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Cochlear Functions • Transduction- Converting acousticalmechanical energy into electro-chemical energy. • Frequency Analysis-Breaking sound up into its component frequencies Transduction• Inner Hair Cells are the true sensory transducers, converting motion of stereocilia into neurotransmitter release. Mechanical Electro-chemical • Outer Hair Cells have both forward and reverse transduction-Mechanical Electro-chemical Mechanical Electro-chemical Hair Cell Activation • Involves Ion Flow into cell • Through channels in the stereocilia • Bending stereocilia causes # of open channels to change. • Toward Modiolus = Fewer channels open • Away from Modiolus = More open Ion Channels are opened by “TIP LINKS” • Tip Links connect tip of shorter stereocilia to the side of a stereocilium in the next taller row • Bending toward taller rows pulls tip links • Bending toward shorter rows relaxes tip links Tip Links Resting (or Membrane) Potentials • Inner Hair Cell = - 45 mV • Outer Hair Cell = - 70 mV Stereocilia bent toward tallest row • Potassium flows into cell • Calcium flows into cell • Voltage shifts to a less negative value • More neurotransmitter is released Sensory Physiology • The basic unit of the nervous system is the Neuron or nerve cell • Neurons undergo voltage changes– SPONTANEOUSLY – IN RESPONSE TO STIMULATION The Neuron Neural Activity • Post-synaptic Potentials-- Local, Variable changes in voltage near synapse • Action Potentials-- Conducted through axon, “all or none,” “spike” • For image of AP’s traveling down an axon: http://bio.winona.msus.edu/berg/ANIMTNS/actpot.htm An Action Potential (or Spike) Action Potentials • Are generated spontaneously – At a slow rate by some neurons – At a faster rate by some neurons • And occur more frequently with STIMULATION • Spike rate increases through a range of about 30 dB Spike Rate (APs/sec) Spike Rate Increases Thru a 30 dB Range 90 80 70 60 50 40 30 20 10 0 Spike Rate 0 5 10 15 20 25 30 35 40 45 50 55 60 Stimulus Level (dB SPL) Cochlear Mechanics • Passive: Bekesy’s Traveling Wave Theory • Active: Outer Hair Cell Motion enhances stimulation of inner hair cells The Traveling Wave BASE APEX Bekesy’s Theory describes Passive Mechanics • Based on work in “dead” cochleae • Highly damped -- not sharply tuned • Active Undamping occurs in live and healthy cochleae • Like pumping on a swing--adds amplitude The Active Component Adds to Bekesy’s Traveling Wave The Active Component • Improves Sensitivity for soft sounds • Improves frequency resolution Frequency Tuning Curves Show these Effects = plots of response threshold as a function of frequency They have a characteristic shape • sharp tip (shows best sensitivity at one freq) • steep high frequency tail • shallow low frequency tail Tuning Curves Passive Only Active + Passive More on Tuning & Tuning Curves: • Frequency of “tip” is called the CHARACTERISTIC FREQUENCY • Can be seen for: basilar membrane, hair cells, nerve cells Tonotopic Organization • Mapping of Characteristic Frequency by place in the auditory nervous system • In the Auditory portion of cranial nerve VIII the map is spiral-shaped with highest CF neurons on the outside, lowest CF neurons at the center. Head-Related Effects • Head-Baffle--the mere presence of your head alters the sound field. • Head Shadow - loss of energy at far ear for frequencies above approx 2000 Hz Signs of Peripheral Activation • Otoacoustic Emissions (OAEs) • Cochlear Potentials Otoacoustic Emissions • Low-level sounds produced by the cochlea and recordable in the external ear canal. • • • • Spontaneous Click-evoked Distortion Product Stimulus Frequency Recording OAEs Spectrum of Sound in Ear Canal OAEs: Measures of Cochlear Health Cochlear Potentials: • Resting Potentials: voltages which exist without external stimulation e.g., Endolymphatic Potential, Cell Membrane Potential • Stimulus-Related Potentials: voltages occurring in response to sounds We’ll talk about 3 of these from the cochlea Cochlear Microphonic • Least valuable from a clinical standpoint. • Is an alternating current (AC) response that mirrors the waveform of low to moderately intense sound stimuli • Appears to arise from outer hair cells in the basal-most turn of the cochlea Summating Potential (SP) • Is a direct current or DC potential • Lasts for duration of stimulus. Compound Action Potential (CAP) • Summation of APs in large number of VIIIth nerve neurons • following onset (and offset) of stimulus