Chapter 7: Audition

September 25
th
, 2014
Introducing Sensory Systems: Bringing the
World into the Mind
https:/.../art_multi_en_cont/art_multi_en
Learning objective
Students will be able to accurately describe
the signal transduction process that occurs
when a sound wave enters the ear, and
describe how that information gets to the
brain.
From: The Christmas Carol
Charles Dickens
You dont believe in me,
observed the ghost.

Why do you doubt your
senses?
From: The Christmas Carol
Because, said Scrooge,
a little thing affects them.
A slight disorder of the
stomach makes them
cheats. You may be an
undigested bit of beef, a
blot of mustard, a crumb
of cheese, a fragment of
underdone potato

Theres more of gravy than of grave about
you, whatever you are!
Lets talk about this.
Do we trust our senses?

What is it that we can sense?

Is this representative of the world around us?

Knowing what we do about the brain, what must
happen for us to be able to sense anything?
We perceive the world as a result of:

Sensory information, transmitted by neurons to the
brain, from specialized receptors that transduce
physical stimuli into neural potentials

Perception occurs only after active processing of that
sensory information by multiple neuronal networks
within the brain.

The study of how we perceive the world is a
subdiscipline of psychology, and can be explored in
Dr. Schirillos class (Psych 329).
The world, as we know it,
is an illusion (well, kind of)
For most sensory systems:
Stimuli in the environment impinge in some way on a
receptor cell.

The receptor alters its membrane potential in
response.
Depolarizations increase NT output
Hyperpolarizations decrease NT output

These changes in NT output directly affect the firing
rates of primary sensory neurons, which code relay
the signal (down axons in the form of action
potentials) from the receptor into the brain


Sensory receptors:
Specialized cells that transduce physical or
electromagnetic stimuli into graded potentials (in
most cases).

In this case, receptor refers to the reception of the
sensory stimulus

These graded potentials most often affect the
neurotransmitter output of the receptor cell,
affecting the activity of post-synaptic sensory
neurons.

We perceive the world as a result of:
1) What our sensory receptors can receive and
transduce, and how it is encoded.

How does primate vision differ from that of
other mammals (such as the cat?


http://www.sciencebuddies.org/science-fair-
projects/project_ideas/HumBio_p011.shtml#procedure
We perceive the world as a result of:
2) How our brains take the coded information
and process it to make sense of our world.

The brain can be tricked, and we call these
tricks perceptual illusions.


The world, as we know it,
is an illusion (well, kind of)
http://www.tonyboon.co.uk/imgs/illusions/wwheels.htm
Audio Illusions:
Infinitely ascending tones:
http://www.youtube.com/watch?v=ugriWSmRxc
M

Try this virtual haircut at home, with
headphones!
http://www.youtube.com/watch?v=IUDTlvagjJA&f
eature=related


For a sensory stimulus to be received by the
brain:
The signal has to impinge on (or affect) a receptor for
that stimulus
This is called reception

The receptor has to detect the stimulus and alter its
NT output as a result
This is called transduction

Primary sensory neurons (which project into the
brain) have to detect the change of NT input and
alter their firing as a result (i.e. increase or decrease
action potential production).
This is called coding
Audition
Physical and Perceptual Dimensions of Sound
Amplitude (Loudness):
= Height of sound wave
(Intensity of sound wave)
Loud
Bolt of lightning
Soft
Whispering
Frequency (Pitch):
= Speed of sound wave
Low
Bass
High
Alarm
Children hear higher
frequencies than adults

Can be in 80 dB or less
indefinitely

If your ears are ringing
after a concert, you did
damage!!!
Audiograms for different species
The basics
A sound wave enters the ear
It is amplified by the middle ear
It is transduced to electrical signals in the inner ear
It is sent to various brain areas


The structure of the
human ear
Outer ear: pinna,
external auditory canal

Middle ear: tympanic
membrane (ear drum),
ossicles (3 tiny bones):
malleus, incus, stapes
Inner ear: oval window,
cochlea (audition),
semicircular canals (vestibular
system)

(malleus)
(incus)
(stapes)
Figure 7.1 p. 206
The function of these structures
Pinna: sound localization
Tympanic membrane: vibrates
same as sound frequency
Ossicles: amplification
Because need greater force
to move wave thru fluid-
filled cochlea than thru air
Malleus, incus, stapes
Oval window: vibrates with
same freq. but more force
Cochlea: transduction

Figure 7.1 p. 206
Together the
tympanic membrane
and the ossicles
convert air vibrations
to fluid vibrations,
which are detected
in the inner ear
Correct terminology
for the ossicles:

Hammer = Malleus
Anvil = Incus
Stirrup = Stapes
Cochlea
3 long canals
Hair cells (auditory receptors) in organ of
corti (btw tectorial and basilar membranes)
Hair cells are mechanoreceptors

Oval window vibrates vibration of fluid in cochlea
hair cells are displaced hair cells are activated
and thereby trigger action potentials in axons of
auditory nerve
And Again
Hair cells
Auditory (8
th
)
cranial nerve
Hair cells
Stereocilia
Notice
shortest to
tallest
Outer H.C.s Inner H.C.s
3 rows
~20,000
Stereocilia elongate
amplify the movement of the
basilar membrane
sharpen frequency response
of IHCs

Actual sensory receptors
1 row
~3500
Most auditory info
90-95% of auditory
nerve fibers come from
IHCs.
The movement of the stapes
upon the cochlear fluid
causes the basilar
membrane to move up or
down

The longest stereocilia of
OHC is attached to the
bottom of the tectorial
membrane, so when the
basilar membrane moves,
so does the Organ of Corti
and the tectorial plate.

This causes a shearing
motion btwn the tectorial
and basilar membrane, thus
displacing the stereocilia of
IHCs.

Endolymph (fluid in
cochlear duct) has very
high K
+
concentration
Auditory nerve
Note: different than neurons
High external K
+
no action potential
K+ channels open and K+
enters the hair cell

The cell becomes
depolarized

The change in voltage
opens VC Ca+ channels,
which allow Ca+ into the
cell

Ca+ allows vesicles
(containing NT) to
release, spilling NT into
the cleft and onto the
auditory nerve


How do we identify the amplitude and
frequency of a sound??
Base: wide basilar membrane: high frequency sounds
Apex: narrow: low frequency sounds
How to we perceive frequency?
Place theory
The place on the basilar
membrane that moves tells our
brain what the frequency is (Best
explains high frequency sounds)

Frequency matching
Basilar membrane vibrates in
synchrony with sound

Action potentials produced at
same frequency as sound wave
(Best explains low frequency
sounds)
Pitch theory uses a
combo of place
theory and
frequency matching.

Perception of high
pitches is not well
understood.
Sound and the Ear (contd.)
Volley principle: auditory nerve as a whole
produces volleys of impulses (for sounds up
to about 4,000 per second)
No individual axon solely approaches that
frequency
Requires auditory cells to precisely time their
responses
Hearing of higher frequencies not well
understood

Sound and the Ear (contd.)
People vary in their sensitivity to pitch
Amusia: the impaired detection of frequency
changes (tone deafness)
Associated with thicker than average
auditory cortex in the right hemisphere but
fewer connections from auditory cortex to
frontal cortex
Sound and the Ear (contd.)
Absolute pitch (perfect pitch) is the
ability to hear a note and identify it
Genetic predisposition may contribute to it
Main determinant is early and extensive
musical training
More common among people who speak tonal
languages
Flow of Auditory information
Auditory nerve
(CN VIII)
Cochlear nucleus
(sound localization)
Superior olivary nucleus
(sound localization)
Inf. colliculus (fine-tuning,
understanding speech,
integration with visual info)
Medial Geniculate Nucleus
of thalamuc (sensory
integration, filter)
Auditory cortex
in temporal lobe
The Auditory Cortex
The primary auditory cortex (area A1) is the
destination for most information from the
auditory system
Located in the superior temporal cortex
Each hemisphere receives most of its
information from the opposite ear
The Auditory Cortex (contd.)
Organization of the auditory cortex
parallels that of the visual cortex
Superior temporal cortex contains area MT
Allows detection of the motion of sound
Area A1 is important for auditory imagery
Requires experience to develop properly
Axons leading from the auditory cortex develop less
in people deaf since birth
The Auditory Cortex (contd.)
The cortex is necessary for the advanced
processing of hearing
Damage to A1 does not necessarily cause deafness
unless damage extends to the subcortical areas
The auditory cortex provides a tonotopic map
in which cells in the primary auditory cortex
are more responsive to preferred tones
Some cells respond better to complex sounds than
pure tones
Tonotopic organization of auditory
cortex
The Auditory Cortex (contd.)
Areas around the primary auditory cortex exist
in which cells respond more to changes in
sound than to prolonged sounds
Cells outside A1 respond to auditory objects
(animal cries, machinery noise, music, etc.)
Because initial response is slow, most likely
responsible for interpreting the meaning of sounds
Impaired hearing
Conductive hearing loss
= middle-ear deafness
Bones of middle ear dont transmit info to cochlea
Disease, infection, tumor
Correction: Surgery, hearing aids




Sensorineural deafness
= inner-ear deafness or nerve
deafness
Damage to cochlea, hair cells
or auditory nerve
Can be confined to certain
part of cochlea
Inherited, prenatal problems,
childhood disorders, noise-
induced
Correction: Hearing aids,
cochlear implant
Hearing Loss (contd.)
Tinnitus is a frequent or constant ringing in
the ears
Experienced by many people with nerve
deafness
Sometimes occurs after damage to the
cochlea
Axons representing other part of the body
innervate parts of the brain previously
responsive to sound
Similar to the mechanisms of phantom limb
Sound Localization
Sound localization depends upon comparing
the responses of the two ears
Three cues:
Sound shadow
Time of arrival
Phase difference
Humans localize low frequency sound by
phase difference and high frequency sound by
loudness differences
Sound Localization (contd.)
Three mechanisms:
High-frequency sounds (2000 to 3000Hz)
create a sound shadow
Difference in time of arrival at the two ears
most useful for localizing sounds with sudden
onset
Phase difference between the ears provides
cues to sound localization with frequencies up
to 1500 Hz
Readjusting sound localization is slow process