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Transcript
Frequency
representation
Part 2
Development of mechanisms
involved in frequency representation
Summary of development of
frequency representation
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Frequency discrimination and frequency resolution
are immature in humans at birth, at least at high
frequencies
Frequency discrimination at high frequencies
becomes mature at about the same time as
frequency resolution, around 6 months
Frequency discrimination at low frequencies
continues to mature to at least school age
What are possible auditory physiological bases
of immaturity? Are there other explanations?
Overview of lecture
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Cochlear development (prenatal events for
humans)
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Anatomy
Physiology
Neural contributions to the development of
frequency representation
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Frequency resolution
Temporal code for frequency
Development of the
cochlea
Frequency resolution is established
in the mature cochlea and the
nervous system maintains that fine
frequency resolution
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Very early cochlear development
From Crossin et al, 1984
Timing of cochlear development
events in humans
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From Northern & Downs, 1991
Morphological development of the
cochlea
Prenatal and postnatal development
of the cochlea
Developmental events happen first
at the base, and progress apically
Hair cell innervation
Development of hair cell
innervation
Some potentially important
developments
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Changes in basilar membrane stiffness and
other mechanical properties of the organ of
Corti.
Differentiation and innervation of hair cells
Development of the stria vascularis and
endocochlear potential.
At the onset of cochlear function, sensitivity
and frequency resolution are poor.
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Frequency gradient in development
of frequency resolution?
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Measuring cochlear frequency
resolution using otoacoustic emissions
f1
f1
Level
Level
f2
Frequency
f2
suppressor
Frequency
2f1-f2
Frequency
Level
Level
f1
f2
2f1-f2
f1
Frequency
f2
Measuring cochlear frequency
resolution using otoacoustic emissions
Suppressor level
to reduce
emission by 3 dB
Suppressor frequency
Cochlear frequency resolution
development in humans
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So if cochlear tuning is mature,why do 3-monthold infants have immature frequency resolution?
You can have mature and immature tuning
in neurons tuned to the same frequency
Cochlea
Tuning quality
Superior olive
Auditory nerve
Frequency
Development of the end bulb of
Held
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Development of neural tuning in
human infants
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Sound frequency to which animals
first respond
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A paradox in development
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Place code shift
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Place code shift at three
frequencies
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Other observations about the place
code shift
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Responses in the nervous system shift with
the responses in the cochlea.
The shift occurs in mammals.
If an animal is trained to respond to a
certain frequency early in life, they will act
like they learned to respond to a lower
frequency later in life.
Possible causes of the place code
shift
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Middle ear response
Outer hair cells
Basilar membrane stiffness
Timing in humans?
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Development of the temporal code
for frequency
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Phase locking takes
longer to develop than
frequency tuning.
Phase locking
develops in the central
nervous system later
than at the periphery.
Development of phase locking in
humans
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Development of phase locking in
humans
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Development of frequency
representation: Conclusions
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Frequency resolution at the level of the cochlea is
mature prior to term birth, but at the level of the
brainstem is still immature until about 6 months.
The cochlear map of frequency shifts during
development; in humans this occurs prenatally.
The development of the temporal code for
frequency has not been studied extensively, but
there is some evidence that it may take longer to
develop than the place code.