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.
Auditory hair cells that survive mechanical injury in culture begin their recovery by reforming the kinocilium. This study is based on cultures of the organ of Corti of newborn mice and two control animals. The axonemal patterns were examined in 165 kinocilia in cross-section. In the immature and regenerating kinocilium, one of the normally peripheral doublets is frequently located inward, forming the modified 8 + 1 (double) form; the distribution of the remaining microtubules is irregular. As the cell matures, the 9 + 0 form predominates. Overall, 34-61% of auditory kinocilia consist of 9 + 0 microtubules. The 9 + 2 (single) form, previously thought to characterize the organelle, occurs only in about 3-14%, whereas the remaining population comprises the modified 8 + 1 (double) form. Normally, the kinocilium lasts only about 10 postnatal days; however, post-traumatic hair cells reform their kinocilia regardless of age. Concomitant with the regrowth of the kinocilium, the basal body and its cilium take a central location in the cuticular plate, stereocilia regrow, and the cytoplasmic area adjacent to the basal body displays pericentriolar fibrous densities, growth vesicles, and microtubules, all surrounded by actin filaments. Pericentriolar bodies nucleate microtubules. Involvement of microtubules is seen in the alignment of actin filaments and in the formation of the filamentous matrix of the cuticular plate. We propose that reformation of the kinocilium in recovering post-traumatic hair cells indicates the possible role of its basal body in the morphogenesis and differentiation of cuticular plates and stereocilia.
Development of the cholinergic enzymes, choline acetyltransferase (ChAT) and AChE, and of the AChE-positive innervation in the cochlea was studied biochemically and morphologically in the postnatal mouse up to 26 days. Both ChAT and AChE are already present at birth in levels comparable to 50 and 20% of near-adult values, respectively. Increases in the enzymatic activities occur mainly during the second postnatal week. ChAT increases primarily in the basal turn; the specific activities in the basal and mid turns become about equal and at least twice of the values found in the apex. AChE increase continues throughout the entire cochlea; at all times its activity is highest in the base and lowest in the apex. In the light microscope, AChE-positive fibres are seen to enter the organ in the intraganglionic bundle during late foetal development and travel upwards via radial bundles. The fibres destined for outer hair cells usually differentiate first and take a separate route. They either cross the prospective tunnel of Corti directly or take a spiral course in front of inner pillar cells to form the inner pillar bundle. The tunnel fibres are radially oriented and provide the innervation to outer hair cells in narrow vertical sectors. In most cases, the outer hair cells are being innervated by the 4th day. Between the 4th and the 6th day, the tunnel fibres reach the outer hair cells in the third row; the first and second outer spiral bundles are formed. The AChE-positive innervation of the inner spiral bundle and plexus forms in short segments, and the bundle may be still discontinuous even by the 6th day. By the 12th day the innervation is complete. In the electron microscope, the stain for AChE may allow identification of growing efferent fibres before their ultrastructural differentiation. Both ChAT and AChE activities are early markers of the differentiating efferent system. An ingrowth of the cholinergic fibres to the entire cochlea occurs before birth. The greatest increase of AChE occurs between the 4th and 10th day, relating in time to efferent synaptogenesis.
The preservation and development of the innervation pattern in the organ of Corti have been studied in culture up to 27 days in vitro. The explants were obtained from the newborn mouse. Segments of the cochlear duct dissected together with the appropriate sectors of the spiral ganglion may retain their structural organization for about two weeks. Maturation of some nonneuronal elements which occurs during that time is followed by a subsequent regression of the organ. Only a fraction of the explanted neurons survive. However, the surviving neurons, if connected with the hair cell region, maintain a complex peripheral innervation pattern that contains all the major fibre components which characterize the normal pattern in a young mouse. The peripheral innervation pattern in culture seems largely composed of preserved fibres, that is, of fibres which at the time of explantation have already ramified within the organ of Corti. Nonetheless, there is evidence for growth or maturation, in culture, of at least some peripheral processes of the spiral neurons. Thus, only in older cultures is the innervation of the apical tip established. Likewise, it is only in older explants that the inner spiral bundle becomes prominent. Spiral neurons survive in culture in several modes. Most frequently, the central process is altogether absent and the neuron is effectively a unipolar cell which maintains only the peripheral process. A distinct minority of neurons is bipolar possessing both the peripheral process and a central axon which grows freely, though no central target is present. A neuron may survive also as a unipolar or, rarely, as a bipolar cell with no processes entering the organ of Corti. The observations imply that (1) most or all major fibre systems in the organ of Corti carry components of spiral neuron origin; (2) a small population of spiral neurons innervating a short segment of the organ contributes importantly not only to the radial but also to the spiral innervation of the segment.
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