Audition

The ear
Physiological processes (from pressure
changes to electrical discharges)
Dr. Stephan Moratti
 The auditive system’s task

• Deliver the sound stimulus to the receptors.

• Transduction from air pressure changes to neural electrical
discharges
• Processing of these signals to determine pitch, loudness,
timbre and localization of the sound.
 Transmission of vibrations

• Transmission of vibrations from one structure to another.

• From the tympanic membrane to the receptors.
• Outer, middle, and inner ear.
 The outer ear

• Pinnae
• Auditory canal
– tube
– 3 cm long in adults
– Protective function
– 3 cm of distance + earwax protect
• Also, constant temperature
– Resonance frequency of the canal:
• amplifies SPLs of frequencies between 1,000 y 5,000 Hz
• See audibility curve!

• Tympanic membrane
 The middle ear

• Sound waves set the tympanic membrane into vibration.

• This vibration is transmitted to the structures of the middle
ear.
– Little cavity(2 cm3)
– Ossicles (smallest bones of the human body)
• Malleus (hammer)
• Incus (anvil)
• Stapes (stirrup)
– Ovel window
 The middle ear

• Why so much transmission?

• Example: If your are diving in a pool, and somebody outside is

trying to talk to you.
 The middle ear

• Why is there so much mechanic transmission?

• Concentrating the vibration from the tympanic

membrane to the small stapes area increases
the pressure by a factor of 20!

• Ossicles lesioned: The sound level has to be

increased by a factor of 10 to 50 in order to
achieve the same loudness perception.

• Lever principle.

• Protect your middle ear!

 The middle ear

• Small muscles of the middle ear (smallest skeletal muscles of the human
body)
• Connected with the ossicles.
• Contract during intense sound stimulation (protection)
• Reduce the effects of low frequency sounds with great amplitude (like a
filter)
• Suppression of interference of sounds from
chewing or swallowing.
The inner ear

• The liquid inside the cochlea vibrates due to the

vibration of the stapes against the oval window
 The inner ear

The cochlear partition contains the structures that transduce the mechanical vibrations into
electrical signals.
 The inner ear
The hair cells and the tectorial membrane
The oval window transmits the vibration to the
cochlear liquid. The basilar membrane starts to
vibrate.

The basilar membrane moves up and down. The

tectorial membrane moves forward and backwards

The cilia of the inner hair cells bend due to their

movement against the surrounding liquid.

The cilia of the outer hair cells bend due to their

movement against the tectorial membrane.
The inner ear
Transduction

The movement of the cilia provokes the opening and closing of ion
channels.

When they open

Ions enter the cell

Liberation of neurotransmitters

Electrical discharges of the auditory nerve fiber

 The inner ear
Frequency coding by exact phase locking of neural discharges

Sound pressure increases > Cilia bends to the right

Sound pressure decreases-> Cilia bends to the left

Cilia bends to the right ->

ion channels open

Cilia bends to left ->

ion channels close

Phase locking between sound wave and

auditory nerve fiber firing
 The inner ear
Temporal coding of sound pressure frequency

One single neuron cannot code high frequencies

because of the refractory time of neurons.

However, many neurons can code higher

frequencies by temporal coding.

Þ Phase locking up to 4000 Hz – 5000 Hz.

Þ However, temporal coding is not all!

 Vibration of the basilar membrane
place theory of frequency codification

Georg von Békésy discovered how the basilar membrane vibrates and received the Novel Price
for that discovery in 1961 (later we will see why).
 Vibration of the basilar membrane
place theory of frequency codification

Békésy bored holes in cochleas of dead animals and human cadavers and observed how the
membrane vibrates to different frequencies. He observed traveling waves as a function of sound
frequency.
Vibration of the basilar membrane
place theory of frequency codification

The place of maximum vibration along the

membrane depends on the frequency of the
sound stimulus.

Place theory of audition!

Base: high frequencies

Apex: low frequencies
Vibration of the basilar membrane
place theory of frequency codification
 Vibration of the basilar membrane
place theory of frequency codification

Tonotopic map determined by placing electrodes

along the basilar membrane of a guinea pig.

Þ Localization of the maximal electrical response.

 Vibration of the basilar membrane
place theory of frequency codification

More precise method: Frequency tuning curves of auditory nerve fibers in cat.

Characteristic frequency of a nerve fiber

Vibration of the basilar membrane
place theory of frequency codification
At the behavioral level: masking (Egan & Hake, 1950).
Vibration of the basilar membrane
place theory of frequency codification
Vibration of the basilar membrane
place theory of frequency codification

The cochlear implant

 Vibration of the basilar membrane
place theory of frequency codification

For complex sounds the cochlea separates the fundamental and harmonic frequencies by its
place code (it is doing a Fourier transform!)
Vibration of the basilar membrane
place theory of frequency codification

Watch this nice summary video!