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R Fettiplace

Publications and source records attributed to R Fettiplace.

At least 37 records · Page 2Linked to original sources
PubMed1995

Confocal imaging of calcium microdomains and calcium extrusion in turtle hair cells.

We have studied spatial Ca2+ distribution in hair cells filled with the low affinity fluorescent indicator Calcium Green 5N using real-time confocal microscopy and whole-cell recording. During depolarizations lasting several hundred milliseconds, Ca2+ fluorescence increased at a number of hotspots around the base of the cell but changed little near the hair bundle. The hotspots required influx of Ca2+ through voltage-dependent channels, and they expanded during the pulse from an initial diameter of < 1 micron. Strong Ca2+ buffers like BAPTA slowed their growth rate. On repolarization, the fluorescence decayed with two time constants: approximately 0.1 s, which may represent Ca2+ diffusion away from the entry sites, and 10 s, probably reflecting Ca2+ extrusion. Extrusion occurs mainly via a CaATPase that can be blocked by vanadate. We suggest the hotspots are microdomains of Ca2+ attaining a concentration of at least 85 microM near assemblies of synaptic release sites.

Animals↗
PubMed1995

The calcium-activated potassium channels of turtle hair cells.

A major factor determining the electrical resonant frequency of turtle cochlear hair cells is the time course of the Ca-activated K current (Art, J. J., and R. Fettiplace. 1987. Journal of Physiology. 385:207-242). We have examined the notion that this time course is dictated by the K channel kinetics by recording single Ca-activated K channels in inside-out patches from isolated cells. A hair cell's resonant frequency was estimated from its known correlation with the dimensions of the hair bundle. All cells possess BK channels with a similar unit conductance of approximately 320 pS but with different mean open times of 0.25-12 ms. The time constant of relaxation of the average single-channel current at -50 mV in 4 microM Ca varied between cells from 0.4 to 13 ms and was correlated with the hair bundle height. The magnitude and voltage dependence of the time constant agree with the expected behavior of the macroscopic K(Ca) current, whose speed may thus be limited by the channel kinetics. All BK channels had similar sensitivities to Ca which produced half-maximal activation for a concentration of approximately 2 microM at +50 mV and 12 microM at -50 mV. We estimate from the voltage dependence of the whole-cell K(Ca) current that the BK channels may be fully activated at -35 mV by a rise in intracellular Ca to 50 microM. BK channels were occasionally observed to switch between slow and fast gating modes which raises the possibility that the range of kinetics of BK channels observed in different hair cells reflects a common channel protein whose kinetics are regulated by an unidentified intracellular factor. Membrane patches also contained 30 pS SK channels which were approximately 5 times more Ca-sensitive than BK channels at -50 mV. The SK channels may underlie the inhibitory synaptic potential produced in hair cells by efferent stimulation.

Animals↗
PubMed1993

The functional morphology of stereociliary bundles on turtle cochlear hair cells.

The stereociliary bundles of hair cells from the basilar papilla of the red-eared turtle were examined with transmission and high resolution scanning electron microscopy to provide a description of their morphology, orientation and inter-ciliary connections for comparison with physiological observations. Bundles on hair cells in the basilar membrane region are of a uniform shape and orientation, but bundles on the apical and basal limbus are distinct in having elongated kinocilia more than twice the length of the tallest stereocilia. Bundles in the basilar membrane region show a roughly two-fold increase in height from 5 to 9 microns from base to apex. Electrical recordings from isolated hair cells indicate that the bundle height is inversely proportional to the cell's characteristic frequency. It is argued that the change in dimensions is insufficient to contribute significantly to the cochlea's frequency selectivity. The cytoplasm adjacent to the kinocilium is filled with microtubules and large vesicles, and there are coated pits in the apical membrane which, it is suggested, may be indicative of rapid turnover of the membrane in this region.

Animals↗
PubMed1993

The effects of low calcium on the voltage-dependent conductances involved in tuning of turtle hair cells.

1. The voltage-dependent conductances of turtle cochlear hair cells of known resonant frequency were characterized by tight-seal, whole-cell recording during superfusion with solutions containing normal (2.8 mM) and reduced (0.1-10 microM) Ca2+. 2. In 1 microM Ca2+, the current flowing through the voltage-dependent Ca2+ channels was increased roughly fivefold and had a reversal potential near 0 mV. This observation may be explained by the Ca2+ channels becoming non-selectively permeable to monovalent cations in low-Ca2+ solutions. Lowering the Ca2+ further to 0.1 microM produced little increase in the current. 3. The size of the non-selective current increased systematically with the resonant frequency of the hair cell over the range from 10 to 320 Hz. This suggests that hair cells tuned to higher frequencies contain more voltage-dependent Ca2+ channels. 4. There was a good correlation between the amplitudes of the non-selective current and the K+ current which underlies electrical tuning of these hair cells. The amplitude of the K+ current also increased systematically with resonant frequency. 5. In cells with resonant frequencies between 120 and 320 Hz, the K+ current was completely abolished in 1 microM Ca2+, consistent with prior evidence that this current flows through Ca2+ activated K+ channels. In a majority of cells tuned between 50 and 120 Hz, the K+ current was incompletely blocked in 1 microM Ca2+, but was eliminated in 0.1 microM Ca2+. In all hair cells the K+ current was abolished by 25 mM tetraethylammonium chloride. 6. In cells tuned to 10-20 Hz, the K+ current was not substantially diminished even in 0.1 microM Ca2+, which argues that it may not be Ca2+ activated. 7. In cells tuned to frequencies above 100 Hz, the K+ current could still be evoked by depolarization during superfusion with 10 microM Ca2+. However, its half-activation voltage was shifted to more depolarized levels and its maximum amplitude was systematically reduced with increasing resonant frequency. 8. These observations are consistent with the notion that in cells tuned to more than 50 Hz, there is a fixed ratio of the number of voltage-dependent Ca2+ channels to Ca(2+)-activated K+ channels, the numbers of each increasing in proportion to resonant frequency. The results also provide indirect evidence that the Ca(2+)-activated K+ channels in cells tuned to higher frequencies may be less sensitive to intracellular Ca2+.

Animals↗
PubMed1991

The actions of calcium on the mechano-electrical transducer current of turtle hair cells.

1. Mechano-electrical transducer currents evoked by deflections of the hair bundle were recorded in turtle isolated hair cells under whole-cell voltage clamp. The outcome of perfusing with solutions of reduced Ca2+ concentration was investigated. 2. The transducer current was roughly doubled by lowering the concentration of divalent cations from normal (2.2 mM-Mg2+, 2.8 mM-Ca2+) to 0 Mg2+, 0.5 mM-Ca2+. No significant effects on the current's kinetics or reversal potential, or on the current-displacement relationship, were noted. 3. If the Ca2+ concentration was lowered to 50 microM (with no Mg2+), there was about a threefold increase in the maximum current but other changes, including loss of adaptation and a decreased slope and negative shift in the current-displacement relationship, were also observed. As a result, more than half the peak transducer current became activated at the resting position of the hair bundle compared to about a tenth in the control solution. 4. The extra changes manifest during perfusion with 50 microM-Ca2+ had also been seen when the cell was held at positive potentials near the Ca2+ equilibrium potential. This supports the view that some consequences of reduced external Ca2+ stem from a decline in its intracellular concentration. 5. With 20 microM-Ca2+, a standing inward current developed and the cell became unresponsive to mechanical stimuli, which may be explained by the transducer channels being fully activated at the resting position of the bundle. 6. The results are interpreted in terms of a dual action of Ca2+: an external block of the transducer channel which reduces the maximum current, and an intracellular effect on the position and slope of the current-displacement relationship; the latter effect can be modelled by internal Ca2+ stabilizing one of the closed states of the channel. 7. During perfusion with 1 microM-Ca2+, the holding current transiently increased but then returned to near its control level. There was a concomitant irreversible loss of sensitivity to hair bundle displacements which we suggest is due to rupture of the mechanical linkages to the transducer channel. 8. Following treatment with 1 microM-Ca2+, single-channel currents with an amplitude of -9 pA at -85 mV were sometimes visible in the whole-cell recording. The probability of such channels being open could be modulated by small deflections of the hair bundle which indicates that they may be the mechano-electrical transducer channels or conductance about 100 pS. 9. Open- and closed-time distributions for the channel were fitted by single exponentials, the mean open time at rest being approximately 1 ms. The mean open time was increased and the mean closed time decreased for movements of the hair bundle towards the kinocilium.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1989

Activation and adaptation of transducer currents in turtle hair cells.

1. Transducer currents were recorded in turtle cochlear hair cells during mechanical stimulation of the hair bundle. The currents were measured under whole-cell voltage clamp in isolated cells that were firmly stuck to the floor of the recording chamber. 2. Stimuli were calibrated by projecting the image of the hair bundle onto a rapidly scanned 128 photodiode array. This technique showed that, while the cell body was immobilized, the tip of the bundle would follow faithfully the motion of an attached glass probe up to frequencies of more than 1 kHz. 3. The relationship between inward transducer current and bundle displacement was sigmoidal. Maximum currents of 200-400 pA were observed for deflections of the tip of the bundle of 0.5 microns, equivalent to rotating the bundle by about 5 deg. 4. In response to a step deflection of the bundle, the current developed with a time constant (about 0.4 ms for small stimuli) that decreased with the size of displacement. This suggests that the onset of the current was limited by the gating kinetics of the transduction channel. The onset time course was slowed about fourfold for a 20 degrees C drop in temperature. 5. For small maintained displacements, the current relaxed to about a quarter of the peak level with a time constant of 3-5 ms. This adaptation was associated with a shift of the current-displacement relationship in the direction of the stimulus. The rate and extent of adaptation were decreased by lowering external Ca2+. 6. Adaptation was strongly voltage sensitive, and was abolished at holding potentials positive to the reversal potential of the transducer current of about 0 mV. It was also diminished by loading cells with 10 mM of the Ca2+ chelator BAPTA. These observations suggest that adaptation may be partly controlled by influx of Ca2+ through the transducer channels. 7. Removal of adaptation produced asymmetric responses, with fast onsets but slow decays following return of the bundle to its resting position; the offset time course depended on both the magnitude and duration of the prior displacement. 8. In some experiments, hair bundles were deflected with a flexible glass fibre whose motion was monitored using a dual photodiode arrangement. Positive holding potentials abolished adaptation of the transducer currents, but had no influence on the time course of motion of the fibre. We have no evidence therefore that adaptation is caused by a mechanical reorganization within the bundle.

Adaptation, Physiological↗
PubMed1987

Variation of membrane properties in hair cells isolated from the turtle cochlea.

1. Hair cells were enzymatically isolated from identified regions of the turtle basilar papilla and studied with the patch-electrode technique. The experimental aim was to relate the resonance properties seen during current injection to the membrane currents measured in the same cell under whole-cell voltage clamp. 2. Solitary hair cells had resting potentials of about -50 mV, and produced a damped oscillation in membrane potential at the onset and termination of a small current step; the resonant frequency varied from 9 to 350 Hz between cells, and was correlated with the region of papilla from which a cell had been isolated. The inferred frequency map was consistent with the tonotopic arrangement described previously in the intact papilla. 3. Depolarizations from the resting potential under voltage clamp activated a large net outward current with a steep voltage dependence, and the steady-state current-voltage relationship was strongly rectified about the resting potential. Input resistances tended to be smaller in cells with higher resonant frequencies, although there was no concurrent variation in membrane area as inferred from the cell capacitance. 4. The kinetics of the outward current evoked by a small depolarizing step depended upon the resonant frequency, fo, of the hair cell, and were slower in low-frequency cells. On repolarization to the resting potential the current decayed exponentially with a time constant that changed from 150 ms in the lowest-frequency cell to less than 1 ms in the highest-frequency one. The time constant was approximately proportional to 1/f0(2). 5. Following repolarization to different membrane potentials, the tail current was found to reverse around -80 mV, indicating that the outward current was due mainly to K+. 6. The outward current was abolished by extracellular application of 25 mM-tetraethylammonium chloride (TEA), or on exchange of Cs+ for K+ in the intracellular medium filling the recording electrode, each experiment supporting the contention that K+ is the major current carrier. Such treatments also removed the oscillations in membrane potential evoked by imposed current steps. 7. Addition of TEA or intracellular perfusion with Cs+ also revealed a fast inward current with an ionic sensitivity consistent with its being carried by Ca2+. Like the K+ current, the Ca2+ current was activated by small depolarization from the resting potential, and over this voltage range it was about five to ten times smaller than the K+ current. Its activation was more rapid than the fastest outward currents in high-frequency cells.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗
PubMed1986

Electrical resonance and membrane currents in turtle cochlear hair cells.

The electrical and mechanical properties of single hair cells from the turtle's cochlea were examined to investigate the basis of their electrical resonance. Receptor potentials were measured with intracellular micropipettes in the isolated basilar papilla. At the onset and termination of a step displacement of the ciliary bundle the receptor potential showed a damped oscillation reflecting the frequency selectivity of the cell. Resonance frequencies increased systematically from apex to base of the cochlea. Similar oscillations could be elicited by a current step injected through the recording electrode. Solitary hair cells enzymatically isolated from the papilla were investigated with the tight-seal whole-cell recording method. Cells retained their properties in response to current steps and had resonance frequencies between 10 and 350 Hz. In voltage clamp such cells displayed a large outward K+ current and an inward Ca2+ current both activated by depolarization from the resting potential. The relaxation time constant of the K+ current was inversely correlated with the resonance frequency of the cell, varying from 150 ms in the lowest frequency cells to less than 1 ms in the highest ones. It is argued that variation in the kinetics of this current is the major factor responsible for the range of resonance frequencies. In preparations of the isolated papilla a flexible glass fibre, attached to the tip of a ciliary bundle, was used to deliver constant force steps to the bundle and to monitor its displacement. Receptor potentials were simultaneously recorded. At the beginning and end of a force step towards the kinocilium, the bundle vibrated at a frequency which coincided with the electrical resonance frequency of the cell.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗
PubMed1985

Efferent modulation of hair cell tuning in the cochlea of the turtle.

Intracellular recordings were made from turtle cochlear hair cells in order to study the changes in their tuning properties resulting from electrical stimulation of the efferent axons. Efferent stimulation caused a reduction in the amplitude of the receptor potential at the hair cell's most sensitive or characteristic frequency, an increased amplitude at frequencies more than an octave below the characteristic frequency, and no change at very high frequencies. These differential effects resulted in a broadening of each cell's tuning curve, which, during maximal efferent stimulation degenerated from a sharply tuned resonance to a critically damped low-pass filter. Efferent alterations in tuning were also inferred from the oscillations in membrane potential produced by acoustic clicks or extrinsic currents. The quality factor (Q) of tuning, derived from the decay of the oscillations, was progressively reduced with synaptic hyperpolarizations up to about 5 mV in amplitude. A consequence of efferent action was that the wave forms of transient pressure changes were more faithfully encoded as changes in hair cell membrane potential. Hyperpolarization of a hair cell by steady current injection resulted in a lowering of its characteristic frequency and quality factor, and an increase in steady-state resistance. By comparison, for a given reduction in quality factor, efferent stimulation was associated with a smaller change in characteristic frequency. This difference is expected if the resonance is also damped by the shunting action of the synaptic conductance. Perfusion with perilymphs containing 0.5-15 mM of the potassium channel blocker, tetraethylammonium bromide (TEA) reduced the hair cell's frequency selectivity, whether assayed acoustically or with extrinsic currents. Lower TEA concentrations abolished the efferent inhibitory post-synaptic potential with only a minor change in tuning. TEA produced other effects different from efferent stimulation including (i) a lowering of the characteristic frequency, and (ii) a highly asymmetric receptor potential. These observations suggest that the efferents do not simply block membrane conductances associated with tuning. We conclude that the efferent modification of the shape of the tuning curve may be a composite result of the synaptic conductance and the hyperpolarization of the hair cell membrane.

Action Potentials↗
PubMed1985

The mechanical properties of ciliary bundles of turtle cochlear hair cells.

The mechanical behaviour of the ciliary bundles of hair cells in the turtle cochlea was examined by deflecting them with flexible glass fibres of known compliance during simultaneous intracellular recording of the cell's membrane potential. Bundle motion was monitored through the attached fibre partially occluding a light beam incident on a photodiode array. The change in photocurrent was assumed to be proportional to bundle displacement. For deflexions of 1-100 nm towards the kinocilium, the stiffness of the ciliary bundles was estimated as about 6 X 10(-4) N/m, with the fibre attached to the top of the bundle. When the fibre was placed at different positions up the bundle, the stiffness decreased approximately as the inverse square of the distance from the ciliary base. This suggests that the bundles rotate about an axis close to the apical pole of the cell and have a rotational stiffness of about 2 X 10(-14) N. m/rad. Step displacements of the fixed end of the flexible fibre caused the hair cell's membrane potential to execute damped oscillations; the frequency of the oscillations in different cells ranged from 20 to 320 Hz. Displacements towards the kinocilium always produced membrane depolarization. The amplitude of the initial oscillation increased with displacements up to 100 nm and then saturated. For small displacements of a few nanometres, the hair cell's mechanoelectrical sensitivity was estimated as about 0.2 mV/nm. Force steps delivered by the flexible fibre caused the bundle position to undergo damped oscillations in synchrony with the receptor potential. The mechanical oscillations could be abolished with large depolarizing currents that attenuated the receptor potential. When placed against a bundle, a fibre's spontaneous motion increased and became quasi-sinusoidal with an amplitude several times that expected from the compliance of the system. It is suggested that the hair bundle drives the fibre. We conclude that turtle cochlear hair cells contain an active force generating mechanism.

Action Potentials↗
PubMed1984

Efferent desensitization of auditory nerve fibre responses in the cochlea of the turtle Pseudemys scripta elegans.

Extracellular recordings were made from single auditory afferents in the isolated half-head of the turtle, and changes in their acoustic sensitivity were examined following electrical stimulation of the efferent fibres to the basilar papilla. Short trains of efferent shocks caused a prolonged elevation of the pure tone thresholds of the auditory afferents and an abolition of their spontaneous activity. These changes could be demonstrated in a majority of recordings and without antidromic firing of the afferent. The amount of desensitization increased steeply with shock number and a train of ten closely spaced shocks could elevate the threshold at the most sensitive or characteristic frequency by four orders of magnitude. Desensitization also occurred with single efferent shocks at repetition frequencies exceeding 25/s. Discharge rate versus sound pressure functions were constructed for a number of afferents. The maximum slope of the functions, and the saturated firing rates were both reduced by efferent stimulation; there was also an over-all shift of the rate-intensity function to higher stimulus levels. Such effects would enable the afferent to signal a wider range of sound pressures. Efferent stimulation caused a broadening of the afferent frequency-threshold curves by removal of the narrowly-tuned region around the characteristic frequency. We suggest that the loss in tuning and concomitant improvement in temporal resolution may be a functionally important consequence of efferent action.

Acoustic Stimulation↗
PubMed1984

Synaptic hyperpolarization and inhibition of turtle cochlear hair cells.

Intracellular recordings were made from turtle cochlear hair cells in order to examine the properties of the post-synaptic potentials evoked by electrical stimulation of the efferent axons. Single shocks to the efferents generated a hair cell membrane hyperpolarization with an average amplitude generally less than 1 mV and lasting for about 100 ms. With short trains of shocks, the size of the post-synaptic potential grew markedly to a maximum of 20-30 mV. The interaction between pairs of shocks separated by a varying interval was studied. For an interval of 4 ms, the response to the second shock was increased on average by a factor of 3 and the conditioning effect of the first shock decayed with a time constant of about 100 ms. We suggest the augmentation in response to trains of shocks may be partly due to facilitation of efferent transmitter release. The efferent post-synaptic potentials could be reversibly abolished by perfusion with perilymphs containing 3 microM-curare or atropine, and infusion of acetylcholine gave a transient membrane hyperpolarization. These observations are consistent with efferent action being mediated via a cholinergic synapse onto the hair cells. The post-synaptic potentials could be reversed in polarity by injection of hyperpolarizing currents through the recording electrode. The reversal potential was estimated as about -80 mV, 30 mV negative to the resting potential. Near reversal, a small brief depolarization was evident and may constitute a minor component of the synaptic response. The value of the reversal potential was unaffected by substitution of the perilymphatic chloride, but was altered in a predictable manner by changes in extracellular potassium concentration indicating that the post-synaptic potentials arise mainly by an increase in the permeability of the hair cell membrane to potassium ions. Throughout the post-synaptic hyperpolarization there was a reduction in the sensitivity of the hair cell to tones at its characteristic frequency. The desensitization, maximal for low sound pressures, varied in different cells from a factor of 1.6 to 28. At the peak of the largest synaptic potentials, the receptor potential remained negative to the resting potential with all but the loudest characteristic frequency tone s. We suggest that there are two factors in efferent inhibition; one a r duction in the receptor potential at the hair cell's characteristic frequency and the other a hyperpolarization of its membrane potential which should reduce the release of excitatory transmitter onto the afferent terminals.

Acetylcholine↗
PubMed1983

Auditory nerve responses to imposed displacements of the turtle basilar membrane.

Impulse activity of single auditory nerve fibres was recorded in the isolated half-head of the turtle in response to displacements of a piezoelectric probe placed on the basilar membrane. The temporal pattern of firing in response to sinusoidal displacements of amplitude 0.1-1.0 nm r.m.s. at a fibre's characteristic frequency could be matched to that generated by low-level tonal stimuli delivered to the tympanum. Frequency-threshold curves for acoustic and mechanical stimuli had similar shapes and differed only at frequencies above 500 Hz where the middle ear should filter acoustic but not direct mechanical stimuli. Step displacements of the basilar membrane gave a transient periodic discharge which resembled the responses to acoustic clicks. Most fibres initially increased their firing rate for rarefaction clicks and displacements towards the scala tympani.

Acoustic Stimulation↗
PubMed1982

Efferent regulation of hair cells in the turtle cochlea.

Intracellular recordings were made from hair cells in the isolated cochlea of the turtle to characterize the inhibition achieved by the cochlea's efferent innervation. A short train of shocks delivered to the efferent axons produced in the hair cells slow hyperpolarizing synaptic potentials which could be reversed by shifting the membrane potential more negative than about -80 mV. Throughout the efferent hyperpolarization, there was a reduction of up to 25-fold in the amplitude of the receptor potential for tones presented at the hair cell's characteristic frequency. Efferent stimulation also was shown to degrade the cell's tuning properties. It is argued that the combined effects of the hyperpolarization and the loss in hair cell sensitivity could account for a threshold elevation of at least 70 dB in the auditory nerve fibres.

Action Potentials↗
PubMed1981

An electrical tuning mechanism in turtle cochlear hair cells.

1. Intracellular recordings were made from single cochlear hair cells in the isolated half-head of the turtle. The electrical responses of the cells were recorded under two conditions: (a) when the ear was stimulated with low-intensity tones of different frequencies and (b) when current steps were injected through the intracellular electrode. The aim of the experiments was to evaluate the extent to which the cochlea's frequency selectivity could be accounted for by the electrical properties of the hair cells.2. At low levels of acoustic stimulation, the amplitude of the hair cell's receptor potential was proportional to sound pressure. The linear tuning curve, which is defined as the sensitivity of the cell as a function of frequency when the cell is operating in its linear range, was measured for a number of hair cells with characteristic frequencies from 86 Hz to 425 Hz.3. A rectangular current passed into a hair cell elicited a membrane potential change consisting of a damped oscillation superimposed on a step. Small currents produced symmetrical oscillations at the beginning and end of the pulse. Larger currents increased the initial ringing frequency if depolarizing and decreased it if hyperpolarizing.4. For small currents the frequency of the oscillations and the quality factor (Q) of the electrical resonance derived from the decay of the oscillations were close to the characteristic frequency and Q of the hair-cell linear tuning curve obtained from sound presentations.5. The hair cell's membrane potential change to small-current pulses or low-intensity tone bursts could be largely described by representing the hair cell as a simple electrical resonator consisting of an inductance, resistor and capacitor.6. When step displacements of 29-250 nm were applied to a micropipette, placed just outside a hair cell in the basilar papilla, an initial periodic firing of impulses could be recorded from single fibres in the auditory nerve. Currents of up to 1 nA, injected through the same micropipette, failed to produce any change in the auditory nerve discharge. The experiment demonstrates that current injection does not produce gross movements of the electrode tip.7. The contribution of the electrical resonance to hair-cell tuning was assessed by dividing the linear tuning curve by the cell's impedance as a function of frequency. The procedure assumes that the electrical resonance is independent of other filtering stages, and on this assumption the resonance can account for the tip of the acoustical tuning curve.8. The residual filter produced by the division was broad; it exhibited a high-frequency roll-off with a corner frequency at 500-600 Hz, similar in all cells, and a low-frequency roll-off, with a corner frequency from 30 to 350 Hz which varied from cell to cell but was uncorrelated with the characteristic frequency of the cell.9. The phase of the receptor potential relative to the sound pressure at the tympanum was measured in ten cells. For low intensities the phase characteristic was independent of the sound pressure. At low frequencies the receptor potential led the sound by 270-360 degrees , and in the region of the characteristic frequency there was an abrupt phase lag of 90-180 degrees ; the abruptness of the phase change depended upon the Q of the cell.10. The calculated phase shift of the electrical resonator as a function of frequency was subtracted from the phase characteristic of the receptor potential. The subtraction removed the sharp phase transition around the characteristic frequency, and in this frequency region the residual phase after subtraction was approximately constant at +180 degrees . This is consistent with the idea that the hair cells depolarize in response to displacements of the basilar membrane towards the scala vestibuli. The high-frequency region of the residual phase characteristic was similar in all cells.11. It is concluded that each hair cell contains its own electrical resonance mechanism which accounts for most of the frequency selectivity of the receptor potential. All cells also show evidence of a broad band-pass filter, the high frequency portion of which may be produced by the action of the middle ear.

Animals↗
PubMed1981

Non-linearities in the responses of turtle hair cells.

1. Intracellular recordings were made from single cochlear hair cells in the isolated half-head of the turtle. Receptor potentials were recorded while the ear was stimulated with high-intensity tones in order to examine the cochlear non-linearities which shape the hair cell responses.2. The size of a hair cell's voltage response to a tone burst was reduced, abolished and then reversed by steady depolarizing currents of increasing strength. The average current needed to produce reversal was about 0.3 nA, the reversal potential being close to zero with respect to the scala tympani.3. Short current pulses injected on the peaks and dips of the receptor potential showed that the membrane resistance and time constant were decreased on the depolarizing phase of the receptor potential. These changes were not due to non-linearity in the hair cell's current-voltage curve in the absence of acoustic stimulation. The results are consistent with the idea that the transducer causes the cell to depolarize by increasing the membrane conductance to ions with an equilibrium potential close to zero.4. Saturated receptor potentials from poorly tuned cells exhibited a pronounced asymmetry, with the maximum depolarizing excursion being several times the maximum hyperpolarizing excursion. This asymmetry was not seen in sharply tuned cells. It is proposed that the asymmetry is present in the transducer conductance change and in sharply tuned cells is reduced in the receptor potential by subsequent filtering.5. For high sound pressures which produced close to a saturated response, the hair cell voltage wave form displayed a number of non-linear features dependent upon the frequency of stimulation relative to the characteristic frequency (c.f.). The most prominent feature occurred at very low frequencies where the potential exhibited damped oscillations on the depolarizations and hyperpolarizations; these ;ringing frequencies' lay above and below the c.f. of the cell respectively.6. The ;ringing frequencies' varied with the c.f. of the cell but for a given cell were largely independent of the frequency of stimulation. The ;ringing frequencies' could be changed by injecting steady currents into the cell during acoustic stimulation; depolarizing currents increased the ringing frequencies and hyperpolarizing currents decreased the frequencies.7. The hair cell's response to a continuous test tone at the c.f. of the cell could be suppressed by simultaneous addition of a second tone whose sound presure was comparable to, or greater than, the test tone. The degree of suppression varied with the intensity and frequency of the second tone, and was maximal close to the c.f. of the cell. The sound pressure required to produce a constant suppression as a function of frequency was sharply tuned, and the tuning of the suppression showed similarities to the frequency selectivity of two-tone suppression described in the auditory nerve.8. An attempt was made to reconstruct the main features of the receptor potential at high intensities.

Animals↗