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In the mammalian vestibular nerve, some afferents have highly irregular interspike intervals and others have highly regular intervals. To investigate whether spike timing is determined by the afferents’ ion channels, we studied spiking activity in their cell bodies, isolated from the vestibular ganglia of young rats. Whole cell recordings were made with the perforated-patch method. As previously reported, depolarizing current steps revealed distinct firing patterns. Transient neurons fired one or two onset spikes, independent of current level. Sustained neurons were more heterogeneous, firing either trains of spikes or a spike followed by large voltage oscillations. We show that the firing pattern categories are robust, occurring at different temperatures and ages, both in mice and in rats. A difference in average resting potential did not cause the difference in firing patterns, but contributed to differences in after hyperpolarizations. A low-voltage-activated potassium current (ILV ) was previously implicated in the transient firing pattern. We show that ILV grew from the first to second postnatal week and by the second week comprised Kv1 and Kv7 (KCNQ) components. Blocking ILV converted step-evoked firing patterns from transient to sustained. Separated from their normal synaptic inputs, the neurons did not spike spontaneously. To test whether the firing-pattern categories might correspond to afferent populations of different regularity, we injected simulated excitatory postsynaptic currents at pseudorandom intervals. Sustained neurons responded to a given pattern of input with more regular firing than did transient neurons. Pharmacological block of ILV made firing more regular. Thus ion channel differences that produce transient and sustained firing patterns in response to depolarizing current steps can also produce irregular and regular spike timing. [ABSTRACT FROM AUTHOR]

Rodent vestibular afferent neurons offer several advantages as a model system for investigating the significance and origins of regularity in neuronal firing interval. Their regularity has a bimodal distribution that defines regular and irregular afferent classes. Factors likely to be involved in setting firing regularity include the morphology and physiology of the afferents' contacts with hair cells, which may influence the averaging of synaptic noise and the afferents' intrinsic electrical properties. In vitro patch clamp studies on the cell bodies of primary vestibular afferents reveal a rich diversity of ion channels, with indications of at least two neuronal populations. Here we suggest that firing patterns of isolated vestibular ganglion somata reflect intrinsic ion channel properties, which in vivo combine with hair cell synaptic drive to produce regular and irregular firing. [ABSTRACT FROM AUTHOR]

Hair bundle structure is a major determi- nant of bundle mechanics and thus of a hair cell’s ability to encode sound and head movement stimuli. Little quantitative information about bundle structure is available for vestibular organs. Here we characterize hair bundle heights in the utricle of a turtle, Trachemys scripta. We visualized bundles from the side using confocal images of utricular slices. We measured kinocilia and stereocilia heights and array length (distance from tall to short end of bundle), and we calculated a KS ratio (kinocilium height/height of the tallest stereo- cilia) and bundle slope (height fall-off from tall to short end of bundle). To ensure that our measurements reflect in vivo dimensions as closely as possible, we used fixed but undehydrated utricular slices, and we measured heights in three dimensions by tracing kinocilia and stereocilia through adjacent confocal sections. Bundle heights vary significantly with position on the utricular macula and with hair cell type. Type II hair cells are found throughout the macula. We identified four subgroups that differ in bundle structure: zone 1 (lateral extra- striola), striolar zone 2, striolar zone 3, and zone 4 (medial extra- striola). Type I hair cells are confined to striolar zone 3. They have taller stereocilia, longer arrays, lower KS ratios, and steeper slopes than do neighboring (zone 3) type II bundles. Models and experiments suggest that these location- and type-specific differences in bundle heights will yield parallel variations in bundle mechanics. Our data also raise the possibility that differences in bundle structure and mechanics will help explain location- and type-specific differences in the physiological profiles of utricular afferents, which have been reported in frogs and mammals. [ABSTRACT FROM AUTHOR]