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P M Sellick

Publications and source records attributed to P M Sellick.

At least 19 recordsLinked to original sources

The continuing search for outer hair cell afferents in the guinea pig spiral ganglion.

Antidromic stimulation of the stump of the VIIIth nerve was combined with microelectrode recording in the spiral ganglion of the guinea pig cochlea in an attempt to identify a sub-population of neurons with long-latency antidromic action potentials that might correspond to the thin unmyelinated afferent neurons emanating from the outer hair cells. The techniques used were similar but not identical to those employed in an earlier study by Brown (1994). By far the largest population of cells contacted had short antidromic latencies (0.58+/-0.12 ms, 76 units) and also responded to acoustic stimulation. These were assumed to be type I afferents emanating from inner hair cells. Eight cells had antidromic latencies larger than 1 ms, all but one of which had a zero spontaneous rate. All eight of these longer-latency cells were unresponsive to acoustic stimulation despite the fact that short-latency neurons in the same cochleas showed robust responses to sound before and after they were contacted. Four of these longer-latency cells had their antidromic thresholds accurately measured and two had significantly higher thresholds to electrical stimulation (0.1 ms duration) than type I cells in the same animal while two had similar electrical thresholds. Attempts to trace the eight long-latency neurons to the outer hair cells using intracellular injection of horseradish peroxidase were unsuccessful. On the basis of present evidence, we cannot conclude definitively that the long-latency neurons found in the spiral ganglion belong to the outer hair cell afferent population.

Animals

The modulation of the sensitivity of the mammalian cochlea by low frequency tones. I. Primary afferent activity.

The neural activity of single cells within the spiral ganglion of the first turn of the guinea pig cochlea has been recorded. Period histograms of the spike activity have been collected. The histograms were synchronised with a loud low frequency tone (40 Hz) presented at a constant intensity below the threshold of the cells to phase locking. The activity evoked by a continuous high frequency tone within the tip of the frequency threshold curve was modulated by the presence of a subthreshold low frequency tone. Suppression of this evoked activity was observed for low frequency displacements of the cochlear partition towards scala tympani, and potentiation was observed for moderate displacements towards scala vestibuli. At higher intensities of the low frequency tone suppression is also observed for peak displacements of the partition towards scala vestibuli. The changes to these histograms with increases in the intensity of the high frequency tone are described. Hysteresis, flattening of the histograms at high intensities and nonmonotonic growth of firing rate at certain phases of the low frequency tone were observed. These changes are believed to be due to fast adaptation of the neural response rather than similar changes to the d.c. receptor potential within the inner hair cells that the neurons innervate.

Acoustic Stimulation

The modulation of the sensitivity of the mammalian cochlea by low frequency tones. III. Basilar membrane motion.

The simultaneous presentation of an intense low frequency tone and a moderate intensity high frequency tone produced modulation of the high frequency motion of the cochlear partition in the first turn of the guinea pig cochlea. This modulation was in synchrony with the displacement caused by the low frequency tone. Maximum reduction in mechanical sensitivity was observed for peak displacements towards scala tympani, whereas a less pronounced reduction was observed for peak displacements towards scala vestibuli. Modulation was reduced or absent for high frequency tones more than an octave below the characteristic frequency of the mechanical tuning curve and was absent post mortem.

Acoustic Stimulation

Comparison between the tuning properties of inner hair cells and basilar membrane motion.

Measurements were made of inner hair cell receptor potentials and basilar membrane motion in the 17-21 kHz region of the guinea pig cochlea. The latter were made using the Mossbauer technique. Isoamplitude curves at 0.9 mV d.c. receptor potential were compared with isovelocity curves at 0.04 mm/s and the corresponding basilar membrane displacement at CF. The Mossbauer source (20 X 60 or 60 X 85 microns) was placed either in the middle of the basilar membrane or on the extreme modiolar edge. These two source positions yielded broad and narrow mechanical tuning curves, respectively. The latter approximated the receptor potential curves most closely but deviated by 10-15 dB on the low frequency side of the tuning curve tip.

Action Potentials

The influence of Mossbauer source size and position on phase and amplitude measurements of the guinea pig basilar membrane.

Phase and amplitude measurements were made from the incus and basilar membrane in guinea pig using the Mossbauer technique. The basilar membrane/incus ratio had a maximum of about 60 dB and a phase accumulation of between 9 and 12 radians to CF. Two source sizes were used (60 X 85 and 20 X 60 microns) and the source was placed either on the modiolar edge of the basilar membrane or in the middle. Notches in plots of the basilar membrane/incus ratio occur at stimulus frequencies that appear to be associated with source size rather than position, suggesting that artefacts could be produced by the presence of the Mossbauer source.

Animals

The alteration of the low frequency response of primary auditory afferents by cochlear trauma.

The response phase of primary auditory afferents in the first turn of the guinea pig cochlea to low frequency sinusoidal stimuli is altered by trauma to the cochlea. Loud sounds of sufficient intensity to produce temporary shifts in the thresholds of these cells at their characteristic frequency produce a reversal of the neural response from increased firing for basilar membrane displacements towards scala tympani before trauma to an increased firing towards scala vestibuli following trauma. These changes can be reversible.

Acoustic Stimulation

Low-frequency characteristics of intracellularly recorded receptor potentials in guinea-pig cochlear hair cells.

Intracellular receptor potentials were recorded from inner and outer hair cells in response to low-frequency tones, from the basal, high-frequency region of the guinea-pig cochlea. The receptor potentials recorded from inner hair cells are asymmetrical about the resting membrane potential with the depolarizing phase, which corresponds to rarefaction in sound pressure, exceeding the phase of hyperpolarization by a factor of about 3. It was found that the relationship between the peak-to-peak voltage responses and sound pressure level could be described by rectangular hyperbolae. When the frequency of the sound stimulus was progressively increased from 100 Hz to 4 kHz, the 'periodic' (a.c.) component of the receptor potential was attenuated with respect to the 'continuous' (d.c.) component. The characteristics of the inner hair cells could be described by two stages of low-pass filtering, with one of the filters having the same corner frequency as the electrical time constants which varied in different cells between 178 and 840 Hz. Receptor potentials recorded intracellularly from two morphologically identified outer hair cells were symmetrical about the resting membrane potential (about -65-70 mV) and had a maximal amplitude of only 5 mV at frequencies and intensities which yield 20-30 mV voltage responses from inner hair cells. No d.c. component receptor potentials were recorded in response to high-frequency tones. Phase and amplitude measurements were made from receptor potentials from inner hair cells, and from 'cochlear microphonic potentials' which were recorded from the organ of Corti and scala tympani. The phase of depolarization in both potentials was associated with displacement of the basilar membrane towards the scala vestibuli. The phase of the intracellular receptor potentials leads the cochlear microphonic by about 90 degrees and the sound pressure by about 180 degrees at frequencies below 100 Hz. Above this frequency the phase lead progressively declines and at higher frequencies becomes a phase lag. These phase relationships indicate that inner hair cells respond to the velocity of the basilar membrane at frequencies below 200-600 Hz, and to its displacement above this, and that the voltage responses of the inner hair cells are limited by their membrane time constants. It is suggested that outer hair cells respond to basilar membrane displacement throughout their frequency range. It is shown that, with respect to frequency, the different growth rates of the cochlear microphonic potentials and inner hair cell receptor potentials, and the dominance of cochlear microphonic potentials in the organ of Corti, result in an effective electrical interaction between inner hair cells and cochlear microphonic potentials.

Action Potentials

A comparison between basilar membrane and inner hair cell receptor potential input-output functions in the guinea pig cochlea.

Intracellular recordings were made from inner hair cells and basilar membrane motion was measured at a similar place, but in different preparations, in the first turn of the guinea pig cochlea. Potential recordings were made using glass microelectrodes and mechanical measurements were made using the Mössbauer technique. Intensity functions of DC receptor potential and basilar membrane velocity in animals with good and poor thresholds are presented. In animals with good thresholds, stimuli at and above the characteristic frequency produce similarly compressive input-output functions for both inner hair cell receptor potentials and basilar membrane motion. However, for frequencies lower than the characteristic frequency, receptor potential input-output functions obtained from animals in good and poor condition show saturation at high stimulus intensities at which basilar membrane motion is linear. This discrepancy is believed to be due to a nonlinear inner hair cell transduction mechanism. We propose that nonlinearity observed in receptor potential input-output functions is a consequence of the simple cascading of a frequency-dependent nonlinear mechanical input and a frequency-independent nonlinear transduction process.

Action Potentials

Modulation of responses of spiral ganglion cells in the guinea pig cochlea by low frequency sound.

Period histograms were generated from single unit data obtained from the spiral ganglion of the first turn of the guinea pig cochlea in response to continuous tones between 40 and 500 Hz. With the lowest intensities used, spontaneous activity was suppressed during basilar membrane displacement (inferred from cochlear microphonic phase) towards scala vestibula and activity was enhanced during displacement towards scala tympani. At higher intensities the response changed to excitation during maximal basilar membrane velocity towards scala vestibuli. These patterns were delayed by about 0.5 ms producing large phase delays at the higher frequencies. We postulate that the displacement response is produced by cochlear microphonic originating from the outer hair cells acting on the inner hair cell membrane. In contrast, the velocity response is produced by the inner hair cell receptor potential. The effect of a 40 Hz tone on activity evoked by tones above, at, and below the characteristic frequency was investigated by generating period histograms synchronous with the 40 Hz tone. We found that activity evoked by tones around the characteristic frequency of the cell was suppressed during displacement of the basilar membrane towards scala tympani and enhanced in the opposite direction at 40 Hz intensities that had no effect on spontaneous activity. Further increase in the 40 Hz intensity produced suppression during scala vestibuli displacement with activity remaining only during the zero crossings. Still further increase produces the 40 Hz tone alone response. Activity evoked by tones in the low frequency 'tails' of the frequency threshold curve was not similarly modulated. This phenomenon is though to be related to basilar membrane nonlinearity for frequencies close to the cut-off. Investigation of the effect of a 40 Hz tone on the threshold of the compound action potential confirmed data obtained from single units.

Animals

Measurement of basilar membrane motion in the guinea pig using the Mössbauer technique.

Basilar membrane motion was measured at the 16-19 kHz place of the guinea pig cochlea using the Mössbauer technique. The threshold of the gross cochlear action potential (CAP) evoked by pure-tone bursts was used as an indication of neural threshold. CAP threshold deteriorated progressively after the cochlea was opened and the Mössbauer source placed on the basilar membrane. A close relationship was found between the amplitude of basilar membrane motion at the source place frequency and CAP threshold. Basilar membrane velocity at CAP threshold SPL was about 0.04 mm/s over a 60-dB range of CAP threshold. Intensity functions for basilar membrane motion were linear for frequencies more than an octave below the source place frequency but demonstrated progressive saturation for frequencies greater than an octave below the CF. This nonlinear behavior was eliminated as the CAP threshold became less sensitive and was absent post mortem. Isovelocity curves at the 0.04 mm/s criterion were remarkably similar to frequency threshold curves from primary afferent fibers innervating a similar place on the basilar membrane. The isovelocity curve was a better fit than the isoamplitude curve suggesting that inner hair cells respond to basilar membrane velocity. As the CAP threshold deteriorated, the isovelocit curves lost sensitivity around the best frequency, whereas sensitivity to frequencies below 10 kHz remained constant even after the animal was killed. We suggested that most of the frequency response and nonlinear behavior of inner hair cells and afferent fibers may be found in basilar motion.

Animals

The responses of inner hair cells to basilar membrane velocity during low frequency auditory stimulation in the guinea pig cochlea.

Intracellular receptor potentials were recorded from inner hair cells in the basal turn of the guinea pig cochlea in response to low frequency tones. These were compared with the cochlear microphonic (CM) recorded from the scala tympani and sound pressure at the tympanic membrance. The CM is symmetrical and behaves as if it responds to basilar membrane displacement. The depolarizing phase of the inner hair cell receptor potential exceeds the hyperpolarizing phase with a ratio of about 3:1 in response to sinusoidal stimulation. Below 100 Hz inner hair cell receptor potentials phase lead sound pressure by 180 degrees and their amplitudes increase at a rate of 12 dB/octave. Above 200 Hz their receptor potentials are in phase with CM. The capacitative impedances of the hair cells delay and attenuate the intracellular receptor potentials at frequencies above 178-873 Hz. We conclude that CM is dominated by the responses of outer hair cells, and that a frequencies below 100-200 Hz inner hair cells respond to basilar membrance velocity. Above this they respond to basilar membrane displacement.

Acoustic Stimulation

Intracellular studies of hair cells in the mammalian cochlea.

1. Intracellular recordings were made from inner hair cells in the first turn of the guinea-pig cochlea, the recording sites being confirmed by the injection of Procion yellow dye and subsequent histology. 2. The receptor potential, in response to a pure tone burst, consisted of an AC response which followed the wave form of the stimulus and was analogous to the extracellularly recorded cochlear microphonic and a depolarizating DC response which followed the envelope of the tone burst and was analogous to the extracellularly recorded summating potential. 3. The DC response was broadly tuned at high sound pressure having a maximal amplitude of 27 mV at a sound pressure level of ca. 100 db; however the bandwidth of the response was reduced at lower sound pressure level. Isoamplitude curves for the DC response were indistinguishable from the threshold curves for auditory nerve fibres. 4. The AC response was tuned in a similar fashion to the DC response except that it was attenuated at 6-9 db/octave with respect to the DC response. It is suggested that this difference was due to the effect of membrane capacitance and resistance on the AC response. In contrast the extracellularly recorded AC component was not subject to this attenuation. 5. The total resistance and capacitance in three cells were found to be 46-61 Momega and 7.8-15.8 muF respectively. 6. Intracellular resistance changes were measured during sound stimulation, the resistance change being proportional to the DC receptor potential, indicating constant current flow through the hair cell. The current varied between 0.37 and 0.81 nA between cells. The time constant for seven cells was found to lie between 0.31 and 0.76 msec. 7. A map of the basilar membrane showing position of hair cells against characteristic frequency corresponded to the cut-off frequencies of the basilar membrane mechanical measurements and the innervation sites of spiral ganglion cells.

Animals

Measurement of potassium and chloride ion concentrations in the cupulae of the lateral lines of Xenopus laevis.

1. Potential measurements were made with double barrel ion selective electrodes from the cupulae of lateral line organs in the aquatic toad Xenopus laevis. 2. A positive endocupular potential (ECP) of 15-50 mV was recorded within the cupula, immediately above the hair cells. 3. Increases in the Cl- and K+ potentials were recorded when the ion selective electrodes touched the cupula. Cupular Cl- and K+ varied between 35 and 70 mM and 24 and 100 mM respectively. This variation existed between, rather than within, different animals. 4. Subcutaneous injections of 0-4 ml. 2 mM ouabain greatly reduced the ECP and cupular K+, whereas 0-4 ml. of Xenopus Ringer had no effect. 5. Changing the bath Cl- a hundredfold had no effect on the ECP. It was concluded that the ECP was produced by an electrogenic K+ pump which maintained high K+ levels within the cupula.

Animals