CHAPTER 5: THE AUDITORY SYSTEM SOUND AND EAR

Audition: The detection of sound and ability to determine its source

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Essential for loacing food, finding mates and avoiding predators.

Audition is a distant sense stimulus can be detected from far away
Audition operations at the fastest time scale

The Physics of Sound

3 Essential Requirements to hear sound:
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Something that creates the sound

Sound must propagate through a medium
o Bell Jar experiment: the sound produced by a bell becomes les audible if enclosed in
a jar in which air is slowly pumped out
Mechanism to translate sound energy into biological signal to generate the experience
known as hearing

The Creation of Sound

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Vibrational Properties of Objects: Inertia + Elasticity = Vibrations

o Inertia an object will initially resist deformation, but once started it will continue
until an opposing force acts on it
o Elasticity The opposing force that brings an object to its original state
o i.e. using a tuning fork, vibrations of the prongs will create a simple harmonic
motion represented by a sinusoidal function.
Impact of a Sound Source on the Medium: Compression and rarefaction of air
o Compression Increased air pressure b/c of prongs moving outward pushing against
it
o Rarefaction Decrease air pressure b/c of prongs moving inwards leaving a space
o Sound wave created by the propagation of the momentary compression /
rarefaction of air

The Properties of Sound

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Amplitude and Sound Intensity:

o Amplitude pressure change from the baseline to the highest peak (measure in Pa
or kPa)
Increase in loudness of sound = Increase in amplitude
o Intensity a word used to describe the loudness of sound
Intensity of an objection is considered in relation to a reference point
Reference point = the lowest sound level that can be heard (20Pa)
Intensity is represented by Decibels (dB). A log scale where 10dB = 1bel
and 1bel = log(Is/Ir)
SPL = Sound pressure level.
dBSPL = the reference value use to calculate intensity is the lowest sound
level that can be heard
Sound Frequency: # of complete cycles / second. Measured in Hertz (Hz)
o Humans can only hear between 20-2000 Hz
o Natural (Resonant) Frequency frequency of an object due to its mass and
stiffness.
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o As stiffness (or tension) increases frequency increases

o As Mass increases frequency decreases
o As Length increases frequency decreases
Speed of Sound: 331.5m/s @ 0C in AIR
o Speed is entirely dependent on the characteristics of the medium inertia and
elasticity
o Inertia (density) of medium is related to speed inversely increase density
decrease speed
o Temperature Increase temp increases speed
Increasing temperature causes expansion of molecules decreases density
increase speed
1C increase = 0.6m/s increase in speed of sound
o Elasticity of medium is related to speed directly increase elasticity increase
speed
i.e. sound actually travesl 16x faster in steel than air b/c of elasticity

Complex Sounds
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Complex Periodic Sounds: When the pattern of pressure change repeats itself at regular
intervals over time
o It does not have a sinusoidal function BUT its formed by adding multiple sinusoidal
function with different frequencies
o Harmonic Series a highly periodic wave characterized by the summation of precise
series of sine waves (i.e. sound produced on a musical instrument)
Complex Aperiodic Sounds: When the pattern of pressure change does not repeat itself at
a regular interval
o Aka NOISE: White noise noise with frequencies within the human hearing range
(i.e. traffic sounds)
Fourier Analysis: Decomposing complex sound into the simple sine functions that formed it
o Fourier Spectrum a way to represent the deconstructed complex sound by using
vertical lines
X = Frequency (location of vertical line) and Y = amplitude (height of the
vertical line)

Removes the need to plot each component of the sine wave functions
Ohms law of Hearing: Human hear sound by deconstructing them into simple tones
similar to Fourier Analysis

Sound Transmission
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The Inverse Square Law: Sound intensity is inversely related to the square of the
distance from the sound source (i.e. If distance r is doubled to 2r intensity decreases to
1/4 of original)
o Only true if there were no objects in the way
Interaction of Sound with Object:
o Reflection when sound bounces off a medium with greater resistance
Echo (aka reverberations) = reflected sound
Reverberant room = a room with highly reflective walls
o Absorption depends on the absorption coefficient (= proportion of sound absorbed
compared to the initial sound) ; Anechoic = a room with highly absorptive walls
o Diffraction when sound waves bend around an object
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High frequencies sound cannot be diffracted as easily b/c of more

compressed undulations of pressure change.
Architectural Acoustics: warmth, brilliance, texture, and blend are important factors
o Reverberation time a factor used to quantified sound reflection
o More reverberation Concert halls will be perceived as having more Fullness
o Less reverberation Concert halls will be perceived as having more Clarity
o Intimacy the impression of being physically close to the sound source ( occurs if
the sound reaches the listener really quickly)

Auditory Processing of Sound Physical Characteristics

Anatomical Components of the Human Ear
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The Outer Ear: Pinna, external auditory canal and tympanic membrane (ear drum)
o Pinna funnels sounds into the ear. Composed of cartilages and skin
o Auditory Canal S-shaped canal lined with wax-secreting glands to protect the
insides of the ear
o Tympanic Membrane thin elastic membrane that transfer vibrations to the middle
ear. Also have protective functions against foreign bodies.
The Middle Ear:
o Eustachian Tube connects the middle ear to the throat to equalize pressure across
the ear drum
o Ossicles (malleus, incus, stapes) transfer the vibration from the ear drum to the
oval window.
Ossicles are suspended by ligaments to allow free vibrations
The Inner Ear: Located in a cavity of the temporal bone known as Bony Labyrinth
o Cochlea fluid filled bony structure that contains the sensory transduction
apparatus of hearing
o Interior of the Cochlea is divided into 2 fluid filled channels
Scala Vestibuli (biggest) uppermost channel. Basal end is connected to
the oval window
Scala tympani lowermost channel. The basal end contains the round
window
Scala media (cochlear duct) middle channel. Smallest.
The Reissners membrane separate this channel from the Scala
vestibule
The Basilar membrane separates this channel from the Scala
tympani
The fluid here is different from the other 2 and they dont mix
o Vestibuli and Tympani are joined together at the apex by a small opening =
Helicotrema
Sound Transmission through the Ear
o Pinna auditory canal tympanic membrane Ossicles oval window fluids
in the scala vestibule and Reissners membrane scala tympani + basilar
membrane Round window

Amplitude Preservation

The Lever Effect: Small amplification of sound as it passes malleus to the incus. Up to an
amplification of 2dB
The Condensation Effect: Large amplification of sound due to transfer of vibration from
large surface area of the tympanic membrane to the small area of the stapes bone. A gain
of 25dB.
The Resonance effect: The Pinna and ear canal has a resonant frequence of 4000-5000Hz
and 2500 Hz respectively. Therefore sound coming in at these frequencies will be be
amplified by 10-15dB.
The Directional effect: the ossicular chain of bones in the middle ear channels all the
incoming sound energy only upon the oval window.
o Without it, both oval and round window will respond equally to incoming pressure
resulting in no net movt of the cochlear fluid.

Frequency Representation
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The Frequency Theory: Neural firing rate = sound frequency; neural firing amplitude =
intensity; BUT
o Neurons fire all-or-nothing therefore no differences in amplitude, all are the same
o Theres actually a time constraint that limits how fast neurons can fire
o Width of basilar membrane is not uniform entire thing cannot vibrate at the same
frequency

The Place Theory: Sounds of different frequencies produce a vibrational pattern whose
maximum amplitude occurs at different places along the basilar membrane
o This is possible because the basilar membrane is narrow at the basal end and gets
wider at the apex
o The tension also decreases from the basal end to the apex
o As a sound travels across, there will be maximum displacement of the basilar
membrane at a site where it matches the incoming frequency.
o Narrow and high tension at the basal end receptive to high frequency
o Wide and low tension at the apex end receptive to low frequency
Tonotopic Organization
o The way the basilar membranes resonant frequency is organized creates a
tonotopic map.
o this allows effective neural coding of frequency
Frequency Analysis of complex Sounds
o The basilar membrane acts as a filter, a frequency analyzer, by separating complex
sounds into simple sinusoidal functions
o As the complex sounds enters the cochlea it creates a virbational disturbance in the
fluids. However b/c the basilar membrane only responds to certain frequency at
specific places, where the incoming sound spectrum matches the resonant
frequency on the membrane there will be greater displacement.

Auditory Processing of Sound Biological Mechanisms

Auditory Transduction
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Organ of Corti Structural Features

o The conversion of vibrations to neural signals occurs within this structure which lies
on top of the basilar membrane . It is entirely within the cochlear duct.
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The Arch of Corti a rigid inverted V structure that divides the organ into an inner
and outer portion
o Hair cells most important cells with regard to auditory signal processing
Inner hair cells located on the inner side of the arch and extends the full
length of cochlea
Outer hair cells located on the outer side of the arch and arranged into 3
rows extending the length of the cochlea
o Sterocilia fine filaments that protrude from the upper surface of hair cells.
o Tectorial membrane gelatinous structure that lies above the sterocillia
Organ of Corti Mechanical Response to Sound Stimulation
o When the basilar membrane moves up and down, the tectorial membrane also
bends creating a shearing force on to the steriocillia in between them
o The sterocillia hairs will bending according to yhr movement of the basilar
memebrane
Stereocillia bending outward upward deflection of the membrane
Stereocillia bending inward downward deflection of the membrane
Transudctional Mechanism in Hair Cells: PASSIVE response to auditory stimulation
o Most of the neural output from the cochlear actually comes from the inner hair cells
o Tip Link a very thin fibre that connects all the sterocillia.
o Bending of sterocillia Bending of the tip link which uncovers the ion gates (i.e. K+
and Ca2+) influx of ions depolarization.
This is important so that we can hear RAPIDLY
o 2nd depolarization + increase in Ca2+ release of neurotransmitter at the base of
the hair cells neurotransmitter (glutamate) depolarizes the cochlear nerve fibres
signal carried to higher centres
Roles of Outer Hair Cells: ACTIVE response to auditory stimulation Mechanical or
Electrical
o Mechanical Response of outer hair cells net effect is to help bend the tectorial
membrane
1st physical change within the contractile elements of the outer hair cells
2nd slower response occurs through nerve stimulation
o Electrical mechanism enhances the passive electrical response functions of the
inner hair cells
Cochlear microphonic an electrical response across the entire organ of Corti
by the movement of outer hair cells in response to basilar membrane
vibration
The ionic properties of the fluid create a steady current that is modulated by
the sterocilia of the outer hair cells. The electrical waveform is exactly the
same as the sound wave
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Neural Capture of Auditory Signals

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Vestibulocochlear Nerve (CN VIII): Main pathway for the transmission of auditory signals
out of the cochlea and into the CNS
Anatomical Organization
o Fibres of the cochlear nerve originate from the cochlear ganglion. The cochlear
ganglion lies just outside of the cochlear and follows the cochlear spiral Spiral
Ganglion
o The neurons are bipolar neurons with the shorter branch innervating the hair cells
o 90% of the cochlear nerve fibres terminate upon the inner hair cells.
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Afferent vs. Efferent nerve Fibres

Afferent Fibres
Receives signals mainly from Inner hair cells
1 neuron innervate 1 hair cell, but each hair cell
may have more than 1 nerve innervating it
Multiple nerves allow the different properties of
sound to be transmitted independently

Efferent Fibres
Sends signals mainly to Outer hair cells
Increase activity of these neurons reduce
firing rate of action potentials of afferent
Function unknown

Neural Coding of Sound Intensity

o Greater amplitude = greater deflection of basilar membrane = greater bending of
sterocilia = greater depolarization of inner hair cells = greater release of
neurotransmitter on to afferent nerve
o The 10 afferent nerves that innervate each hair cell have different intensity
threshold ranging from 0dB to 100dB and beyond. This encodes the entire audible
range of intensity.
o Hair cells can control how much neurotransmitter it releases to each afferent nerve
difference in neurotransmitter release means that it can trigger different levels of
neural activity among a parallel set of fibres
Neural Coding of Sound Frequency Place Theory Mechanisms
o Since the basilar membrane shows maximum vibrational amplitudes at different
places along its length in response to different sound frequencies, therefore a
particular afferent fibre will only carry signals that was depolarizedby the particular
sound frequency associated with its place in the membrane .
o Tuning Curve displays the minimum sound intensity required to obtain a neural
response as a function of frequency
Characteristic Frequency the lowest point on the curve
o Frequency Response Curve Shows the neural response to frequency and intensity
Neural Coding of Sound Frequency Frequency Theory Mechanisms
o Phase Lock Response All the afferent nerves produce an action potential at the
same frequency as the incoming sound. (i.e. an action potential is produced during
each cycle). Only works up to 4000Hz

Subcortical Auditory Structures

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2 Functions
o Serve as a relay mechanisms that transmits information from one site to another
o They enhance, modify, and further process the auditory signals
Ascending Pathways: Afferent nerves come together forming the Cranial Nerve VIII enter
brain stem and synapse at Cochlear nucleus Superior Olive + Inferior Colliculus on both
sides Inferior Colliculus will project to the Medial geniculate Nucleus on both side +
inferior colliculus of the opposite side From MGN it goes to the Auditory Cortex
o Cochlear Nucleus only receive inputs from their corresponding side. There is no
cross over at this level (monaural neurons). 2 types of cells Relay cells +
enhancing + sharpening signals cells
o Superior Olive Auditory Processing
o Inferior Colliculus Auditory relay + Coordination of acoustic reflexes (midbrain)
o Medial Geniculate Nucleus largely a relay signals to the cortex
Characteristics of Subcortical Structures
o Response Modification by way of inhibitory + excitatory signals in the cochlear
nucleus
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sharpens the neural representation important for sound localization +

detecting sounds that are present in a noisy background
o Presence of a Tonotopic Map prominent in the inferior colliculus in which each
layer codes for one frequency (Isofrequency sheets) ensures that frequency
information is retained
o Laterality the extent to which the subcortical neurons can be separately driven by
the 2 ears
Binaural Neurons excited by contralateral sound stimulation
This is consistent with the fact that neurons on one side of the brain
represents the opposite side of the body
Monaural Neurons can be excited or inhibited by ipsilateral sound input
Descending Pathway: Modulates auditory response to sound in 2 different way
o inhibition of outer hair cell by Olivocochlear neurons to allow the auditory system to
encode a higher range of sound intensities by preventing early saturation of
discharge rates.
o Activation of the 2 small muscles in the middle ear, Tensor Tympani and Stapedius
Muscular contractions lead to decreased sound transmission to protect the
middle and inner ear component from sudden burst of loud sound.

The Auditory Cortex

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Anatomical Organization:
o Area A1 The primary Auditory cortex in the temporal lobe which receives signals
from the MGN
o Area A2 Surrounds A1 and processes higher order acoustic information
o Wernickes Area located in the left hemisphere ONLY and is important for speech
comprehension
Functional Organization:
o Tonotopic Arrangement of the Auditory Cortex
Low Frequencies towards the anterior end of A1
Higher frequencies towards the posterior end of A1
o Columnar Organization of Neural response resulting from Binaural inputs
Summation Response inputs from both ears are excitatory and therefore the
neurons can be driven by sound stimulation of either ear
Suppression Response the neurons is excited by sound input from the
opposite side but is inhibited by stimulation on the same side
Parallel Auditory Pathways: It appears that neural circuits involved in sound pattern
analysis are separate from those involved in sound localization analysis

Auditory Dysfunction
Causes of Hearing Loss
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Conductive Loss: Occurs when the outer or middle ear is affected reduced sound
transmission to the cochlea
o Middle ear infection (aka otitis media) most common cause
o Otosclerosis inherited diseases that produces abnormal development and function
of the ossicles
caused by build up of calcium which restricts the movement of the bones
reducing the transfer of sound energy by 30dB
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o Other Causes blockage of the outer ear by ear wax or perforation of the ear drum
Sensorineural Loss: occurs when there is damage to the cochlea or to the nerves of the
inner ear
o Ingestion of Ototoxic drugs some antibiotics and aspirin b/c they destroy hair cells
leads to Tinnitus the perception of ringing in the ear in absence of
external sound
o Traumatic injury fracture of the temporal bone
o Tumours especially in cochlear nerve acoustic neurinomas
o Diseases rubella in pregnant mother, Menieres Diseases excessive cochlear
fluid production
o Intense Environmental Noise causes injury to the hair cells and the transduction
mechanism
aka Noise-induced hearing loss. >80dB will cause damage. >120dB its
painful
Deaf-mutism: the absence of language vocalization ability due to hearing loss in prelingual children
Hereditary Factors and Aging:
o Genetic Causes
Usher Syndrome autosomal recessive gene that causes variable degrees of
deafness either at birth or later in life. Often accompanied by visual
problems.
Waardenburg Syndrome causes hearing loss and changes in skin and hair
pigmentation.
2 different eye colour, hearing loss can vary from moderate to severe;
Dominant gene
o Presbycusis hearing loss due to aging. High pitch sounds are first to go.

Diagnosis and Treatment

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Bone Conduction test: use to distinguish b/w conductive and sensorineural loss by
applying a vibrating tuning fork somewhere on the skull
o Normal hear sound whether it is touching the skull or not
o Conductive can only hear sound when the tuning fork touches the skull
o Sensorineural cannot hear sound with tuning fork touching the skull or not.
Hearing Aids: amplifies incoming sound and delivers them to the ear. Only works if
cochlear is working
o Contains a microphone, amplifier and speaker
Cochlear implant: contains a small microphone and processor that converts the sound
signal into an electric current that is delivered to the inner ear via fine wires
o Single Channel Inputs one wire to the cochlear nerve. No good.
o Multi-channel Inputs 20+ wires that delivers different frequency to the different
nerve along the basilar membrane