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Acoustic Sensitivity
Francisco E. Olucha Bordonau
Universitat de València

Content

Acoustic Sensitivity ................................................................................................................... 1

Acoustic Sensitivity ................................................................................................................... 1

Inner ear................................................................................................................................ 1

Bony labyrinth .................................................................................................................... 1

Membranous labyrinth ........................................................................................................ 2

The auditory brainstem....................................................................................................... 2

Thalamocortical auditory system ........................................................................................ 4

Acoustic Sensitivity
Along the auditory pathway, the neural systems that manage acoustic information performs a
discrimination of the components of the sound in order to enhance all aspects that help to
identify and locate its source. Thus main aspects to be detected in a sound are its features and
sequences and its location in an azimutal world. We can define two components in the auditory
pathway, the brainstem nuclei and the thalamocortical system. In addition to the ascending
system there is a descending component that from the auditory cortex reach the cochlear
organs.

Inner ear
The inner ear is the organ derived from embryonic statoacoustic placode. It is carved in the
petrous part of the temporal bone forming the bony labyrinth. Within the bony labyrinth is a
membranous structure, the membranous labyrinth which contains the mecanorreceptor. The
membranous labyrinth contains endolymph and the space between the bony and membranous
labyrinth contains perilymph. The mecanorreceptor are sensitive to sound waves (auditory
system) or the endolymph movement (vestibular system).

Bony labyrinth
It consists of three components: cochlea, vestibule and semicircular ducts. The three
components are in bony continuity with each other.

The cochlea is located medially. This is a coiled tube in a spiral of 2.5 turns around an axis
called spiral modiolus. The base of the modiolus is oriented towards the internal auditory canal
which opens in the posterior aspect of the petrous part of the temporal bone. The first round
makes a prominence on the medial side of the middle ear chamber called the tympanic
promontory. The modiolus is perforated along its axis by small ducts parallels to its axis which
contain the axonic ganglionar processes and arrive to the spiral lamina which runs all along the
cochlea attached to the modiolar axis.

The vestibulum is an oval chamber that is placed laterally to the cochlea from where the
semicircular ducts are attached. The verstibulum contains the organs of balance, the utriculum
and the saculum.
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Membranous labyrinth
Within the bony labyrinth there is a membranous labyrinth that contains mecanorreceptors. The
sensitive elements are placed along the cochlea, in the maculae of the utriculum and saculum
and in the crests of the ampullae of the semicircular ducts. The membranous labyrinth is
continuous in the different elements that make up the statoacoustic system. The membranes of
the cochlear duct are continuous to the saculum by the ductus reunions, also there is a
utriculum-saculum duct which contacts both structures and from where there is a endolymphatic
duct which ends in the closed endolymphatic sac. The semicircular ducts are in direct contact
with the utricle.

In the cochlea, the sensitive hairy cells appear along the membranous duct in a sensitive organ
called spiral organ (Corti’s organ). The cochlear duct containing the endolymph is confined
between the vestibular membrane, the outer wall of the cochlear duct and the basilar
membrane. This outer wall of the cochlear duct is attached by the spiral ligament to the bony
laberynth and from the point of confluence between the basilar and vestibular membranes to the
spiral lamina. This configuration leaves three spaces in a transverse section: the vestibular
scala, the cochlear duct and the tympanic scala. The space between the vestibular membrane
and the bone is occupied by perilymph that is in continuity with the perilymph of the saculum
and is called the vestibular scala. The cochlear duct appears in the middle and contains
endolymph. Finally the tympanic space appears between the basilar membrane and the bony
wall. It contains perilymph and starts in the round window which contacts with the middle ear. In
the apex of the modioulus, the vestibular and the tympanic scalas are in continuity to each
other.

The Corti’s organ is placed on the basilar membrane. It a row of internal hair cells and three
rows of outer hair cells. Hair cells are surrounded by supporting cells. Two of these cells leave a
triangular space called Corti’s tunel which separates the inner and outer group of cells. All the
set is covered by a gelatinous membrane tectoria which embeds the ciliae. The movement of
the stapes on the oval window acts as a piston over the perilyph of the scala vestibula and this
movement is transmitted to the membrane tectoria which in case activates the underneath hair
cells. Activation of the hair cells is transmitted to the terminal processes of the ganglion cells.
These terminals penetrate the spiral lamina and arrive to the spiral ganglion where the ganglion
cells are located. Axons of the ganglion neurons pass through the modiolus and join the
vestibular fibers to form the VIII cranial pair.

The auditory brainstem

The first station of the auditory pathway is the cochlear nuclei that are situated laterally at the
junction between the pons and the medulla. The cochlear complex is composed of the dorsal
cochlear nucleus (DCN) and the anterior ventral (AVCN) and anterior dorsal (ADCN) cochlear
nuclei. The axons reaching the cochlear complex arise from the spiral ganglion. When arriving
to the cochlear complex divides into "T" by giving one ascending and one descending branches.
The ascending branch targets the anterior dorsal cochlear nucleus while the descending branch
divides again giving rise to two additional branches one for the dorsal cohelar and the other for
the posteroventral cochlear nucleus. The acoustic part of the VIII cranial nerve displays a
tonotopic organization in which the high frequencies stimulate the base of the cochlea while
high frequencies stimulates the apex. This distribution is multiplied by three. That is, each of the
three cochlear nuclei display a tonotopic organization in which high frequencies are represented
in the ventral aspects of each nucleus while progressively more dorsal levels are targeted by
progressively lower frequencies.

The cochlear nuclei originate three acoustic bundles: dorsal acoustic stria, intermediate acoustic
stria and ventral acoustic stria (also known as the trapezoid body). The dorsal stria crosses
contralaterally just below the floor of the IV ventricle and in the lateral aspects of the tegmentum
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originates the lateral lemniscus. The intermediate stria is a bundle of fibers that gives rise to
collaterals to the ipsilateral olivary complex and the main bulk crosses the midline at
intermediate levels to target the contralateral superior. Part of the fibers of the intermediate stria
joints the dorsal stria to make up the lateral lemniscus. olivary complex. is projected on the
ipsilateral superior olive complex, but some fibers cross the midline and projected onto the
contralateral superior olive complex, some of the stress fibers and intermediate still rostralment
lemnisc added to the side. Finally, the trapezoid body contains fibers arising from the
anteriorventral cochlear which projects to both ipsilateral and contralateral superior olivary
complex.

The superior olivary complex is group of nuclei located in the ventral part of the brainstem
between the medulla and the pons is composed of the nucleus of the trapezoid body, medial
and lateral superior olivary nuclei and a set of periolivary nuclei. Most of the neurons in the
superior olivary complex receive information from the two ears at the same time. Thus it is the
first step in which information from both ears can be compared. This circumstance allows
locating the source of a sound in space. The superior olive complex evaluates differences in the
intensity of the sound and the phase of the sound that is the difference between arriving to each
ear. A sound that is emited by a source on the right side will arrive first to the rtight ear than to
the left one, and a sound that is being emited by a source just in the fromt will arrive at the same
time to both ears and no differences in phase will be detected between right and left ears. In the
case of the lateral supraolivary nucleus, its bipolar neurons are arranged as an “S” so that each
dendrite receives axons from each ear. This convergence friom both sources allows locating the
source of the sound.

The superior olivary complex and cochlear nuclei originate fibers passing through the lateral
walls of the brainstem forming the so-called lateral lemniscus. This is the ascending pathway
from the cochlear and superior olivary nuclei arriving to the inferior colicles. From this and
subsequent steps thew auditory information runs through two parallel pathways: the main or
lemniscal pathway and the accessory or extrategmental pathway. The main, lemniscal, pathway
is where an accurate evaluation of the cognitive aspects, such as frequency, intensity duration..,
of the sound is being done. The extrategmental pathway is less sensitive to the formal aspects
of the sound but is quicker in its emotional value (coming from a predator, a pup…). The lateral
lemniscus is not only a tract, among its bundles some rows of cells may relay ascending
auditory information to the inferior colliculus, these groups of cells are called the nuclei of the
lateral lemniscus. Medial to the lateral lemniscus and its nuclei there are also some groups of
cells that are also involved in auditory processing and are called as the paralemniscal nucleus.

The inferior colliculus is an oval structure containing two main divisions: the central nucleus and
the pericentral region. The fibers of the lateral lemniscus target the central nucleus. This
nucleus displays a tonotopic organization in which obliquus bands of isofrequency can be
detected. According to this tonotopy, lower frequencies are represented in more superficial
isofrequency bands, while progressively higher frequencies are progressively represented in
deeper isofrequency bands. The central nucleus represents the lemniscal pathway which, in
turn projects to the ventral nucleus of the medial geniculate body. The central nucleus also
projects to pericentral nuclei which also receive information coming from the paralemniscal
nuclei. The pericentral nuclei makes up the starting point of the extralemniscal auditory pathway
which is being send to the deep layers of the superior colliculus. In addition, projections of the
pericentral nuclei also reach the nuclei surrounding the ventral nucleus of the medial geniculate
body that are part of the indirect extrategmental auditory pathway.

Along the brainstem pathways crossing occurs at different levels. The cochlear nuclei receive
information primarily ipsilateral. The superior olivary complex receives information from both
ears. Many of the fibers coming from the acoustic striae and superior olive complex run by
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contralateral lateral lemniscus. The isofrequency bands of each central nucleus of the inferior
colliculus project to each other through the inferior colliculus commissure. At the end most of the
information of one ear is being projected to the contralateral inferior colliculus. Upper to this
point the auditory information does not cross to the contralateral side.

In the same way as auditory information progress through the brainstem to the thalamus and
them to the cortex there is a descending pathway that from the cerebral cortex project to the
inferior colliculus and from there to the cochlear nuclei and nuclei of the superior olivary
complex. Fibers arising from the cuperior olivary complex make up the olivo-cochlear bundle
which target the Corti’s organ in the cochlea thus regulating the flow of information from the
spiral ganglion to the cochlear nuclei.

Thalamocortical auditory system

The projection from the thalamus to the inferior colliculus is direct that means not crossed.
There are two parallel pathways, i.e. the main and the accessory pathway. The main pathway
starts at the central nucleus of the inferior colliculus and project to the ventral medial geniculate
body. In contrast, the accessory pathway arise from the pericentral nuclei of the inferior
colliculus and is projected over the ventral and medial nucleis of the medial geniculate body.
Thus the medial geniculate body is composed of the ventral, the dorsal and medial geniculate
nuclei. In all the three divisions there is a representation of the tonotopìc map, however the
tonotopy in the ventral nucleus is much more accurate. The projection collicular projections over
the thalamus is glutamatergic and GABAergic.

The medial geniculate body projects onto the auditory cortex. The ventral nucleus projects to
primary auditory cortex. The dorsal nucleus projects mainly to the accessory auditory cortex.
Finally, the medial nucleus of the medial geniculate body projects to both the primary and the
associative auditory cortex. The difference between primary and secondary auditory cortex is
accuracy and latency. The accuracy of sound occurs in terms of frequency and interaurality.
Primary cortex neurons respond only to very narrow frequencies coming mainly from the
contralateral cochlea. By contrast, secondary cortex’s neurons respond to wider frequencies
and coming from both ears. Furthermore, neurons in the primary cortex fire at very short
latencies, around 10 msec while the secondary cortex neurons respond at latencies around 100
msec. These differencies means that the primary cortex neurons are sensitive to a direct
accurate projection, while secondary cortex neurons receive projection from more elaborated
and delayed circuits.

The primary cortex is represented by areas 41 and 42 which are included in the temporal lip of
the Sylvian fissure in the top half of the superior temporal gyrus. The secondary cortex is
around the primary cortex corresponds to areas 24 and 22. The area 22 in the dominant (left)
hemisphere is related to oral language comprehension (Wernicke area). The association areas
receive projections from the thalamus and from the auditory primary and from visuakl
associative cortical areas, some of these pathways are involved in understanding written
language. The association areas project to the amygdala where the sound may acquire
emotional value. Also cortico cortical projections from the Wernicke area are directed towards
the speech motor areas corresponding to area 44 (Broca's area) located in the opercular gyrus
just anterior to the area controlling motor muscles of the face and mouth.
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Diagram representing the flow of information along the auditory pathway up to the auditory
cortex where the information becomes conscious. It also represents the efferent pathway that
targets the superior olivary complex and from there to the cochlea. a) cochlea, b) cochlear
nuclei, c) superior olivary complex, d) nuclei of the lateral lemniscus e) tectal longitudinal
column, f) inferior colliculus, g) medial geniculate body.
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Diagram representing the elements that makes up the membranous labyrinth and its
sensitive parts. a) superior vestibular ganglion, b) inferior vestibular ganglion ( together the
superior and inferior vestibular ganglia makes up the ganglion of Scarpa), c) spiral ganglion
and auditory nerve, d) cochlear duct, e) saculum with its sensitive side, f) utriculum with its
sensitive part, g) ampulla of the superior semicircular duct, h) ampulla of the lateral
semicircular canal, i) ampulla of the posterior semicircular canal
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Horizontal section of the human cochlea in which it can be observed the 2 ½ turns around the
modiolus. a) cochlear component of the statoacoustic nerve, b) spiral duct occupied by spiral
ganglion of Corti, c) tympanic scala, d) and cochlear duct, e) vestibular scala, f) modiolus g)
helicotrema (apex of the cochlea where vestibular and tympanic scalae connect to each other).
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Microphotography of a rat cochlea in which it is represented the main structures of the cochlear
labyrinth. a) tympanic scala (perilymph), b) vestibular scala (perilymph), c) cochlear duct
(endolymph), d) membrane tectoria, e) vestibular border of the limbus, f) spiral lamina, g), spiral
eminence, h) vascular stria, i) spiral ligament, j) basilar membrane, k) vestibular membrane
(Reissner).

Diagram representing the cochlear organ (of Corti) framed in the previous figure. a) membrane
tectoria, b) inner hair cells, c) external hair cells, d) inner pillar, e) outer pillar, f) cochlear tunnel
(tunnel of Corti), g) supporting cells (of Hensen), h) fibers of the ganglion cells.
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Diagram representing the location of the vestibular and cochlear nuclei in a dorsal view of the
brain stem (A), a transverse section at the pons-medulla transition (B) and a transverse section
at the same level in the rat stained for acetyl choline esterase (C). a) superior cerebellar
peduncle, b) middle cerebellar peduncle, c) inferior cerebellar peduncle, d) anterior ventral
cochlear nucleus, e) posterior ventral cochlear nucleus, f) dorsal cochlear nucleus, g) superior
vestibular nucleus, h) medial vestibular nucleus, i) lateral vestibular nucleus, j) inferior vestibular
nucleus, k) spinal trigeminal tract, l) nucleus of the trapezoid body, m), medial nucleus of the
superior olivary, n) lateral nucleus of the superior olivary complex, o) olivo-cochlear tract, p)
cochlear fibers, q) vestibular fibers, r) principal trigeminal nucleus, s), spinal trigeminal nucleus,
t) reticular formation.
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Microphotographs of transverse sections of superior (A) and inferior (B) colliculus of the rat
brain stained with toluidine blue. a) periaqueductal gray substance, b) surface layer of superior
colliculus, c) intermediate gray layer of superior colliculus d) deep gray layer of superior
collicullus, e) tectal longitudinal column, f) central nucleus of the inferior colliculus, g) pericentral
nuclei of the inferior colliculus, h)commissure of the inferior colliculus.
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