Frequency Threshold Curves Research Articles

Nonlinear phenomena as observed in the ear canal and at the auditory nerve

We report here several measures of nonlinear effects in the mammalian ear made in the external auditory meatus and in single neurons of the auditory nerve. We have measured the 2f1-f2 and the f2-f1 distortion products and we have found that the neural distortion product threshold curve for 2f1-f2 mirrors the low-frequency side of the frequency threshold curve, when the neural distortion product threshold curve of 2f1-f2 is plotted versus log(f2/f1) its slope is about 50 dB/oct and its intercept is 10-20 dB above the frequency threshold at the characteristic frequency CF, substantial 2f1-f2 distortion was seen in all animals studied while the f2-f1 distortion product was only rarely found at substantial levels, and the distortion product pressure observed in the ear canal was at a level equal to that detected at threshold by the neural units under study. We have also made measurements of two-tone rate suppression thresholds using two new and consistent threshold paradigms. We find that for high and intermediate characteristic frequency neural units the suppression threshold is independent of frequency and at a level of about 70 dB SPL, the suppression above CF is much less than below CF, and the tip of the frequency tuning curve can be suppressed by up to 40 dB by a low-frequency suppressor.

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.

Frequency threshold curves and simultaneous masking functions in high-threshold, broadly-tuned, fibres of the guinea pig auditory nerve.

Tuning curves for simultaneous masking were measured electrophysiologically, in single fibres of the auditory nerve. The recordings were made in guinea pigs with cochlear hearing losses. The masking paradigm was an analogy of that used in the determination of psychophysical tuning curves in man. Below the probe frequency, the slope of the masking function was similar to that of the frequency-threshold curve. Here, changes in the slope of the frequency-threshold curve with hearing loss were closely mirrored by changes in the slope of the masking function. Above the probe frequency, however, the masking function for low-threshold fibres had a shallower slope than the frequency-threshold curve. When the high-frequency slope of the frequency-threshold curve became shallower with hearing loss, the changes were not mirrored in the masking function, until the threshold was raised to 70-80 dB SPL. The results are discussed in terms of the influence of cochlear nonlinearity on frequency resolution, and the vulnerability of the nonlinearity.

Frequency threshold curves and simultaneous masking functions in single fibres of the guinea pig auditory nerve

Tuning curves for simultaneous masking were measured electrophysiologically, in single fibres of the guinea pig auditory nerve. The masking paradigm used was an analogy of that used in the determination of psychophysical tuning curves in man. The resulting tuning curves ran nearly parallel to the corresponding neural frequency threshold curve, over all except the high-frequency portion of the tuning curve. There, the masking functions had a shallower slope than the excitatory frequency threshold curve. The frequency at which the slope became shallower had a close and consistent dependence on fibre characteristic frequency, reaching a value of 1.2 times fibre characteristic frequency in high-frequency fibres. Analysis of firing rates during the threshold determination gave information about the mechanism of the masking, and the results were supported by theoretical analysis. The results give information useful for the interpretation of psychophysical tuning curves, determined by simultaneous masking, in man.

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.

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.

Functional significance of dendritic swelling after loud sounds in the guinea pig cochlea.

Exposure of the guinea pig cochlea to loud pure tones caused a dramatic swelling of afferent dendrites beneath the inner hair cell (IHC). This swelling occurred in a restricted region of the cochlea basalward of the exposure frequency location. For a 110 dB tone swelling was just detectable in 1 micron sections for a 18 3/4 min exposure and was clearly visible after a 22 1/2 min exposure. Swelling was reversible. Exposures which caused swelling produced a loss in sensitivity of the flat low frequency 'tail' of the frequency-threshold curves of single auditory neurons whose most sensitive frequency was a 1/2 octave higher than the exposure frequency. The findings are consistent with the notion that dendritic swelling causes a non-selective decrease in sensitivity to all frequencies of sound.

Electrophysiological characteristics of representation of auditory and somatosensory systems in the turtle midbrain

Experiments with unanesthetized tubocurarine-immobilized turtles show that the auditory representation is localized in the medio-dorsal part of tegmentum, torus semicircularis, which contains monomodal (auditory) and bimodal (auditory and somatosensory) units. The somatosensory system is wider represented, overlapping the auditory system in the medial parts of tegmentum. The focus of auditory representation is localized in lateral parts of the dorsal tegmentum (n. intercollicularis) where monomodal somatic units were recorded. The predominance of contralateral somatic projections is revealed. Frequency-threshold curves based on threshold of evoked potentials were V-shaped but flattened. Range of perceived frequencies was equal to 40-6000 Hz, range of optimal frequencies was 100-400 Hz. Auditory mesencephalic cells may be attributed to three types of discharge patterns: phasic, sustained and burst firing. Tuning to a single optimal frequency was typical of phasic cells but most of cells with sustained firing were sensitive to two or even three frequencies.

A mathematical model of the Pacinian corpuscle.

This article describes a mathematical model of the Pacinian corpuscle based on the analysis of the available experimental data and on previous theoretical research. The model includes the main anatomofunctional constituents of the corpuscle: the capsula and the mechano-to-neural transduction; its structure accounts for the formation of the receptor potential and of the spikes on the nerve terminal. Comparison of the theoretical predictions with the experimental results, in response to different types of stimuli provides a substantial validation of the model and an explanation for the basic aspects of the transduction, in particular for: a) the receptor potential time course for isolated stimuli; b) the frequency response, in terms of receptor potential; c) the frequency threshold curve for the spikes; d) the firing rate, I.S.I. and P.S.T. histograms and the synchronization coefficient, in response to sustained sinusoidal inputs. Possible lines for future experimental research are suggested from the model predictions.

The effects of intracochlear and systemic furosemide on the properties of single cochlear nerve fibres in the cat.

1. Tuning properties and spontaneous discharge rate of single cochlear fibres in the anaesthetized cat were determined during short- and long-term poisoning of the cochlea by locally and systemically applied furosemide.2. With intra-arterial administration of furosemide, short-term reversible elevation occurred of the low threshold sharply tuned ;tip' segment of the frequency threshold (;tuning') curve (f.t.c.) by up to 40 db, without substantial changes in the threshold of the low frequency ;tail' segment of the f.t.c. These changes could occur in part without changes in the spontaneous activity and entirely without changes in the maximal evoked activity. These effects were observed in all fibres examined, the characteristic frequencies of which ranged from 3.5 to 31 kHz.3. Intracochlear administration of furosemide in 0.9 mM concentrations produced similar changes, but these were not reversible.4. The changes correlated with the depression of the amplitude of the gross cochlear action potential. The cochlear microphonic potential, however, was either unchanged, or only slightly reduced.5. In long-term furosemide poisoning of the cochlea, fibres with anomalous response properties were found alongside fibres having normal tuning. The former exhibited either reduced excitability of the low threshold tip segment, or a tip segment attenuated in both excitability and threshold.6. It is concluded that the selective effects of furosemide on the tip segment of cochlear fibre f.t.c.s offer further evidence for a physiologically vulnerable ;second filter' in the cochlea. The selective influence of the furosemide on the low threshold tip segment provides support for the hypothesis that the normal f.t.c. is generated by two largely independent processes: one vulnerable, low threshold and sharply tuned, and the other less vulnerable, but high threshold and more broadly tuned.7. The findings, obtained with an agent known to produce reversible impairment of hearing in man, provide direct physiological evidence in support of the hypothesis that in sensorineural hearing loss of cochlear origin the frequency selectivity of cochlear nerve fibres is impaired.

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.

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.

Reverse correlation study of cochlear filtering in normal and pathological guinea pig ears.

The filtering properties of single cochlear fibres have been determined in normal and kanamycin-treated guinea pigs using the reverse correlation technique. This method allows investigation of filtering over a wide dynamic range. For normal guinea pig fibres, the near threshold filter functions obtained with this method correspond to the tone derived frequency threshold curves ((FTCs). The 10 dB bandwidth of the filter functions increased monotonically with increasing noise levels above threshold. Thus with noise levels at approximately 50 dB above threshold, the 10 dB bandwidth had increased by a factor of 1.3--3. The changes in 3 dB bandwidth with increasing levels were, for some fibres, different from those of the 10 dB bandwidths. For the pathological fibres, the derived filter functions corresponded to their tone determined FTCs, and were therefore comparatively broadly tuned. Their tuning (Q10dB) approximated to those of normal fibres when the latter were measured 60 dB or more above threshold (i.e., at similar levels of stimulus), and did not increase further with increase in level. The findings in the normal guinea pig are consistent with those obtained by others in rodents, but are not consistent with those from the cat, where normal filtering is more robust to high levels of stimulus noise.

Location-specific components of the gross cochlear action potential: an assessment of the validity of the high-pass masking technique by cochlear nerve fibre recording in the cat.

The auditory high-pass masking technique has been used in attempts to define the origin, along the cochlear partition, of the gross cochlear action potential (CAP) and the gross brain stem potential. Theoretically, the high-pass masking paradigm should be frequency and location specific at the cochlear level, and some indirect evidence does point to this specificity. However, this hypothesis has not yet been directly substantiated. In the present experiment, click-evoked cochlear nerve activity was recorded simultaneously from the round window and from single fibres of the cochlear nerve, with and without high-pass maskers spaced in octaves from 0.5 to 16 kHz, at three intensities, in the anaesthetized cat. The "derived' CAPs were computed and compared with the mapping of single cochlear fibre responses under the same conditions. With one main exception, the conclusions drawn on the origin of the frequency components of the "derived' potentials were found to be valid in the normal cat. The exception concerned fibres with characteristic frequencies below 1-2 kHz, where the substantial spread towards the high frequencies of their frequency threshold curves, and the effects of lateral suppression or of other "remote masking' phenomena rendered the high-pass masking less location specific. From these results and certain assumptions, we would predict the high-pass masking technique to be valid in electrophysiological investigations in normal humans for frequencies down to 0.5-1 kHz.

Chronic effects of loop diuretics on the guinea-pig cochlea.

Groups of pigmented guinea pigs (weighing 480 to 650 g at the start of the experiment) were given daily intraperitoneal injections of 16 mg/kg bumetanide or 10, 20 or 40 mg/kg furosemide over a period of 4 weeks. Two weeks following the last diuretic dose the animals were anaesthetized with urethane and the responses of single auditory nerve fibres to pure-tone and click stimuli were recorded. The tuning of individual units, as measured by Q10 dB values of frequency threshold curves, tended to be more affected than threshold at characteristic frequency by both loop diuretics. Light microscope examination of treated cochleas suggested greater effect of the drugs on the organ of Corti than on stria vascularis.

The frequency selectivity of the normal and pathological human cochlea.

Human AP tuning curves (tone on tone simultaneous and forward masking curves) were measured during transtympanic electrocochleography. For subjects with near normal thresholds, average Q10 dB values (simultaneous masking) were 2.3 at 2 kHz, 3.6 at 4 kHz, and 4.7 at 8 kHz. Patients with threshold elevations of more than 40 dB, resulting from sensorineural hearing loss of cochlear origin, had tuning curves less sharply tuned by a factor of 2-3, with Q10 dB values of 1-2 at 2 kHz and at 4 kHz, and 1-2.3 at 8 kHz. AP tuning curves and single fibre tuning curves (frequency threshold curves) were measured in normal guinea pigs; cochlear fibre tuning is sharper than AP tuning (simultaneous masking) by a factor of 1.8 (2-20 kHz). Assuming that this factor can be applied to the human cochlea, estimates of normal human cochlear fibre Q10 dB values are 4.2 at 2 kHz, 6.5 at 4 kHz, and 8.5 at 8 kHz.

Auditory nerve fibre activity in the Tokay gecko

We have recorded from single auditory nerve fibres in the lizard,Gekko gecko, while changing the animal's temperature within the range 19–30 °C. Small steps of 0.5–2 °C (as measured next to the otic capsule or within the contralateral inner ear) cause a small but reliable and reversible change in the tuning characteristics of a fibre: its tuning curve (frequency-threshold curve) shifts to higher frequencies with warming and lower frequencies with cooling (Figs. 1, 2 and 3). The sharpness of tuning and overall sensitivity of the fibres are relatively unaffected by these procedures. The mean shift in characteristic frequency (CF) for 34 fibres was 0.06 octaves per °C (Fig. 4A). A more accurate estimate is given by the mean shift in ‘centre frequency’: a point midway between the low-frequency and high-frequency sides of the tuning curve at 10 dB above CF threshold. This shift was 0.05 octaves per °C (Fig. 4B), which is equivalent to a thermalQ 10 of approximately 1.4 for the frequency change with temperature. Comparison of these results and analogous frequency shifts with temperature reported for other auditory organs suggests that tuning in the mammalian cochlea is less temperature-dependent than it is in nonmammalian inner ears; this may reflect a basic difference in how tuning is achieved. The observed temperature sensitivities are discussed in terms of what might be expected for tuning by basilar and/or tectorial membranes and for tuning of conductances within the hair cell membrane.

Threshold elevation at high frequencies of the auditory nerve action potential in acute versus chronic recordings in guinea pigs

Thresholds of the auditory nerve compound action potential (CAP) responses to filtered clicks from 0.5 to 40 kHz, defining a CAP frequency threshold curve, have been determined in the same guinea pigs: (1) in acute conditions, the animal still anesthetized at the end of the operation for permanent implantation of a round-window electrode, and (2) several days later in the implanted awake and unrestrained animal. The results show higher CAP thresholds for high frequencies in acute conditions as compared to chronic recordings. This difference appears above about 12.5 kHz and increases progressively to around 40 dB at 40 kHz. Similar effects, although somewhat less pronounced, were observed during anesthesia alone in already implanted guinea pigs. These observations could explain the discrepancies which appear between electrophysiological thresholds, acutely recorded either from single fibers or whole-nerve responses, and behavioral audiograms. Moreover, in the accurate evaluation of the cochlea, slight impairments may be masked by this phenomenon as illustrated in guinea pigs treated with low doses of ototoxic antibiotic.

Psychophysical frequency resolution in the cat as determined by simultaneous masking and its relation to auditory-nerve resolution.

Critical bandwidths were measured behaviorally at 1 and 2 kHz by simultaneous masking in four cats. Three methods were used. They were (i) the masking of a tone by noise of variable bandwidth, (ii) the masking of a narrow-band signal by two tones, and (iii) the masking of a tone by noise of rippled spectrum. The three methods agreed closely and gave a mean critical bandwidth of 410 Hz at 1 kHz and 690 Hz at 2 kHz. These values were about three times as great as the electrophysiologically-determined effective bandwidths of single fibers of the auditory nerve at the same frequencies, both as measured in other animals, and in one of the animals that had been tested behaviorally as well. Psychophysical tuning curves were also determined behaviorally; in contrast, they agreed closely with auditory-nerve fibers in both bandwidth and slope. The results suggest that the critical band as measured by simultaneous masking is not a close relation of the frequency-threshold curve of auditory-nerve fibers, but that the psychophysical tuning curve possibly may be. Possible reasons and implications are discussed.

Forward masking of auditory nerve fiber responses.

1. Responses of single fibers were obtained from the auditory nerve of chinchillas. Tone-burst stimuli consisted of a masking stimulus followed by a probe stimulus. Forward masking of a fiber's response is defined as a reduction in the magnitude of the probe-evoked response caused by the addition of the masking stimulus. 2. The recovery of probe response magnitude as a function of the time interval between masker offset and probe onset (delta T) follows an exponential time course. A relationship between the time course or magnitude of poststimulus recovery and the characteristic frequency (CF) of a fiber was not detected. 3. The iso-forward masking contour near the threshold of the masking effect across masker frequencies approximates a fiber's frequency threshold curve (FTC). In other words, forward masking tuning curves are essentially the same as frequency threshold curves. 4. The frequency dependence of forward masking is compared to that of two-tone suppression. Tonal stimuli outside the boundaries of a fiber's FTC that produce two-tone suppression are ineffective forward maskers. Certain frequency/intensity combinations within the FTC may produce both suppression and forward masking and tones within the remaining area of the FTC produce no suppression but are effective forward maskers. 5. Both the time course and the magnitude of the forward masking effect are dependent on the discharge rate evoked by the masker regardless of the masker's absolute level or spectral content. An increase in masker-evoked excitation causes an increase in time constant and a greater reduction in probe response magnitude, rd. The function relating rd to masker level parallels the firing rate/masker level function up to 40 dB above response threshold. 6. A decrease in masker duration from 100 ms leads to a decrease in both rd and the time constant of recovery. There is no significant difference between the 100 and 200 ms duration conditions. 7. Forward masking in single fibers is related to the period of poststimulus recovery of spontaneous activity, a component of a fiber's response pattern to the masker, and this component is tentatively identified as a period of recovery from short-term adaptation.