Sound Waves: Stimulus for Audition

Sounds are created when objects vibrate

Vibrations of an object cause molecules in the objects surrounding medium to
vibrate as well, which causes pressure changes in the medium.

Sound waves travel at a particular speed depending on the medium.

Examples:
Speed of sound through air is about 340 meters/second.
Speed of sound through water is 1500 meters/second.

Physical qualities of sound waves

Amplitude: The magnitude

(intensity) of air molecules
moving through the air.
Frequency: The number of
times per second that a pattern
of pressure repeats.
Units for measuring sound:
Hertz (Hz): A unit of measure
for frequency. One Hz equals
one cycle per second.
Decibel (dB): A unit of measure
for the physical intensity of
sound. Larger decibel number
translates into a more intense
sound.

The frequency range of hearing varies in different species

We hear between 20
and 20,000 Hz.
Whales, dolphins
and dogs can hear
a wider range
of sounds than we
canat both the low
and high frequency
ends.
Bats, seals
and rats can hear
higher frequencies but
not lower.
Fish can hear lower
but not higher.
Birds and frogs have
a limited range of
hearing.

Sounds that we hear in our daily environments vary greatly in intensity

Pain threshold
High risk
threshold

What Is Sound?
The simplest kind of sounds: sine (sinusoidal) waves, or pure tone.

Sine waves are not common in everyday sounds because not many vibrations in
the world are so pure (exception is the tuning fork).
Most sounds in the world are complex sounds.
For example: Human voices, bird songs, music.
All sound waves can be described as some combination of sine waves.

Example: waveform of sound coming from a clarinet

Harmonic spectrumthe complicated
waveforms produced by a resonating
source, such as a string of a violin or the
reed of a woodwind. Corresponds to our
perception of timbre or uniqueness.
Fundamental frequency: The lowestfrequency component of a complex
periodic sound. The rate at which the
complex waveform pattern repeats.
Overtones: Set of higher frequency sound
waves that vibrate at whole-number
(integer) multiples of the fundamental
frequency.

Fourier Analysis-mathematical method of breaking down frequency

components of a complex sound.
1

A: Waveform on left can be described in a spectrum (on right) as an amplitude (height of bar)
and a frequency (location of bar (about 1000 Hz)).
B: Smaller amplitude and higher frequency sound described by a shorter bar at a higher
frequency.
A and B: Complex sound can be described by a combination of two sine waves, A and B.
Combining these sine waves will reproduce the original sound.
As well see later, the inner ear conducts a form of Fourier analysis of sound waves.

Figure 9.8 Harmonic sounds with the same fundamental frequency can sound different

Middle C as a pure tone only has

one frequency.
None of these is the same as the
pure tone, but they all are
harmonics of itwith different
overtones.

Timbre is the complex quality of

sound that allows one to hear the
difference between a middle C
note played on a tenor
saxophone, a trombone, or a
piano.

External and Internal Structures of the Human Ear

Air vibrations, a mechanical force, are transduced into neural activity.
The external earthe pinna and the ear canal (or auditory meatus)
collects sound waves.
The spiral-shaped cochlea of the inner ear
converts vibrations into neural activity.

The middle ear concentrates

sound energies.
Three ossicles (bones), the
malleus, incus, and stapes,
connect the tympanic
membrane (eardrum) to the
oval window.
http://www.tutis.ca/Senses/L9Auditory/L9Auditory.swf

The organ of Corti is the part of the

cochlea that converts sound to neural activity.

Structure of the Ear

Pinna
Funnel-like external structure designed to catch sound waves in the surrounding
environment and deflect them into the ear canal.

Pinna size and shape vary

greatly among mammals.

Pinnae and ear canal make up the outer ear.

Tympanic membrane (eardrum) and three tiny bones, the
ossicles, make up the middle ear.
Cochlea, otolith organs and semicircular canals
(vestibular system) make up the inner ear.

The middle ear relays sound information from the outer ear to the inner ear.
Ossicles: The smallest bones in the body.
Malleus (also called the hammer): Receives vibrations from the
tympanic membrane.
Incus (anvil): The middle ossicle connects malleus to the stapes.
Stapes (stirrup): Connected to the incus on one end and the oval
window of the cochlea on the other.
Amplification of vibrational energy provided by the
ossicles is essential to the ability to hear faint sounds.
Ossicles have hinged joints that work like levers
to amplify sounds energy.
Stapes has a smaller surface than the eardrum,
so sound energy is concentrated (easy to
squash a grape with a sheet of plywood).
Inner ear consists of fluid-filled chambers.
Takes more energy to move liquid than air.

http://www.tutis.ca/Senses/L9Auditory/L9Auditory.swf

The middle ear acts to protect against loud sounds.

Tensor tympani and stapedius:

Two muscles in the middle ear that
decrease ossicle vibrations by
stiffening when contracted.
Muffles loud sounds and protects
the inner ear.
However, acoustic reflex follows
onset of loud sounds by 200 msec.
so cannot protect against abrupt
sounds (e.g., gun shot).

Back-and-forth movement of the stapes moves the oval window which moves
the fluid inside the cochlea.

Back-and-forth movement of the stapes moves the oval window which moves
the fluid inside the cochlea.
Fluid flowing past the basilar membrane causes it to move up and down like
a board floating on a lake after a speedboat goes by.

A shearing force bends the hairs of

hair cells because the tips of the
hairs are embedded in the tectorial
membrane which moves less.

Detecting loudness: More intense sound waves trigger bigger waves of fluid in
the cochlea and more intense movements of the basilar membrane.
This results in greater shearing action of the hair cells.

Coding of amplitude and frequency in the cochlea

Place code: Different parts of the cochlea are tuned to
different frequencies.
The basilar membrane changes
Basilar membrane performs a type
Fourier
transform
fromof
narrow
and thick
at the oval
on the sound wave.
window to wider and thinner at
the cochlear apex.
Wider portions of the basilar
membrane vibrate at lower
frequencies and narrower portions
vibrate at higher frequencies.
Like snapping a rope that is wide and thin
at the end, narrow and thick near your
hand.
Hair cells at different locations along the
basilar membrane relay frequency
information to the brain (another example
of a labeled line).

Another look at the frequency selectivity of the basilar membrane

The sound wave moves the eardrum and

attached ossicular chain. The stapes footplate,
in the oval window, transfers the vibrations to
the inner ear structures. Depending on the
frequency, the vibration will have a maximum
effect (resonance) at different points along the
basilar membrane
Here, a high frequency sound affects a basal portion of the cochlea

A low frequency sound affects a more apical part of the cochlea

Video 6.1 Inside the Ear

[Click image to open in browser.]

Mechanism of sound transduction

The tip of each stereocilium is connected to the side of its neighbor by a tiny
filament called a tip link.
The bending of stereocilia
opens an ion channel, allowing
an influx of potassium (K+) ions
into the hair cell.

The K+ ions cause depolarization which allows Ca2+ to enter the base of the hair
cell, causing release of neurotransmitter into the synapse between the hair cell
and an afferent auditory neuron.
Thus, when a hair cell is displaced by movement of the basilar membrane it
releases more transmitter.

Animation 6.3 Sound Transduction

Two kinds of hair cells: Inner and outer hair cells

Inner hair cells: Convey almost all information about
sound waves to the brain (using afferent fibers).
Outer hair cells: Convey information from the brain (using
efferent fibers).
They are involved in a feedback system that modulates
the inner hair cell response by pushing and pulling on the
tectorial membrane.
Voltage signal from brain makes outer hair cells lengthen
and shorten. When depolarized, they shorten and pull
the basilar and tectorial membranes closeramplifying
the inner hair cells depolarization. Allows us to better
hear faint sounds.
Rock around the Clock Video:
http://www.youtube.com/watch?v=bB3x7kp9El8

There are four kinds of neural connections between the brain and hair cells.
IHC afferents convey action
potentials that provide sound
perception to the brain.
IHC efferents lead from the
brain to the IHCscontrol
responsiveness.

OHC afferents convey

information to the brain about
the state of the basilar
membrane.

OHC efferents lead from the brain

to OHCsthe brain can command
changes in their length, which
affects the basilar membrane.

Auditory brain structures:

Cochlear nucleus: The first
brain stem nucleus at which
afferent auditory nerve
fibers synapse.
Superior olive: An early
brain stem region in the
auditory pathway where
inputs from both ears
converge.
Inferior colliculus: A
midbrain nucleus in the
auditory pathway.
Medial geniculate nucleus:
The part of the thalamus
that relays auditory signals
to the auditory cortex.

Animation 6.4 Mapping Auditory Frequencies

Primary auditory cortex has tonotopic organization that matches the frequency
profile of the basilar membrane.

Cues for Binaural Hearing: two kinds of binaural cues locate a sound source.
1. Intensity differencesdifferences in loudness
at the two ears.

2. Latency differencesdifferences
between the two ears in the time of arrival
of sounds.
Onset disparitydifference in hearing at
the beginning of a sound.
Ongoing phase disparitycontinuous
difference between ears in arrival of parts
of a sound wave.

Lateral Superior Olive (LSO) and Medial Nucleus of the Trapezoid Body (MNTB)
encode sound localization through interaural intensity differencesworks best for high
frequency sounds.

High frequency sounds are

short wave and dont bend
around head very well.

Medial Superior Olive (MSO) encodes the location of a sound by interaural

time differences

Medial superior olive (MSO) has cells that are coincidence detectors. They receive coincident innervation
from the right and left cochlear nuclei.
Cells within the MSO are located at different distances from the respective cochlear nuclei.
Length of the axonal connections determine which MSO cell receives coincident activation by action
potential.

Types of Hearing Loss

Conduction deafness
disorders of the outer or
middle ear that prevent
sounds from reaching the
cochlea

Sensorineural deafness hair cells

fail to respond to movement of the
basilar membrane.
Ototoxic (ear-damaging) effects may
be due to drugs, noise pollution, or
loud sounds.
Damage to hair cells can result in
tinnitus, a persistent ringing in the
ears.

http://www.tutis.ca/Senses/L9Auditory/L9Auditory.swf

Central deafnesshearing loss

caused by damage to auditory
brain areas, such as by stroke,
tumors, or traumatic brain injury
Word deafnessunable to
recognize spoken words.
Cortical deafnessdifficulty in
recognizing all complex sounds or
speech.

Sensorineural Hearing Loss

Start out with about 3,500 inner hair cells
and 12,000 outer hair cells.
Damage to the hair cells in the cochlea,
frequently the result of loud sounds or
ototoxic drugs (some antibiotics, NSAIDs
and heavy metals).

Residents of quiet Easter Island suffered less hearing loss at all frequencies than
their peers who lived off the island.
Hearing loss (especially high frequencies) is correlated with age.

Cochlear implants can be useful for those with sensorineural deafness.

Different frequencies of sound detected by the electronic circuitry stimulate

the nerves innervating the different parts of the basilar membrane. These
devices completely bypass the eardrum and hair cells.

Watch from start through direction:

http://www.tutis.ca/Senses/L9Auditory/L9Auditory.swf
Learn about Fourier Analysis:
http://sites.sinauer.com/wolfe3e/chap1/fourierF.htm
Learn about hearing and the auditory system (Parts 9.1 and 9.2):
http://sites.sinauer.com/wolfe3e/chap9/startF.htm
Learn about missing fundamental frequencies:
http://sites.sinauer.com/wolfe3e/chap10/missingfundF.htm