The Inner Ear and Hair Cells (CH 12, pg. 283-300) I. Sound – changes in pressure A) Stimulus Parameters – Frequency (pitch) & Amplitude (loudness) 1. Humans – frequency range 20 Hz to 20kHz, pressure in dB B) Tympanic Membrane –> Ossicles (amplify sound pressure) -> Cochlea Fig. 12-3 C) The Cochlea Uncoiled 1. Ossicles transmit pressure to cochlea through Oval Window 2. Basilar Membrane – a) tapered, narrower (and stiffer) at oval window (base) b) wider, less stiff at apex c) each point on basilar membrane tuned to a particular frequency Amplitude of vibration Fig. 12-5 3 Text http://www.neurophys.wisc.edu/animations/ http://www.youtube.com/watch?v=yADT4gvzwE4 Endolymph (high K+), +80 mV Perilymph (0 mV) Fig. 12-4 Fig. 12-6 3) Vibration of basilar membrane causes bending of hair cells II. Hair Cells A. Kinocilium , Stereocilia – contain actin filaments B. Tip Links – stretching opens cation selective channels – 1. Channels located near the tips of the cilia C. Steps in the process of mechanotransduction 1. K+ enters at distal ends of cilia (endolymph) 2. Depolarizes Cell 3. Voltage sensitive Ca+2 channels open 4. Calcium entry causes exocytosis of vesicles containing synaptic transmitter 5. Transmitter release --> depolarization & action potential in postsynaptic cell (vestibular or auditory afferent) D. Evidence – record extracellular currents from saccular hair cells 1. EM of hair cells & locations of recording electrode Hudspeth, J. Neurosci. 2: 1-10, 1982 2. Some important controls Recording electrode In Contact Adjacent to it Stimulus probe Toward & away from kinocilium Perpendicular to it E. Frequency Tuning (Selectivity) 1. Frequency response of two hair cells in turtle cochlea 2. A pulse of sound produces resonant oscillations 3. Injecting current (depolarizing cell) produces oscillations at same frequency Fettiplace, TINS, 10: 421-425, 1987 Mechanism of Frequency tuning (CH 13, Box B) Endolymph (high K+) a) K+ enters at tip links b) Depolarization c) Voltage-gated Ca2+ opens d) Opens Ca2+ - gated K+ channel e) K+ leaves repolarization f) Ca2+ channel closes g) K+ channel closes h) Depolarization from tip link channels Perilymph (low K+) F. Transient signaling – Adaptation (Purves, Ch 14, Box B) 1. Maintain sensitivity to transient stimuli Hair cell bent by a constant amount Probability of K+ Channel Being Open Fig. 13-Box B Basilar Membrane Filtering City Traffic Conversation Fan at the back of this room Mosquito (Sellick, Patuzzi, and Johnstone, 1983) Prestin in action Outer Hair Cells are “Pre-Amps”