Hearing and Language

Chapter 9

HEARING

LANGUAGE
 Hearing

A receptor is a cell, often a specialized neuron, that is

suited by its structure and function to respond to a
particular form of energy, such as sound.
 A receptor’s function is to convert that energy into a neural
response.
An adequate stimulus is the energy form for which the
receptor is specialized.
The pattern of the information contained in the
sensory stimulus makes information meaningful.
Many people consider audition, along with vision, to
be the most important senses.
 Hearing

Sensation is the acquisition of sensory information.

Perception is the interpretation of this information.
The adequate stimulus for audition is vibration in a
conducting medium, such as air.
Frequency refers to the number of cycles or waves of
compressions and decompressions
 Frequency provides the perception of pitch.
 A pure tone, produced by striking a tuning fork for
example, has only one frequency.
 Complex sounds, such as that produced by a clarinet, are
composed of multiple frequencies.
 Hearing

The intensity of a sound is perceived as loudness.

 Intensity refers to the amplitude or size of the wave. It
represents the physical energy of a sound.
 Loudness is influenced by the frequency of the sound.
 For example, we are most sensitive to sounds between
2000 and 4000 Hz, the range in which most
conversation occurs.
 Sounds outside this range would seem less loud.

 Also, the intensity of the sound influences the perception

of pitch.

◊
 Hearing

The outer ear, or pinna:

 captures the sound and amplifies it by funneling it into
the smaller auditory canal.
 selects for sounds in front and to the side of us, partially
blockinh those behind.

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 Hearing

The Middle Ear

 The tympanic membrane, or eardrum, collects the
vibrations and transmits them to the ossicles.
 The lever action of the ossicles transfers the vibration to
the cochlea.
 Amplification as the vibration passes from the eardrum to
the smaller base of the stirrup compensates for the loss of
energy in passing from air into the liquid inside the
cochlea.
 We can detect sound when the eardrum vibrates as little
as the diameter of the hydrogen atom.
◊
The Outer, Inner, and Middle Ear
Figure 9.4
 Hearing

The inner ear includes

the cochlea.
 The cochlea is divided
into the fluid-filled
vestibular canal,
tympanic canal,
cochlear canal.
 The stirrup sends
vibrations throughout the
cochlea and to the sound-
analyzing structure.

◊ Figure 9.5
 Hearing

The organ of Corti rests on the basilar membrane.

 It consists of four rows of hair cells and the tectorial
membrane above the hair cells.
 Vibration bends the hair cells, opening potassium and
calcium channels.
 This depolarizes the cells and sets off signals in the neurons.
 The less numerous inner hair cells provide the majority of
information about the auditory stimulus.
 Lengthening and shortening of the outer hair cells against
the tectorial membrane adjusts the organ of Corti’s rigidity.
 This amplifies weak signals and provides adjustable
frequency selectivity.
 Hearing

The Auditory Pathway

 Auditory neurons form part of the auditory (8th) cranial
nerves.
 The neurons travel to the inferior colliculi, to the medial
geniculate nucleus of the thalamus, and to the auditory
cortex.
 Neurons from each ear go to both temporal lobes, but
there are more connections to the opposite side than the
same side.
 The auditory cortex is topographically organized:
Neurons from adjacent receptors project to adjacent cells
in the cortex, forming a map of the unrolled cochlea.
◊
Fig. 7-5, p. 194
 Hearing

 Secondary auditory areas are involved in analyzing

complex sounds and understanding their meaning.
 The dorsal stream travels from the auditory cortex to the
parietal lobes (for spatial location of sounds) and to the
frontal lobes (for directing eye movements and planning
movements). It is the auditory “where” system.
 The ventral stream passes from the temporal to the
frontal lobes. It is involved in identifying sounds and is
the auditory “what” system.

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 Hearing

Frequency Analysis
 Frequency theory assumes that the auditory mechanism
transmits the actual sound frequencies to the auditory
cortex for analysis there.
 Telephone theory: Individual neurons in the auditory
nerve fire at the same frequency as the rate of vibration
of the sound source.
 Volley theory: Groups of neurons follow the frequency
of a sound at higher frequencies.
 Even volleying fails to follow sounds beyond about
5200 Hz, so the frequency theory is inadequate.
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 Hearing

 Place theory: The frequency of a sound is identified

according to the location of maximal vibration on the
basilar membrane and, therefore, which neurons are
firing most.
 Higher frequencies cause the base end to vibrate most
and low frequencies cause the apex end to vibrate most.
 The auditory cortex is topographically organized, in the
form of a tonotopic map.
Thus, each successive area responds to successively
higher frequencies.

◊
Tonotopic Map
Figure 9.12
 Hearing

 However, place theory alone is inadequate.

 The basilar membrane vibrates about equally at the
lowest frequencies.
 Frequency-specific neurons have not been found below
200 Hz.
 Frequency-place theory:
 Neurons follow the sound’s frequency below about

200 Hz.
 Higher frequencies are detected by place analysis.

◊
 Hearing

Analyzing Complex Sounds

 The basilar membrane does a Fourier
analysis of a complex sound, separating
it into its sine wave components.

Figure 9.14: Fourier analysis

of a clarinet note
 Hearing

 We also must sort out meaningful sounds embedded in a

confusing background of sounds. This is known as the
cocktail party effect.
 Selective attention enhances some sounds and
suppresses others.
 These separated sounds become auditory objects
 We must then identify a sound.

 This occurs in the ventral “what” area.

 Voices are identified in the superior temporal area.

 Environmental sounds are identified primarily in

posterior temporal areas and to some extent in
frontal areas.
 Hearing

Locating Sounds With Binaural Cues

 Binaural cues permit us to locate sounds quickly and
accurately.
 When a source is to one side or the other, the head blocks
some of the sound energy; thus, there is a difference in
intensity at the two ears.
 A sound directly to the left or the right of the listener
takes about 0.5 ms to travel the additional distance to the
second ear. Thus, there is also a difference in time of
arrival at the two ears.
◊
Differential Intensity & Time of Arrival Cues
(sound shadow)
 Hearing

 At low frequencies, a sound arriving from one side of the

body will be at a different phase of the wave at each ear.
 As a result, the rising or falling pressure will be
different at the two eardrums.

Figure 9.18: Phase differences at the two ears

Fig. 7-7, p. 196
Fig. 7-8, p. 197
 Hearing

Brain Circuits for Locating Sounds

 The “time of arrival” circuit (studied most extensively)
contains coincidence detectors.
 In this circuit, a longer pathway from one ear
compensates for the delay in sound reaching the other
ear.
 A coincidence detector fires most when it receives input
from both ears at the same time.
 As a result, each detector is specialized for sounds
arriving at a particular angle to the body.

◊
Difference in Time of Arrival Circuit
Figure 9.19
 Hearing

Next, this directional information must be integrated

with
 information from the visual environment
 and information about the position of the body in
space.
 This involves the parietal lobes, part of the dorsal “where”
stream.

◊
 Audition

 Two categories of hearing impairment include:

1. Conductive or middle ear deafness.
2. Nerve deafness.
 Audition

Conductive / middle ear deafness occurs if

bones of the middle ear fail to transmit sound
waves properly to the cochlea.
Caused by disease, infections, or tumerous bone
growth.
Can be corrected by surgery or hearing aids that
amplify the stimulus.
Normal cochlea and normal auditory nerve
allows people to hear their own voice clearly.
 Audition

 Nerve or inner-ear deafness results from

damage to the cochlea, the hair cells, or the
auditory nerve.
 Can vary in degree
 Can be confined to one part of the cochlea.
 people can hear only certain frequencies.
 Can be inherited or caused by prenatal
problems or early childhood disorders (rubella,
syphilis, inadequate oxygen to the brain during
birth, repeated exposure to loud noises, etc).
 Audition

 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.
 Audition

 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 difference.
 Audition

 Three mechanisms:
1. High-frequency sounds (2000 to 3000Hz) create a
“sound shadow”, making the sound louder for the
closer ear.
2. The difference in the time of arrival at the two ears is
most useful for localizing sounds with sudden onset.
3. Phase difference between the ears provides cues to
sound location for localizing sounds with frequencies
up to 1500 Hz.
 Language

Language includes the generation and

understanding of written, spoken, and gestural
communication.
 Broca’s area was discovered in 1861 when Paul Broca
studied a stroke patient with injury in the frontal area.
 Symptoms of Broca’s aphasia include:
 Nonfluent speech;
 Anomia, or trouble finding words;
 articulation problems;
 lack of grammatical, or function, words.
 Reading and writing are impaired as much as speech.
 Comprehension is impaired when the meaning depends
on grammatical words.
 Language

 Wernicke’s area is in the posterior portion of the left

temporal lobe.
 Wernicke’s Aphasia
 The individual has trouble understanding spoken and
written language.
 However, the term receptive aphasia is misleading,
because the patient has as much difficulty producing
language as understanding it.
 Speech, for example, is fluent but meaningless (and is
often referred to as “word salad”).

◊
Language-Related Areas of the Cortex
Figure 9.20
 Language

The Wernicke-Geschwind model is an effort to explain

how Broca’s area and Wernicke’s area interact to
produce language.
 Answering a Spoken Question:
AUDITORY CORTEX ➞ WERNICKE’S AREA ➞ BROCA’S AREA
Broca’s area then communicates with the facial area of the
motor cortex to produce speech.
 Reading Aloud: The visual information is first transformed
into an auditory form in the angular gyrus; then
ANGULAR GYRUS ➞ WERNICKE’S AREA ➞ BROCA’S AREA
 What would happen if the response is to be
written?
 This model is generally accurate but oversimplified; for
example, functions are not so localized.
Wernicke-Geschwind Model of Language
Figure 9.21
 Language

Reading, Writing, and Their Impairment

 Alexia, the inability to read (cannot recognize letters),
and agraphia, the inability to write, are presumably due
to disruption of pathways in the angular gyrus.
 These pathways connect the visual projection area with
the auditory and visual association areas.
 Dyslexia, an impairment of reading, can be acquired
through damage, but is more often developmental.

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 Language

Brain Differences in Dyslexia

 The left planum temporale, where Wernicke’s area is
located, left is typically larger than on the right; in
dyslexics it is larger on the right or equal in size.
 Neurons in the left planum temporale lack orderly
arrangement.
 The most reliably identified genes are involved in
neuron guidance and migration.

◊
Anomalies in the Dyslexic Brain
Figure 9.24

Left planum temporale in a normal brain (left)

and in the brain of a person with dyslexia (right)
 Language

 People with dyslexia have both auditory and visual-

perceptual difficulties.
 They have trouble detecting the frequency and
amplitude changes that distinguish letter sounds.
 Words are read backwards, mirror-image letters

(b and d) are confused, and words appear to move

around on the page.
 According to the magnocellular hypothesis, dyslexia
involves deficiencies in auditory and visual magnocellular
cells.

◊
 Language

 According to the phonological hypothesis, the major

problem is an impairment of phoneme processing.
 A phoneme is a small sound unit that distinguishes one
word from another (as in book versus cook).
 fMRI indicates the problem is in an auditory word
analysis area, not in the area that recognizes words by
their visual form.
 Dyslexia occurs much less frequently in countries
where the languages are phonologically simpler

◊
 Language

Recovery From Aphasia

 The right hemisphere can take over language functions
following left-hemisphere damage, as long as the injury
occurs early in life.
 If damage occurs later in life, language control is more
likely to shift into bordering areas in the left hemisphere.
 The ability of other areas to take over language functions
may be due to their normal participation in language.
For example:
 Language

 The right hemisphere contributes prosody to speech;

prosody is the use of intonation, emphasis and rhythm
to convey meaning.
 The right hemisphere also is important in
understanding information from language that is not
specifically communicated by the meaning of the
words:
 when meaning must be inferred from an entire
discourse;
 when the meaning is figurative rather than literal, as
in the moral of a story.
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 Language

A Language-Generating Mechanism?
 Children have a remarkable readiness to learn language;
on average, they learn a new word every 90 waking
minutes.
 This readiness led researchers to believe there is a
language acquisition device, a part of the brain that is
dedicated to learning and producing language.
 Other researchers agree that there are biological
mechanisms that make language acquisition so easy.
 For example, speaking and signing children follow the
same sequences in learning language.
 Their interpretation: Language has co-opted areas
specialized for abilities that language requires.
 Language

Specializations in the brain suggest that it is innately

well fitted for creating and learning language.
 The left hemisphere is dominant for language in 90% of
right-handed people and most left-handed.
 Broca’s area is larger and the lateral fissure and planum
temporale are longer on the left than on the right.
 Even newborns show left-hemisphere response to speech.

 Sign language activates the left hemisphere, even in

individuals deaf from birth.
 Language

Animal language research has the potential to reveal the

roots of language.
 Because chimps lack an adequate larynx for forming word
sounds, researchers have attempted to communicate with
them using various sign and symbol languages.
 Washoe learned to use 132 symbols, but critics said the
behavior was not complex enough to represent true
language.
 Washoe and three other chimps taught her son 47 signs and
they regularly carried on conversations among themselves.
Chimp Using American Sign Language
Figure 9.30
 Language

Other animals also share some of the brain

organization associated with human language.
 Chimps, monkeys, dolphins, and canaries show left-
hemisphere dominance for meaningful sounds or
gestures.
 Similar structures in these animals likely provide
prelanguage communicative abilities.
We share apparent genetic antecedents, such as
FOXP2.
However, the key to language probably lies in slight
modifications of those genes and in the genes that are
turned on or turned off.
◊
 Language

We do share mirror neurons with other species, and

they may be critical to the development of language.

QuickTime™ and a
Sorenson Video 3 decompressor
are needed to see this picture.

Figure 9.32
Brown areas are active during
imitation of others’ actions.
They overlap Broca’s and
Wernicke’s areas (yellow).