Vestibular receptors in mammals: afferent discharge characteristics and efferent control.
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Selective expression of the NOS II isoform during mouse vestibulocochlear receptorgenesis.
Based on in vitro studies, nitric oxide (NO) is reported to be involved in initial neuronal differentiation. In order to compare this finding with the situation in vivo, we have looked for the expression of the three NO synthase isoforms in the developing mouse vestibulocochlear system. From these isoforms only the inducible NOS II is expressed during inner ear development. Examination of a series of embryonic and early postnatal animals, up to postnatal day 6, reveals a maturation-dependent, monophasic expression of this isoform. Initial expression is observed by day 10 of gestation in nerve cells of the vestibolocochlear ganglion and on their fibres. By day 14 of gestation, these afferent fibres penetrate the epithelium of the prospective receptor fields making contact with early, differentiating immunoreactive cochlear hair cells and receptor cells of the macula and crista ampullaris. This receptor-cell-derived immunoreactivity vanished in differentiated sensory hair cells by postnatal day 6, when both the constitutive isoforms and subsequent activated members of the down stream second messenger cascade (guanylate cyclase/cGMP) of the adult mouse were not then detectable. The strict phasic expression of NOS-II, independent of the second messenger system mentioned above, implies that there is a unique role for the inducible NOS isoform in nerve cell differentiation, independent of the NO/guanylate cyclase/cGMP pathway.
Aging↗
Self-motion magnitude estimation during linear oscillation: changes with head orientation and following fatigue.
Two types of experiments concerning estimated magnitude of self-motion during exposure to linear oscillation on a parallel swing are described in this paper. Experiment I examined changes in magnitude estimation as a function of variation of the subject's head orientation, and Experiments II a, II b, and II c assessed changes in magnitude estimation performance following exposure to sustained, "intense" linear oscillation (fatigue-inducing stimulation). The subjects' performance was summarized employing Stevens' power law (R = k . Sn, where R is perceived self-motion magnitude, k is a constant, S is amplitude of linear oscillation, and n is an exponent). The results of Experiment I indicated that the exponents, n, for the magnitude estimation functions varied with head orientation and were greatest when the head was oriented 135 degrees off the vertical. In Experiments II a-c, the magnitude estimation function exponents were increased following fatigue. Both types of experiments suggest ways in which the vestibular system's contribution to a spatial orientation perceptual system may vary. This variability may be a contributing factor to the development of pilot/astronaut disorientation and may also be implicated in the occurrence of motion sickness.
Acceleration↗
Inhibition of vestibular and neck reflexes in forelimb extensor muscles during the episodes of postural atonia induced by an anticholinesterase in decerebrate cat.
In precollicular decerebrate cats, the multiunit EMG activity of forelimb extensor muscles (e.g. the triceps brachii) was recorded during sinusoidal stimulation of labyrinth and neck receptors at the frequencies of 0.026-0.15 Hz, +/- 10 degrees and the resulting responses were tested during tonic activation of a cholinergic mechanism. In agreement with previous findings, the first harmonic component of the EMG responses to roll tilt of the animal leading to selective stimulation of labyrinth receptors was characterized by an increased activity during side-down tilt and a decreased activity during side-up tilt (labyrinth responses); on the other hand just the opposite changes were elicited for the same directions of neck rotation (neck responses). For the parameters of stimulation reported above, the responses were always related to position and not to velocity of displacement. Intravenous injection of an anticholinesterase (eserine sulphate, 0.10-0.15 mg/kg) which produced a state of postural atonia, associated with bursts of rapid eye movements (REM), similar to that occurring spontaneously in unrestrained cats during desynchronized sleep or REM sleep, also decreased the tonic activity of the triceps brachii and abolished the EMG responses of this muscle to sinusoidal stimulation of labyrinth and neck receptors. This suppression persisted throughout the episode of postural atonia associated with REM bursts. The abolition of the labyrinth and neck reflexes acting on forelimb muscles was not only dependent on the dose of anticholinesterase, but also on the state of the animal. In fact, somatosensory or acoustic stimuli applied during the REM episodes abolished the rhythmic oculomotor activity and determined the prompt recovery of both the decerebrate rigidity and the EMG responses of the triceps brachii to labyrinth and neck stimulation. The postural atonia as well as the tonic depression of the vestibular and neck reflexes acting on forelimb extensor muscles can in part at least be attributed to cholinergic activation of medullary reticulospinal neurons exerting a postsynaptic inhibitory influence on extensor motoneurons. However, since these inhibitory reticulospinal neurons collaborate with excitatory vestibulospinal neurons to the motoneuronal responses during stimulation of labyrinth and neck receptors, we cannot exclude that the suppression of the vestibular and neck reflexes may also depend on occlusion of the unit responses at reticular level.
Animals↗
Mechanisms for sensory adaptation in frog vestibular organs.
The effects of external low-Ca, high-Mg solutions were tested both on frog isolated semicircular canals and on single cells isolated from these sensory organs. Our results showed that these media were able to cancel slow adaptation of the ampullar microphonic current in the whole organ and to abolish a Ca-dependent K current (IK(Ca)) in single hair cells, suggesting that IK(Ca) is involved in vestibular sensory adaptation.
Adaptation, Physiological↗
[Structure of the vestibular apparatus and ionic composition of the body of Xenopus laevis larvae as affected by weightlessness].
Clawed frog larvae (stages 45-46) that developed for 9 and 8 days beginning with the blastula and tail bud stages, respectively, in the weightless state were investigated. Scanning microscopy of the larval labyrinths did not reveal significant qualitative changes in the receptor and supporting cells of the maculae and in the otolith membrane. Determination of the electrolyte composition (Na, K, Ca, Mg) of the larval body showed no significant changes in the relative content of these elements. The morphometric examinations indicated an increase of the size of the utricular otoliths by 1.3 times as a result of exposure to weightlessness. There was a tendency for a greater asymmetry between the left and right otoliths in the same larva.
Animals↗
Hair cell replacement in avian vestibular epithelium: supporting cell to type I hair cell.
Previous investigations have demonstrated that the sensory epithelium of the avian vestibular system possesses the capacity to replace hair cells both on an ongoing basis and following severe damage. Supporting cells, within the sensory epithelium, are believed to be the progenitors of the regenerated hair cells. In the present study we describe the series of events leading to the formation of a regenerated vestibular hair cell in post-hatched birds. Young chickens received injections of streptomycin sulfate in order to damage the sensory epithelium of the vestibular system. These injections were followed by injections of the cell proliferation marker tritiated-thymidine. At predetermined intervals, the animals were killed, and the vestibular organs were processed for tissue autoradiography. Our results confirm that hair cells originate from supporting cells. The data also indicate that postmitotic cells migrate towards the lumen of the epithelium where they differentiate into Type II hair cells. At a later time, some of the new Type II hair cells further differentiate into Type I hair cells. These results suggest that both types of avian vestibular hair cells have a common ancestor. The data also provide evidence in support of the hypothesis that calyx enclosed Type I hair cells, only present in birds and mammals, are a more differentiated stage of Type II hair cells.
Animals↗
Apoptosis during chick inner ear development: some observations by TEM and TUNEL techniques.
In order to clarify the occurrence, distribution and possible role of apoptosis during inner ear development, the ultrastructural aspects (by TEM) (at 9-19 incubation day and 1 day after hatching) and the distribution of the apoptotic phenomenon (by the TdT-mediated dUTP nick end-labeling technique), were studied in the crista ampullaris of chick embryo at 5-19 days of incubation to hatching and of postnatal 1-day old chick. We found, in the sensorial epithelium, dark supporting cells in chick embryos and mainly dark hair cells in postnatal chicks, both with ultrastructural features consistent with those of apoptosis. The presence of apoptotic phenomena was confirmed by the TUNEL technique. According to our findings, it is hypothesized that apoptosis in the inner ear may be involved: 1) at first, in macroscopic remodelling of the membranous labyrinth in early developmental stages, 2) later, in the correct differentiation of the hair and of the supporting cells, leading to characteristic cellular pattern formation and 3) finally, in physiological cell turnover of the postnatal chicken sensorial epithelium of the crista.
Animals↗
Carbohydrates of the vestibular end organs. An ultrastructural study using gold-labeled lectins.
The presence of carbohydrates in the glycocalyx of the vestibular end organs of the guinea pig was investigated at the ultrastructural level using lectin-gold solutions. The glycocalyx of both the sensory and the supporting cells has variable sugar components. The glycocalyx of the sensory cells including the ciliary interconnections contains N-acetyl-glucosamine, galactose and mannose. In contrast, the glycocalyx of the supporting cells had a lower amount of N-acetylglucosamine. The ciliary interconnections, which have been considered to be a part of the glycocalyx, have larger amounts of galactose and mannose than the surface glycocalyx. These findings indicate that the sugar components may be closely related to the functional significance of the inner-ear glycocalyx and that the functional properties of the glycocalyx may differ between the sensory and the supporting cells of the vestibular end organs.
Animals↗
Morphometric analysis and fine structure of the vestibular epithelium of aged C57BL/6NNia mice.
The vestibular organs of young and very old C57BL/6NNia (B6) mice were compared by light and electron microscopy. Hair cell density decreased an average of 14% in the utricle, 19% in the saccule and posterior crista, 23% in the horizontal crista, and 24% in the anterior crista. Hair cell size remained the same throughout the mouse's life span as did the ratio of Type I to Type II hair cells. The most apparent sign of advanced age was dense inclusions found in sensory and supporting cells. Although small inclusions were present at five weeks, by 29 months, additional, larger forms appeared. An unusual melanin-like form was characteristic of old Type I hair cells. Synaptic morphology and synaptic bodies were well preserved even in very old B6 mice. Elongated bars were common in Type I hair cells and spheroid synaptic bodies were the most common form in Type II hair cells. Large clusters of synaptic bodies occurring in both young and old mice were seen only in Type I hair cells. Although the B6 strain suffers from genetically determined early cochlear degeneration, it does not experience early degeneration of the peripheral vestibular organs.
Aging↗
Identification of hair cell progenitors and intermitotic migration of their nuclei in the normal and regenerating avian inner ear.
Postembryonic production of sensory hair cells occurs in both normal and aminoglycoside-damaged avian inner ears. The cellular source and mechanism that results in new differentiated hair cells were investigated in the avian vestibular epithelia using three distinct cell-cycle-specific labeling methods to identify proliferating sensory epithelial cells. First, immunocytochemical detection of the proliferating cell nuclear antigen, an auxiliary protein of DNA polymerase, allowed labeling of cells in late G1, S, and early G2 phases of the cell cycle. Second, a pulse-fix tritiated thymidine autoradiographic protocol was used to identify cells in S phase of the cell cycle. Finally, Hoechst 33342, a fluorescent DNA stain, was used to identify epithelial cells in mitosis. The distribution of cells active in the cell cycle within the normal and ototoxin-damaged vestibular epithelium suggests that supporting cells within the sensory epithelia are the cellular precursors to the regenerated hair cells. Differences between the proliferation marker densities in control and damaged end organs indicate that the upregulation of mitotic activity observed after streptomycin treatment is due primarily to an increase in the number of dividing progenitor cells. The differences between the extent of ototoxic damage and the level of reparative proliferative response suggest a generalized stimulus, such as a soluble chemical factor, plays a role in initiating regeneration. Finally, after DNA replication is initiated, progenitor cell nuclei migrate from their original location close to the basement membrane to the lumenal surface, where cell division occurs. This pattern of intermitotic nuclear migration is analogous to that observed in the developing inner ear and neural epithelium.
Aminoglycosides↗
Nature of optokinetic response and zonal organization of climbing fiber afferents in the vestibulocerebellum of the pigmented rabbit. I. The flocculus.
In pigmented rabbits anesthetized with N2O (70%) and halothane (2-4%), Purkinje cells were extracellularly recorded in the flocculus. A large central visual field (60 degrees x 60 degrees) was used to optokinetically stimulate either the ipsi- or contralateral eye, and the direction and velocity selectivities of complex spike responses were examined. For optokinetic stimulation (OKS) delivered to the ipsilateral eye (n = 129), the preferred direction was forward (F, n = 57) or upward (U, n = 37), while the remaining cells (n = 35) showed no response (N). For OKS delivered to the contralateral eye (n = 107), the preferred direction was backward (B, n = 11), downward (D, n = 42) or upward (U, n = 2), and the rest (n = 52) showed N. Cells tested with both eyes (n = 89) fell into five categories based on the preferred direction to ipsi- and contralateral OKS: (1) ipsi-F and contra-B (F/B type, n = 9), (2) ipsi-F but contra-N (F/N type, n = 28), (3) ipsi-U and contra-D (U/D type, n = 13), (4) ipsi-U but contra-N (U/N type, n = 17), and (5) ipsi-N but contra-D (N/D type, n = 22). The optimum velocity was within 0.1-2.0 degrees/s for all cells. On the average, the best response was obtained at 0.2-0.5 degrees/s stimulation. All ipsi-F cells responded to electrical stimulation of the optic tract (OT), while most cells preferring ipsi-U, contra-B and contra-D directions did not respond. No characteristic feature was found in cells innervated with collateralized climbing fiber branches to the nodulus. In the flocculus, cells preferring horizontal orientation (H cells, preferring ipsi-F and/or contra-B directions) were localized in a narrow dorsoventral zone (less than 1.0 mm) along the caudal border of the rostral one third, while those preferring the vertical orientation (V cells, preferring ipsi-U and/or contra-D directions) were in two distinct narrow zones located rostral and caudal to the H cell zone. H and V cells were intermingled in the central portion of the ventral flocculus. These four zones are in good agreement with previously defined H, anterior V, posterior V and R zones, respectively. The results indicate that the subdivision of the flocculus which controls horizontal (vertical) eye movements receives information regarding movements of the visual surround in the horizontal (vertical) orientation through visual climbing fiber afferents, thus being organized in olivo-cortico-nuclear functional units for control of eye movements.
Action Potentials↗
Pattern of gentamicin-induced cochlear degeneration in the guinea pig. A morphological and electrophysiological study.
Gentamicin-induced cochlear degeneration in the guinea pig was studied by complete hair-cell counting (cytocochleograms) and phase-contrast and interference microscopical examination of the stria vascularis and Reissner's membrane. Gentamicin (100 mg/kg/day) was administered over a period of 7-17 days. The first loss of hair cells (OHC) occurred in a region 6-8 mm from the round window. From this 'degeneration point', the loss of haircells progressed towards the round window (fast) and the apex (slowly). The stria vascularis showed no signs of degeneration. Reissner's membrane, on the other hand, showed intracellular vacuolization of the endolymphatic cells over the complete length of the cochlea after 12 or more days intoxication. Hearing loss was measured by electrocochleography with skin electrodes. The histologic findings were compared with the objective audiograms.
Animals↗
Effects of electrical stimulation and reversible lesions of the olivocerebellar pathway on Purkinje cell activity in the flocculus of the cat.
The activity of Purkinje cells (P-cells) in the flocculus of 8 lightly anesthetized cats and one alert cat was recorded for periods of up to several hours each. The resting simple spike (SS) rate in the anesthetized cats was 37 +/- 21 Hz (mean +/- standard deviation), similar to that observed in the alert cat. Complex spikes (CSs) were evoked by an electrode placed in the climbing fiber (CF) decussation or inferior olive (IO). For each P-cell, SS activity was suppressed completely at or above a cut-off frequency of evoked CSs; median cut-off rate was 5 Hz (range 1-10 Hz, 15 cells). Reversible lesions of the CF pathway were made by microinjection of 1-10 microliter of saturated lidocaine into the IO or CF decussation. This abolished spontaneous CS activity and produced two reversible effects on SS discharge: (1) an increase in mean SS rate of 98%, from 23 +/- 13 to 40 +/- 18 Hz (11 cells); and (2) a decrease of 50% in the variability of SS firing rate. Similar effects were observed in two P-cells whose CF axons were lesioned mechanically. These results show that electrical stimulation and reversible lesions in areas previously shown to alter the gain of the vestibulo-ocular reflex (VOR) also alter CF input to the flocculus, suggesting that the gain changes were caused by changes in CS rate. This study confirms and extends the observation that a reciprocal relationship exists between CS rate and SS background rate, and therefore further suggests that the changes in the gain of the VOR might be due to changes in SS background rate.
Animals↗
Atrophy of middle and short stereocilia on outer hair cells of guinea pig cochleas with experimentally induced hydrops.
Scanning electron microscopy was employed to investigate hair cell morphology at different stages in the development of experimentally induced hydrops in the guinea pig. A particular form of morpho-pathology, never before described, was identified as characteristic of hydropic cochleas. The pathology was characteristically identified as atrophy of the short and middle stereocilia on the outer hair cells while the inner hair cell stereocilia did not have such a pathology. The atrophy was restricted to the upper cochlear turns in remarkable correspondence with the low/middle frequency sensitivity loss and was detected only at the end of the period of fluctuating thresholds. These stereocilia perturbations appear therefore to be linked with the threshold fluctuations and represent the first evidence for a clear correlation between hair cell morphology and physiology in the experimental model of endolymphatic hydrops. Such a morphopathology might also be expected to occur in cochleas of Menière's patients but may have been overlooked in the past because of the discrete nature of the pathology.
Animals↗
A role of climbing fibers in regulation of flocculonodular lobe protein kinase C expression during vestibular compensation.
The behavioral recovery from unilateral labyrinthectomy (UL) in rats is accompanied by asymmetric expression of Protein kinase C (PKC) in parasagittal regions of the flocculonodular lobe within 6 h after UL, which resolves to the control, symmetric pattern within 24 h. These changes consist of a regionally selective increase in the number of PKC-immunopositive Purkinje cells contralateral to the lesion. This study tested the hypotheses (1) that climbing fiber innervation inhibits PKC expression and (2) that climbing fibers are essential for the observed changes in PKC expression within 6 h after UL. The patterns of flocculonodular lobe Purkinje cell PKCdelta expression were analyzed 6 h post-operatively in both UL and sham-operated that had been treated previously with 3-acetylpyridine to destroy the inferior olive. These data were compared with previous results from rats with an intact olive. The results suggest that at least two signals regulate the zonal distribution of Purkinje cell PKCdelta expression in the flocculonodular lobe during the early period of compensation from UL. Climbing fiber activation appears to reduce PKC expression, while extraolivary mechanisms appear to up-regulate PKC expression. It is suggested that the climbing fiber signals may act as a molecular 'filter' or 'automatic gain control' which adjusts the contributions of these kinases to synaptic plasticity within the context of the background activity of climbing fibers.
Animals↗
Role of primate flocculus during rapid behavioral modification of vestibuloocular reflex. I. Purkinje cell activity during visually guided horizontal smooth-pursuit eye movements and passive head rotation.
1. Extracellular recordings were obtained from 124 Purkinje cells (P-cells) in the flocculus of alert monkeys. P-cell simple spike-firing rate was analyzed quantitatively during various combinations of smooth-pursuit eye movement and passive head rotation. 2. During sinusoidal smooth eye movements, 80% of the P-cells displayed increased firing rate during ipsilateral and 20% during contralateral eye movement. Over the frequency range 0.3--1.4 Hz, firing-rate modulation was proportional to and in phase with maximum eye velocity. During the steady state of triangle-wave tracking, firing rate increased monotonically as a function of eye velocity. Since firing rate was uncorrelated with retinal-error velocity, one component of P-cell firing rate was related to eye velocity. 3. During the transient phase of triangle-wave tracking, when an instantaneous change in the direction of target movement caused a large retinal-error velocity, 40% of the P-cells were related only to eye velocity. Sixty percent of the P-cells displayed an overshoot or undershoot in firing rate, indicating a relationship to either retinal-error velocity or eye acceleration as well as to eye velocity. 4. During the vestibuloocular reflex (VOR), evoked by head rotation in the dark, P-cell firing rate was only weakly modulated. In contrast, when the monkey suppressed the VOR by fixating a target that rotated with him, P-cell rate was deeply modulated. Since the modulation was proportional to and in phase with maximum head velocity, another component of P-cell firing rate was related to head velocity. 5. Of 36 P-cells tested, 35 displayed firing-rate modulation during both suppression of the VOR and smooth-pursuit eye movement. P-cells that reached peak firing rate during ipsilateral head rotation also reached peak firing rate during ipsilateral smooth eye rotation. Average population sensitivitites to head velocity and eye velocity were equal. In three conditions in which eye and head velocity were elicited simultaneously, P-cell firing rate could be predicted by the linear, vector addition of the separate eye and head velocity components of firing rate. Therefore, the relatively weak modulation of P-cell firing rate during the VOR in the dark can be accounted for by the cancellation of equal but opposite head and eye velocity components. 6. The connections of flocculus P-cells to interneurons in the brain stem VOR pathways have been established in other mammals. In the context of those connections, P-cell firing patterns were appropriate to facilitate the eye movements the monkey was required to make. We conclude that the flocculus is important for sustaining any smooth eye movements that are different from those evoked by head rotation in the dark. The eye velocity component may represent an efference copy signal that sustains ongoing eye velocity during smooth pursuit.
Animals↗
Inner ear therapy for neural preservation.
A gradual loss of auditory neurons often occurs following sensorineural hearing loss. Since the cochlear implant must stimulate the remaining auditory neuron population, it would be beneficial to preserve as many auditory neurons as possible. Neurotrophic factors protect auditory neurons from degradation after sensorineural hearing loss in experimental animals, but have not yet been translated into the clinical setting. Current experimental and clinical techniques for drug delivery to the inner ear are examined in this review, covering the routes for drug delivery to the cochlea and the delivery systems used to introduce them. Duration of treatment, drug diffusion, effectiveness and safety are discussed with references to how they may be translated to the implementation of neurotrophic factor treatment for neural preservation.
Animals↗