to wall of duct) and the
endolymph
If, for example, the head is rotated toward the right, the
endolymph in the lateral semicircular canal remains stationery
at the onset of the movement because of its inertia, and the
result is a relative movement in the opposite direction
(hydrodynamic inertia); both contralateral cupulae are
displaced to the left. Then the endolymph slowly overcomes
the intertia and follows the rotation of the head. However,
when the rotation ceases, the endolymph continues to move
for a while in the same direction, and the contralateral cupulae
are bent to the right. The semicircular ducts are particularly
concerned with reflex control of visual movements. The jerky
eye movements caused by rotation of the head (rotatory
nystagmus) are dependent on the defections of the cupulae.
The slow component of the nystagmus is always in the
direction of the cupula deflection.
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(b) Maculae:
sensory epithelium per se essentially the same as for cristae in cell
structure, but are not linear (they are curving or hook-shaped), and
sensory cells are oriented relative to an imaginary longitudinal axis
(striola)
utricular and saccular maculae oriented at right angles to one
another
when standing, macula utriculi is horizonal; macula sacculi is
vertical
orientation of sensory cells relative to striola is opposite
cells of macula utriculi have kinocilium toward striola
cells of macula sacculi have kinocilium away from striola
In the place of the cupulae, maculae have an overlying gelatinous
otolithic membrane which contains 3-5 crystaline particles of
calcium carbonate and protein (otoliths or otoconia) in its outer
surface (opposite that in which the sensory hairs are embedded).
Sensory hairs are bent by effect of gravity on and inertia
(consequent to linear acceleration) of otoliths.
2. Sensory receptor of the acoustic apparatus (the spiral organ of Corti)
contained within cochlear duct (scala media)
suspended in cochlear canal by attachments to the spiral lamina and
the outer wall of the canal via the spiral ligament
lateral or outer wall: stria vascularis
highly vascular, thick, pseudostratified epithelium (source of
endolymph?)
upper or apical wall (separating it from scala vestibuli) = vestibular
membrane (Reissners)
lower wall or floor (separating it from scala tympani) = spiral or
basilar membrane
width of spiral membrane increases as it ascends to apex
spiral organ resides on the basilar membrane
at the acute angle formed by the junction of the vestibular and spiral
membranes is a dense tissue layer covered by a tall epithelium, the
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spiral limbus, which extends peripherally as two lips lying on either
side of the internal spiral sulcus:
a tympanic lip, which extends along the spiral membrane to the
spiral organ
a vestibular lip, to which the tectorial membrane (which overlies
the spiral organ) is attached.
(a) Spiral organ (of Corti)
Complex neuroepithelium
Lies on basilar membrane at margin of spiral lamina, from basal turn
to apex
A triangular inner canal (tunnel of Corti, actually an intercellular
space) lies at the center, filled with cortilymph (extracellular fluid),
bound by:
vertical inner pillar cells, resting on the tympanic lip of the
spiral limbus
oblique outer pillar cells, resting on the spiral membrane
have a wide basal part that includes the nucleus
contain long bundles of supporting tonofibrils
On either side, phalangeal (Dieters supporting) cells underlie
rows of hair cells:
(i) inner row of hair cells
single row of piriform cells (like Type I) throughout turns of
cochlea
phalangeal cells support and surround
(ii) outer row of hair cells
row gradually widens from three ranks of cylindrical hair cells
(like Type II) in basal turn to five at the apex
each outer phalangeal cell supports a sensory hair cell on a
cupped lower part of the cell
phalangeal processes ascend between sensory cells and
end in flat head plates which together form a superficial
perforated reticular membrane or lamina surrounding the
apical ends of the hair cells sealing the endolymph
compartment from the intercellular spaces of the spiral organ
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hair cells have thick cuticular plates in which sensory hairs are
anchored, usually in three V- or W-shaped rows of increasing
height
hair cells do not have kinocilium, but retain a basal body on the
outer aspect
a gelatinous tectorial membrane extends from the vestibular
lip of the spiral limbus to overlie the hair cells
the tips of the sensory hairs of the outer hair cells are
embedded in this
the variations in the perilymphatic fluid pressure produced by the
vibrations of the footplate of the stapes at the oval window cause
traveling waves displacing the spiral membrane and thus distorting
the spiral organ; thus the sensory hairs of the sensory cells are
deflected against the tectorial membrane. High frequency sounds
cause maximal displacement of the spiral membrane near the base
of the cochlea, where spiral membrane is narrowest and outer row
of hair cells has three ranks; low-frequency sounds cause maximal
displacement near the apex, where spiral membrane is widest and
outer row of hair cells has five ranks [tonotopic localization].
Perception of loudness depends on the degree of displacement of
the spiral membrane at any given frequency range.
Wow!!