Audition

Josefina C. Ochoa, PhD, RPm, LPT

Evolution has been described as “thrifty.”
After it has solved one problem, it
modifies that solution for other problems
instead of starting from scratch. For
example, imagine a gene for visual
receptors in an early vertebrate. Make a
duplicate of that gene, modify it slightly,
and presto: the new gene makes receptors
that respond to different wavelengths of
light, and color vision becomes possible.
 Sound and the Ear

Under optimum conditions, human hearing is sensitive to sounds that vibrate the eardrum by less
than one-tenth the diameter of an atom, and we can detect a difference between two sounds as little
as 1/30 the interval between two piano notes (Hudspeth, 2014).

Ordinarily, however, we attend to hearing in order to extract useful information. If you hear
footsteps in your home or a snapped twig in the forest, you know you are not alone. If you hear
breathing, you know some person or animal is close. Then you hear the sound of a familiar friendly
voice, and you know that all is well.
 Physics and Psychology of Sound

Sound waves are periodic compressions of air, water,

or other media. When a tree falls, the tree and the
ground vibrate, setting up sound waves in the air that
strike the ears.

Sound waves vary in amplitude and frequency. The

amplitude of a sound wave is its intensity. In
general, sounds of greater amplitude seem louder,
but exceptions occur. For example, a rapidly talking
person seems louder than slow music of the same
physical amplitude.
The frequency of a sound is the number of
compressions per second, measured in hertz
(Hz, cycles per second).

Pitch is the related aspect of perception.

Sounds higher in frequency are higher in
pitch. The figure on the right illustrates the
CONCEPT 2
amplitude and frequency of sounds. The Venus has a beautiful name and
is the second planet from the
height of each wave corresponds to Sun
amplitude, and the number of waves per
second corresponds to frequency.
In addition to amplitude and pitch, the third
aspect of sound is timbre (TAM-ber), meaning
tone quality or tone complexity. Two musical
instruments playing the same note at the same
loudness sound different, as do two people
singing the same note at the same loudness.

For example, any instrument playing a note at 256

Hz will simultaneously produce sound at 128 Hz,
512 Hz, and so forth, known as harmonics of the
principal note. The amount of each harmonic
differs among instruments.
Interestingly:
People communicate emotion by alterations
in pitch, loudness, and timbre.

For example, the way you say “that was

interesting” could indicate approval (it really
was interesting), sarcasm (it really was
boring), or suspicion (you think someone
was hinting something).

Conveying emotional information by tone of

voice is known as prosody.
 Structures of the Ear

Rube Goldberg (1883–1970) drew cartoons of complicated, far-fetched inventions. For example, a person’s tread on the
front doorstep might pull a string that raised a cat’s tail, awakening the cat, which then chases a bird that had been resting
on a balance, which swings up to strike a doorbell.
The functioning of the ear is complex enough to resemble a Rube Goldberg device, but unlike Goldberg’s inventions, the
ear actually works.

Anatomists distinguish the outer ear, the middle ear, and the inner ear. The outer ear includes the pinna, the familiar
structure of flesh and cartilage attached to each side of the head. By altering the reflections of sound waves, the pinna helps
us locate the source of a sound. We have to learn to use that information because each person’s pinna is shaped differently
from anyone else’s. Rabbits’ large movable pinnas enable them to localize sound sources even more precisely.
“This is a quote, Words full of
wisdom that someone important said
and can make the reader get
inspired.”

—SOMEONE FAMOUS—
 Pitch Perception

Your ability to understand speech or enjoy music depends on your ability to differentiate among sounds of different
frequencies. How do you do it?

According to the place theory, the basilar membrane resembles the strings of a piano, with each area along the
membrane tuned to a specific frequency. If you sound a note with a tuning fork near a piano, you vibrate the piano
string tuned to that note.

According to this theory, each frequency activates the hair cells at only one place along the basilar membrane, and the
nervous system distinguishes among frequencies based on which neurons respond. The downfall of this theory is that
the various parts of the basilar membrane are bound together too tightly for any part to resonate like a piano string.
 Place Theory

60% 30% 25%

mercury saturn mars

Mercury is the closest It is composed of Mars is actually a very
planet to the Sun hydrogen and helium cold place
According to the frequency theory, the entire basilar membrane vibrates in synchrony with a sound,
causing auditory nerve axons to produce action potentials at the same frequency. For example, a
sound at 50 Hz would cause 50 action potentials per second in the auditory nerve.

The downfall of this theory in its simplest form is that the refractory period of a neuron, though
variable among neurons, is typically about 1/1,000 second, so the maximum firing rate of a neuron is
about 1000 Hz, far short of the highest frequencies we hear.
The current theory is a modification of both theories. For low-frequency sounds (up to about 100 Hz
—more than an octave below middle C in music, which is 264 Hz), the basilar membrane vibrates in
synchrony with the sound waves, in accordance with the frequency theory, and the auditory nerve
axons generate one action potential per wave.

Soft sounds activate fewer neurons, and stronger sounds activate more. Thus, at low frequencies, the
frequency of impulses identifies the pitch, and the number of firing cells identifies loudness.
 The Auditory Cortex

As information from the auditory system passes

through subcortical areas, axons cross over in the
midbrain to enable each hemisphere of the forebrain
to get most of its input from the opposite ear
(Glendenning, Baker, Hutson, & Masterton, 1992).
The information ultimately reaches the primary
auditory cortex.
The organization of the auditory cortex parallels that of the visual cortex (Poremba et al., 2003). For
example, just as the visual system has separate pathways for identifying objects and acting upon
them, the auditory system has a pathway in the anterior temporal cortex specialized for identifying
sounds, and a pathway in the posterior temporal cortex and the parietal cortex specialized for locating
sounds (Lomber & Malhotra, 2008).

Just as patients with damage in a brain area become motion blind, patients with damage in parts of the
superior temporal cortex become motion deaf. They hear sounds, but they do not detect that a source
of a sound is moving (Ducommun et al., 2004).
 Sound Localization

You are walking alone when suddenly you hear a loud noise. You want to know what
produced it (friend or foe), but equally, you want to know where it came from. Sound
localization is less accurate than visual localization, but nevertheless impressive.
Owls localize sounds well enough to capture mice in the dark.

Determining the direction and distance of a sound requires comparing the responses
of the two ears. One method is the difference in time of arrival at the two ears. A
sound coming directly from one side reaches your closer ear about 600 microseconds
(μs) before the other. A smaller difference in arrival times, indicates a sound source
nearer to your midline. Time of arrival is useful for localizing sounds with a sudden
onset. Most birds’ alarm calls increase gradually in loudness, making them difficult
for a predator to localize.
For people with amusia, the auditory cortex appears to be approximately
normal, but it has fewer than average connections to the frontal cortex (Hyde et
al., 2007; Loui et al., 2009; Norman-Haignere et al., 2016). That is, the deficit
is evidently not in hearing itself. Rather, these people have poor memory for
pitch (Tillman, Leveque, Fornoni, Abouy, & Caclin, 2016), and perhaps poor
attention to pitch.

Transcranial alternating current applied to the scalp is a non-invasive way to

stimulate the underlying area of the brain. When this procedure was applied to
part of the right prefrontal cortex of people with amusia, their ability to
remember pitch improved to almost normal levels (Schaal, Pfeifer, Krause, &
Pollok, 2015). The implication is that amusia results from either an impairment
of the prefrontal cortex, or input to it from the auditory cortex.
 Deafness

Although few people are totally unable to hear, many people have enough impairment to prevent speech comprehension.
The two categories of hearing loss are conductive deafness and nerve deafness.

Diseases, infections, or tumorous bone growth can prevent the middle ear from transmitting sound waves properly to the
cochlea. The result, conductive deafness or middle-ear deafness, is sometimes temporary.

If it persists, it can be corrected by surgery or by hearing aids that amplify sounds. Because people with conductive
deafness have a normal cochlea and auditory nerve, they readily hear their own voices, conducted through the bones of the
skull directly to the cochlea, bypassing the middle ear. Because they hear themselves clearly, they may accuse others of
mumbling or talking too softly.
Nerve deafness, or inner-ear deafness, results from damage to the cochlea, the hair cells, or the
auditory nerve. If it is confined to one part of the cochlea, it impairs hearing of certain frequencies
and not others.

Nerve deafness can be inherited, it can result from disease, or it can result from exposure to loud
noises. For example, many soldiers, construction workers, and fans of loud rock music expose
themselves to noise levels that damage the synapses and neurons of the auditory system. Gradually
they begin to notice ringing in the ears or impaired hearing (Kujawa & Liberman, 2009).
Tinnitus (tin-EYE-tus) is frequent or constant ringing in the ears.
In some cases, tinnitus may be due to a phenomenon similar to
phantom limb. Damage to part of the cochlea is like an amputation:
If the brain no longer gets its normal input, axons representing
other parts of the body may invade part of the brain area that
usually responds to sounds.

In many cases, people who have lost their hearing in a particular

range report ringing in the ears in the same range, suggesting that
some other input is activating part of the auditory cortex. However,
many people have tinnitus without hearing loss or reorganization of
the cortex (Elgoyhen, Langguth, De Ridder, & Vanneste, 2015).
Evidently tinnitus can result from more than one cause.
 Hearing, Attention, and Old Age

Many older people have hearing problems despite wearing hearing aids. The hearing aids make the sounds
loud enough, but people still have trouble understanding speech, especially in a noisy room or if someone
speaks rapidly.

Part of the explanation is that the brain areas responsible for language comprehension have become less
active (Peelle, Troiani, Grossman, & Wingfield, 2011). This trend might be just a natural deterioration, or
it might be a reaction to prolonged degradation of auditory input. That is, if someone delays getting
hearing aids, the language cortex doesn’t get its normal input and it begins to become less responsive.
 APPLICATIONS

We spend much of our day listening to language, and we sometimes forget that the original, primary
function of hearing has to do with simpler but extremely important issues: What do I hear? Where is
it? Is it coming closer? Is it a potential mate, a potential enemy, potential food, or something
irrelevant? The organization of the auditory system is well suited to answering these questions.
Thank You for
Listening!
Josefina C. Ochoa, PhD, RPm, LPT