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IHC-TM connect-disconnect in relation to sensitization and masking of a HF-tone burst by a LF tone. IV.

Evidence continues to accumulate that although the outer hair cells (OHCs) of the cochlea are firmly bonded to the tectorial membrane (TM), the inner hair cells (IHCS) are not. This is the fourth in a series of papers that explores how the idea of a set of disconnected hair cells that "impact" the TM is consistent with psychophysical data. The paper extends the exploration to the masking of high-frequency (HF) tone bursts by low-frequency (LF) tones and shows that the model can explain the important features of these complex data.

Auditory Threshold↗

ABR measurements in the cat using a forward-masking paradigm.

Probe-elicited wave V amplitudes of the auditory brainstem response (ABR) were measured using a forward-masking paradigm. Subjects were anesthetized cats. For individual experiments, probe frequency and intensity were fixed and masker frequencies and intensities were varied. For each masker frequency, the extent to which the probe-elicited wave V amplitude was reduced by the preceding masker was plotted as a function of masker intensity. The rising segments of the masking functions were fitted with straight lines, using a least-squares procedure, to obtain estimates of their slopes. Masking grew most rapidly for masker frequencies below probe frequency, becoming progressively less steep as masker frequency increased. ABR tuning curves were constructed by using the linear fits to define the masker intensity that caused a 50% reduction in probe-elicited wave V amplitude. The shapes of these tuning curves were comparable to whole-nerve action potential (AP) tuning curves obtained under similar stimulus conditions. These results indicate that ABR amplitude measurements in a forward-masking paradigm can be used to estimate the growth of response to masking stimuli and frequency selectivity in a manner similar to AP amplitude measurements.

Action Potentials↗

Place specific influences on the wave I to V interpeak latency of the auditory brain-stem response.

There is controversy over whether the wave I to V interpeak latency (I-V IPL) of the auditory brain-stem response can be manipulated by cochlear processing. In this study, a forward masking paradigm was used to test the predictions of two contrasting models of I-V IPL generation. The paradigm was designed to determine if the I-V IPL can be affected by masking selected portions of the cochlear response region. The results from ten normal hearing subjects suggest that: (1) waves I and V can be masked semi-independent of each other, and (2) the I-V IPL can be shortened or prolonged by masking the basal or apical portion of the cochlear response region respectively. These findings support the hypothesis that, at least in normal hearing subjects, wave V is biased to reflect more apical cochlear events than wave I. Additionally, they offer tentative support for anecdotal reports of shortened I-V IPLs in the presence of high-frequency hearing loss.

Adult↗

Effects of aspirin on psychophysical measures of frequency selectivity, two-tone suppression, and growth of masking.

Three psychophysical measures of nonlinearity were evaluated before and during a course of aspirin ingestion to investigate the role of outer hair cells (OHCs) in these measures, as aspirin is thought to alter the functioning of OHCs. Six normal-hearing individuals received a moderate dose (3.9 g/day) of aspirin for four days, producing essentially flat, temporary hearing losses that ranged from 5-20 dB. The losses were about 2 dB greater for a 300-ms signal than for a 15-ms signal, indicating reduced temporal integration with aspirin. On the final three days of aspirin use, three experiments were completed; each was designed to measure one aspect of nonlinear behavior: (1) the effects of level on frequency selectivity in simultaneous masking using notched-noise maskers, (2) two-tone suppression using forward maskers at the signal frequency (fs) and suppressor tones above fs, and (3) growth-of-masking functions in forward masking using a masking tone below fs. Signal frequencies of 750 and 3000 Hz were used to evaluate the effects of aspirin at relatively low- and high-frequency regions of the cochlea. In experiment 1, aspirin broadened the auditory filters and reduced the effect of level on frequency selectivity. In experiment 2, aspirin reduced or eliminated two-tone suppression. And, in experiment 3, aspirin reduced the slopes of the growth-of-masking functions. Thus, the aspirin was effective in reducing nonlinearity in all three experiments, suggesting that these measures reflect the same (or a similar) active, nonlinear mechanism, namely the compressive nonlinearity provided by the OHCs. In all experiments, aspirin tended to have larger detrimental effects on the nonlinear measures at 3000 Hz than at 750 Hz, which can be explained in terms of greater involvement of nonlinear processing at higher frequencies. Finally, these effects of aspirin were found to be similar to those observed in preliminary measurements in two subjects with mild, permanent hearing loss.

Adult↗

Inter-relationship between different psychoacoustic measures assumed to be related to the cochlear active mechanism.

The active mechanism in the cochlea is thought to depend on the integrity of the outer hair cells (OHCs). Cochlear hearing loss is usually associated with damage to both inner hair cells (IHCs) and OHCs, with the latter resulting in a reduction in or complete loss of the function of the active mechanism. It is believed that the active mechanism contributes to the sharpness of tuning on the basilar membrane (BM) and is also responsible for compressive input-output functions on the BM. Hence, one would expect a close relationship between measures of sharpness of tuning and measures of compression. This idea was tested by comparing three different measures of the status of the active mechanism, at center frequencies of 2, 4, and 6 kHz, using subjects with normal hearing, with unilateral or highly asymmetric cochlear hearing loss, and with bilateral loss. The first measure, HLOHC, was an indirect measure of the amount of the hearing loss attributable to OHC damage; this was based on loudness matches between the two ears of subjects with unilateral hearing loss and was derived using a loudness model. The second measure was the equivalent rectangular bandwidth (ERB) of the auditory filter, which was estimated using the notched-noise method. The third measure was based on the slopes of growth-of-masking functions obtained in forward masking. The ratio of slopes for a masker centered well below the signal frequency and a masker centered at the signal frequency gives a measure of BM compression at the place corresponding to the signal frequency; a ratio close to 1 indicates little or no compression, while ratios less than 1 indicate that compression is occurring at the signal place. Generally, the results showed the expected pattern. The ERB tended to increase with increasing HLOHC. The ratio of the forward-masking slopes increased from about 0.3 to about 1 as HLOHC increased from 0 to 55 dB. The ratio of the slopes was highly correlated with the ERB (r = 0.92), indicating that the sharpness of the auditory filter decreases as the compression on the BM decreases.

Adult↗

Auditory edge detection: a neural model for physiological and psychoacoustical responses to amplitude transients.

Primary segmentation of visual scenes is based on spatiotemporal edges that are presumably detected by neurons throughout the visual system. In contrast, the way in which the auditory system decomposes complex auditory scenes is substantially less clear. There is diverse physiological and psychophysical evidence for the sensitivity of the auditory system to amplitude transients, which can be considered as a partial analogue to visual spatiotemporal edges. However, there is currently no theoretical framework in which these phenomena can be associated or related to the perceptual task of auditory source segregation. We propose a neural model for an auditory temporal edge detector, whose underlying principles are similar to classical visual edge detector models. Our main result is that this model reproduces published physiological responses to amplitude transients collected at multiple levels of the auditory pathways using a variety of experimental procedures. Moreover, the model successfully predicts physiological responses to a new set of amplitude transients, collected in cat primary auditory cortex and medial geniculate body. Additionally, the model reproduces several published psychoacoustical responses to amplitude transients as well as the psychoacoustical data for amplitude edge detection reported here for the first time. These results support the hypothesis that the response of auditory neurons to amplitude transients is the correlate of psychoacoustical edge detection.

Animals↗

On the relationship between speech- and nonspeech-evoked auditory brainstem responses.

Auditory brainstem response (ABR) reflects activation of the neural generators along the ascending auditory pathway when a sound is heard. In this study, we explored the relationship between brainstem encoding of click and speech signals in normal-learning children and in those with language-based learning problems. To that end, ABR was recorded from both types of stimuli. We found that the normal pattern of correlation between click- and speech-evoked ABRs was disrupted when speech-evoked ABRs were delayed. Thus, delayed responses to speech were not indicative of clinically abnormal responses to clicks. We conclude that these two responses reflect largely separate neural processes and that only processes involved in encoding complex signals such as speech are impaired in children with learning problems.

Acoustic Stimulation↗

Audiological assessment in Ramsay Hunt syndrome.

Ramsay Hunt syndrome is known to cause symptoms and signs of vestibulocochlear dysfunction, including sensorineural hearing loss. The present study investigates the audiological features of a group of 15 patients with this syndrome. A complete otolaryngological, neurologic, and audiological workup was performed in every patient, including auditory brain stem response measurements and recording of transiently evoked otoacoustic emissions. In most patients, some degree of hearing loss was evident, and abnormal latencies and interpeak latencies of the auditory brain stem response, or even absence of the waves, were observed. Transiently evoked otoacoustic emissions were present in only 6 cases, and caloric tests showed unilateral weakness in the majority of the patients. In all of the performed tests, abnormalities were present only on the affected side. The audiological data suggested cochlear or retrocochlear involvement or involvement at more than one site along the auditory pathway.

Adolescent↗

Safe level for noise exposure?

The safety of long, continuous exposure to several different levels of noise was determined. Chinchillas were exposed for two or nine days to an octave band of noise with a center frequency of .5 kHz or 4 kHz. By examining the inner ear for evidence of acute cellular degeneration, exposures which produced minimal permanent damage could be identified easily. Differences were found between the two bands of noise in the level required to produce injury, and the type, location and extent of damage which occurred as the level was increased. The findings help explain why exposure to industrial noise, which is generally low frequency, first produces measurable auditory changes for high frequency tones.

Acoustic Stimulation↗

Otoacoustic emissions: an approach for monitoring aminoglycoside-induced ototoxicity.

Ototoxic drugs, such as aminoglycosides, affect outer hair cell integrity in the inner ear. Transiently evoked otoacoustic emission (TEOAE) characteristics are related to outer hair cell function and can be expected to reflect the influence of ototoxic agents. Transiently evoked otoacoustic emissions were measured during amikacin sulfate therapy in nine patients. The duration of treatment for individual patients ranged from 9 to 33 days. A reversible decrease of overall TEOAE level, occurring after a treatment period longer than 16 days, was found in the majority of patients. The monitoring of TEOAEs is proposed as a method for early identification and, as a result, prevention of aminoglycoside-induced ototoxicity.

Adolescent↗

Interaural phase and level difference sensitivity in low-frequency neurons in the lateral superior olive.

The lateral superior olive (LSO) is believed to encode differences in sound level at the two ears, a cue for azimuthal sound location. Most high-frequency-sensitive LSO neurons are binaural, receiving inputs from both ears. An inhibitory input from the contralateral ear, via the medial nucleus of the trapezoid body (MNTB), and excitatory input from the ipsilateral ear enable level differences to be encoded. However, the classical descriptions of low-frequency-sensitive neurons report primarily monaural cells with no contralateral inhibition. Anatomical and physiological evidence, however, shows that low-frequency LSO neurons receive low-frequency inhibitory input from ipsilateral MNTB, which in turn receives excitatory input from the contralateral cochlear nucleus and low-frequency excitatory input from the ipsilateral cochlear nucleus. Therefore, these neurons would be expected to be binaural with contralateral inhibition. Here, we re-examined binaural interaction in low-frequency (less than approximately 3 kHz) LSO neurons and phase locking in the MNTB. Phase locking to low-frequency tones in MNTB and ipsilaterally driven LSO neurons with frequency sensitivities <1.2 kHz was enhanced relative to the auditory nerve. Moreover, most low-frequency LSO neurons exhibited contralateral inhibition: ipsilaterally driven responses were suppressed by raising the level of the contralateral stimulus; most neurons were sensitive to interaural time delays in pure tone and noise stimuli such that inhibition was nearly maximal when the stimuli were presented to the ears in-phase. The data demonstrate that low-frequency LSO neurons of cat are not monaural and can exhibit contralateral inhibition like their high-frequency counterparts.

Acoustic Stimulation↗

Frequency analysis of transiently evoked otoacoustic emissions in sensorineural hearing disturbance.

Transiently evoked otoacoustic emissions (TEOAE) are present in nearly all normal hearing ears and recognized as useful for objective auditory screening. In the spectral analysis of TEOAE with click stimuli, the normal power spectrum of TEOAE has yet to be obtained. In this study, the normal range of the power spectrum was determined for 42 adults with normal hearing and compared with sensorineural hearing disturbance data. The power spectrum of TEOAE in normal hearing adults sloped down at high frequencies and was the same for right and left ears in both males and females. For comparison with sensorineural hearing disturbance, a significant correlation between the audiogram and power spectrum of TEOAE was sought. Frequency specificity was noted in the power spectrum of TEOAE with click stimulation.

Adult↗

Use of frequency modulation combined with amplitude change to elicit frequency-specific whole-nerve response.

The averaged whole-nerve response to frequency modulation combined with amplitude change was recorded from the round window of the guinea pig. A downward shift of 20% in the frequency of a 500-Hz, 1-kHz or 2-kHz pure tone with an amplitude increase elicited the responses. The latency for each frequency was investigated at different intensities. The latency is greater for lower-frequency tones and this latency increase agrees well with the traveling wave delay for the intensity range used in this study. A downward shift with an amplitude decrease failed to yield the response. These findings suggest that the response to a downward shift of frequency with an amplitude increase results from new activation due to an apical extension of the envelope of the traveling wave and thus represents the activity in a restricted area of the basilar membrane. Thus, the frequency-modulated tone in downward direction with increased amplitude can be a frequency-specific stimulus.

Acoustic Stimulation↗

Attenuation of auditory-evoked potentials during voluntary movement in man.

Long-latency auditory evoked potential (AEP) amplitudes were reduced during voluntary thumb movement. The cerebral component N100 did not change during the movement. By contrast, the cerebral potential P200 and N100-P200 amplitudes were attenuated during the motor task compared to control. This report cannot clearly distinguish if the attenuation obtained in the AEP during the performance of the movement is related to motor events in the cerebral cortex causing 'gating' or to other factors such as habituation or vigilance changes.

Auditory Threshold↗

Evoked-response forward-masking functions in chinchillas with noise-induced permanent hearing loss.

Evoked-response forward-masking functions were measured by chronic electrodes in the inferior colliculus of the chinchilla before and after exposure to an intense tone that produced a permanent hearing loss. Before exposure, the forward-masking time constants ranged from 50 to 90 ms. After exposure, the forward-masking time constants increased significantly in the region of hearing loss, but not in regions where hearing was normal. The effect of the hearing loss on the time course of forward masking was most pronounced once the hearing loss exceeded 20-25 dB. These physiological changes in the evoked-response forward-masking functions appear to parallel those observed psychophysically in human listeners.

Animals↗

Dependence of auditory brainstem response on click polarity and high-frequency sensorineural hearing loss.

An investigation is presented on the effect of click polarity upon the auditory brainstem response (ABR) in cases of simulated high-frequency hearing loss. A high-frequency hearing loss was stimulated in 12 normal subjects by applying a high-pass noise masker, the noise cutoff frequency being varied in half-octave steps through the range of audiometric frequencies. With this paradigm, the effects of click polarity on the ABR are similar to those reported in patients with high-frequency hearing loss, including changes in waveform morphology and substantial peak latency shifts. To evaluate the role of individual cochlear frequency channels in a more direct way, half-octave narrow-band responses were determined by pair-wise subtraction of the high-pass-noise-masked responses. As far as ABR peak latencies are concerned, stimulus polarity effects tend to increase towards the lower frequency bands. Thus, the selective elimination of the contribution to the ABR of cochlear high-frequency channels can be held responsible for the increased dependence of ABR on click polarity observed in high-frequency hearing loss. However, statistical evaluation of peak I, III and V latencies, both in high-pass-noise-masked and in narrow-band ABRs, reveals that the latency effects are not systematic between subjects. Mechanisms producing the observed rarefaction-condensation differences, possible explanations for the large intersubject variability and the consequences for clinical interpretation of ABR are discussed.

Acoustic Stimulation↗

A study of the effects of cochlear loss on the auditory brainstem response (ABR) specificity and false positive rate in retrocochlear assessment.

The auditory brainstem response (ABR) test is widely used as a screen for retrocochlear dysfunction in individuals suffering idiopathic auditory and vestibular disturbance. Although its sensitivity for lesions of the lower brainstem is well established, hearing loss is known to have significant effects upon the test's specificity and false positive rate. This study analysed the effects of aspects of cochlear hearing loss including level, slope and general audiogram shape on these properties of the ABR test in a large clinical group. The study comprised 306 patients (153 male, 153 female) referred for ABR screening to exclude retrocochlear dysfunction. Of 566 ears meeting the inclusion criteria, 85 (15 per cent) failed the test. On follow-up eight ears (3 per cent) were confirmed to have some form of retrocochlear abnormality, seven of which were pathological. All had abnormal ABR results. No cases of retrocochlear abnormality were found in the ABR pass group. In those patients diagnosed as free of retrocochlear problems, the importance of the I-V interval as a primary index of function was confirmed. It displayed a high specificity (>90 per cent) which was largely independent of the level or slope of high-frequency hearing loss. In contrast wave V specificity decreased with increasing loss and increasing slope. Its value as a functional index was limited with losses greater than 70 dB HL in whom specificity fell below 70 per cent. The study concluded that, despite reducing interval availability above 60 dB HL, wave V latency cannot be used as an alternative index because it displays weakening specificity over this range of loss. With the problems inherent in applying latency corrections, the author advocates the use of improved procedures for wave I identification in these cases.

Adult↗