2

THE ANATOMY AND PHYSIOLOGY OF THE

EAR AND HEARING
Peter W.Alberti
Professor em. of Otolaryngology Visiting Professor University of Singapore
University of Toronto Department of Otolaryngology
Toronto 5 Lower Kent Ridge Rd
CANADA SINGAPORE 119074
[email protected]

2.1. INTRODUCTION

Hearing is one of the major senses and like vision is important for distant warning and
communication. It can be used to alert, to communicate pleasure and fear. It is a conscious
appreciation of vibration perceived as sound. In order to do this, the appropriate signal must
reach the higher parts of the brain. The function of the ear is to convert physical vibration into
an encoded nervous impulse. It can be thought of as a biological microphone. Like a
microphone the ear is stimulated by vibration: in the microphone the vibration is transduced
into an electrical signal, in the ear into a nervous impulse which in turn is then processed by
the central auditory pathways of the brain. The mechanism to achieve this is complex. This
chapter will deal mainly with the ear, first its structure and then its function, for it is the ear
that is mainly at risk from hazardous sounds.
The ears are paired organs, one on each side of the head with the sense organ itself, which
is technically known as the cochlea, deeply buried within the temporal bones. Part of the ear
is concerned with conducting sound to the cochlea, the cochlea is concerned with transducing
vibration. The transduction is performed by delicate hair cells which, when stimulated,
initiate a nervous impulse. Because they are living, they are bathed in body fluid which
provides them with energy, nutrients and oxygen. Most sound is transmitted by a vibration of
air. Vibration is poorly transmitted at the interface between two media which differ greatly in
characteristic impedance (product of density of the medium and speed of sound within it, c),
as for example air and water. The ear has evolved a complex mechanism to overcome this
impedance mis-match, known as the sound conducting mechanism. The sound conducting
mechanism is divided into two parts, an outer and the middle ear, an outer part which catches
sound and the middle ear which is an impedance matching device. Let us look at these parts
in detail (see Figure 2.1).

2.2. SOUND CONDUCTING MECHANISMS

2.2.1. The Outer Ear

The outer ear transmits sound to the tympanic membrane. The pinna, that part which protrudes
from the side of the skull, made of cartilage covered by skin, collects sound and channels it into
54 Anatomy and physiology of the ear and hearing

the ear canal. The pinna is angled so that it catches sounds that come from in front more than
those from behind and so is already helpful in localizing sound. Because of the relative size
of the head and the wavelength of audible sound, this effect only applies at higher
frequencies. In the middle frequencies the head itself casts a sound shadow and in the lower
frequencies phase of arrival of a sound between the ears helps localize a sound. The ear canal
is about 4 centimetres long and consists of an outer and inner part. The outer portion is lined
with hairy skin containing sweat glands and oily sebaceous glands which together form ear
wax. Hairs grow in the outer part of the ear canal and they and the wax serve as a protective
barrier and a disinfectant. Very quickly however, the skin of the ear canal becomes thin and
simple and is attached firmly to the bone of the deeper ear canal, a hard cavity which absorbs
little sound but directs it to the drum head (eardrum or tympanic membrane) at its base. The
outer layer of the drumhead itself is formed of skin in continuity with that of the ear canal.

Figure 2.1. The pinna and external auditory canal form the outer ear, which is separated
from the middle ear by the tympanic membrane. The middle ear houses three ossicles, the
malleus, incus and stapes and is connected to the back of the nose by the Eustachian tube.
Together they form the sound conducting mechanism. The inner ear consists of the cochlea
which transduces vibration to a nervous impulse and the vestibular labyrinth which houses
the organ of balance. (from Hallowell and Silverman, 1970)

In life, skin sheds and is continuously renewing. Ear canal skin grows like a fingernail
fromthe depths to the exterior so that the skin is shed into the waxy secretions in the
outer part
 Anatomy and physiology of the ear and hearing 55

and falls out. This is the reason for not using cotton buds to clean the ear canal because very
frequently they merely push the shed skin and wax deep into the canal, impacting it and
obstructing hearing. The ear canal has a slight bend where the outer cartilaginous part joins the
bony thin skinned inner portion, so that the outer part runs somewhat backwards and the inner
part somewhat forwards. This bend is yet another part of the protective mechanism of the ear,
stopping foreign objects from reaching the tympanic membrane. However it means that to inspect
the tympanic membrane from the outside, one must pull the ear upwards and backwards. The
tympanic membrane separates the ear canal from the middle ear and is the first part of the sound
transducing mechanism. Shaped somewhat like a loudspeaker cone (which is an ideal shape for
transmitting sound between solids and air), it is a simple membrane covered by a very thin layer
of skin on the outside, a thin lining membrane of the respiratory epithelium tract on the inner
surface and with a stiffening fibrous middle layer. The whole membrane is less than a 1/10th of
millimetre thick. It covers a round opening about 1 centimetre in diameter into the middle ear
cavity. Although the tympanic membrane is often called the ear drum, technically the whole
middle ear space is the ear drum and the tympanic membrane the drum skin.

2.2.2. The Middle Ear

The middle ear is an air filled space connected to the back of the nose by a long, thin tube
called the Eustachian tube. The middle ear space houses three little bones, the hammer, anvil
and stirrup (malleus, incus and stapes) which conduct sound from the tympanic membrane to
the inner ear. The outer wall of the middle ear is the tympanic membrane, the inner wall is the
cochlea. The upper limit of the middle ear forms the bone beneath the middle lobe of the
brain and the floor of the middle ear covers the beginning of the great vein that drains blood
from the head, the jugular bulb. At the front end of the middle ear lies the opening of the
Eustachian tube and at its posterior end is a passageway to a group of air cells within the
temporal bone known as the mastoid air cells. One can think of the middle ear space shaped
rather like a frying pan on its side with a handle pointing downwards and forwards (the
Eustachian tube) but with a hole in the back wall leading to a piece of spongy bone with
many air cells, the mastoid air cells. The middle ear is an extension of the respiratory air
spaces of the nose and the sinuses and is lined with respiratory membrane, thick near the
Eustachian tube and thin as it passes into the mastoid. It has the ability to secret mucus. The
Eustachian tube is bony as it leaves the ear but as it nears the back end of the nose, in the
nasopharynx, consists of cartilage and muscle. Contracture of muscle actively opens the tube
and allows the air pressure in the middle ear and the nose to equalize.
Sound is conducted from the tympanic membrane to the inner ear by three bones, the
malleus, incus and stapes. The malleus is shaped like a club; its handle is embedded in the
tympanic membrane, running from its centre upwards. The head of the club lies in a cavity of
the middle ear above the tympanic membrane (the attic) where it is suspended by a ligament
from the bone that forms the covering of the brain. Here the head articulates with the incus
which is cone shaped, with the base of the cone articulating with the head of the malleus, also
in the attic. The incus runs backwards from the malleus and has sticking down from it a very
little thin projection known as its long process which hangs freely in the middle ear. It has a
right angle bend at its tip which is attached to the stapes(stirrup), the third bone shaped with
an arch and a foot plate. The foot plate covers the oval window, an opening into the vestibule
of the inner ear or cochlea, with which it articulates by the stapedio-vestibular joint.
56 Anatomy and physiology of the ear and hearing

2.3. THE SOUND TRANSDUCING MECHANISM

2.3.1. The Inner Ear

2.3.1.1. Structure

The bony cochlea is so called because it is shaped like a snail shell It has two and a half turns
and houses the organ of hearing known as the membranous labyrinth surrounded by fluid
called the perilymph. The cochlea has a volume of about 0.2 of a millilitre. In this space lie
up to 30,000 hair cells which transduce vibration into nervous impulses and about 19,000
nerve fibres which transmit the signals to and from the brain. It is easiest to think of the
membranous labyrinth by imagining the cochlea to be straightened out as a bony tube closed
at the apex and open at the base with the round and oval windows and a connection to the
vestibular labyrinth (see Figure 2.2). It is in continuity with the vestibular labyrinth or organ
of balance which in technical terms acts as both a linear and angular accelerometer, thus
enabling the brain to know the position of the head in relationship to gravity and its
surroundings. The organ of balance will not be dealt with any further.
( Koklea tulang yang disebut demikian karena berbentuk seperti cangkang siput ini memiliki dua setengah
berbalik dan rumah organ pendengaran dikenal sebagai labirin membran dikelilingi oleh cairan yang disebut
perilymph tersebut. koklea memiliki volume sekitar 0,2 mililiter. Dalam ruang ini berbohong hingga sel 30.000
rambut yang transduce getaran menjadi impuls saraf dan sekitar 19.000 serabut saraf yang mengirimkan sinyal
ke dan dari otak. Hal ini paling mudah untuk berpikir dari labirin membran dengan membayangkan koklea yang
akan diluruskan sebagai tabung tulang ditutup pada puncak dan terbuka di dasar dengan putaran dan jendela oval
dan koneksi ke labirin vestibular (lihat Gambar 2.2). Hal ini dalam kontinuitas dengan labirin vestibular atau
organ keseimbangan yang dalam istilah teknis bertindak baik sebagai linear dan accelerometer sudut, sehingga
memungkinkan otak untuk mengetahui posisi kepala dalam hubungan dengan gravitasi dan sekitarnya. Organ
keseimbangan tidak akan dibahas lebih lanjut.)

Figure 2.2. The cochlea is a bony tube, filled with perilymph in which floats the
endolymph filled membranous labyrinth. This separates the scala vestibuli from the
scala media. (from Hallowell and Silverman, 1970)

Vibration of the foot plate of the stapes vibrates the perilymph in the bony cochlea. This
fluid is essentially incompressible. Therefore, there has to be a counter opening in the
labyrinth to allow fluid space to expand when the stapes foot plate moves inwards and in turn
to move inwards when the stapes foot plate moves outwards. The counter opening is provided
by the round window membrane which lies beneath the oval window in the inner wall of the
middle ear. It is covered by a fibrous membrane which moves synchronously but in opposite
phase with the foot plate in the oval window.
The membranous labyrinth is separated into three sections, by a membranous sac of
triangular cross section which run the length of the cochlea. The two outer sections are
the scala vestibuli which is connected to the oval window, and the scala tympani which is
connected to the round window. The sections are filled with perilymph; they connect at
the apex by a small opening known as the helicotrema which serves as a pressure
equalizing mechanism at frequencies well
 Anatomy and physiology of the ear and hearing 57

below the audible range. They also connect at the vestibular end with the fluid surrounding the
brain, through a small channel known as the perilymphatic aqueduct. The membranous labyrinth,
also known as the cochlear duct, is filled with different fluid called endolymph. On one side it is
separated from the scala vestibuli by Reissner's membrane, and on the opposite side from the
scala tympani by the basilar membrane (see Figure 2.3). The basilar membrane is composed of a
great number of taut, radially parallel fibres sealed between a gelatinous material of very weak
shear strength. These fibres are resonant at progressively lower frequencies as one progresses
from the basal to the apical ends of the cochlea. Four rows of hair cells lie on top of the basilar
membrane, together with supporting cells. A single inner row is medial, closest to the central core
of the cochlea. It has an abundant nerve supply carrying messages to the brain. The three outer
rows, which receive mainly an afferent nerve supply, are separated from the inner row by tunnel
cells forming a stiff structure of triangular cross section known as the tunnel of Corti (see Figure
2.3). Any natural displacement of the cochlear partition results in a rocking motion of the tunnel
of Corti and consequently a lateral displacement of the inner hair cells.

Figure 2.3. A cross section of one turn of the cochlea showing details of the
membranous labyrinth. (from Hallowell and Silverman, 1970)

The hair cells derive their name from the presence at their free ends of stereocilia which
are tiny little stiff hair like structures of the order of a few micrometers long (Figure 2.4). The
stereocilia of the hair cells are arranged in rows in a very narrow cleft called the subtectorial
space formed by the presence above the hair cells of the radially stiff tectorial membrane. The
58 Anatomy and physiology of the ear and hearing

cilia of the outer hair cells are firmly attached to the tectorial membrane while the cilia of the
inner hair cells are either free standing are loosely attached to the tectorial membrane.
In summary then, anatomically, the ear consists of a sound conducting mechanism and a
sound transducing mechanism. The sound conducting mechanism has two parts, the outer ear
consisting of the pinna and ear canal, and the middle ear consisting of the tympanic
membrane. The middle ear air space is connected to the nose by the Eustachian tube and to
the mastoid air cells housing the ossicular chain, the malleus, stapes and incus. The inner ear,
or cochlea, transduces vibration transmitted to the perilymph via the ossicular chain into a
nervous impulse which is then taken to the brain where it is perceived as sound.

Figure 2.4. A surface view looking down on the top of the hair cells; note the three rows
of outer hair cells and the one row of inner cells.

2.3.1.2. Function

Transduction of vibration in the audible range to a nervous impulse is performed by the inner hair
cells; when the basilar membrane is rocked by a travelling wave, the cilia of the inner hair cells
are bent in relation to the body of the cell, ion passages are opened or closed in the body of the
cell and the afferent nerve ending which is attached to the hair cell base is stimulated (Transduksi
getaran di kisaran terdengar untuk impuls saraf dilakukan oleh sel-sel rambut bagian dalam;
ketika membran basilar adalah diguncang oleh gelombang berjalan, silia dari sel-sel rambut
bagian dalam yang bengkok dalam kaitannya dengan tubuh sel, bagian ion dibuka atau ditutup
dalam tubuh sel dan akhir aferen saraf yang melekat dasar sel rambut dirangsang.).
As mentioned earlier, the basilar membrane responds resonantly to highest frequencies at
the basal end nearest the oval window and to progressively lower frequencies as one
progresses toward the apical end. At the apical end the basilar membrane responds resonantly
to the lowest frequencies of sound. A disturbance introduced at the oval window is
transmitted as a wave which travels along the basilar membrane with the remarkable property
that as each frequency component of the travelling wave reaches its place of resonance it
stops and travels no further. The cochlea is thus a remarkably efficient frequency analyser
(Seperti disebutkan sebelumnya, membran basilar merespon resonantly ke frekuensi tertinggi
pada akhir basal terdekat jendela oval dan frekuensi untuk semakin rendah sebagai salah satu
berlangsung menjelang akhir apikal. Pada akhir apikal membran basilar merespon resonantly
dengan frekuensi terendah suara. Gangguan diperkenalkan pada jendela oval ditransmisikan
sebagai gelombang yang bergerak sepanjang membran basilar dengan properti yang luar biasa
bahwa setiap komponen frekuensi gelombang perjalanan mencapai tempatnya resonansi
berhenti dan perjalanan tidak lebih. koklea demikian analisa frekuensi sangat efisien.)
The cochlea has an abundant nerve supply both of fibres taking impulses from the cochlea to
the brain (afferent pathways) and fibres bringing impulses from the brain to the cochlea (efferent
fibres). When stimulated the inner hair cells trigger afferent nervous impulses to the brain. Like
virtually all neural-mechanisms there is an active feedback loop. The copious nerve supply to the
outer hair cells is overwhelmingly efferent, although the full function of the efferent
(koklea memiliki pasokan saraf yang melimpah baik dari serat mengambil impuls dari koklea
ke otak (jalur aferen) dan serat membawa impuls dari otak ke koklea (serat eferen). Ketika
dirangsang sel-sel rambut bagian dalam memicu impuls aferen saraf ke otak. Seperti hampir
semua saraf-mekanisme ada umpan balik yang aktif. Pasokan saraf berlebihan pada sel-sel rambut
luar adalah sangat eferen, meskipun fungsi penuh eferen yang)
 Anatomy and physiology of the ear and hearing 59

pathways is not yet fully understood. It has been suggested that the purpose of the active
feedback system which has been described is to maintain the lateral displacement of the
stereocilia in the sub tectorial space within some acceptable limits (jalur belum sepenuhnya
dipahami. Ia telah mengemukakan bahwa tujuan dari sistem umpan balik aktif yang telah
dijelaskan adalah untuk menjaga perpindahan lateral Stereosilia di ruang tectorial sub dalam
beberapa batas yang dapat diterima)

2.4. THE PHYSIOLOGY OF HEARING (How does this all work?)

2.4.1. The Outer and Middle Ears

Let us deal first with the sound conducting mechanism. The range of audible sound is
approximately 10 octaves from somewhere between 16 and 32 Hz (cycles per second) to
somewhere between 16,000 and 20,000 Hz. The sensitivity is low at the extremes but
becomes much more sensitive above 128 Hz up to about 4,000 Hz when it again becomes
rapidly less sensitive. The range of maximum sensitivity and audibility diminishes with age.
(Mari kita berurusan pertama dengan suara mekanisme melakukan. Kisaran suara yang dapat
didengar adalah sekitar 10 oktaf dari suatu tempat antara 16 dan 32 Hz (siklus per detik) ke
suatu tempat antara 16.000 dan 20.000 Hz. sensitivitas rendah pada ekstrem tapi menjadi jauh
lebih sensitif atas 128 Hz sampai sekitar 4.000 Hz ketika lagi menjadi cepat kurang sensitif.
Kisaran sensitivitas maksimum dan kemampuan mendengar berkurang dengan usia.

The head itself acts as a natural barrier between the two ears and thus a sound source at
one side will produce a more intense stimulus of the ear nearest to it and incidentally the
sound will also arrive there sooner, thus helping to provide a mechanism for sound
localization based on intensity and time of arrival differences of sound. High frequency
hearing is more necessary than low frequency hearing for this purpose and this explains why
sound localization becomes difficult with a high frequency hearing loss. The head in humans
is large in comparison to the size of the pinna so the role of the pinna is less than in some
other mammals. Nonetheless, its crinkled shape catches higher frequency sounds and funnels
them into the ear canal. It also blocks some higher frequency sound from behind, helping to
identify whether the sound comes from the front or the back.
(Kepala itu sendiri bertindak sebagai penghalang alami antara dua telinga dan dengan
demikian sumber suara di satu sisi akan menghasilkan stimulus lebih intens dari telinga
terdekat untuk itu dan kebetulan suara juga akan tiba di sana lebih cepat, sehingga membantu
untuk menyediakan mekanisme untuk suara lokalisasi berdasarkan intensitas dan waktu
perbedaan kedatangan suara. pendengaran frekuensi tinggi adalah lebih penting daripada
rendah pendengaran frekuensi untuk tujuan ini dan ini menjelaskan mengapa lokalisasi suara
menjadi sulit dengan gangguan pendengaran frekuensi tinggi. Kepala pada manusia besar
dibandingkan dengan ukuran pinna sehingga peran pinna kurang dari di beberapa mamalia
lainnya. Meskipun demikian, bentuk berkerut yang menangkap suara frekuensi yang lebih
tinggi dan saluran mereka ke dalam liang telinga. Hal ini juga blok beberapa tinggi suara
frekuensi dari belakang, membantu untuk mengidentifikasi apakah suara itu berasal dari
bagian depan atau belakang)
The ear canal acts as a resonating tube and actually amplifies sounds at between 3000 and
4,000 Hz adding to the sensitivity (and susceptibility to damage) of the ear at these frequencies.
(Liang telinga bertindak sebagai tabung beresonansi dan benar-benar menguatkan suara di antara
3000 dan 4000 Hz menambah sensitivitas (dan kerentanan terhadap kerusakan) dari telinga pada
frekuensi ini)
The ear is very sensitive and responds to sounds of very low intensity, to vibrations which
are hardly greater than the natural random movement of molecules of air. To do this the air
pressure on both sides of the tympanic membrane must be equal. Anyone who has their ear
blocked even by the small pressure change of a rapid elevator ride knows the truth of this.
The Eustachian tube provides the means of the pressure equalization. It does this by opening
for short periods, with every 3rd or 4th swallow; if it were open all the time one would hear
one's own every breath (telinga sangat sensitif dan merespon suara intensitas sangat rendah,
getaran yang hampir tidak lebih besar dari gerakan acak alami molekul udara. Untuk
melakukan hal ini tekanan udara di kedua sisi membran timpani harus sama. Siapa saja yang
memiliki telinga mereka diblokir bahkan oleh perubahan tekanan kecil naik lift yang cepat
tahu kebenaran ini. Tabung Eustachian menyediakan sarana pemerataan tekanan. Hal ini
dilakukan dengan membuka untuk jangka pendek, dengan setiap 3 atau 4 walet; jika itu
terbuka sepanjang waktu satu akan mendengar sendiri setiap nafas)

Because the lining membrane of the middle ear is a respiratory membrane, it can absorb
some gases, so if the Eustachian tube is closed for too long it absorbs carbon dioxide and
oxygen from the air in the middle ear, thus producing a negative pressure. This may produce
pain (as experienced if the Eustachian tube is not unblocked during descent of an aeroplane).
The middle ear cavity itself is quite small and the mastoid air cells act as an air reservoir
cushioning the effects of pressure change. If negative pressure lasts too long, fluid is secreted
by the middle ear, producing a conductive hearing loss.
The outer and middle ears serve to amplify the sound signal. The pinna presents a fairly
large surface area and funnels sound to the smaller tympanic membrane; in turn the surface of
the tympanic membrane is itself much larger than that of the stapes foot plate, so there is a
hydraulic amplification: a small movement over a large area is converted to a larger
movement of a smaller area. In addition, the ossicular chain is a system of levers which serve
to amplify the sound. The outer and middle ears amplify sound on its passage from the
exterior to the inner ear by about 30 dB (Karena membran selaput telinga tengah adalah
membran pernapasan, dapat menyerap beberapa gas, jadi jika tabung Eustachio ditutup terlalu
lama menyerap karbon dioksida dan oksigen dari udara di telinga tengah, sehingga
menghasilkan tekanan negatif. Hal ini dapat menghasilkan nyeri (seperti yang dialami jika
tabung Eustachio tidak diblokir selama keturunan dari pesawat terbang). Rongga telinga
tengah itu sendiri cukup kecil dan sel-sel udara mastoid bertindak sebagai reservoir udara
bantalan efek perubahan tekanan. Jika tekanan negatif berlangsung terlalu lama, cairan
disekresi oleh telinga tengah, menghasilkan gangguan pendengaran konduktif.)
60 Anatomy and physiology of the ear and hearing

2.4.2. The Inner Ear

The function of the inner ear is to transduce vibration into nervous impulses. While doing so,
it also produces a frequency (or pitch) and intensity (or loudness) analysis of the sound.
Nerve fibres can fire at a rate of just under 200 times per second. Sound level information is
conveyed to the brain by the rate of nerve firing, for example, by a group of nerves each
firing at a rate at less than 200 pulses per second. They can also fire in locked phase with
acoustic signals up to about 5 kHz. At frequencies below 5 kHz, groups of nerve fibres firing
in lock phase with an acoustic signal convey information about frequency to the brain. Above
about 5 kHz frequency information conveyed to the brain is based upon the place of
stimulation on the basilar membrane. As an aside, music translated up into the frequency
range above 5 kHz does not sound musical.
As mentioned above each place along the length of the basilar membrane has its own
characteristic frequency, with the highest frequency response at the basal end and lowest
frequency response at the apical end. Also any sound introduced at the oval window by
motion of the stapes is transmitted along the basilar membrane as a travelling wave until all
of its frequency components reach their respective places of resonance where they stop and
travel no further. For example, a 1 kHz tone induces resonance at about the middle of the
basilar membrane. Any frequency components lower than 1 kHz must travel more than half
the length of the basilar membrane, whereas high frequency components, greater than 1 kHz
must travel less than half the length of the basilar membrane. Evidently the brain must
suppress high frequency information in favour of low frequency information as the travelling
wave on the basilar membrane passes through places of high frequency resonant response. An
explanation is thus provided for the observation that low frequency sounds, for example
traffic noise, are very effective in masking high frequency sounds, for example the fricatives
of speech, making telephones near busy streets difficult to use.
How does the brain cope with intensity? The physiological range of intensity of the normal
ear is huge. As a matter of interest it is the same as that of the eye when the responses of the cones
and rods are considered together; thus the visual analogue is appropriate. It is as wide as seeing a
candle flicker on a dark night at a hundred meters to looking indirectly into a bright sun. The
range is so great that only the logarithmic response characteristic of variable rate processes and
thus favoured by anatomical systems, is capable of encompassing it. The normal range of human
hearing is from 0 to 100 dB(A), before sound becomes uncomfortably loud.
Mounted on the basilar membrane close to the end nearest the central core of the cochlea are a
single row of inner hair cells followed by three rows of outer hair cells which are separated from
the single row of inner hair cells by a stiff structure of triangular cross section known as the
tunnel of Corti. Any natural displacement of the cochlear partition results in a rocking motion of
the tunnel of Corti and consequently a lateral displacement of the inner hair cells.
The ear has evolved a very intriguing mechanism to cope with the large range in
sound intensity encountered in the environment. Only the inner hair cells initiate nervous
impulses which are heard as sound. They are not particularly sensitive but they are
rugged and they are placed at the inner edge of the basilar membrane which is relatively
immobile . The point where the basilar membrane vibrates most is about its middle so
that the inner hair cells are spared the most violent vibration of very intense sound. The
question then arises, how do the inner hair cells respond to slight or moderate amounts
of stimulation? Here the outer hair cells play a major role. When they are stimulated by
the travelling wave they respond actively and physically contract. They have muscle
proteins in their wall and literally shorten. Because they are attached
 Anatomy and physiology of the ear and hearing 61

both to the Reissner's membrane and the basilar membrane, this produces an additional shear
movement of the membranous labyrinth, which amplifies the travelling wave at the point of
maximal stimulation. This amplified movement is transmitted to the inner hair cells which then
respond. If the amount of movement of the basilar membrane is slight, the amount of outer hair
cell contracture adds significantly to the basilar cell movement; if the amount of movement is
large the contracture adds nothing to the already great displacement of the membranous labyrinth.
If the outer hair cells are damaged they no longer contract in response to slight sounds and the
inner hair cells are not stimulated. This produces a hearing loss for low intensity sound. If the
sound is more intense, the inner hair cells are stimulated directly and they respond normally so
that the ability to hear louder sounds remain unimpaired. This is a common phenomenon known
as loudness recruitment. The inner hair cells are much "tougher" than outer hair cells and much
less likely to be damaged by ageing, noise or most ototoxic drugs, so ageing, noise and ototoxic
drugs usually only produce hearing loss but not deafness. It was noted earlier that the ear is most
sensitive to sounds between approximately 3000 and 4000 Hz, in part because of the amplifying
mechanism of the ear canal. Thus, the most intense stimulus is produced at these frequencies and
the outer hair cells which respond to these frequencies are most at risk from damage. Prolonged
exposure to loud sounds damages these hair cells and thus explains the hearing loss from noise
which occurs first at 3 to 4 kHz.

2.5. CENTRAL AUDITORY PROCESSING

The nervous impulses are carried along the 8th (statico-acoustic nerve) from the cochlea to
the brain stem. Here the nerve fibres reach nuclei where they relay with other nerve fibres.
The fibres from each auditory nerve split, some passing to one side of the brain, others
remaining on the same side. Thus, as auditory stimuli pass up each side of the brain from both
ears, unilateral hearing loss cannot be caused by a brain lesion. The fibres pass up the hind
brain to the mid brain and the cerebral cortex. There are many central functions, some of
which will be examined but most of which lie outside the scope of this chapter.( Impuls saraf
dilakukan di sepanjang 8 (statico-akustik saraf) dari koklea ke batang otak. Berikut serabut
saraf mencapai inti di mana mereka menyampaikan dengan serabut saraf lainnya. Serat dari
setiap perpecahan saraf pendengaran, beberapa melewati salah satu sisi otak, orang lain yang
tersisa pada sisi yang sama. Dengan demikian, sebagai rangsangan pendengaran melewatkan
setiap sisi otak dari kedua telinga, gangguan pendengaran unilateral tidak dapat disebabkan
oleh lesi otak. Serat melewatkan otak belakang ke pertengahan otak dan korteks serebral. Ada
banyak fungsi sentral, beberapa di antaranya akan diperiksa tetapi sebagian besar berada di
luar lingkup bab ini.)

2.5.1. The Ability to Block Out Unwanted Sounds.

In a crowded noisy room a young person with normal hearing can tune in and out
conversations at will. This is known technically as the cocktail party effect. The brain quite
automatically adjusts time of arrival and intensity differences of sound from different signal
sources so that the one which is wanted passes to the cortex and all others which do not meet
these criteria are suppressed by feedback loops. This requires both good high frequency
peripheral hearing, two ears and an additional central mechanism. Even in the presence of
normal bilateral peripheral hearing, the elderly lose part of the central mechanism and find it
difficult to listen in crowded rooms. This is compounded if there is some hearing loss. (Di
ruang bising ramai orang muda dengan pendengaran normal dapat mendengarkan dan keluar
percakapan di akan. Hal ini dikenal secara teknis sebagai efek pesta koktail. otak cukup
otomatis menyesuaikan waktu kedatangan dan intensitas perbedaan suara dari sumber sinyal
yang berbeda sehingga salah satu yang ingin lolos ke korteks dan semua orang lain yang tidak
memenuhi kriteria tersebut ditekan oleh loop umpan balik. Ini membutuhkan baik baik
frekuensi tinggi pendengaran perifer, dua telinga dan mekanisme sentral tambahan. Bahkan di
hadapan sidang perifer bilateral normal, orang tua kehilangan bagian dari mekanisme pusat
dan merasa sulit untuk mendengarkan di kamar ramai. Hal ini diperparah jika ada beberapa
gangguan pendengaran.)

2.5.2. Spatial Localization.

A normal human can localize quite accurately the source of the sound. One knows from what
direction the sound is coming; one knows where to turn one's head to look for a speaker; as
one knows where to look for an aeroplane or a bird. There are specific neurones which deal
with this in the mid brain.
(Seorang manusia normal dapat melokalisasi cukup akurat sumber suara. Yang tahu dari arah
mana suara itu berasal; satu tahu ke mana harus berpaling kepala seseorang untuk mencari
pembicara; sebagai salah satu tahu di mana untuk mencari pesawat terbang atau burung. Ada
neuron khusus yang menangani hal ini di otak pertengahan
62 Anatomy and physiology of the ear and hearing

2.5.3. On and Off Sounds

Hearing has an alerting function especially to warning signals of all kinds. There are brain
cells which respond only to the onset of a sound and others which respond only to the
switching off of the sound, i.e. a change. Think only of being in an air conditioned room
when the air conditioner turns on, one notices it. After a while it blends into the background
and is ignored. When it switches off, again one notices it for a short time and then too the
absence of sound blends into the background. These cells allow the ear to respond to acoustic
change - one adjusts to constant sound - change is immediately noticeable. This is true too
with machinery and a trained ear notices change (Mendengar memiliki fungsi menyiagakan
terutama untuk sinyal peringatan dari segala jenis. Ada sel-sel otak yang merespon hanya
untuk timbulnya suara dan lain-lain yang merespon hanya untuk switching off suara, yaitu
perubahan. Pikirkan hanya berada di ruang ber-AC ketika AC menyala, satu pemberitahuan
itu. Setelah beberapa saat itu memadukan ke latar belakang dan diabaikan. Ketika nonaktif,
lagi satu pemberitahuan itu untuk waktu yang singkat dan kemudian juga adanya suara
memadukan ke latar belakang. Sel-sel ini memungkinkan telinga untuk menanggapi
perubahan akustik - satu menyesuaikan dengan suara konstan - perubahan segera terlihat. Hal
ini berlaku juga dengan mesin dan terlatih perubahan pemberitahuan telinga.)

2.5.4. Interaction of Sound Stimuli with Other Parts of the Brain

Sound stimuli produce interaction with other parts of the brain to provide appropriate responses.
Thus, a warning signal will produce an immediate general reaction leading to escape, a
quickening of the heart rate, a tensing of the muscle and a readiness to move. A baby's cry will
alert the mother in a way it does not alert others. The sound of martial music may lead to bracing
movement of those to whom it is being played and induce fear and cowering in the hearts and
minds of those at whom it is being played. Certain sounds can evoke anger, others pleasure. The
point is that the sensations produced by hearing are blended into the body mechanism in the
central nervous system to make them part of the whole milieu in which we live (rangsangan suara
menghasilkan interaksi dengan bagian lain dari otak untuk memberikan tanggapan yang tepat.
Dengan demikian, sinyal peringatan akan menghasilkan reaksi umum langsung mengarah ke
melarikan diri, gejolak denyut jantung, sebuah menegang otot dan kesiapan untuk bergerak.
menangis A bayi akan mengingatkan ibu dengan cara itu tidak mengingatkan orang lain. Suara
musik bela diri dapat menyebabkan menguatkan gerakan mereka kepada siapa ia sedang
dimainkan dan menginduksi rasa takut dan meringkuk di hati dan pikiran orang-orang di
antaranya yang sedang dimainkan. suara-suara tertentu dapat membangkitkan kemarahan, orang
lain senang. Intinya adalah bahwa sensasi yang dihasilkan oleh pendengaran dicampur ke dalam
mekanisme tubuh dalam sistem saraf pusat untuk membuat mereka bagian dari seluruh
lingkungan di mana kita hidup.)

REFERENCES

Hallowell, Davis and S. Richard Silverman (Ed.), (1970). Hearing and Deafness, 3rd ed.,
Holt, Rinehart and Winston.
3

THE PATHOPHYSIOLOGY OF THE EAR

Peter W.Alberti
Professor em. of Otolaryngology Visiting Professor University of Singapore
University of Toronto Department of Otolaryngology
Toronto 5 Lower Kent Ridge Rd
CANADA SINGAPORE 119074
[email protected]

Things can go wrong with all parts of the ear, the outer, the middle and the inner. In the
following sections, the various parts of the ear will be dealt with systematically.

3.1. THE PINNA OR AURICLE

The pinna can be traumatized, either from direct blows or by extremes of temperature. A hard
blow on the ear may produce a haemorrhage between the cartilage and its overlying
membrane producing what is known as a cauliflower ear. Immediate treatment by drainage of
the blood clot produces good cosmetic results. The pinna too may be the subject of frostbite,
a particular problem for workers in extreme climates as for example in the natural resource
industries or mining in the Arctic or sub-Arctic in winter. The ears should be kept covered in
cold weather. The management of frostbite is beyond this text but a warning sign, numbness
of the ear, should alert one to warm and cover the ear.

3.2. THE EXTERNAL CANAL

3.2.1. External Otitis

The ear canal is subject to all afflictions of skin, one of the most common of which is
infection. The skin is delicate, readily abraded and thus easily inflamed. This may happen
when in hot humid conditions, particularly when swimming in infected water producing what
is known as swimmer's ear. The infection can be bacterial or fungal, a particular risk in warm,
damp conditions.
The use of ear muffs particularly in hot weather may produce hot, very humid conditions
in the ear canal leaving it susceptible to infection, and similarly insertion and removal of ear
plugs may produce inflammation. Although this is surprisingly rare; it does occur particularly
in those working with toxic chemicals. These people should take care to wash their hands
before inserting or removing ear plugs or preferably use ear muffs. The soft seal of a muff
should be kept clean and if reusable plugs are used, they should also be regularly washed.
Inflamed or infected ear canals should be treated by a physician.