Period histograms, which display the distribution of spikes throughout the period of a periodic stimulus, were computed for discharges recorded from single auditory nerve fibers of the squirrel monkey when low-frequency tones were employed. For frequencies up to about 4 kHz, where phase locking is observed, the average phase angle of the discharges was tracked as frequency was varied in small steps from low to high. To a first approximation, the cumulative shift in average phase angle is a linear function of frequency as would be observed for an ideal delay line. The slopes of these phase-versus-frequency lines were found to be related through a power law to the best frequencies of the fibers. Thus, it seems possible to estimate the travel time of a mechanical disturbance between the oval window and any point on the cochlear partition. A more detailed examination revealed that average phase angle is also sensitive to intensity, depending upon the relation of the stimulating frequency to the best frequency. For stimulating frequencies below best frequency, discharges tend to occur progressively later in the cycle as intensity increases. Above best frequency, an opposite tendency often prevails if a change in phase angle occurs. At or near best frequency, little change in average phase angle is noted as intensity varies.
Studies of synaptogenesis in the developing organ of Corti in the intact mouse and in culture indicate that the inner and outer hair cells contain three populations of synaptic ribbons, i.e. ribbons adjacent to nerve fibres, free intracellular ribbons and misplaced ribbons apposed to non-neuronal elements. Ribbons adjacent to nerve fibres can be further classified into: ribbons synaptically engaged, ribbons participating in formation of presynaptic complexes only and ribbons that are not engaged to the hair cell membrane. In the developing innervated cultures the ribbon distributions are similar to those in the normal animal. Inner and outer hair cells differ in distribution of the ribbons. In the inner hair cells the ribbons adjacent to the nerve fibres are dominant (over 90%) and most of them (88%) are synaptically engaged. In the outer hair cells the presynaptic ribbons dominate the population (up to 60%) during the first postnatal week when the cells acquire afferent synaptic connections. This stage is followed by a marked reduction in the number of all ribbons. In the intact animal the rapid decrease results in a relative increase of misplaced and free ribbons. These changes are presumably due to the loss of some of the afferents. In the denervated hair cells the distribution of ribbons indicated the presence of conspicuous scatter. In the areas of incomplete denervation, however, the ribbons are apposed to the preserved fibres. Despite denervation, most of the ribbons develop the entire presynaptic complex in apposition to non-neuronal structures. The different populations of synaptic ribbons appear to reflect different stages in synapse formation. Possibly, the synaptic body originates in the interior of the hair cell and subsequently migrates to the cell membrane. In any case, a nerve fibre appears critical in influencing the location of the synaptic ribbon. At the apposition of the ribbon to the hair cell membrane, presynaptic densities are formed and the ribbon appears to become anchored. Typically, the nerve fibre membrane apposed to the presynaptic complex responds with the formation of postsynaptic densities.
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