4/11/11 Inner Ear Physiology 1 Inner Ear Physiology 2 Inner Ear Physiology • Transduction • Tympanic membrane • Acoustical/mechanical • Oval window • Mechanical/hydraulic • Basilar & tectorial membrane • Hydraulic/mechanical • Hair Cells (stereocilia) • Mechanical/electric • Hair Cells (base) • Electrical/chemical • Auditory Neuron • Chemical/electrical 3 1 4/11/11 Block diagram of the cochlea 4 Frontal view of unrolled cochlea 5 Effect of stapes insertion in cochlea 6 2 4/11/11 Effect of outward motion of stapes. 7 Effect of outward movement of stapes 8 9 3 4/11/11 Theories of Hearing • Place Coding Theories • Resonance theory (Helmholtz, 1885) • Traveling wave theory (Georg von Bekesy 1928-1958- Nobel Prize 1961) • Many other theories including standing wave theory, pressure pattern theory, frequency analytic theory, etc. See Zemlin for details. • Temporal Coding Theories • Telephone theory (Rutherford, 1886) • Volley theory (Wever, 1949) 10 Place Theory: Impedance characteristics of the basilar membrane Resistance Mass reactance Stiffness reactance • Apical end is 5x wider and 100x more flaccid than basal end. 11 Resonance Theory The BM acts as a series of filters or tuned rods (analogy of piano strings). It performs a frequency analysis (Fourier Analysis) of the incoming signal. 12 4 4/11/11 Traveling Wave Theory • Most popular TW theory developed by Bekesy (1928-1958). • Used cochlear models and direct observation of cochlea. 13 Traveling wave 14 Traveling wave for complex tones Basilar Membrane Traveling Wave 8 kHz + 2 kHz 16 kHz + 2 kHz + 1 kHz 16 kHz + 8 kHz + 4 kHz + 2 kHz 15 5 4/11/11 Traveling wave and nonlinearities 16 Traveling wave and upward spread of masking • Excitation patterns and can be used to represent traveling wave. • Notice that low frequencies can mask high frequencies (e.g., 1000 masking 2000 Hz) better than the high frequencies mask lows (e.g., 1000 masking 500 Hz). 17 Telephone Theory • Problem is that the absolute refractory period is about 1 msec. • That is, a hair cell can fire only once per millisecond (1000 times per second or 1000 Hz). • Therefore, it would not be possible to transmit sounds to the CNS greater than 1000 Hz. • Another problem is that damage to certain parts of the basilar membrane (basal end) helps confirm a place theory. 18 6 4/11/11 Volley Theory • Modification of Telephone theory. • Receptor cells fire in groups: first one, then another, then a third • Resetting: the first group of cells to fire are resetting while the second and third groups fire and so on 19 Combining Place and Volley Theories Frequencies < 300, 400 Hz Mid Frequencies > 4000, 5000 Hz Predominate Comments Theory Volley The traveling wave is too broad to allow for specificity. Both theories apply Traveling Temporal coding not Wave supported in high frequencies 20 Electrical potential of the cochlea 21 7 4/11/11 22 Role of Hair Cells on Hearing Thresholds • Normal behavioral thresholds are due to amplification of motion by OHCs • Slight to moderate hearing loss is due to a breakdown of OHCs, and not IHCs • Severe hearing loss is due to the breakdown of OHCs and IHCs 23 Nerve Cell Anatomy Synapses Soma Terminal Boutons Axon & Myelin Dendrites Neurotransmitter 8 4/11/11 Auditory Neurons are Bipolar 25 Block diagram of auditory neuron 26 Sensitivity (Detection) vs. Specificity (Identification) Afferent Innervation of Inner and Outer Hair Cells Specificity: Many-to-one arrangement Nerve Fiber IHC Sensitivity: One-to-many arrangement Nerve Fiber OHC 9 4/11/11 Innervation 28 Summary 29 10