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Tuning in the basilar membrane  Vibration of basilar membrane can be described by a ‘tuning curve’  Amplitude required for the basilar membrane to vibrate at a certain constant displacement, as a function of the frequency of the input sound Neural connections in the cochlea  Afferent and efferent fibers of the VIIIth cranial nerve (auditory nerve)  Afferent: From organ of Corti to brain  Efferent: From brain to organ of Corti  Peripheral processes of auditory nerve neurons enter the cochlea through small openings on the edge of the osseous spiral lamina  These openings are called ‘habenulae perforata’  These fibers are then gathered in the modiolus Organization of the auditory nerve bundle in the modiolus  Fibers from the apex in the middle  Fibers from the base on the outside  This nerve bundle then goes to the cochlear nucleus in the brain Afferent fibers  Around 30,000 neurons in man  Only 5-15% of these innervate the OHC   These are called Type II or outer spiral fibers  One neuron connects to one OHC (one-to-many) Rest innervate the IHC  These are called Type I or radial fibers  Many neurons connect to one IHC (many-to-one) Efferent fibers  Originate from the olivocochlear bundle in the auditory brainstem  Efferent fibers synapse on the afferent nerves innervating the IHC  Efferent fibers synapse directly on the OHC Discharge pattern of a neuron  Neural spike has an initial large potential shift  Following this, refractory or rest period of around 1 msec Other terms  Spontaneous discharge rate: Neuron’s discharge rate without a stimulus  Threshold: Minimum stimulus level needed to cause an increase in the discharge rate above the spontaneous discharge rate Spontaneous rate and thresholds  Neurons with high spontaneous rate: Low threshold  Neurons with low spontaneous rate: High threshold Rate-Level function  Also called input-output or intensity function  Increase the level of the acoustic stimulus and measuring changes in the discharge rate of a single neuron Response area  Also called isolevel or isointensity curve  Iso: “Same”  Plotting how a neuron fires in response to sounds of different frequencies at a fixed intensity Tuning curve  Define a certain threshold for a neuron  Plot the level of the tone required for this neuron to discharge at this threshold, as a function of the frequency of the tone Encoding of frequency Two theories: 1. Place theory Tonotopic organization 2. Temporal theory Based on the periodic nature of nerve firing Place theory Different neurons in the nerve respond to different frequencies Frequency of input determined by which neuron(s) in the auditory nerve fires at the greatest rate Temporal theory  For lower frequencies (< 5000 Hz), discharge rates of neurons are proportional to the period of the input stimulus.  So for lower frequencies, rate of discharge of the neuron also provides information about the frequency of the stimulus. Encoding of intensity  Increase in discharge rate with increase in stimulus intensity  However, for most neurons, increase in discharge rate only occurs for a limited range of input intensity  Possible that discharge rate of many neurons may be combined to account for the 140 dB dynamic range of the ear