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Auditory System


         1) Physical properties of sound

         2) Peripheral mechanisms in audition
                ear structure
                transduction

         3) Coding




                                                             1




                Functions of the Auditory System

    1) Sound identification (What is it? )
           Frequency
           Intensity

    2) Sound localization (Where is it?)
          Interaural timing differences
          Interaural frequency differences

    3) Communication




                                                             2




10% of the population has hearing disorders

1 in 1000 children have congenital deafness

1 in 3 people over the age of 60 has hearing loss

40% of the cells that detect sound are destroyed by age 65


Health cost in America is $56 billion/year



                                                             3




                                                                 1
Physical Properties of Sound

                                        Movement of a single
     Sound is energy produced           pulse of sound
     by vibrating bodies, which
     produces a disturbance in
     air molecules.

     The disturbance travels as
     a longitudinal wave, which
     has a region of condensation
     (close together) and a region
     of rarefaction (far apart).




                                   Region of       Region of
                                   rarefaction     condensation
                                                                         4




                     Properties of Sound
Speed:
In air, the speed of sound is 344m/s.

Frequency (wavelength/s):
The number of sound pulses that travel past a fixed point in a second.
                Species - Frequency Range
                Humans      20 - 20,000 Hz
                Bats        1000-100,000 Hz
Pitch:
Perception of frequency. High frequency sound is heard as a high pitch.

Intensity (amplitude):
Amount of air pressure.
                Normal breathing is about 10 dBs
                A typical conversation is 60 dB
                A jet taking off is 140 dB
Loudness
Perception of intensity. Loudness proportional to log (Intensity)

                                                                         5




               Sound Illusion: Doppler Shift
The frequency of sound that reaches our ears changes when
the sound source is moving.

The change in perceived pitch due to movement is known as
the Doppler shift.




                                                                         6




                                                                             2
THE EAR


Outer--- funnels sound; converts sound into physical vibration

Middle---transmits vibrations to the inner ear

Inner---converts fluid movements into neural firing




                                                                     7




                           The EAR

   OUTER                     MIDDLE                   INNER




                     Tympanic
External Auditory    Membrane
Meatus                                                               8




      Function of Middle Ear

                                            Malleus       Incus
 1. Three bones in the middle ear
    transmit the signal
         malleus- hammer
         incus-anvil
                                                            Stapes
         stapes-stirrup               Tympanic Membrane


 2. The middle ear amplifies signal by
    increasing the pressure on the oval
    window

  3. Inner ear muscles provide adjustable
     intensity control (dampen sounds)



                                                                  10




                                                                         3
INNER EAR




                                                       11




        Inner Ear: Chambers of the Cochlea




Scala vestibuli-contacts oval window
Scala media-enclosed by other 2, contains endolymph
Scala tympani-in contact with round window
                                                       12




               The Cochlea Unwound




        Oval Window    Scala Vestibuli
                                         Helicotrema

     Scala Media




                              Scala Tympani
       Round Window




                                                       13




                                                            4
Fluid movement in “unwound” cochlea
  •Fluid flows from oval window to round window




 Fluid path taken by wavelength
 outside of audible frequency




 Fluid path taken by wavelength
 within audible frequency

                                                                      14




        Cochlear Representation Of Sound - Theories


                                  Sound Frequency Cochlear Representation
Place theory:
Frequency is represented
by the location of the mechanical
response. Low frequency sounds
are closer to the helicotrema.
High frequency are closer to the
oval window


Frequency Theory:
This theory postulates that the
cochlea vibrates in synchrony
with the sound



                                                                      15




       Georg Von Bekesy
       1961 Nobel prize



 Von Bekesy studied
 cochlear response by
 opening a window to
 the cochlea and observing
 the effect of frequency on
 movement of the basilar
 membrane.




                                                                      16




                                                                            5
Discovery One: Basilar membrane has very tiny vibrations




                                                                            17




 Discovery Two: Different sounds produce vibrations
                   at different places

   Sound caused a traveling wave, with a maximum point of
   deflection that varied as a function of frequency.

                  Tuning Curves for various frequency signals
     Deflection




                                                                Low
                                                                Frequency




High                       Distance from Stapes
Frequency

                                                                            18




                                                                            19




                                                                                 6
Resonance properties of basilar membrane
                tune it to different frequencies


        Basal                                       Apical
        Thick                                       Thin
        Stiff                                       Floppy
        High frequency                              Low frequency




  stapes                 Basilar membrane                    helicotrema




                      A TONOTOPIC MAP!!!
                                                                         20




             Shape of Cochlear Response to Sound


                      Actual Response



                                                 Difference between
      Response seen by                           actual response and
      Von Bekesy                                 response predicted by
                                                 von Bekesy is the active
                                                 response.




        Active Response is caused by outer hair cells, which
        inject energy back into vibration of cochlear membranes

                                                                         21




                 Transduction of Sound by Hair Cells
                     Structure of Organ of Corti

               Tectorial membrane

                                                                  Inner
                                                                  Hair Cells
Outer
Hair Cells




                           Basilar Membrane
                                                                         22




                                                                               7
The hair bundle of hair cells




                                                           23




 How do hair cells sense mechanical stimulation?




               60 dB causes 10 nm deflection
                                                           24




       Tip Link Hypothesis of Activation and Adaptation




1) The stereocilia are connected together through tip
   links. The tip links end on gating springs.
2) Fluid movement in one direction puts pressure on
   gating links, leading to ion channels opening.

3) Fluid movement in opposite direction reduces pressure
   on gating links, leading to ion channels closing.

                                                           25




                                                                8
Tip Link Hypothesis




       • Channel connected to a sliding motor

       • Motor moves channel, changes tension on tip link

       • Allows the channel to rapidly open and rapidly adapt
                                                                26




                The channel is a potassium channel!!!!



                    “normal” cell               hair cell



                        Na                           K
extracellular           0 mV                       +80 mV



                        K                          K
intracellular
                      -45 mV                      -45 mV



                                                                27




                 Hearing aids or cochlear implants?

                 What’s the difference?




                                                                28




                                                                     9
Beyond the Ear…How sound is encoded in the brain


   •Basics of the auditory pathways

   •Properties of higher order auditory cells




                                                                       29




           Auditory Pathways- The highlights




                                                                       30




             Coding of Auditory Information


Lower Levels (cochlea, auditory nerve, cochlear nuclei)
       Tonotopic Organization


Superior Olive---Sound location

        receives input from both ears

        Interaural timing differences---sound reaches one ear first,
        determine sound location by the difference in time

        Interaural frequency differences--- sound frequency gets
        distorted by head, another way to determine location




                                                                       31




                                                                            10
Processing of Location in Inferior Colliculus



Cells show varying kinds of binaural interactions.

   • Some show an excitatory response to input to both ears (EE cells)
   •
   • Others show excitation to input to one ear, and inhibition
        to input to the other ear.

   • Interaural phase difference cells fire maximally when the
        input to the two ears is out of phase by a specified amount.

   • Cells sensitive to temporal order of input to two ears.




                                                                       32




                Major concepts for hearing

    1. Ear is specialized to detect sounds
    2. Cochlea has a tonotopic map
    3. Hair cells transduce sound information quickly
       - Direct gating of ion channels
       - Rapid adaptation
    4. Brain retains maps of different frequencies
    5. Brain computes sound location by differences in
       frequency and timing between the two ears




                                                                       33




                                                                            11

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Audition Ppt

  • 1. Auditory System 1) Physical properties of sound 2) Peripheral mechanisms in audition ear structure transduction 3) Coding 1 Functions of the Auditory System 1) Sound identification (What is it? ) Frequency Intensity 2) Sound localization (Where is it?) Interaural timing differences Interaural frequency differences 3) Communication 2 10% of the population has hearing disorders 1 in 1000 children have congenital deafness 1 in 3 people over the age of 60 has hearing loss 40% of the cells that detect sound are destroyed by age 65 Health cost in America is $56 billion/year 3 1
  • 2. Physical Properties of Sound Movement of a single Sound is energy produced pulse of sound by vibrating bodies, which produces a disturbance in air molecules. The disturbance travels as a longitudinal wave, which has a region of condensation (close together) and a region of rarefaction (far apart). Region of Region of rarefaction condensation 4 Properties of Sound Speed: In air, the speed of sound is 344m/s. Frequency (wavelength/s): The number of sound pulses that travel past a fixed point in a second. Species - Frequency Range Humans 20 - 20,000 Hz Bats 1000-100,000 Hz Pitch: Perception of frequency. High frequency sound is heard as a high pitch. Intensity (amplitude): Amount of air pressure. Normal breathing is about 10 dBs A typical conversation is 60 dB A jet taking off is 140 dB Loudness Perception of intensity. Loudness proportional to log (Intensity) 5 Sound Illusion: Doppler Shift The frequency of sound that reaches our ears changes when the sound source is moving. The change in perceived pitch due to movement is known as the Doppler shift. 6 2
  • 3. THE EAR Outer--- funnels sound; converts sound into physical vibration Middle---transmits vibrations to the inner ear Inner---converts fluid movements into neural firing 7 The EAR OUTER MIDDLE INNER Tympanic External Auditory Membrane Meatus 8 Function of Middle Ear Malleus Incus 1. Three bones in the middle ear transmit the signal malleus- hammer incus-anvil Stapes stapes-stirrup Tympanic Membrane 2. The middle ear amplifies signal by increasing the pressure on the oval window 3. Inner ear muscles provide adjustable intensity control (dampen sounds) 10 3
  • 4. INNER EAR 11 Inner Ear: Chambers of the Cochlea Scala vestibuli-contacts oval window Scala media-enclosed by other 2, contains endolymph Scala tympani-in contact with round window 12 The Cochlea Unwound Oval Window Scala Vestibuli Helicotrema Scala Media Scala Tympani Round Window 13 4
  • 5. Fluid movement in “unwound” cochlea •Fluid flows from oval window to round window Fluid path taken by wavelength outside of audible frequency Fluid path taken by wavelength within audible frequency 14 Cochlear Representation Of Sound - Theories Sound Frequency Cochlear Representation Place theory: Frequency is represented by the location of the mechanical response. Low frequency sounds are closer to the helicotrema. High frequency are closer to the oval window Frequency Theory: This theory postulates that the cochlea vibrates in synchrony with the sound 15 Georg Von Bekesy 1961 Nobel prize Von Bekesy studied cochlear response by opening a window to the cochlea and observing the effect of frequency on movement of the basilar membrane. 16 5
  • 6. Discovery One: Basilar membrane has very tiny vibrations 17 Discovery Two: Different sounds produce vibrations at different places Sound caused a traveling wave, with a maximum point of deflection that varied as a function of frequency. Tuning Curves for various frequency signals Deflection Low Frequency High Distance from Stapes Frequency 18 19 6
  • 7. Resonance properties of basilar membrane tune it to different frequencies Basal Apical Thick Thin Stiff Floppy High frequency Low frequency stapes Basilar membrane helicotrema A TONOTOPIC MAP!!! 20 Shape of Cochlear Response to Sound Actual Response Difference between Response seen by actual response and Von Bekesy response predicted by von Bekesy is the active response. Active Response is caused by outer hair cells, which inject energy back into vibration of cochlear membranes 21 Transduction of Sound by Hair Cells Structure of Organ of Corti Tectorial membrane Inner Hair Cells Outer Hair Cells Basilar Membrane 22 7
  • 8. The hair bundle of hair cells 23 How do hair cells sense mechanical stimulation? 60 dB causes 10 nm deflection 24 Tip Link Hypothesis of Activation and Adaptation 1) The stereocilia are connected together through tip links. The tip links end on gating springs. 2) Fluid movement in one direction puts pressure on gating links, leading to ion channels opening. 3) Fluid movement in opposite direction reduces pressure on gating links, leading to ion channels closing. 25 8
  • 9. Tip Link Hypothesis • Channel connected to a sliding motor • Motor moves channel, changes tension on tip link • Allows the channel to rapidly open and rapidly adapt 26 The channel is a potassium channel!!!! “normal” cell hair cell Na K extracellular 0 mV +80 mV K K intracellular -45 mV -45 mV 27 Hearing aids or cochlear implants? What’s the difference? 28 9
  • 10. Beyond the Ear…How sound is encoded in the brain •Basics of the auditory pathways •Properties of higher order auditory cells 29 Auditory Pathways- The highlights 30 Coding of Auditory Information Lower Levels (cochlea, auditory nerve, cochlear nuclei) Tonotopic Organization Superior Olive---Sound location receives input from both ears Interaural timing differences---sound reaches one ear first, determine sound location by the difference in time Interaural frequency differences--- sound frequency gets distorted by head, another way to determine location 31 10
  • 11. Processing of Location in Inferior Colliculus Cells show varying kinds of binaural interactions. • Some show an excitatory response to input to both ears (EE cells) • • Others show excitation to input to one ear, and inhibition to input to the other ear. • Interaural phase difference cells fire maximally when the input to the two ears is out of phase by a specified amount. • Cells sensitive to temporal order of input to two ears. 32 Major concepts for hearing 1. Ear is specialized to detect sounds 2. Cochlea has a tonotopic map 3. Hair cells transduce sound information quickly - Direct gating of ion channels - Rapid adaptation 4. Brain retains maps of different frequencies 5. Brain computes sound location by differences in frequency and timing between the two ears 33 11