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Spontaneous, click-, and toneburst-evoked otoacoustic emissions from normal ears.

Evoked and spontaneous otoacoustic emissions were recorded bilaterally in a group of normal subjects (n = 14) using clicks and tonebursts at four frequencies (0.5, 1, 1.5, and 3 kHz). All ears (n = 28) demonstrated evoked emissions, but not to every stimulus type. The 0.5-kHz toneburst evoked emissions in only 10 (36%) ears, the 1.5-kHz toneburst in all ears, and the remaining stimuli in at least 80% of ears. Two distinct patterns of evoked emissions were identified. Five (18%) ears showed short, broadband click-evoked emissions lasting less than 20 ms after stimulus onset. In these ears, toneburst-evoked emissions were often more prominent than click-evoked emissions and no spontaneous emissions were detected. Twenty-three (82%) ears showed click-evoked emissions lasting longer than 20 ms poststimulus onset. Spectral analysis of these emissions demonstrated several (2-10) narrow frequency peaks. Highly similar peaks were present in the spectra of toneburst-evoked emissions within the range of toneburst spectra. Spontaneous emissions were recorded in 12 of the 23 ears. In these ears, at the frequencies of spontaneous emissions, prominent peaks in both click- and toneburst-evoked emission spectra were always present. Otoacoustic emission characteristics correlated significantly between the ears of individual subjects inferring that a symmetrical cochlear mechanism generates otoacoustic emissions.

Acoustic Stimulation↗

Speculations about noise and the evolution of vertebrate hearing.

Two of the most conspicuous and celebrated properties of human hearing--the abilities to discriminate pitch and to localize sound sources, may have been secondary derivations to which the ear was preadapted by having evolved in the inevitable presence of noise.

Adaptation, Physiological↗

Neural temporal coding of low pitch. I. Human frequency-following responses to complex tones.

The neural basis of low pitch was investigated in the present study by recording a brainstem potential from the scalp of human subjects during presentation of complex tones which evoke a variable sensation of pitch. The potential recorded, the frequency-following response (FFR), reflects the temporal discharge activity of auditory neurons in the upper brainstem pathway. It was used as an index of neural periodicity in order to determine the extent to which the low pitch of complex tones is encoded in the temporal discharge activity of auditory brainstem neurons. A tone composed of harmonics of a common fundamental produces a sensation of pitch equal to that of the 'missing' fundamental. Such signals generate brainstem potentials which are spectrally similar to FFR recorded in response to sinusoidal signals equal in frequency to the missing fundamental. Both types of signals generate FFR which are periodic, with a frequency similar to the perceived pitch of the stimuli. It is shown that the FFR to the missing fundamental is not the result of a distortion product by recording FFR to a complex signal in the presence of low-frequency bandpass noise. Neither is the FFR the result of neural synchronization to the waveform envelope modulation pattern. This was determined by recording FFR to inharmonic and quasi-frequency-modulated signals. It was also determined that the 'existence region' for FFR to the missing fundamental lies below 2 kHz and that the most favorable spectral region for FFR to complex tones is between 0.5 and 1.0 kHz. These results are consistent with the hypothesis that far-field-recorded FFR does reflect neural activity germane to the processing of low pitch and that such pitch-relevant activity is based on the temporal discharge patterns of neurons in the upper auditory brainstem pathway.

Acoustic Stimulation↗

Tonotopic evolution during development.

The evolution of cochlear tonotopy can be observed by the study of the modification of tuning curves as obtained from auditory nerve fibers in the kitten. The differential development of the two components of the tuning curves, i.e. the tail, which is restricted to lower frequencies that appear at first during ontogenesis, and the tip which is restricted to higher frequencies that appear later, can explain the ontogenetic variations of tonotopy observed in the past. An hypothesis is presented where the tail of the tuning curve is related to the first functioning of the basal inner hair cells during development, whereas the delayed appearance of the tip is related to the basal outer hair cells. The base-apex gradient of maturation of receptors, along with their lateral gradient of development can explain the tonotopic shift observed during the cochlear ontogenesis.

Animals↗

Temporal response patterns of single auditory nerve fibers elicited by periodic electrical stimuli.

Single auditory nerve fibers exhibit firing synchronized to one or both phases of periodic AC stimulus currents. Responses to biphasic pulses depend on order and excitation sites of the two phases. Sine and triangle stimuli between 100 Hz and 500 Hz elicit similar response patterns. Responses to square waves are sometimes more synchronized and generally shifted in phase with respect to sine wave responses. Preferred firing phase(s): (1) are largely independent of stimulus intensity; (2) vary among fibers; (3) may shift continuously or discontinuously over several seconds before steady state is achieved. Responses to an unprocessed synthetic vowel stimulus were dominated by pitch period, first formant, and 'spurious' components.

Acoustic Stimulation↗

Sensitivity of neurons in the dorsal medullary nucleus of the grassfrog to spectral and temporal characteristics of sound.

The responses of 58 dorsal medullary nucleus units to a set of spectrally and temporally structured stimuli were investigated. Responses to tonepips and noise indicated monomodal spectral sensitivities, with diverse response patterns. Phase-locking was strong for frequencies from 0.1 to 0.2 kHz, and in one unit extended up to 0.6 kHz. To clicks, amplitude modulated tonebursts and natural and artificial versions of the mating call various responses were found. Most low-frequency units fired tonically. They showed a non-selective or low-pass rate response to increasing modulation frequency, and a low-pass synchronization behavior to the envelope. A group of mid-frequency units fired phasically and exhibited a band-pass rate characteristic of amplitude modulated tonebursts. Frequently this was combined with a low-pass rate characteristic of click trains. These units hardly responded to the time-reversed mating call, but fired in a time-locked fashion to the pulses of the original mating call, up to a signal-to-noise ratio of 0 dB. This suggests that aspects of pulse envelope and interpulse interval are coded in the dorsal medullary nucleus.

Animals↗

Physiological and psychophysical correlates of temporal processes in hearing.

Discharges of auditory nerve fibers are synchronized to stimulus frequencies below 4-5 kHz. The phase-locking phenomenon has been studied in considerable detail in several animal species. Although strikingly close correspondences exist between phase-locking behavior in animals and human perceptual performance on certain tasks, there is still no clear evidence that the human brain actually bases perceptual decisions on temporally encoded frequency information. The alternative to temporal coding is rate-place coding, in which frequency is assigned on the basis of peaks in cochlear excitation patterns. This paper reviews pertinent physiological, psychophysical and modeling data in three classes of experiment whose results are explanable in terms of both rate-place and temporal processing of neural responses. The experiments deal with the pitch of complex tones, vowel identification, and pure-tone frequency discrimination. The data described here suggest that temporal models of frequency coding compete well with and in some cases offer a more parsimonious explanation of perceptual performance than rate-place codes do, particularly at low and middle frequencies. A potentially important implication of the analyses conducted here is that humans may not code frequency information in synchronized activity as well as other species. The data suggest that within limits the human ear is capable of using either temporal and rate-place frequency codes, and that the specific code employed by the perceptual processor is task-dependent.

Animals↗

Synergistic interactions of gentamicin and pure tones causing cochlear hair cell loss in pigmented guinea pigs.

The effects of single and repeated combinations of gentamicin and sound on Preyer reflex and cochlear hair cells in pigmented guinea pigs have been examined. Gentamicin at 50 mg/kg for 10 days and an 8 kHz pure tone (116 dB for 60 min) cause little or no hair cell loss when given alone. In combination, hair cell loss occurs with the gentamicin and sound acting synergistically. If the pure tone is given on the first of a 10-day gentamicin course there is considerable loss of hair cells in the basal coil; if given on the 10th day the loss is several times less. Repeating the treatments after 3 weeks increases the hair cell loss and the synergistic effect. A possible explanation for the differential synergistic effect is discussed.

Animals↗

Influence of envelope rise time on neural responses in the auditory system of anurans.

The influence of envelope rise time on neural responses was investigated in the central auditory pathway of frogs. Single unit and evoked potential recordings were made from the dorsal medullary nucleus (DMN) and thalamus, respectively. It was found that phasic neurons (15% of the population) in the DMN responded preferentially to stimuli with rapid (less than 25 ms) rise times. Acoustically evoked potentials (AEPs) recorded from the thalamus, specifically the Nucleus of Bellonci, also showed more pronounced responses to stimuli with rapid, rather than slower, envelope rise times. Interestingly, the leopard frog mating call, which has a rapid onset, elicited strong neuronal discharge both within the DMN and thalamus. In contrast, the mating call of bullfrogs, a species sympatric with leopard frogs, has a characteristically slow (greater than 100 ms) envelope rise time and elicited little, if any, response. These findings indicate the presence of neural specializations within the frog's auditory pathway for the optimal detection of the conspecific mating call. The relevance of these findings for stimulus coding in the auditory pathway of other vertebrates is discussed.

Animals↗

Off-responses in the auditory system in relation to the signal end phase and neuronal characteristic frequency.

Off-responses of single neurons from the inferior colliculus (IC) as well as summed off-responses (evoked potentials) from the IC, cochlear nucleus and auditory nerve were studied while varying the end phase of tonal signals. It was found that with tonal signals higher than the characteristic frequency, off-responses at all the auditory levels studied were greatest at the end phases near 0 and 180 degrees and were of minimal value at the end phases near 90 and 270 degrees. On the contrary, with tonal signals lower than the characteristic frequency, the greatest off-responses corresponded to the end phases near 90 and 270 degrees, and responses of the lowest value were registered at the end phases near 0 and 180 degrees. The observed phenomena did not result from transient 'off'-responses of the acoustic system. It is suggested that they reflect certain cochlear processes connected with different effectiveness of oscillation phases at frequencies below and above the resonance.

Animals↗

Interferometric measurement of the amplitude and phase of tympanic membrane vibrations in cat.

The amplitude and phase of the tympanic membrane and malleus vibrations were measured over a wide frequency range with a homodyne interferometer. When sound pressure was maintained constant near the tympanic membrane, the malleus frequency response followed the typical pattern up to 10 kHz as measured by previous investigators. At higher frequencies the response changes dramatically. Instead of decreasing with frequency, between 10 and 20 kHz the vibration amplitude oscillates around a value which is only about 20 dB lower than the low frequency plateau level. Measurements of malleus vibration at several points along its length indicate that its mode of vibration changes at high frequencies, and no longer consists of a simple rotational component. All points on the tympanic membrane vibrate in phase with the malleus up to a frequency of 1 kHz. Above 5 kHz discrete resonances are observed, and the response varies strongly with position on the tympanic membrane.

Animals↗

Evoked otoacoustic emissions as cochlear Bragg reflections.

The reflection of cochlear waves is mathematically described for various assumptions about the dependence of the echo-generating mechanism on place and frequency, without requiring knowledge of the physical details of the mechanism. Any 'wave-related' reflections, caused by intrinsic features of the resonance or active-feedback mechanism, cannot explain long group delays in an approximately shift-similar basilar membrane (BM). Only 'BM related' scattering due to inhomogenities is a possible explanation then. Reflection at a fixed point of the BM could cause only narrow-band evoked otoacoustic emissions (EOAEs). Long group delays in a wider frequency band can be obtained by assuming Bragg reflection at an approximately periodic inhomogenity of BM parameters, possibly due to spatial variations of active undamping. This explains the long delays, about inversely proportional to frequency, the decreasing instantaneous frequency, the often found multi-packet or modulated structure of echoes (vanishing at higher SPL), the spectra of simultaneous and delayed EOAEs and their relation to the threshold of hearing. These phenomena do not necessarily require multiple reflection due to middle/inner-ear mismatch.

Auditory Threshold↗

Rapid assessment of sound-evoked olivocochlear feedback: suppression of compound action potentials by contralateral sound.

The compound action potential (CAP) measured at the round window of anesthetized cats in response to low-level tone pips can be significantly suppressed by addition of tones or noise to the opposite ear. This contralateral-sound suppression disappears upon transection of the olivocochlear bundle. The frequency and level dependence of the suppression phenomenon are well explained by known sound-evoked discharge properties of single olivocochlear neurons. Thus, the contralateral-sound suppression of cochlear CAP should prove useful as a rapid measure of the magnitude of the sound-evoked efferent feedback to the cochlea.

Animals↗

Intonation of musical intervals by musical intervals by deaf subjects stimulated with single bipolar cochlear implant electrodes.

Some subjects with cochlear implants have been shown to associate electrical stimulus pulse rates with the pitches of musical tones. In order to clarify the role of these pitch sensations in a musical context, the present investigation examined the intonation accuracy achieved by implant subjects when adjusting pulse rates in the reconstruction of musical intervals. Using a method of adjustment, the subjects altered a variable pulse rate, relative to a fixed reference rate, on one electrode, in the tuning of musical intervals abstracted from familiar melodies. At low pulse rates, subjects generally tuned the intervals to the same frequency ratios which define tonal musical intervals in normal-hearing listeners, with error margins comparable to musically untrained subjects. Two subjects were, in addition, able to transpose these melodic intervals from a standard reference pulse rate to higher and lower reference rates (reference and target pulse rates with geometric means of the intervals ranging from 81 to 466 pulses/s). Generally, the intervals were adjusted on a ratio scale, according to the same frequency ratios which define analogous acoustical musical intervals. These results support the hypothesis that, at low pulse rates, a temporal code in the auditory nerve alone is capable of defining musical pitch.

Acoustic Stimulation↗

Auditory enhancement at the absolute threshold of hearing and its relationship to the Zwicker tone.

Auditory enhancement describes an improvement in the detection of a tonal signal in a broad-band masker with a spectral gap at the signal frequency if the signal is delayed in its onset relative to the masker. This auditory enhancement may be based on an increase of the effective signal level instead of a decline in the effective masker level. In order to evaluate whether this signal enhancement also exists at the threshold of hearing, we measured the absolute threshold for pure-tone pulses of different frequencies with and without preceding band-rejected noise. Such noise also causes the sensation of the Zwicker tone-a faint pure tone lasting for a few seconds immediately after the noise presentation. The pitch of this sensation is a complex function of the noise parameters but always lies at a frequency within the rejected band. During the Zwicker tone sensation, auditory sensitivity for tone pulses at frequencies adjacent to the Zwicker tone was improved by up to 13 dB instead of being reduced which might be expected due to the presence of the simultaneously audible Zwicker tone. The failure to influence this threshold shift with low-frequency tones and measurements of the ear's acoustical response indicate that this threshold improvement may be produced through neuronal disinhibition rather than through a release from mechanical suppression in the cochlea.

Acoustic Stimulation↗

Binaural fusion and the representation of virtual pitch in the human auditory cortex.

The auditory system derives the pitch of complex tones from the tone's harmonics. Research in psychoacoustics predicted that binaural fusion was an important feature of pitch processing. Based on neuromagnetic human data, the first neurophysiological confirmation of binaural fusion in hearing is presented. The centre of activation within the cortical tonotopic map corresponds to the location of the perceived pitch and not to the locations that are activated when the single frequency constituents are presented. This is also true when the different harmonics of a complex tone are presented dichotically. We conclude that the pitch processor includes binaural fusion to determine the particular pitch location which is activated in the auditory cortex.

Acoustic Stimulation↗

Assessment of outer hair cell function and blood antioxidant status of rabbits exposed to noise and metal welding fumes.

OBJECTIVES: To investigate the interaction between welding fumes and noise in causation of hearing impairment. METHODS: Groups of rabbits (n=6) were exposed to noise, welding fumes or combination of both prior to Distortion Product Otoacoustic-Emissions (DPOAEs) analysis. The function of outer hair cells (OHCs) was examined by DPOAE assessment over a broad range of frequencies. Variations in DPOAE amplitude were compared between control (n=6) and exposed (n=18) groups. RESULTS: The DPOAEs levels measured at different frequencies (1379-6299 Hz) were found to decrease significantly (P<0.05) in rabbits exposed to 110 dB sound pressure level (SPL) broadband noise (8h/day, 12 days). In rabbits, exposed to carbon-steel welding fumes alone (157 mg/m(3)), the threshold shift was limited to the high frequencies (2759-6299 Hz), whereas, mixed exposure to noise and fumes resulted in reduction of DPOAEs at all the frequencies. Changes in DPOAEs were associated with increased susceptibility of erythrocytes to oxidation (P<0.05). Exposure to noise or fumes alone or simultaneously, suppressed total antioxidant ability of plasma as measured by ferric reducing ability of plasma (FRAP). Noise alone or in combination with fumes resulted in depletion of blood glutathione (GSH). Despite suppression of FRAP in the exposed groups, GSH was found to remain unchanged due to welding fumes suggesting that antioxidants other than GSH are affected by toxicants present in metal welding fumes. CONCLUSION: Exposure to very high levels of welding fumes can increase noise-related effects on OHC function by extending hearing threshold shift to wide band frequencies.

Air Pollutants, Occupational↗