The auditory system

Subject

Status Completed

Week 5

Property Chapter 13

Sound

refers to pressure waves - generated by vibrating air molecules.

refers to an auditory percept.

The amplitude & freq. of sound pressure - change at ear roughly ~ correspond to
that listener’s exp. of loudness and pitch.

A sine wave - produced by the vibrating tines of a tuning fork.

Sinusoidal cycles of compression and rarefaction - form of circular motion w. 1

complete cycle equivalent to 1 full revolution.

Phase diff. have corresponding time diff. → appreciating how the auditory
system locates sounds in space.

Sounds in speech - contain energy distributed across a broad freq. spectrum.

Complex waveforms - have a periodic quality can be modelled ~ sum of

sinusoidal waves of varying amp., freq., and phases.

Fourier transform - decomposes complex signal into its sinusoidal

components.

The inner ear acts as a sort of acoustical prism - decomposing complex sounds into
a myriad of constituent freq.

The auditory system’s ability - encode these temporal features

⇒ imp. to speech & melody perception + the perceptual grouping of auditory ob. in
complex acoustical scenes.

The auditory system 1

 The audible spectrum

Human w. normal hearing - able to detect sounds that fall w/in a freq. range 20HZ
→ 20 kHz ~ the upper limit dropping off in adulthood.

Diff. species - tends to emphasise certain freq. bandwidths in both their vocalisation
+ range of hearing.

vocalisation distinguished from the noise ‘barrier’ created by environmental

sounds.

Echolocated animals - rely on very high freq. vocal sounds → resolve spatial
features of the target.

Animals avoiding predation have auditory system - tuned to low-freq. vibrations

→ approach predators transmit thru. the substrate.

A synopsis of auditory function

The auditory system transform sound into distinct patterns of neural activity →
integrated w. info. from other sensory systems/ brain regions imp. to movement,
attention and arousal.
→ guide behaviour - incl. orienting to acoustical stimuli ~ engaging in intraspecies
communication ~ distinguishing self-generated sounds from other sounds in the
environment.

Transformation - 1st occur at the external & middle ears → collect sound waves and
amplify their pressure.

sound energy in the air can be transmitted to the fluid-filled cochlea of the inner
ear.

inner ear - enable the freq., amp., and phase of the original signal →
transduced by the sensory hair cells → encoded by electrical act. of the
auditory nerve fibers.

The systematic representation of sound freq. along the length of the cochlea
(tonotopy) → imp. organisational feature preserved thru. the central auditory
pathways.

The auditory system 2

 The peripheral auditory info. - diverges into several parallel central pathways.

The superior olivary complex - info. from 2 ears interacts → the site of initial
processing of the cues - allows listeners to localise sound in space.

The cochlear nucleus - projects to inferior colliculus of the midbrain, major

integrative center → auditory info. can interact w. the motor system.

The inferior colliculus - relays auditory info. to the thalamus & cortex ~ plays a
prominent role in the perception of speech and music.

Both the peripheral & central auditory system - tuned to conspecific

communication vocalisations → pointing to the interdependent evolution of the
entire system for perceiving these signals.

The external ear

consist of the pinna, concha, and auditory meatus → gathers sound energy and
focuses it on the eardrums (tympanic membrane).

meatus - selectively boosts the sound pressure 30-100 fold for freq. around 3 kHz
via passive resonate effects.

This amplification - sensitive to freq. in the range of 2-5 kHz → explains. why
hearing loss near this freq. following exposure to high-intensity broadband noise.

⇒ In human - this range directly related to speech perception.

⇒ selective hearing loss in the range of 2-5 kHz disproportionately degrades
speech recognition.

2nd imp. function of the pinna and concha - selectively filter diff. sound freq. →
provide cues about the elevation of the sound source.

convolutions of the pinna - shaped the external ear transmits more high-freq.
components from an elevated source.

⇒ effect demonstrated by recording identical sounds from diff. elevations after

passing thru. artificial external ear.

The middle ear

The auditory system 3

 The sound induced vibrations are converted to neural impulses.

The major function of the middle ear - match relatively low-impedance airborne
sounds to the higher-impedance fluid of the inner ear.

A low-impedance medium (air) ~ higher-impedance medium (water) → all of the

acoustical energy in reflected.

The middle ear overcomes this prob. - ensure transmission of the sound energy
across the air-fluid boundary ~ by boosting the pressure measured at the tympanic
membrane almost all of the acoustical energy is reflected.

2 mechanical processes - occur w/in the middle ear to achieve this large pressure
gain.

The first and major boost is achieved by focusing the force - impinging on the
relatively large-diameter tympanic membrane → smaller-diameter oval window.

2nd and related process - relies on the mechanical advantage gained by the level
action of 3 small, interconnected middle ear bones (ossicles).

💡 Conductive hearing loss - invol. damage to the external or middle ear →

lower the efficiency at which sound energy is transferred to inner ear.

can be overcome by artificially boosting sound pressure levels w. an

external hearing aid.

Sensorineural hearing loss - due to conductive prob./ damage either to the

hair cells of inner ear ~ auditory nerves.

Contraction of these muscles - triggered auto. by loud noises/ during self- genrated
vocalisation

⇒ counteracts the movements of the ossicles ~ reduce the sound energy

transmitted to the cochlea, serving to protect the inner ear.

Hyperacusis - trigger painful sensitivity to moderate/ even low-intensity sounds.

Bony & soft tissues - have impedance values close to that of water → be transferred
directly thru. the bones & tissues of the head to the inner ear.

The auditory system 4

 The inner ear

Cochlea - energy from sonically generated pressure waves is transformed into

neural impulses.

not only amplifies sound waves → converts them into neural signals ~ act as a
mechanical freq. analyzer → decomposing complex acoustical waveforms into
simpler elements.

⇒ imp. to consider this structure in some detail.

uncoiled - form a tube about 35mm long.

bisected from its basal end almost to its apical end by the cochlear partition.

⇒ support the basilar membrane & tectorial membrane.

The oval & round window + other regions - bone is absent surrounding the
cochlea at the basal end of this tube.

Scala vestibuli & scala tympani - fluid filled chambers on ea. side of the cochlear
partition.

Scala media - runs w/in the cochlear partition.

⇒ The helicotrema - joins the scala vestibuli to the scala tympani → allow the fluid
perilymph to mix.

💡 The inward movement of the oval window - displaces the fluid of the inner ear
→ causing the round window to bulge out sightly ~ deform the cochlear
partition.

Basilar membrane - vibrates in response to sound → discharge rates of indv.

auditory nerve fibers - terminate along its length ~ showing both features tuned BUT
respond most intensely to spec. freq.

Freq. tuning w/in inner ear - attributable in part to the geometry of the basilar
membrane

The auditory system 5

 → wider & more flex. at the apical end ~ narrower & stiffer at the basal end.

💡 Georg - shows a membrane that varies systematically in its width & flex. →
vibrates max. at diff. positions as a function of the stimulus freq.

Acoustical stimulus - initiates a travelling wave → propagates from the

base toward the apex of the basilar membrane ~ growing in amp. +
slowing in velocity till max. displacement is reached.

⇒ max. displacement point - respond to high freq. at the base of basilar

membrane and low freq. at the apex ~ giving rise to a topographical mapping
of freq.

The indv. tones of complex sound - generated the spectrally complex stimuli that
causing the vibration equivalent to the superposition of the vibrations.
→ process of spectral decomposition - imp. strategy for detecting the various
harmonic combinations → distinguish natural sounds that have a periodic character.

The traveling wave - initiates sensory transduction by displacing the sensory hair
cells ~ sit atop the basilar membrane.

Stereocilia - protrude from the apical ends of the hair cells → leading to volt.
changes across the hair cell membrane.

Hair cells & the mechanoelectrical transduction of sound waves

The inner hair cells - sensory receptors ~ 95% of the fibers of the auditory nerve →
project to the brain arise from this subpopulation.

The outer hair cells - imp. role in modulating basilar membrane motions → imp.
component of the cochlear amplifier.

Ea. hair bundle - contains anywhere from 30 to few 100 stereocilia (kinocilium).

Kinocilium - true ciliary structure & in mammalian cochlear hair cells →

disappears a shortly after birth.

The auditory system 6

 The stereocilia - simpler → containing only an actin cytoskeleton.

Ea. stereocilium tapers - inserts into the apical membrane → form a hinge
about which ea. stereocilium pivots.

Fine filamentous structures - tip links → runs in parallel to the plane of bilateral
symmetry ~ connecting the tips of adjacent stereocilia.

provide the means for rapidly translating hair bundle movement into a receptor
potential.

⇒ displacement of the hair bundle parallel to plane of bilateral symmetry in direction

of tallest stereocilia - stretches the tip links → opening cation-selective
mechanoelectrical transduction (MET, hcMET) channels located at the end of the
link ~ depolarising the hair cell.
~ opp. direction - compresses the tip links → closing the hcMET channels ~
hyperpolarising the hair cell.

The tension on tip link varies - modulate the iconic flow → result in graded
receptor potential ~ follow the movement of stereocilia.

→ leads to transmitter release from the basal end of the hair cell → triggers
action potentials in cranial nerve VIII fibers ~ follow the up- down vibration of
basilar membrane at low freq.

Hair cells - convert the displacement of the stereociliary bundle into an electrical
potential in as little 10us.

The need for microsecond resolution - requires direct mechanical gating of the
transduction channel.

The exquisite mechanical sensitivity of the stereocilia - presents substantial risks.

eg. high intensity sounds - break tip links ~ destroy the hair bundle result in
profound hearing deficits.

💡 Imp. goal of current research - identify stem cells & factors → contribute to
the regeneration of human cells ~ affording a possible therapy for some forms
of sensorineural hearing loss.

The auditory system 7

 The ionic basis of mechanotransduction in hair cells

At the resting potential - only small fraction of the transduction channels are open.

→ cause depolarisation as K+ & Ca2+ enter the cell.

Depolarisation opens volt-gated Ca channels in hair-cell membrane

→ the resultant Ca2+ influx - cause transmitter release from the basal end of the
cell onto the auditory nerve endings.

The receptor potential - biphasic

1. movement toward the tallest stereocilia - depolarise the cell.

2. opp. direction of movement - hyperpolarise the cell → allows hair cell to

generate a sinusoidal receptor potential in response to sine stimulus.
⇒ preserving the temporal info. present in the original signal (freq. around 3
kHz).

The cell’s membrane time-constant - filters the asymmetrical

displacement-receptor current function of the hair cell bundle → produce a
tonic depolarisation of the soma ~ augmenting transmitter release + exciting
auditory nerve terminals.

K+ serves both depolarise & repolarise the cell → enabling the hair cell’s K+
gradient to be largely maintained by passive ion movement alone.

The apical end - protrudes into the scala media → exposed to the K+ rich & Na+
poor endolymph produced by dedicated ion-pumping cells in the stria vascularis.

The auditory system 8

 The basal end - bathed in perilymph ~ same fluid that fills the scala tympani →
resembles other extracellular fluids in that K+ poor & Na+ rich.

⇒ endolymph is about 80mV is more (+) than perilymph. (endocochlear potential)

~ inside hair cells about 45mV more (-) than perilymph/ about 12mV more (-) than
endolymph.

The electrical gradient across the membrane of the stereocilia - drives K+ thru.
open transduction channels into the hair cell

→ depolarise the hair cell ~ opening volt-gated K+ & Ca2+ channels located in the
membrane of the hair cell soma.

⇒ hair cell operates 2 distinct compartments - ea. dominated by its own Nernst
equilibrium potential for K+ → ensures that hair cell’s ionic gradient does not run down.

💡 A sensorineural hearing deficit - rupture of Reissner’s membrane

(presence of ethacrynic acid) - poison the ion-pumping cells of the stria
vascularis.
⇒ the hair cell exploits the diff. ionic milieus of apical & basal surface -
provide fast & energy-efficient repolarisation.

The auditory system 9

 The hair cell mechanoelectrical transduction channel

A single hair bundle - possess few as 200 functional channels → represent a tiny
fraction of all hair bundle proteins ~ factors that have rendered biochemical
purification impractical.

The complexity of the transduction apparatus - indicate the pore-forming molecule

→ must interact w. a variety of other accessory proteins to enable
mechanotransduction.

The mechanotransduction in hair cells - is the product of a multi-molecular machine

comprising these and others incl. associated tip links.

The cochlear amplifier

Recent studies - indicate the normal hearing depends on the act. of biological
amplifier located w/in the cochlea.

The tuning of auditory periphery - too sharp to be explained by passive mechanics

alone.

At low sound intensities - the basilar membrane vibrates 100-fold more than
predicted by linear extrapolation ~ from the motion measured at high intensities.

Otoacoustic emissions - the ear generate sounds under certain conditions →

detected by placing a sensitive microphone at the eardrum & monitoring the
response after briefly presenting a tone/ click.
→ provide a useful means to assess cochlear function in newborn.
(Cochlear microphonics)

The outer hair cells - essential component of the cochlear amplifier.

high sensitivity ‘notch’ of auditory nerve tuning curves - lost when hair cells
slectively inactivated.

isolated outer hair cells - contract + expand in response to small electrical

currents ⇒ provide a potential source of energy to drive an active process w/in

The auditory system 10

 the cochlear.

The opening & closing of hcMET channels - provides another source of energy
driving basilar membrane motion.
→ raising the possibility that inner hair cells also contribute to amplification.

Tuning & timing in the auditory nerve

The rapid response time of transduction - allows the membrane potential of hair
cells ~ follow deflection of hair bundle up to moderately high freq. of oscillation
(~3kHz).

the temporal modulation in sound - fall below 3kHz → encoded by the temporal
patterns of act. of hair cells & their associated nerve fibers.

Some mechanisms - used to transmit auditory info. at high freq.

Tonotopically organised basilar membrane - provided an alternative to temporal

coding (labeled line coding mechanism).

nerve fibers related to apical end - respond to low freq.

~ related to basal end - respond to high freq.

⇒ tuning curves - threshold function/ characteristic freq. - lowest threshold of the

tuning curves.

💡 Cochlear implants - effectively bypass the impaired transduction apparatus

→ restore some deg. of auditory function.

Most prominent feature of hair cells - follow the waveform of low-freq. sounds
→ imp. in other, more subtle aspects of auditory coding.

⇒ hair cells release transmitter only when depolarised - nerve fibers fire only during
(+) phases of low-freq. sounds.

‘phase locking’ - provides temporal info. from 2 ears to neural centers ~ compare
interaural time diff. for freq. up to 3kHz.

The auditory system 11

 How info. from Cochlear reaches targets in the brainstem

The auditory nerve - along w.

vestibular nerve constitutes cranial
nerve VIII ~ comprise the central
process of bipolar spiral ganglion
cells in the cochlear.

⇒ ea. cell sends a peripheral process - contact few inner hair cells + central process to
innervate the cochlear nucleus.

The tonotopic organisation of the cochlea - maintained in 3 parts of the cochlear

nucleus

ea. contains diff. populations of cells w. quite diff. properties.

Integrating info. from the 2 ears

The auditory system 12

 💡 Monaural hearing loss - strongly implicates unilateral peripheral damage -
either middle/ inner ear. auditory nerve itself.

Sound localisation - localise horizontal position of sound source ~ depend on the

freq. in the stimulus.

interaural time diff. - used to localise the source.

→ the longest time diff. detected by sounds arising directly lateral to one ear ~
are on the order of only 700us.

interaural intensity diff. - used as cues.

The neural circuitry - computes these tiny interaural time diff. ~ consist of binaural
inputs to the medial superior olive(MSO) → arise from left to right anteroventral
cochlear nuclei.

MSO contains cells w. bipolar dendrites - extend both medially (receive inputs
from the contralateral anteroventral cochlear nucleus) & laterally (receive
inputs from the ipsilateral anteroventral cochlear nucleus).

MSO - coincidence detectors → respond when excitatory signals arrive at the

same time (max. sensitive to diff. interaural time delays).
→ sound localisation - perceived on the basis of interaural time diff. → require
phase-locked info. from the periphery ~ avail. to humans only for freq. below 3
kHz.

eg. at frequencies higher than about 2 kHz, the human head

begins to act as an acoustical obstacle, because the
wavelengths of the sounds are too short to bend around it.
→ an acoustical ‘shadow’ of lower intensity is created at the
far ear.

The circuits - compute the position of a sound source on this basis found in lateral
superior olive (LSO) + medial nucleus of the trapezoid body (MNTB).

The auditory system 13

 excitatory axons - project directly from the ipsilateral anteroventral cochlear
nucleus to LSO

LSO receives inhibitory input from the contralateral ear via neuron in the MNTB.

⇒ excitatory-inhibitory interaction - results in a net excitation of the LSO on the same

side of the head ~ sound source.

sounds arising directly lateral to the listener’s head - firing rates will be highest in
the LSO on that side.

~ closer to the listener’s midline - elicit lower firing rates in the ipsilateral LSO.

(Figure 13.15)
⇒ ea. LSO encodes only sounds arising in the ipsilateral hemifield - both LSOs
represent the full range of horizontal positions.

The spectral ‘notches’ created by the shape of the pinnae - detected by neurons in
the dorsal cochlear nucleus.

binaural cues - imp. in localising the azimuthal position of sound sources.

spectral cues - used to localise the elevation of sound sources.

Monaural Pathways from the

Cochlear Nucleus to the Nuclei of
the Lateral Lemniscus (pg.299)

Integration in the Inferior Colliculus

The convergence of binaural inputs in the midbrain - produces sth. new ~ related to
the periphery (topographical representation of auditory space).

Humans have clear perception of both the elevational & azimuthal components of a
sound’s location.
→ indicating that our brains contain a neural representation of auditory space.

Its ability to process sounds w. complex temporal patterns.

→ many neurons in inferior colliculus - respond only to freq.-modulated sounds.

The auditory system 14

 ⇒ High deg. of convergence in IC inputs - convey info. about simpler cues ~ results in
more integrative + complex response properties (imp. to the representation of
auditory ob).

The auditory thalamus

The medial geniculate complex (MGC) - an obligatory relay for all ascending
auditory info. destined for the cortex.

several division - projects to the core region of the auditory cortex + medial &
dorsal divisions → organised like a belt around the ventral division.
→ project to the belt regions - surround the core region of the auditory cortex.

some mammals - MGC strictly maintained tonotopy of the lower brainstem

areas

→ exploited by convergence onto MGC neurons ~ generating specific

responses to certain spectral combinations.

Bat sonar illustrates 2 imp. points about function of the auditory thalamus.

1. MGC display pronounced selectivity for freq. combinations.

2. Cells in MGC - selective not only for freq. combinations ~ BUT also spec. time
intervals btwn the 2 freq.

⇒ speech sound - change continuously over the range of a few milliseconds →

suggest that MGC neurons in human capable of integrating acoustical info. over
millisec. - facilitate speech perception.

The auditory cortex

major target of the ascending fibers from MGC → play essential role in our
conscious perception of sound + the recognition of speech and music.

The core region in macaque monkeys - comprises 3 divisions (A1, R and RT)
→ located in lower bank of the lateral sulcus in the medial & posterior part of the
superior temporal gyrus (STG) in temporal lobe.

The auditory system 15

 A1 and belt areas - contain neurons that strongly responsive to spectral
combinations → characterise certain vocalisations.

Human implicate 2nd regions og auditory cortex in perception of pitch → imp. to

musical sense + to vocal communication - enable to hear 2 speech sounds as
distinct.

filled in a missing freq. further underscores - the auditory cortex is doing more
than faithfully representation what auditory periphery provides as input.

The belt & parabelt regions of the auditory cortex - receive more diffuse input from
belt division of the MGC ~ and prim. auditory cortex.

The auditory cortex obviously does much more than provide a

tonotopic map and respond differentially to ipsilateral and
contralateral stimulation.

Auditory cortical act. - strongly infl.both by linguistic features + cognitve context

→ consistent w. infl. of exp. + task demands on auditory cortical processing of
speech.
⇒ discriminate to complex sound simultaneously.

💡 patients w. bilateral damage to auditory cortex - reveal severe prob. in

processing the temporal order of sound.

Specific regions of human auditory cortex - specialised for processing

elementary speech sounds + temporally complex acoustical signals.

The auditory system 16

 Wernicke’s area - critical to the comprehension of human language ~ contiguous
w. the 2nd auditory area.

The auditory system 17