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Effects of acoustic overstimulation on spectral and temporal processing in the amphibian auditory nerve.

Activity of isolated auditory-nerve fibers in tree frogs (Eleutherodactylus coqui) exposed to continuous 3-min tones of different intensities at their characteristic frequencies (CFs) was recorded. Period histograms show a retardation in the preferred phase of discharge during and after the cessation of the exposure. Postexposure phase shift is concomitant with an elevation in CF thresholds and related to the level of tone exposure above threshold. Vector strength does not show comparable trends of change; postexposure shifts are related to preexposure CF thresholds. Recovery of phase retardation is rapid; units exposed to successive 3-min tones of the same intensities with intervals of 10-14 min between exposures experienced similar changes in their patterns of temporal discharge. Micromechanical changes affecting stereocilia stiffness or structural alterations in the tectorial membrane of the amphibian papilla may underly the transitory phase shifts observed in traumatized anuran auditory fibers.

Acoustic Stimulation↗

Temporal resolution in frog auditory-nerve fibers.

Sinusoidally amplitude-modulated (SAM) noise was monaurally presented to the neotropical frog, Eleutherodactylus coqui, while recording intracellularly from auditory-nerve fibers. Neuronal phase locking was measured to the SAM noise envelope in the form of a period histogram. The modulation depth was changed (in 10% steps) until the threshold modulation depth was determined. This was repeated for various modulation frequencies (20-1200 Hz) and different levels of SAM noise (34-64 dB/Hz). From these data, temporal modulation transfer functions (TMTFs) were produced and minimum integration time (MIT) for each auditory fiber was calculated. The median MIT was 0.42 ms (lower quartile 0.32, upper quartile 0.68 ms). A noise level-dependent effect was noted on the shape of the TMTF as well as the minimum integration time. The latter results may be explained as a loss in spectral resolution with increasing noise level, which is consistent with the correlation that was found between minimum integration time and bandwidth.

Animals↗

Detection and discrimination of synthetic English vowels by Old World monkeys (Cercopithecus, Macaca) and humans.

Abilities to detect and discriminate ten synthetic steady-state English vowels were compared in Old World monkeys (Cercopithecus, Macaca) and humans using standard animal psychophysical procedures and positive-reinforcement operant conditioning techniques. Monkeys' detection thresholds were close to humans' for the front vowels /i-I-E-ae-E), but 10-20 dB higher for the back vowels /V-D-C-U-u/. Subjects were subsequently presented with groups of vowels to discriminate. All monkeys experienced difficulty with spectrally similar pairs such as /V-D/, /E-ae/, and /U-u/, but macaques were superior to Cercopithecus monkeys. Humans discriminated all vowels at 100% correct levels, but their increased response latencies reflected spectral similarity and correlated with higher error rates by monkeys. Varying the intensity level of the vowel stimuli had little effect on either monkey or human discrimination, except at the lowest levels tested. These qualitative similarities in monkey and human vowel discrimination suggest that some monkey species may provide useful models of human vowel processing at the sensory level.

Adolescent↗

The effect of varying the amplitude-frequency response on the masked speech-reception threshold of sentences for hearing-impaired listeners.

In an evaluation of frequency-dependent automatic gain-control systems in hearing aids, the effect of varying the amplitude-frequency response on the speech-reception threshold (SRT) for sentences in noise is studied for 20 hearing-impaired listeners. The noise has a spectrum identical to the long-term average spectrum of the sentences. Speech and noise are shaped by the same amplitude-frequency response; their spectra are varied relative to the bisector of the individual's dynamic range. In four experimental conditions, the effect of a steady-state amplitude-frequency response is studied. Steepening the negative spectral slope of speech and noise appears to cause an increase of masked SRT, possibly due to increased effect of upward spread of masking. The effect of a single transition of the amplitude-frequency response between 10 and -10 dB/oct halfway through the sentence seems to be related to the effect for the fixed -10-dB/oct condition. Two transition times are tested. For a transition time of 0.25 s, the SRT is only a little higher than for 1 s. The results suggest that the amplitude-frequency response may be varied in time without having a detrimental effect on the masked SRT of sentences for hearing-impaired listeners as long as strongly negatively sloping spectra are avoided.

Adult↗

Intensity discrimination as the driving force for loudness. Application to pure tones in quiet.

Loudness functions and their associated neural-count functions are derived for steady-state tones at 250 and 1000 Hz from measurements of intensity discrimination obtained under gated and continuous conditions. The calculations are based on a multichannel generalization of the McGill-Goldberg counting model [J. Acoust. Soc. Am. 44, 576-581 (1968)]. Using the data for just noticeable differences (jnd) in intensity as input, the generalized version gives an integral relation between the neural-count function N(x) and the intensity-jnd function, where x = I/I0 and I0 is the reference intensity. Loudness functions are generated through the prescription L(x) = AN(x)--B. To determine how the detailed shapes of the intensity-jnd functions affect the form of the loudness function within the model, integration was performed over the intensity-jnd functions with and without a power-function approximation. Over a range of intensity levels from 20-95 dB, the best agreement between the calculated and measured loudness functions is obtained from the unaltered intensity-jnd functions. Consistent with psychophysical evidence and several models of intensity coding, the results predict that the output of the whole auditory nerve is unnecessary to maintain the large dynamic range observed for loudness and intensity discrimination.

Animals↗

A population study of auditory-nerve fibers in unanesthetized decerebrate cats: response to pure tones.

Population responses of single auditory-nerve (AN) fibers to 1-kHz pure tones were investigated in unanesthetized decerebrate cats. Driven rate (spikes/s), Fourier component amplitude (spikes/s), and d'e = (driven rate)/(standard deviation) were examined. These results extend the presently available information. First, the AN population responses, previously observed only in anesthetized animals, are now observed in unanesthetized animals. The anesthetized and unanesthetized animal responses are qualitatively similar. Second, a detectability measure d'e was applied, à la signal detection theory, to populations of AN fibers. A previously unknown behavior of the low-spontaneous-rate (SR) (less than 15 spikes/s) fibers was observed: the d'e increased to large values at moderate and high stimulus levels, giving rise to a large, sharply tuned peak in the d'e-place profile, even when the driven rate was saturated. This behavior was absent in the high-SR (greater than 15 spikes/s) fibers. This observation suggests that the d'e of the low-SR fibers is the best code for stimulus intensity among the different response measures examined, and that the d'e-place profile may be a more precise spatial code for stimulus frequency than the commonly considered rate-place code.

Animals↗

A comparison of maximum length and Legendre sequences for the derivation of brain-stem auditory-evoked responses at rapid rates of stimulation.

Experiments were performed in which brain-stem auditory-evoked responses (BAERs) were elicited by two types of pseudorandom pulse trains: maximum length sequences (MLS) and Legendre sequences (LGS). In experiment 1, each pulse sequence was presented at 50 dB nHL with minimum pulse intervals varying from 1 to 10 ms. Wave V latency increased and wave V amplitude decreased with decreasing minimum pulse intervals, with no significant effect of the type of sequence (MLS vs LGS), and no significant interaction between sequence and minimum pulse interval in terms of wave V amplitude or latency. In a second experiment, the minimum pulse interval was held constant at 4 ms, while MLS and LGS levels were varied from 20 to 60 dB nHL. With increasing click intensity, there is a decrease in wave V latency and an increase in wave V amplitude. There was no significant effect of type of sequence (LGS vs MLS) or interaction between type of sequence and stimulus intensity for wave V amplitude or latency. Despite the obvious violation of the assumptions (linearity and stationarity) underlying the application of maximum length sequence analysis and Legendre sequence analysis, both techniques produced reliable responses remarkably similar in morphology to evoked responses obtained by conventional averaging. The results of these experiments support the possibility that analysis methods based on pseudorandom pulse sequences may prove more efficient in data collection and provide a more thorough description of the electrophysiologic behavior of the auditory system compared to conventional averaging.

Adult↗

Across-channel masking and comodulation masking release.

These experiments on across-channel masking (ACM) and comodulation masking release (CMR) were designed to extend the work of Grose and Hall [J. Acoust. Soc. Am. 85, 1276-1284 (1989)] on CMR. They investigated the effect of the temporal position of a brief 700-Hz signal relative to the modulation cycle of a 700-Hz masker 100% sinusoidally amplitude modulated (SAM) at a 10-Hz rate, which was either presented alone (reference masker) or formed part of a masker consisting of the 3rd to 11th harmonics of a 100-Hz fundamental. In the harmonic maskers, each harmonic was either SAM with the same 10-Hz modulator phase (comodulated masker) or with a shift in modulator phase of 90 degrees for each successive harmonic (phase-incoherent masker). When the signal was presented at the dips of the envelope of the 700-Hz component, the comodulated masker gave lower thresholds than the reference masker, while the phase-incoherent masker gave higher thresholds, i.e., a CMR was observed. No CMR was found when the signal was presented at the peaks of the envelope. In experiment 1, we replicated the experiment of Grose and Hall, but with an additional condition in which the 600- and 800-Hz components were removed from the masker, in order to investigate the role of within-channel masking effects. The results were similar to those of Grose and Hall. In experiment 2, the signal was added at the peaks of the envelope of the 700-Hz component, but in antiphase to the carrier of that component and at a level chosen to transform the peaks into dips. No CMR was found. Rather, performance was worse for both the comodulated and phase-incoherent maskers than for the reference masker. This was true even when the flanking components in the maskers were all remote in frequency from 700 Hz. In experiment 3, the masker components were all 50% SAM and the signal was added in antiphase at a dip of the envelope of the 700-Hz component, thus making the dip deeper. Performance was worse for the phase-incoherent than for the reference masker and was worse still for the comodulated masker. The results of all three experiments indicate strong ACM effects. CMR was found only when the signal was placed in the dips of the masker envelope and when it produced an increase in level relative to that in adjacent bands.

Adult↗

The combination of interaural time and intensity in the lateralization of high-frequency complex signals.

In an effort to examine the rules by which information arising from interaural differences of time (IDT) and interaural differences of intensity (IDI) is combined, d"s were measured for trains of high-frequency clicks (4000 Hz, bandpass) possessing various combinations of IDT and IDI. The number of clicks was either 1 or 8, with the interclick interval either 2 or 10 ms. A 2-IFC task was employed in which the paired values of IDT and IDI favored one side during one interval and the other side during the other interval. Data obtained with the combined cues are compared to those obtained with IDTs or IDIs alone in order to determine the degree to which processing of the two cues is done independently. Results suggest that lateralization with such stimuli is based on the sum of the temporal and intensive differences and not on independent evaluations of their separate presences.

Adult↗

Binaural forward and backward masking: evidence for sluggishness in binaural detection.

The threshold of a short interaurally phase-inverted probe tone (20 ms, 500 Hz, S pi) was obtained in the presence of a 750-ms noise masker that was switched after 375 ms from interaurally phase-inverted (N pi) to interaurally in-phase (No). As the delay between probe-tone offset and noise phase transition is increased, the threshold decays from the N pi S pi threshold (masking level difference = 0 dB) to the No S pi threshold (masking level difference = 15 dB). The decay in this "binaural" situation is substantially slower than in a comparable "monaural" situation, where the interaural phase of the masker is held constant (N pi), but the level of the masker is reduced by 15 dB. The prolonged decay provides evidence for additional binaural sluggishness associated with "binaural forward masking." In a second experiment, "binaural backward masking" is studied by time reversing the maskers described above. Again, the situation where the phase is switched from No to N pi exhibits a slower transition than the situation with constant interaural phase (N pi) and a 15-dB increase in the level of the masker. The data for the binaural situations are compatible with the results of a related experiment, previously reported by Grantham and Wightman [J. Acoust. Soc. Am. 65, 1509-1517 (1979)] and are well fit by a model that incorporates a double-sided exponential temporal integration window.

Adult↗

NoSo and NoS pi detection as a function of masker bandwidth in normal-hearing and cochlear-impaired listeners.

NoSo and NoS pi detection thresholds for a 500-Hz pure-tone signal were measured as a function of masking noise bandwidth in normal-hearing and cochlear hearing-impaired subjects. NoSo and NoS pi critical bands were derived from the bandlimited noise functions. A notched noise measure of the monaural critical band was also obtained for each ear. One hypothesis tested was that an asymmetrical monaural critical band would result in a relatively steep improvement of the NoS pi detection threshold as a function of decreasing masker bandwidth and would, therefore, be associated with a wider binaural critical band. This was hypothesized because the outputs of the left and right auditory filters would be more decorrelated the greater the interaural difference in the monaural critical band. However, as the noise bandwidth was narrowed, the decorrelation would lessen, resulting in a relatively steep improvement in NoS pi detection. Results indicated that the masking level difference (MLD) was smaller and that the monaural critical bands were generally wider in cochlear-impaired listeners. NoSo and NoS pi critical bands were somewhat larger in the cochlear hearing-impaired listeners having relatively wide monaural critical bands. There was a significant correlation between monaural critical band asymmetry and the NoS pi critical band; however, this correlation was insignificant when a control was employed for the critical band in the worse ear. Therefore, the present results did not support a strong association between monaural critical band asymmetry and the width of the NoS pi critical band.

Attention↗

Detection of frequency modulation (FM) in the presence of a second FM tone.

A series of three experiments was undertaken to investigate detection of sinusoidal frequency modulation (FM) in the presence of FM at a separate frequency. The first experiment measured detection of modulation for an FM tone with a modulation frequency (fm) of 6 Hz as a function of carrier frequency (fc) under three conditions: (1) in quiet, (2) in the presence of a 2500-Hz pure tone, and (3) in the presence of a 2500-Hz FM tone with fm = 6 Hz, modulating in phase with the signal. Detection of FM in the presence of the second FM tone was worse than for either the signal presented in quiet or in the presence of the unmodulated tone. Threshold varied as an inverse function of frequency separation between the signal and the masker. In the second experiment, FM detection for a signal with fc = 1900 Hz and fm = 6 Hz was measured as a function of the modulation frequency (fm = 2-18 Hz) of the 2500-Hz masker tone. FM detection improved significantly with increasing difference between the modulation frequencies of the signal and the masker. The final experiment measured detection of FM for a signal (fc = 1900 Hz, fm = 6 Hz) in the presence of a second FM tone (fc = 2500 Hz, fm = 6 Hz) as a function of the relative phase of the 6-Hz modulators. Detection of FM improved monotonically as a function of increasing phase difference between the two modulators. The results are discussed in terms of modulation detection interference and perceptual grouping.

Adult↗

On the dependence of (f2-f1) difference tones on subject and on additional masker.

For a group of eight subjects showing stronger than usual irregularities in the level dependence of the quadratic distortion product, the level and phase of the (f2-f1) difference tone were measured using the method of cancellation for three sets of primary frequencies as a function of the primary levels. An additional masker seems to "linearize" the level dependence toward regular behavior. Using data sets produced with and without the additional masker, it is possible to separate two sources of quadratic nonlinearity, one with regular behavior presumably located in the middle ear and another with irregular behavior (similar to that of cubic distortion) presumably located in the characteristics of outer hair cells. Through the "subtraction" of empirically determined patterns from idealized patterns, it is possible to approximate patterns stemming from the inner-ear source alone.

Attention↗

Discrimination of modulation depth of sinusoidal amplitude modulation (SAM) noise.

The detection of sinusoidal amplitude modulation (SAM) provides a lower bound on the degree to which temporal information in the envelope of complex waveforms is encoded by the auditory system. The extent to which changes in the amount of modulation are discriminable provides additional information on the ability of the auditory system to utilize envelope fluctuations. Results from an experiment on the discrimination of modulation depth of broadband noise are presented. Discrimination thresholds, expressed as differences in modulation power, increase monotonically with the modulation depth of the standard, but do not obey Weber's law. The effects of carrier level and of modulation frequency are consistent with those observed in modulation detection: Changes in carrier level have little effect on modulation discrimination; changes in modulation frequency also have little effect except for standards near the modulation detection threshold. The discrimination of modulation depth is consistent with the leaky-integrator model of modulation detection for standards below--10 dB (20 log ms); for standards greater than--10 dB, the leaky integrator predicts better performance than that observed behaviorally.

Attention↗

Cochlear action potential tuning curves recorded with a derived response technique.

Previous action potential (AP) tuning curve methods have used a reduction in amplitude of the probe-elicited AP as an indication of tone-induced masking. The reduction criterion used in different studies has varied from 25% to 100%. For low level probe stimuli, which elicit a low-amplitude AP, this is a sensitive indicator. In contrast, for high-amplitude AP responses elicited by high-level stimuli, the required reduction in absolute terms is large, making it an insensitive indicator. AP tuning curves have been recorded using a sensitive method for detecting masker/probe interaction with a fixed criterion, unrelated to the unmasked AP amplitude. For each masking condition, a derived response was obtained by digitally subtracting the tone-masked AP waveform from the unmasked response. Derived responses are generated if there are ANY changes in the AP waveform induced by the masker, including amplitude changes, latency changes, or even changes in AP morphology not necessarily associated with the major peaks. A fixed criterion (10 microV) of tone-derived (TD) response was used as an indication of interaction of the responses to the masker and probe. Tuning curves generated by this method were compared with those generated by conventional amplitude reduction (AR) methods. TD tuning curves show different characteristics, especially with respect to increasing probe levels. They appear to give a good representation of the array of afferent fibers responding to a probe stimulus. In addition, frequency regions making minor contributions to the AP are better represented in TD tuning curves.

Animals↗

Response timing constraints on the cortical representation of sound time structure.

The precision of spike response timing of 94 primary auditory cortex neurons was studied using conventional extracellular recording techniques in barbiturate-anesthetized cats to which tone- and/or noise-burst stimuli were presented using sealed sound delivery systems. Precision of spike timing was indexed using the standard deviation of the first-spike latent period in responses evoked by repeated presentation of tonal stimuli systematically varied in frequency, amplitude, and/or repetition rate. Within a neuron, variability of first-spike timing was usually proportional to the mean first-spike latency, in agreement with previous reports. In cases where there was a systematic relation between the precision of response timing and the mean latency, a linear correlation accounted for up to 90% of the data variance. Across the 94 neurons, standard deviations seen in responses of minimum latency were related to minimal mean latencies, and were typically in the range from 0.15-1.5 ms. The data suggest that responses to transients in the cortex show a precision of spike timing which is only slightly worse than that seen in cochlear-nerve fibers. This, however, is in dramatic contrast to previous evidence on the steady-state temporal response of cortical cells, which is at least an order of magnitude poorer than that seen in auditory-nerve fibers and many cochlear nucleus cells. These observations may be directly relevant to the known consequences of auditory cortex pathology in man.

Animals↗

Time-domain analysis of auditory-nerve-fiber firing rates.

Time-domain analysis of firing-rate data from over 200 fibers from the auditory nerve of cat has been used to estimate the formants of the synthetic-syllable stimuli. Distinct groups of fibers are identified based on intervals between peaks in the fiber firing rates. The large extent of some of these groups--over an octave in terms of characteristic frequency--and the lack of short intervals in the longer-interval groups suggest that the behavior of the nonlinear cochlear filters for these signals is effectively wideband with steep high-frequency cutoffs. The measured intervals within each group are very similar, and correspond to the period of the formant that dominates the group's response. These intervals are used to estimate the dynamic speech formants. The overall formant estimates are better than those of the previous spectral analyses of the neural data, and the details of lower-formant dynamics are tracked more precisely. The direct temporal representation of the formant in contrasted with the diffuse spectral representation, the dependence of spectral peaks on nonformant parameters, and the distortion of the spectrum by rectification. It is concluded that a time-domain analysis of the responses to complex stimuli can be an important addition to frequency-domain analysis for neural data, cochlear models, and machine processing of speech.

Animals↗

Temporal responses of primarylike anteroventral cochlear nucleus units to the steady-state vowel /i/.

It has previously been shown that a population of units classified as "primarylike" and located in the ventral cochlear nucleus of the anesthetized guinea pig was unable to signal the position of the higher formant-related peaks (greater than 1.5 kHz) of steady-state vowels in terms of a temporal-place representation [Palmer et al., J. Acoust. Soc. Am. 79, 100-113 (1986)]. In this paper, it is demonstrated that units characterized by a prepotential in their spike waveform and a primarylike post-stimulus time histogram shape can encode the relative position of the formant peaks present in the spectra of steady-state vowels in terms of a temporal-place code. The possible reasons for the differences in the present results and the previous report are discussed.

Animals↗