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major components of outer ear
auricle, helix, lobule, external acoustic meatus
major components of middle ear
tympanic membrane, malleus, incus, stapes,
major components of inner ear
semicircular canals, cochlea, maculae
what membrane separates middle ear from outer and inner?
tympanic membrane
frequency
corresponds to pitch, Hertz (Hz)
amplitude
corresponds to loudness/intensity, Decibels (dB)
fourier analysis
complex sounds broken down into their “pure tone” components
fourier synthesis
complex sound can be created by adding “pure tone” components
psychoacoustics
study of psychological correlates of physical dimensions and acoustics
energy does not equal perception
purpose of the pinna
capture sound waves to direct to inner ear, sound localization
function of the ossicles
collect and amplify sound waves from tympanic membrane, joints in between shake, send vibration to cochlea
ossicles
malleus, incus, stapes
how are sound waves visualized
waveforms, spectrogram, spectrum
differences in sound waves
waveform: pressure changes over time
spectrogram: plot a spectrum over time
spectrum: plot of amplitude vs. frequency
muscles of middle ear
tensor tympani and stapedius
acoustic reflex
muscles tense with loud sends and self-generate sounds such as chewing and swallowing
anatomy of cochlea
3 canals, vestibular canal, cochlear duct, tympanic canal
membranes separating layers of cochlea
between vestibular and cochlear: Reissner’s (vestibular) membrane
between cochlear and tympanic: basilar membrane
oval window
where stapes connects to cochlea, vibrations push stapes to oval window, transmits pressure into fluid
round window
pressure release in ear, bulges out in response to sound waves
helicotrema
located at apex of cochlea, connects vestibular and tympanic membrane, pressure equalization and low-frequency sound transmission
what makes up cochlear partition
tectorial membrane, organ of Corti, basilar membrane
inner hair cells
1 row, afferent
arranged front to back, shortest to longest
most auditory info (90-95% of audio info)
outer hair cells
3 rows, mainly efferent innervation
stereocilia on top of OHCs in V or W shape
improve sensitivity and frequency selectively
what happens in cell when hair cells are stimulated
stereocilia bend due to fluid movement
gated ion-channels open
potassium rushes in from surrounding endolymph, hair cell depolarizes
calcium enters, releases NTs
NTs stimulate auditory nerve, sending information to the brain
does stimulation of hair cell lead to action potential
no, but they produce graded receptor potential
equal loudness curves
frequency vs. loudness of sound
ears are most sensitive between 2-4 kHz
low and high frequencies have to be much louder than midrange
why does frequency not equal pitch
can discriminate between 1 Hz difference
500 > 1000 Hz is bigger change than 5000 > 5500, 100% vs. 10% difference
why does decibels not equal loudness
we are sensitive to changes in loudness, can discriminate changes of less than 1 dB
intensity leads to more action potentials and more action potentials are generally interpreted as higher volume
response comprehension
how we perceive and interpret loudness
amplitude coding in cochlea
rate of hair cell depolarization
firing rate of auditory nerve fibers
number of hair cells activated
frequency coding in cochlea
place coding and frequency matching
place code
different frequencies displace different regions of cochlear partition
higher frequencies: closer to base (more accurate)
lower frequencies: closer to apex
frequency matching
phase locking and volley principal
phase locking
each AN fiber fires at particular point in sound wave
works for low frequency, fiber can only fire so fast
volley principal
multiple auditory nerves divide and conquer high frequency sounds
each neuron locks on particular part of sound wave
2-tone suppression
two sounds of different frequencies enter ear, one tone can suppress vibration of basilar membrane
helps cochlea sharpen frequency selectivity
rate saturation
limit of how fast an auditory neuron can fire in response to increasing sound intensity
once saturation is reached, firing rate is peaked and cannot get faster
how many auditory nerves innervate a single hair cell
IHC: Many:1
OHC: 1:Many
characteristic frequency
each IHC and ANF has characteristic frequency based on its location on basilar membrane
base responds to higher frequencies, apex responds to lower frequencies
aspects of stimulus affecting rate of firing on auditory nerve
amplitude, frequency, duration
labeled-line coding
cross-fiber patterning
each ANF is tuned to specific characteristic frequency
brain interprets identity of active neuron to determine frequency of the sound
why is labeled-line coding better at telling us frequency of quiet sounds than loud sounds
as only a few ANF’s respond, they tune closes to the sound’s frequency
at high sounds, many fibers respond, incuding those tuned to nearby frequencies
spiral ganglion
transmits electrical signals from IHCs to cochlear nucleus in brainstem
type I neurons
myelinated, innervate 1 IHC
type II neurons
non-myelinated, connect to multiple OHCs
brain structures involved in auditory pathway in order
pinna, IHC, spiral ganglion, cochlear nucleus, olives, inferior colliculus, medial geniculate nucleus, A1, A2, association cortex
why is the superior olive a good place for sound localization to occur
first point where brain compares input from both ears, allows it to detect tiny differences in timing and loudness
what is the auditory cortex
part of the brain responsible for processing sound information
where is the auditory cortex
temporal lobe, superior temporal gyrus
primary auditory cortex
any sound elicits activity, bilateral activation
secondary auditory cortex
more complex sounds elicit activity
association cortex
more complex sounds elicit activity, also processes other sensory information
tonotopic organization
base of cochlea responds to high frequencies, apex responds to low frequencies
in temporal lobe, each area corresponds to a different frequency spot in cochlea
Broca’s area
in frontal lobe, important for motor speech production
Broca’s aphasia
“broken speech”
non-fluent aphasia, difficulty with motor production, speak slowly and inarticulately
trouble with writing and gestures
Wernicke’s area
in temporal lobe, important for language comprehension
Wernicke’s aphasia
“word salad”
fluent aphasia, smooth, but nonsense speech
impaired ability to remember names of objects, difficulty finding right word
parts of the brain important for language processing
Broca’s area, Wernicke’s area, A1, A2
difference between left and right hemispheres for language/speech processing
language is usually in left hemisphere, lateraliized
how things are said in right hemisphere, emotion
Jeffress model of sound localization
how the brain localizes sound using ITDs to explain differences in the time a sound takes to arrive at each ear
ex: sounds arriving at right ear first will activate right leading neurons first, then signal from left will activate same neuron
what brain region does interaural time difference
medial superior olive
interaural time difference (ITD)
difference in time it takes for sound to reach each ear
works best in low frequencies below 1500 Hz
helps us determine which side a sound is originating from
how do we determine interaural level difference
based on intensity of sound reaching ear (how loud)
helps ears localize sounds for high frequencies (above 1500 Hz)
what brain region determines interaural level difference
lateral superior olive
why is interaural level difference different from front to back
head shadowing, one side blocking sound waves from the other
cones of confusion
3D surfaces around head wehre sounds produce same ITD and ILD
use pinna spectral cues, head movement, and other cues
how does the pinna help with localizing sounds
funnels sound energy into the ear canal, better at funneling some frequencies over others
direct transfer function (DTF)
function that describes how the pinna, ear canal, head, and torso change intensity of sounds with different frequencies
are all frequencies funneled to the eardrum equally
no, depends on pinna and ear canal shapes
certain frequencies get amplifed (2000-5000 Hz)
methods of distance perception
relative intensity of sound
spectral composition of sounds
relative amount of direct energy vs. reverbant energy
conductive hearing loss
caused by problems with ossicles
ex: ear infections, otitis media
otosclerosis
abnormal growth of middle ear bones, can be fixed by surgery
sensorineural hearing loss
most common
defects in cochlea and auditory nerve, hair cells are injured
result of antibiotics or cancer drugs
natural consequences of aging
young people: 20-20000 Hz
college age: 20-15000 Hz
cochlear implants
flexible coils with miniature electrode contacts
signals activate electrode to appropriate positions along cochlear implant