THE EAR

PYSIOLOGICAL ANATOMY
• The ear is organ of hearing & equalibrium
• It is divided into three parts:
. External ear
. Middle ear
. Internal ear
 The external ear/ outer ear
• The pinna
• The external auditory meatus/outer ear canal
• The pinna:
. cartilagenous expanded portion covered with

skin projecting from side of head

. Helps to collect sound waves
. Helps to localize the source of sound
• external auditory meatus
. 2.5 cm long
. Runs forwards & inwards (medially)
. Helps in transporting the sound waves to the
middle ear
. Its lining with skin which secretes wax & oil which by
trapping the foreign bodies keep the ear canal clear
. It teminates at the tympanic membrane which
covers the opening of the middle ear
 The middle ear
• It is an air filled chamber
• Consists of following parts:
. Tympanic membrane / ear drum
. Ear ossicles /auditory ossicles
.pharyngotympanic tube or eustachian tube
. Two skeletal muscles
• tympanic membrane / ear drum
. Consists of connective tissue covered with skin on outside &
mucous membrane on inside
. It has the shape of shallow funnel with the centre umbo
.umbo is pointing inwards
. Umbo is kept in position by handle of malleus
. The presence of air on both sides of it to vibrate freely when
sound waves strike it
. These are then passed to the ear ossicles
• Ear ossicles
.3 minute bony ossicles: malleus(hammer
shaped), incus(anvil shaped), stapes(stirrup
shaped)
. Extend across the cavity from the tympanic
memb to the oval window
• Malleus
.its handle is attached to the inner side of the
tympanic membrane
. Its tip is situated at the umbo
. Its head is firmly bound by ligaments to the
incus
• Incus
. Shape like an iron block on which a smith
hammers metal into shape
. At one side it is attached to the head of
malleus
. On the other side its long process passes
downwards to articulate with the head of
stapes
• stapes
. Stirrup shaped , a support for a horse rider’
foot
. Its foot plate is attached by annular ligament
to the margins of the oval window
.functions : the 3 ossicles function to magnify
the intensity of sound by 1.2-1.3 times
• opposite wall of middle ear contains 2
membrane –covered openings
• they face the internal ear
• an oval window above & a smaller round
window just below it
• Pharyngotympanic tube/ eustachian tube
. 4-5 cm in length
. Connects middle ear cavity with pharynx
. normally the pharyngngeal opening is closed
. It can be opened by swallowing , chewing, or
yawn
• Two skeletal muscles: tensor tympani &
stapedius
. Tensor tympani: is attached to the neck of
malleus
. Is innervated by trigeminal nerve
. Its contraction increases the tension of
tympanic memb by pulling the handle of the
malleus medially
. It keeps the tympanic memb firmly attached
• Stapedius : the smallest skeletal muscle in the
body
. It is attached to the neck of the stapes & to
the post wall of the middle ear
. It is innervated by facial nerve
. On contraction it pulls the foot plate of the
stapes out from the oval window
• Function:
. Both muscles can be reflexely activated by
loud sounds
. This reflexely decreases the amplitude of
sound vibration of tympanic memb
. Thus they serve a protective function by
protection of internal ear from loud sounds
 The internal ear
. Bony labyrinth
. Membranous labyrinth
.bony labyrinth is a series of channels in the
temporal bone which enclose the
membranous labyrinth
. Membranous labyrinth comprises of
vestibule, cochlea & 3 semicircular canals
• Cochlea
. Coiled tube, 35 mm long, 2.5 -2.75 turns
.lamina is divided throughout the length by 2
membranes ( Reissner ‘ membrane & basilar
membrane) into 3 compartments/ scalae
. scala vestibuli ,scala [Link] media
. Helicotrema
. Sec tympanic membrane
• Receptor of hearing : the organ of corti
• tunnel of corti
• inner hair cells
• outer hair cells
• stercocilia
• reticular lamina: tough memb
• tectorial membrane thin but stiff gelatinous
elastic structure made of glycoprotein material
• Innervation
.afferent innervation : cochlear divison of 8th
nerve ,having cell bodies in the spiral ganglion,
27000 fibres
.efferent innervation: efferent cholinergic fibres
arise from both ipsilateral & contralateral
superior olivary nucleus via olivocochlear bundle
. Eff causes inhibition of aff fibres by releasing
Ach, plays in impt role in auditory transduction
• function of hair cells:
• Inner hair cells: are primary sensory cells that
generate A.P in auditory nerve
. stimulus is the fluid moving between tectorial
memb & hair cells
. These cells detect sound & are responsible
for fine auditory discrimination
• outer hair cells:
. Detect presence of sound
. Improve hearing by influencing the vibration
patterns of basilar membrane
 Auditory pathways
• The cochlear divison of 8th nerve is formed by
axons of spiral ganglion
• auditory nerve ends in ventral & dorsal cochlear
nuclei in medulla oblongata ,the site of first synapse
• second order neurons from cochlear nuclei end in
superior olive & trapezoid body on both sides of
brain stem
• from superior olive & trapezoid third order neurons
take origin
• Third order neurons pass via variety of pathways in
the lateral leminiscus to the inferior colliculi on both
sides (centre for auditory reflexes)
• some of these fibres also send collaterals to the
reticular formation & medial geniculate bodies in
thalamus
• from the inferior colliculi ,many fibres project &
relay in the medial geniculate bodies
• medial geniculate neurons finally project to the
primary auditory cortex(area 41)
• At area 41. nerve impulses are perceived as
sound .i.e it receives & perceives auditory
information , such as loudness, pitch, source,
direction of sounds
• auditory association areas (area 22,area21, &area
20)
• area 22 : WERNICKE’ AREA: located in superior
temporal gyrus ,is concerned with comprehension
• area 21 & area20 : are located in middle & inferior
temporal gyrus
• Area 21 &20 are concerned with
interpretation & integration of auditory
impulses
• lesion of area 21 &20 impair auditory short-
term memory without impairing visual
memory
• each ear is bilaterally represented in the
auditory pathway from medulla upwards &
projects equally to two cerebral hemispheres
• Deafness is hardly ever produced by cortical lesions
• The auditory pathways are very [Link] are
modified by experience
• individuals who become deaf before language
skills are fully developed , viewing sign language
activates auditory association areas
• individuals who become blind early in life are
better at localizing sound than individuals with
normal eye sight
 Physical properties
• The vibrating objects cause alternating phases
of compression & rarefaction (presure
changes) in the medium ,spread out as sound
waves
• sound travels in air at a speed of 330 m/sec
at sea level at 0o C ,INC at 20o C, in water it
travels much faster ,at 20o C
• the speed of sound slightly increases with
temp & altitude
• Amplitude (intensity) of sound wave
determines the loudness of sound, greater the
ampitude –louder the sound
• frequency of sound waves determines the
pitch of sound , greater the frequency – higher
the pitch
• the wave form of any periodic vibration can
be resolved
• lowest freq is called fundamental and
multiples of fundamental are called
harmonics/octaves
• quality/timbre of many sounds are of
complicated but of repeating pattern,these
are perceived as musical sounds
• if sound contains many components but not
simple multiples of fundamental ,called noise
• Intensity of sound /amplitude is measured in
terms of max pressure
• 1 bel= log intensity of a given sound/intensity
of a reference sound
• 1 bel = 2 log pressure of a given sound/pressure
of a standard sound
• 1 bel= 0.1 bel
• zero dB=reference sound
• 40-60dB =NORMAL CONVERSATION
• PITCH OF SOUND
• HUMAN EAR can peceive 16-20,000cycles/sec
• pitch of normal male voiceis 120 Hz
• pitch of normal female is 250 Hz
• no. Of pitchs that can be distinguished is 2000
 Mechanism of hearing
• ear converts sound waves in the external
environment into action potentials in the auditory
nerve
• the sound waves are changed greately by the
tympanic membrane & ear ossicles into movements
of the foot plate of the stapes
• these movements set up waves in the fluid present
in the inner ear
• the action of waves on organ of corti generates
action potentials in the nerve fibers
 Role of middle ear
• Tympanic membrane;
. Acts as pressure receiver i.e extremely
sensitive to pressure changes produced by
sound waves on its external surface
.it acts as a resonator i.e. It starts vibrating (in
& out movements) freely when the sound
waves strike
.it critically dampens ( stops the vibration of)
the sound waves
• Ear ossicles
. The ear ossicles function as alever system that converts
the resonant vibrations of tympanic membrane into
movements of the stapes against the perilymph filled scala
vestibuli
. Middle ear contains air ,inner ear contains fluid so sound
is transmitted from the air to the fluid
.As fluid having inertia, sound is not transmitted so easily
into the inner ear
.so it is transmitted by increasing the pressure in the
middle ear
• reason of increasing pressure
. these ossicles move as a single unit
. Magnification of sound intensity by 1.2-1.3
times
.reduction of the area is by about 17 times
.so there is corresponding increase in pressure
at the oval window
.impedence matching : increase of pressure
within middle ear by 22 times
• thus when the stapes is pressed into oval
window, this pressure is transferred to the
perilymph in the scala vestibuli
• amplification of sound intensity is greatest
between 1000-3000 Hz
• sounds below 16 Hz or 20000 Hz are not
amplified at all
• minimum audibility curve i.e ability of the middle
ear to magnify some sounds better than others
• Effective transfer of sound energy from air to
fluid medium is difficult because most of
sound is reflected
• middle ear functions as an impedence
matching device by amplifying pressure of
sound
• 70% of sound energy which falls on the
tympanic membrane gets transmitted to the
fluid in the cochlea
 Tympanic reflex
• It is protective reflex against loud sound
• Loud sound initiates reflex contraction of middle ear
muscles
• Contraction of tensor tympani by tensening of the
tympanic membrane ,decreases its vibrations
• Contraction of stapedius muscle pulls the foot plate of
the stapes out of the window
• Thus excessive sound is prevented from going to oval
window
• Reaction time of this reflex is 40-160 msec
 Role of internal ear
.Vibration of basilar membrane
. Stimulation of hair cells
. Functions of hair cells
. Mechanism of pitch discrimination
 [Link] of basilar membrane
mechanism
. The movements of footplate of the stapes into the
oval window set up pressure in perilymph of scala
vestibuli
. The bony walls are rigid but basilar memb is flexible
. Also the basilar memb is not under tension
. Therefore basilar memb is readily depressed into
scala tympani by the pressure
.finally the pressure is transmitted to round window
• it causes secondary tympanic membrane to bulge
outwards into the middle ear
• conversely an outward movement of stapes & oval
window cause upward movement of basilar membrane
• secondary tympanic membrane plays an impt role in
proper vibration of basilar membrane
• as stapes rocks to and fro in the oval window it sets up
wave motion in the membranous labyrin
• the site of memb at which these vibrations are
maximal will be determined by freq sound wave
 2. Stimulation of hair cells
• the tops of hair cells in corti are held rigid by
reticular lamina & their cilia are embedded in the
tectorial memb
• the tectorial & basilar memb are attached to the
limbus at different points
• therefore, when basilar memb moves ,there is
sharp motion between two stiff structures, tectorial
memb & reticular lamina
• this bending results in generation of action pot in
8th nerve
Mechanism of pitch discrimination
• place theory/ Bekesy travelling wave theory
. He showed the basic pattern of movement of the
basilar membrane ,is that of a travelling wave
. As the wave moves up the cochlea ,its height increases
to a maximum & then drops off rapidly
. The distance from the stapes to the point of max
height varies with freq of vibrations
. High pitched sounds generate waves that reach max
height near the base of cochlea
.
• Low pitched sounds generate waves reach
max ht near apex
• this theory is responsible for discrimination of
sound frequencies only between 2000-
20000Hz
• below 2000 are discriminated by Volley
principle
 VOLLEY PRINCIPLE
• The apex of cochlea contains units which
respond to the lower octaves
• the pure tones upto 2000 Hz produce clear
synchronous volleys of action potential in the
cochlear nuclei of 8th nerve
• this is called Volley principle of freq
discrimination
 Factors affecting pitch of a sound
• freq of sound wave
• freq affects loud sounds : low tones sound
lower & HIGH TONES SOUND HIGHER
• DURATION: PITCH OF A TONE CANNOT BE
PERCEIVED UNLESS IT LASTS FOR MORE THAN
0.01 SEC
• Wave form :the pitch of complex wave of a
given freq can still perceived even when the
fundamental freq is absent
 Endolymphatic potential/electrical activity
of cochlea
• When 2 electrodes ( one inserted into the scala
media containing endolymph & other inserted into
the scala vestibuli containing perilymph) are
connected through a suitable amplifier to CRO
• Then a steady potential difference of +50 - +100
millivolt (av:80 millivolt) is recorded ,called
endolymphatic potential/ endocochlear potential
• It is written with a positive sign ,signifying that
scala media is positive to the scala vestibule
• The interior of the cells of the Reissner ‘
membrane & of the cells of organ of corti &
the cells of stria vascularis is approx 30
mv ,negative to the perilymph of the scala
vestibule
• there is no pot difference between the two
chambers ,scala vestibuli & scala tympani,both
containing perilymph
• CF genesis :
. Cell membrane separates 2 fluid
compartments (ICF & ECF having widely
different composition with inside –ve relative
to outside(ECF)
. Likewise , Reissner ‘ memb separates two
fluids having widely different composition
across across the membrane
• Endolymph is formed by stria vascularis & has an
electrolyte concentration very similar to that of ICF
• Perilymph is formed mainly from plasma with
composition like ECF
• as endolymph is +ve wrt to perolymph, so it
cannot be explained on the basis of Na and k ion
conc. Difference
• The stria vascularis that covers the lateral wall of
the scala media is the source of endolymphatic pot
• The cells in the stria vascularis have high conc of Na+- K+ ATP ase
• there is unique electrogenic K+ pump that accounts for high
conc of K+
• FACTORS:
. Movements of basilar membrane affect endolymphatic pot
. It can be increased by a downward movement of the basilar
membrane & reverse of it
• ringer sol (same composition as that of ECF)
into scala media ( same composition as that of
ICF) abolishes the E.P
• RINGER sol has no effect when injected into
scala tympani
• the injection of a potassium rich ,sodium poor
solution into the scala media does not alter
E.P ,but abolishes it if the injection is made
into the scala tympani or vestibuli
 Cochlear microphonic potentials
• one of the electrical responses to sound
• it is potential fluctuation that can be recorded
between an active electrode & indifferent electrode
• an active electrode is placed on or near the cochlea
• an indiferent electrode is placed anywhere on body
• it is called the microphonic cochlear
potential ,because if these pot are amplified,the
loudspeaker records the pure tones fed into ear as
sound waves upto frequencies of 20,000hz
• these pot are similar to the generator potential they show no latency,do
not obey all or none law, are resistant to ischaemia & anaesthesia
• these pot are produced by transformation of mechanical energy into
electrical energy
• These potentials can be recorded optimally by placing one electrode in
scala media & one in scala tympani
• these pot are developed as a modification of the endolymphatic pot
• like E.P these pot are also altered by movement of basilar memb
• the base of cochlea responds to all freq of sound while apex responds to
only low freq
• IF local degeneration of the organ of cortii occurs by prolonged
exposure to a loud tone, cochlear microphonic potential produced by
this particular band of frequency is abolished
Action pot of auditory nerve fibers
• The processes of hair cells project into endolymph ,
whereas the bases are bathed into the perilymph , this
arrangement is necessary for production of generator pot
• tunnel of corti & the bases of hair cells get filled with
perilymph
• as Reissner’ memb is impermeable to Na & K ions ,
reticular lamina is a barrier to diffusion
• Therefore perilymph from the scala tympani diffuses
across basilar membrane into the tunnel of corti & bases
of hair cells
• hair cell depolarization intiates the release of chemical
transmitter glutamine that activates receptor sites on the
terminals of the afferent neuron
• It causes the generation of action potential in the neuron
• As the nerve fibres from the organ of cortii traverse the
tunnel of cortii, they would be unable to conduct impulses if it
were filled with endolymph which has the same electrolyte
composition as ICF
• The freq of action pot of single first order neuron of 8 th nerve
is directly prop to loudness ( intensity ) of sound
• At low sound intensities ,each axon discharges to sound of
only one freq
• At higher sound intensities, the individual
axons discharge to a wider range of sound
freq ,specially to freq lower than at which
threshold stimulation occurs
• The dotted line shows the boundary area of
nerve fiber
• Thus the low sound intensities will displace
only a limited area of the basilar membrane to
the max,an individual nerve ending may signal
• with high sound intensities, displace larger area
of basilar membrane
• & corresponding other nerve endings are
activated
• at higher sound intensities ,the single second
order neurons show responses to sounds of a
wider range of freq
• there is a sharper cut-off of action pot discharge
to freq below that causing the maximal response
• Pitch discrimination
.remarkable power is to discriminate between sounds
in 64-2000Hz
.according to duplex theory ,diff freq of sound
activates diff areas of basilar memb,causing max
receptor stimulation at one point
.since pathways from the various parts of cochlea to
the brain are separate
. So each auditory nerve fiber get max stimulated by
particular freq
 Intensity discrimitation
• The freq of firing in an auditory nerve fiber
increases as the intensity of sound wave
increases
• a large portion of basilar memb is vibrated as
sound intensity increases,more auditory nerve
fibers get activated
• sound intensityis discrminated by freq of 8th
nerve discharge & no. Of 8th nerve fibres get
activated
 Auditory cortex
• In primary auditory ,there is an orderly tonotropic
representation
• anterolateral part receives impulses arising from
apex of cochlea
• postero medial part receives impulses from base of
cochlea
• this tonotrophic representation is represented
through the auditory pathway
• selective destruction of small portion of MGB
causes deafness for restricted bands of freq
• auditory cortex is concerned with recognition of tonal
patterns , analysis of properties of sound & sound
localization
• sound localization depends on difference of time(as little
as 0.02 msec) & on the difference between the intensity
of sound in two
• difference of time best operates best when freq of
sound is less than 3000Hz
• difference between intensity of sound in two ears sound
localization operates best when freq is more than
3000Hz
• sound localization is markedly interrupted by
lesions of auditory cortex
• many neurons in the auditory cortex receive
input from both ears & they respond
maximally or minimally when the time of
entrance of sound at one ear is delayed by a
fixed time period relative to the time of
entrance of sound at the other ear
 Deafness
• It means inability on a part of an individual to
hear either wholly or partially
• 2 types : conductive deafness & nerve
deafness
• causes of conductive deafness-wax,foreign
body,thickness of tympanic memb,otitis
media, otosclerosis,blockage of eustachian
tube
• causes for nerve
deafness:presbycusis ,hereditary,injury to 8th
nerve,hazards of industrial noise,streptomycin
toxic degenration,tumour, vascular
degeneration
• tinnitus : ringing sensation in ears by irritative
stimulation of int ear or 8 th nerve
 Tests for hearing
• use of human voice
• tuning fork tests
• audiometry
• Treat the cause
• as treatment hearing aids are used
 Theories of hearing
• How does the ear discriminate such a wide
range of sound frequencies(pitch) & also
appreciate their intensities(loudness)?
• A no. Of conceptual frameworks have evolved
like telephone theory, travelling wave theory,
place theory, volley theory etc.
• But each falls short in explaining one or the
other element in the ability to appreciate
sound
 Studied under two variables
• Determination of the frequency(pitch) of
sound
• Determination of sound intensity(loudness)
 Determination of the frequency (pitch) of
sound
• PLACE THEORY
 was proposed in the 19th century by Helmholtz, assumes that
different regions of the basilar membrane are stimulated to
different frequencies
 discrimination of frequencies is a function of a number of
spikes in a fibre along with recruitment of fibres
 The ear has the highest senstivity for pitch discrimination for
frequencies <2000Hz
 It is performed at the apical turn of cochlea
 Therefore , doubt arises as to how be it possible in such a small
loop to have such high selectivity; this is main drawback of this
theory
 TRAVELLING WAVE THEORY
• It is observed that for lower frequencies, the
whole of basilar membrane vibrates(travelling
wave) & as the frequency increases ,only part
of it vibrates
• Therefore due to the travelling wave,
resonance point differs for different frequencies
• But as the frequency increases , the area of
maximum vibration increases & the afferent
neural spikes become irregular
• It appears that if frequency of tones
presented are not too close together,they are
discriminated pretty well by the ear itself
• At high frequency ,there also seems to be
inhibition of low tone