Functional Anatomy of the Cochlea

INTRODUCTION

Is represented by osseous structures, membranous structures, fluids, and specialized sensory and supporting
cells.

The bony morphology of the cochlea provides the foundation upon which the membranous and cellular
structures rest either directly or indirectly. The cochlea is housed within the temporal bone and an
understanding of the anatomy of this bone is important to the understanding of the structure and function of
the cochlea. The position of the cochlea in the temporal bone and the fact that the temporal bone is composed
of a hard and rigid bony foundation provides protection for this extremely delicate and complex organ of
hearing.

OVERVIEW OF THE TEMPORAL BONE

The temporal bone is an osseous structure that is part of the cranium and houses a portion of the external ear,
as well as the middle ear, the inner ear including the cochlea and the vestibular apparatus, and the seventh and
eighth cranial nerves.

The temporal bone is a hard, rigid bone located within the skull and has a myriad of cavities, channels, and
canals that subserve the end organs of hearing and balance; hence, the term labyrinth is used to describe this
bony structure. The main purpose of the bony labyrinth is to provide support and protection for the delicate
anatomical structures of hearing and balance. Within the bony labyrinth are membranous structures that line
the channels and canals of the temporal bone and form the inner ear. These membranous structures, referred
to as the membranous labyrinth, contain the fluids that bathe the sensory cells for hearing and balance.
Within the membranous labyrinth are a number of structures, including the cochlea, or the end organ for
hearing, and five sensory structures that are part of the vestibular end organ (three semicircular canals, the
utricle, and the saccule).

The temporal bone helps form the base of the human skull. It also forms the middle and posterior fossa of the
cranial vault. The posterior fossa houses the cerebellum and the brainstem and is the area that is approached
by the surgeon to remove acoustic tumors and other brainsteam tumors, such as meningiomas of the
cerebellopontine angle. The middle fossa accommodates the anterior-inferior aspect of the temporal lobe.

The four main parts of the temporal bone are the squamous, mastoid, petrous, and tympanic portions
(Figures 4–1, 4–2, and 4–3). Using the external auditory meatus as a reference point, the squamous portion is
superior, the mastoid portion is posterior, the petrous segment is medial, and the tympanic segment is anterior
and inferior to this structure (Anson & Donaldson, 1967).

The squamous portion of the temporal bone is positioned superior to the ear canal and is semicircular, or fan-
shaped, in its orientation relative to the ear canal (see Figure 4–1). Its lateral surface is smooth, and its
superior surface serves as the attachment for the temporal muscle. The superior part has a small sulcus, which
accommodates the middle temporal artery. The inferior, lateral aspect of the squamous portion is
characterized by the zygomatic process, which is a bony arch running posteriorly and anteriorly (see Figure
4–1). The masseter muscle attaches to the zygomatic arch. The medial surface of the squama helps form the
lateral aspect of the middle cranial fossa. Also, on the medial surface of this structure is a sulcus for the
middle meningeal artery (Anson & Donaldson, 1967). The mastoid part of the temporal bone is posterior to
the ear canal and the squamous bone (see Figure 4–1). The lateral aspect of the mastoid portion serves as the
attachment for the occipital and postauricular muscles. The mastoid has a sulcus that accommodates a branch
of the occipital artery. Located on the mastoid part of the temporal bone is an eminence that projects outward.
This part of the bone, which lies directly posterior to the pinna, is where the bone conduction oscillator is
frequently placed on the skin for bone conduction testing. The medial aspect of the mastoid has a notable
sulcus. This sulcus houses the sigmoid venous sinus and is called the sigmoid sulcus.
The petrous portion of the temporal bone is probably the most anatomically complex of the four portions of
this bone. It houses most of the peripheral hearing mechanisms (see Figures 4–2 and 4–3). The petrous part of
the temporal bone is triangular in shape, allowing an anterior and posterior surface. The anterior surface forms
part of the middle cranial fossa. Laterally the petrous bone fuses with the squama. The anterior surface has a
semicanal structure for the tensor tympani muscle, which is innervated by the fifth cranial nerve. It also
contains a semicanal for the Eustachian tube. This surface of the temporal bone is characterized by two small
openings toward the apex of the petrous segment. The first opening is for the facial nerve and the second is
for the petrosal nerve and superior tympanic artery (Anson & Donaldson, 1967).

The temporal bone, in particular the petrous portion, is susceptible to fractures from falls and other types of
head injuries, which often present with audiological consequences. The posterior surface of the petrous bone
makes up part of the posterior fossa (Anson & Donaldson, 1967; Gulya, 1997) (see Figures 4–2 and 4–3).
Midway between the base and apex of the petrous bone is the opening of the internal auditory meatus (IAM)
(see Figure 4–3). This structure is important for hearing, balance, and facial function. Running through the
IAM are the auditory, vestibular, and facial nerves. The IAM can be divided into four quadrants that
accommodate these important nerve routes. If one were facing the opening of the IAM, the upper left area
would contain the facial nerve, while directly inferior to the facial nerve would be the auditory nerve. Across
from the facial and auditory nerves are the vestibular nerves. The superior quadrant houses the nerve fibers
from the superior and horizontal semicircular canals, while the inferior quadrant of the IAM provides the
route of the nerve fibers from the saccule, the utricle, and the posterior semicircular canal. The singular
foramen is a small opening in the inferior lateral aspect of the IAM that provides the route for the fibers of the
posterior semicircular canal. Also located on the posterior surface of the petrous part of the temporal bone is
the bony accommodation for the vestibular aqueduct (VA) (see Figure 4–3). This sulcus is located about
halfway between the opening of the IAM and the sigmoid sinus area (part of the mastoid). The VA is a
channel that contains endolymph and communicates with the endolymphatic sac and duct. Both of these
structures are part of the vestibular end organ. The inferior surface of the petrous bone in combination with
the occipital bone forms the jugular foramen. Also located on the inferior surface of the petrous bone is the
jugular fossa and jugular bulb. (The clinical significance of the jugular tumor will be discussed in Chapter
14).

Osseous (Bony) Cochlea

The cochlea is perhaps the hardest bony structure in the body. It is located in the petrous portion of the
temporal bone, immediately medial to the tympanum, or middle ear cavity. The cochlea’s position in the head
is sometimes difficult to realize. The apex, or cupola, of the cochlea (representing the apical turn) is oriented
anterior and slightly lateral within the head pointing toward the cheekbone (Figure 4–5). The portion of the
cochlea that is opposite the apex points medially and posteriorly toward the back of the head. The bony
cochlea resembles a spiral cone with auditory nerve fibers emanating from inside the cone in a spiral fashion.
The bony cochlea resembles a snail shell, and in humans, it has between 2.2 and 2.9 turns (Buser & Imbert,
1992). These turns are smaller at the apex than at the base of the cochlea (see Figure 4–5). The entire bony
cochlea measures about 1 cm wide and about 5 mm from base to apex (Pickles, 1988). The modiolus is a
central, perforated bony core that accommodates nerve fibers from the hair cells, as well as blood vessels
(Figures 4–6, 4–7, and 4–8). At its medial end the modiolus is continuous with the IAM. The modiolus is also
the central structure around which the cochlea is wrapped. Specifically, the osseous spiral lamina is a shelf-
like structure that winds around the modiolus from base to apex (see Figures 4–6, 4–7, and 4–8). The osseous
spiral lamina is composed of two thin plates between which auditory nerve fibers pass from the hair cells in
the cochlea to eventually connect to nuclei within the brainstem. It is narrower at the apical end of the cochlea
than at the base of the cochlea (Zemlin, 1998). It is interesting to note that the human bony cochlea is wider at
the base and flatter than the bony cochleae found in many small animals.

The spiral lamina’s lower shelf also serves as the support point and connector for the inner aspect of the
basilar membrane (BM) and the scala media or cochlear duct; this duct divides the cochlea into the scala
vestibuli (superior) and the scala tympani (inferior) (see Figure 4–8). The upper shelf of the spiral lamina is
continuous with a structure known as the spiral limbus and serves as an attachment and support point for the
tectorial membrane. The medial aspect of the tectorial membrane along with the end of the spiral limbus,
which is concave in shape, forms the internal spiral sulcus. The bony cochlea has two windows located at the
basal turn on its lateral aspect. The more superior of the two windows is the oval window, which interacts
with the stapes of the middle ear and opens into the scala vestibuli. The oval window in humans ranges from
about 1.2 × 3 mm to 2.0 × 3.7 mm (for a review of inner ear structure measurements, see Yost, 2000).

The round window is inferior to the oval window and opens into the scala tympani. In humans the round
window’s dimensions are approximately 2.25 × 1.0 mm. Although it has been termed the round window
(most likely to differentiate it from the oval window described above), it actually has an ovoid shape that
doesn’t change in size after birth (Rask-Andersen et al., 2012). The anatomy of the round window has
received much attention recently because it is the insertion point for cochlear implants (see Rask-Andersen et
al., 2012).

Lying between the oval and round windows is a bony eminence known as the promontory. The promontory
is positioned to protect the oval and round windows in that it directly faces the middle ear cavity. Foreign
bodies entering the middle ear would first strike the promontory. The promontory is the anatomical site for
needle placement in transtympanic electrocochleography. For this procedure, a transtympanic needle
electrode is placed in and through the lower portion of the eardrum, which allows for alignment with the
promontory. The round window is covered by a membrane that retains cochlear fluids within the cochlea.

Membranous Cochlea and Related Structures

Scalae and Cochlear Fluids

Following the spiral shape of the bony cochlea from its base to its apex is the membranous cochlea (Figure 4–
10). The membranous cochlea is highly elastic, allowing it to move easily within the bony cochlea. There are
three ducts within the cochlea: the scala vestibuli, which is superior; the scala media (often called the
cochlear duct), which is located in the middle; and the scala tympani, which is the most inferior of the three
ducts (Figures 4–11 and 4–12). The oval window is located at the opening of the scala vestibuli, which is the
site of the stapes footplate. The round window membrane is at the basal-most aspect of the scala tympani. The
three scalae are divided by two membranes: the basilar membrane and Reissner’s membrane. The basilar
membrane (BM) is connected to the spiral ligament on the outer wall of the bony cochlea and to the osseous
spiral lamina to form the floor of the scala media or cochlear duct (see Figures 4–11 and 4–12). Reissner’s
membrane forms the roof of the cochlear duct and the floor of the scala vestibuli. The three scalae spiral the
length of the cochlea and are separated from one another, with a single exception. At the apex of the cochlea
the scala vestibuli and scala tympani communicate, and this point of communication is termed the
helicotrema (see Figure 4–11). The cochlear duct (scala media) communicates at its basal region with the
vestibular region (specifically the saccule) of the inner ear by way of the ductus reuniens of Hensen (see
Figures 4–10 and 4–13). This is a narrow passageway, which may be obliterated in the adult — although this
point remains controversial.

The three scalae are filled with fluids (see Figure 4–12). The scala tympani and scala vestibuli are filled with
perilymph (16 to 23 μL in humans) and the scala media with endolymph and cortilymph (2.7 μL in humans)
(Yost, 2000). Perilymph has essentially the same chemical composition as cerebral spinal fluid (CSF). It is
low in potassium (K+) and high in sodium (Na+). Interestingly, however, it appears that the sodium and
potassium concentrations in the perilymph are not uniform throughout the cochlea (Wangemann & Schacht,
1996). The scala vestibuli and scala tympani have slightly different Na+ and K+ concentrations. Cortilymph
is similar to perilymph in its chemical composition, as it is high in sodium and low in potassium (Engstrom,
1960). The cortilymph bathes the hair cells and fills the spaces of Nuel, the external tunnel, and the main
tunnel of Corti. Cortilymph is segregated from the endolymph by the reticular lamina.

Vestibular and Cochlear Aqueducts

Contributing to the fluid systems of the cochlea are two fluid pathways: the vestibular aqueduct (VA)
(endolymphatic) and cochlear aqueduct (CA). The VA (containing endolymph) is the bony channel that
encompasses the endolymphatic duct (ED). The ED courses from the posterolateral wall of the vestibule to
the endolymphatic sac, which is located on the posterior surface of the petrous pyramid of the temporal bone
(see Figure 4–3). The endolymphatic sac’s opening in the temporal bone is 10 mm posterolateral to the
opening of the IAM (Gulya, 1997). The ED is thought to help regulate endolymphatic pressure to avoid the
effects of overproduction or under-absorption of endolymph. Because of the role it may play in the
hydrodynamic function of the inner ear, dysfunction of the ED has been implicated in endolymphatic hydrops
(see Figure 4–13).

The CA (containing perilymph) courses from the basal turn of the scala tympani only a few millimeters from
the round window to the subarachnoid space near the jugular fossa (see Figure 4–13). It runs parallel to the
inferior margin of the IAM (Gulya, 1997). The CA allows transfer of CSF. The CA is patent early in life, but
in many individuals the channel becomes closed in later life. At 50 years of age, it is estimated that only 50%
of individuals will have a patent CA (Wlodyka, 1978). The endolymph has a high potassium (K+)
concentration and is low in sodium (Na+). It is believed that this chemical relationship is maintained by an
active process in the stria vascularis (Buser & Imbert, 1992). Endolymph is most likely produced in a
complex manner, but the stria vascularis is certainly the main player (see Pickles, 1988). The high
concentration of K+ in the endolymph is especially important to cochlear function because a considerable
portion of the transduction current in the hair cells is carried by the K+ ions (Wangemann & Schacht, 1996).
This is related to the high K+ and low Na+ concentrations in the endolymph and the K+ selectivity of ion
channels in the membranes of the hair cells. The low K+ concentration in the perilymph also contributes to
this ion action. Changing the K+ ion concentration in the inner ear fluids influences the cochlear potentials
(Marcus, Marcus, & Thalmann, 1981).

Basilar and Reissner’s Membranes

The BM is from 25 to 35 mm in length in the adult human. Its breadth is greater at the apex than at the base,
with the base measuring about 0.04 to 0.1 mm and the apex 0.36 to 0.5 mm in the adult with some variability
(Buser & Imbert, 1992; Harrison, 2012). The thickness of the BM seems to be a difficult issue to resolve.
However, to the best of our knowledge it appears that the BM is thicker at the base compared with apex
(Harrison, 2012) (Figure 4–16a). However, there are those who report just the opposite (Rask-Andersen et al.,
2012). It should be noted that along the BM’s length there is some variability in its thickness; i.e., slight
changes in the thickness of the BM may be observed randomly along its length (Ehret & Frankenreiter, 1977).
The BM is composed of approximately 24,000 fibers and is oriented transversely (Figures 4–12, 4–15a, and
4–16a, b). It has shorter fibers at the base than at the apex and is stiffer at the base than at the apex. This
allows better tuning for high frequencies at the base and low frequencies at the apex (Harrison, 2012). The
BM has two sections. The inner section, which courses from the spiral lamina to the outer pillar cells, is thinly
structured and is termed the arcuate zone (see Figure 4–12). The second section extends to the spiral
ligament and is structurally much thicker and is termed the pectinate zone (Raphael & Altschuler, 2003). The
arcuate zone, because it is partly enclosed in the osseous spiral lamina, vibrates little, while the pectinate area
moves more freely. The stiffness of the BM decreases and the mass increases as it progresses from the base to
apex of the cochlea (Harrison, 2012; Zwislocki, 2002) (Figure 4–16c).

Reissner’s membrane forms the upper boundary of the scala media (see Figures 4–12 and 4–17). Its
structure includes two layers of cells, termed epithelial and mesoepithelial cells, and a basement membrane.
The epithelial cells are located on the endolymph side of the membrane (i.e., the scala media side), while the
mesothelial cells are situated on the perilymph side (i.e., the scala vestibuli side). These two cell layers are
separated by the basement membrane that lies in between the two cell layers (Rask-Andersen et al., 2012).
This membrane is avascular. It is permeable, by most accounts, but it is responsible for maintaining a
separation of endolymph and perilymph, which in turn maintains the endocochlear potential.

The BM (and Reissner’s membrane) has some interesting relationships with the cochlea in general.
Progressing from the base to the apical region of the cochlea, the scala media becomes smaller, the spiral
lamina narrower, and the BM (and hence Reissner’s membrane) wider (Duvall & Rhodes, 1967; Flock &
Flock, 2003).