Noise susceptibility and immunity of phase locking in amphibian auditory-nerve fibers

Recordings from auditory-nerve fibers in the anesthetized frog revealed that addition of broadband noise results in a reduction in the ability of a fiber to phase lock to a continuous pure tone. In particular, our results suggest that: (i) there is a threshold below which masking noise has little or no effect on vector strength (VS); then with increasing masking noise level, VS appears to decrease monotonically for all test frequencies (TFs); (ii) there exist subpopulations of auditory-nerve fibers in the frog for which the deterioration of phase locking to tones in wideband noise depends critically on the relationship of the TF to the fiber's CF. Specifically, in one subpopulation (43% of the fibers studied), the rate of VS decrease with increasing levels of masking noise is greater for CF tones than it is for TFs greater than CF. The net result is a "crossing" of the VS versus masking noise functions (e.g., Fig. 6); (iii) there exists a small subpopulation of amphibian papillar (a.p.) fibers for which the rate of VS decrease with increasing levels of masking noise is less for TFs less than CF than it is for CF tones (e.g., Fig. 5); (iv) there is a pronounced noise-induced phase lead for TFs greater than CF, whereas, for stimulus tones at or below CF, the preferred firing phase is nearly noise-level independent; (v) the remainder of the sample consists of fibers in which the VS-falloff rates appear to be test-frequency independent; (vi) addition of wideband masking noise to a CF tone, and increasing the CF-tone level in the absence of noise, produced (qualitatively) similar effects on the preferred firing phase of auditory-nerve fibers (e.g., Figs. 1 and 7). Thus amphibian auditory-nerve fibers appear to be energy detectors, i.e., exhibit phase shifts corresponding to the total energy within the filter passband defined by the frequency-threshold curve.     

Responses to amplitude-modulated tones in the auditory nerve of the cat.

Sinusoidally amplitude-modulated (AM) tones are frequently used in psychophysical and physiological studies, yet a comprehensive study on the coding of AM tones in the auditory nerve is lacking. AM responses of single auditory-nerve fibers of the cat are studied, systematically varying modulation depth, frequency, and sound level. Synchrony-level functions were nonmonotonic with maximum values that were inversely correlated with spontaneous rate (SR). In most fibers, envelope phase-locking showed a positive gain. Modulation transfer functions were uniformly low pass. Their corner frequency increased with characteristic frequency (CF), but changed little for CFs above 10 kHz. The highest modulation frequencies to which phase locking occurred were more than 0.8 oct lower than the highest frequencies to which phase locking to pure tones occurs. Cumulative, or unwrapped, phase increased linearly with modulation frequency: The slope was inversely related to CF, and slightly higher than group delays reported for pure tones. High SR, low CF fibers showed the poorest envelope phase locking. In some low CF fibers, phase locking increased at high levels, associated with "peak-splitting" phenomena. Changes in average rate due to modulation were small, and could be enhancement or suppression.     

Asymmetry of masking between complex tones and noise: partial loudness

This experiment examined the partial masking of periodic complex tones by a background of noise, and vice versa. The tones had a fundamental frequency (F0) of 62.5 or 250 Hz, and components were added in either cosine phase (CPH) or random phase (RPH). The tones and the noise were bandpass filtered into the same frequency region, from the tenth harmonic up to 5 kHz. The target alone was alternated with the target and the background; for the mixture, the background and target were either gated together, or the background was turned on 400 ms before, and off 200 ms after, the target. Subjects had to adjust the level of either the target alone or the target in the background so as to match the loudness of the target in the two intervals. The overall level of the background was 50 dB SPL, and loudness matches were obtained for several fixed levels of the target alone or in the background. The resulting loudness-matching functions showed clear asymmetry of partial masking. For a given target-to-background ratio, the partial loudness of a complex tone in a noise background was lower than the partial loudness of a noise in a complex tone background. Expressed as the target-to-background ratio required to achieve a given loudness, the asymmetry typically amounted to 12-16 dB. When the F0 of the complex tone was 62.5 Hz, the asymmetry of partial masking was greater for CPH than for RPH. When the F0 was 250 Hz, the asymmetry was greater for RPH than for CPH. Masked thresholds showed the same pattern as for partial masking for both F0's. Onset asynchrony had some effect on the loudness matching data when the target was just above its masked threshold, but did not significantly affect the level at which the target in the background reached its unmasked loudness. The results are interpreted in terms of the temporal structure of the stimuli.     

Transient-evoked stimulus-frequency and distortion-product otoacoustic emissions in normal and impaired ears

Transient-evoked stimulus-frequency otoacoustic emissions (SFOAEs), recorded using a nonlinear differential technique, and distortion-product otoacoustic emissions (DPOAEs) were measured in 17 normal-hearing and 10 hearing-impaired subjects using pairs of tone pips (pp), gated tones (gg), and for DPOAEs, continuous and gated tones (cg). Temporal envelopes of stimulus and OAE waveforms were obtained by narrow-band filtering at the stimulus or DP frequency. Mean SFOAE latencies in normal ears at 2.7 and 4.0 kHz decreased with increasing stimulus level and were larger at 4.0 kHz than latencies in impaired ears. Equivalent auditory filter bandwidths were calculated as a function of stimulus level from SFOAE latencies by assuming that cochlear transmission is minimum phase. DPOAE latencies varied less with level than SFOAE latencies. The ppDPOAEs often had two (or more) peaks separated in time with latencies consistent with model predictions for distortion and reflection components. Changes in ppDPOAE latency with level were sometimes explained by a shift in relative amplitudes of distortion and reflection components. The pp SFOAE SPL within the main spectral lobe of the pip stimulus was higher for normal ears in the higher-frequency half of the pip than the lower-frequency half, which is likely an effect of basilar membrane two-tone suppression.     

Effects of flanking noise bands on the rate of growth of loudness of tones in normal and recruiting ears

Five subjects with unilateral cochlear hearing impairments and three normally hearing subjects made loudness matches between tones presented alternately to two ears, as a function of the intensity of the tone in the impaired ear (or the left ear of the normal subjects). The impaired ears showed recruitment; the rate of growth of loudness with increasing intensity was more rapid in the impaired ear than the normal ear. Presenting the tone in the impaired ear with two noise bands on either side of the tone frequency, at a fixed signal-to-noise ratio, did not abolish the recruitment. This suggests that recruitment is not caused by an abnormally rapid spread of excitation in the peripheral auditory system. At low signal-to-noise ratios, a continuous background noise reduced the loudness of the tone more than a noise gated with the tone, suggesting that the continuous noise induces adaptation to the tone. The noise had a greater effect on the loudness of the tone in normal ears than in impaired ears. It is possible that the loudness reduction of the tone in noise is mediated by suppression; suppression is weak or absent in impaired ears, and so the loudness reduction is smaller.     

Asymmetry of masking between complex tones and noise: the role of temporal structure and peripheral compression

Thresholds for the detection of harmonic complex tones in noise were measured as a function of masker level. The rms level of the masker ranged from 40 to 70 dB SPL in 10-dB steps. The tones had a fundamental frequency (F0) of 62.5 or 250 Hz, and components were added in either cosine or random phase. The complex tones and the noise were bandpass filtered into the same frequency region, from the tenth harmonic up to 5 kHz. In a different condition, the roles of masker and signal were reversed, keeping all other parameters the same; subjects had to detect the noise in the presence of a harmonic tone masker. In both conditions, the masker was either gated synchronously with the 700-ms signal, or it started 400 ms before and stopped 200 ms after the signal. The results showed a large asymmetry in the effectiveness of masking between the tones and noise. Even though signal and masker had the same bandwidth, the noise was a more effective masker than the complex tone. The degree of asymmetry depended on F0, component phase, and the level of the masker. The maximum difference between masked thresholds for tone and noise was about 28 dB; this occurred when the F0 was 62.5 Hz, the components were in cosine phase, and the masker level was 70 dB SPL. In most conditions, the growth-of-masking functions had slopes close to 1 (on a dB versus dB scale). However, for the cosine-phase tone masker with an F0 of 62.5 Hz, a 10-dB increase in masker level led to an increase in masked threshold of the noise of only 3.7 dB, on average. We suggest that the results for this condition are strongly affected by the active mechanism in the cochlea.     

Neuronal responses to amplitude-modulated and pure-tone stimuli in the guinea pig inferior colliculus, and their modification by broadband noise

Neuronal responses were recorded to pure and to sinusoidally amplitude-modulated (AM) tones at the characteristic frequency (CF) in the central nucleus of the inferior colliculus of anesthetized guinea pigs. Temporal (synchronized) and mean-rate measures were derived from period histograms locked to the stimulus modulation waveform to characterize the modulation response. For stimuli presented in quiet, the modulation gain at low frequencies of modulation (approx less than 50 Hz) was inversely proportional to the neuron's mean firing rate in response to both the modulated stimulus and to a pure tone at an equivalent level. In 43% of units the mean discharge rates in response to the AM stimuli were greatest for those modulation frequencies that generated the largest temporal responses. These discharge-rate maxima occurred at signal intensities corresponding to the steeply sloping part of the neuron's pure-tone rate-intensity function (RIF). The change in mean-rate response to modulated stimuli, as a function of intensity, was qualitatively similar to the pure-tone RIF. Adding broadband noise to the modulated stimulus increased the neuron's temporal response to low modulation frequencies. This increase in modulation gain was correlated with mean firing rate in response to the modulation but did not bear a simple relationship to the noise-induced shift in the RIF measured for a pure tone.     

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.     

Rate responses of auditory nerve fibers to tones in noise near masked threshold

The rate responses of auditory nerve fibers were measured for best frequency (BF) tone bursts in the presence of continuous background noise. Rate functions for BF tones were constructed over a 32-dB range of levels, centered on the behavioral masked thresholds of cats. The tone level at which noticeable rate changes are evoked by the tones corresponds closely to behavioral masked threshold at all noise levels used (-10- to 30-dB spectrum level). As the noise level increases, the response rate to the background noise approaches saturation, and the incremental rate response to tones decreases. At high noise levels, the rate responses to tones of low and medium spontaneous rate fibers are larger than those of high spontaneous rate fibers. Empirical statistics of auditory nerve fiber spike counts are reported; these differ from those expected of a Poisson process in that the variance is smaller than the mean. A new measure of discharge rate is described that allows rate changes to be expressed in units of a standard deviation. This measure allows tone-evoked responses to be interpreted in terms of their detectability in a signal detection task. Rate responses of low and medium spontaneous rate fibers are more detectable than those of high spontaneous rate fibers, especially at high noise levels. There appears to be sufficient information in the rate response of a small number of auditory nerve fibers to support behaviorally observed levels of detection performance.     

Intensity discrimination, increment detection, and magnitude estimation for 1-kHz tones

Intensity difference limens (DLs) were measured over a wide intensity range for 200-ms, 1-kHz gated tones and for 200-ms increments in continuous 1-kHz tones. Magnitude estimates also were obtained for the gated tones over a comparable intensity range. The discrimination data are in general agreement with those from earlier studies but they extend them by showing: (1) good discrimination for gated tones over at least a 115-dB dynamic range; (2) a slight increase in the relative DL (delta I/I) as intensity increases above 95 dB SPL; (3) smaller DLs for increments than for gated tones, with the difference approximately independent of intensity; (4) negligible "negative masking" when thresholds are expressed as intensity differences (delta I). For two of the three subjects, magnitude estimates do not conform to a single-exponent power law for suprathreshold intensities. Over the middle range of intensities where a single exponent is appropriate, the value of the exponent is less than 0.1 for all subjects.     

Across-frequency interference effects in fundamental frequency discrimination: questioning evidence for two pitch mechanisms

Carlyon and Shackleton [J. Acoust. Soc. Am. 95, 3541-3554 (1994)] presented an influential study supporting the existence of two pitch mechanisms, one for complex tones containing resolved and one for complex tones containing only unresolved components. The current experiments provide an alternative explanation for their finding, namely the existence of across-frequency interference in fundamental frequency (F0) discrimination. Sensitivity (d') was measured for F0 discrimination between two sequentially presented 400 ms complex (target) tones containing only unresolved components. In experiment 1, the target was filtered between 1375 and 15,000 Hz, had a nominal F0 of 88 Hz, and was presented either alone or with an additional complex tone ("interferer"). The interferer was filtered between 125-625 Hz, and its F0 varied between 88 and 114.4 Hz across blocks. Sensitivity was significantly reduced in the presence of the interferer, and this effect decreased as its F0 was moved progressively further from that of the target. Experiment 2 showed that increasing the level of a synchronously gated lowpass noise that spectrally overlapped with the interferer reduced this "pitch discrimination interference (PDI)". In experiment 3A, the target was filtered between 3900 and 5400 Hz and had an F0 of either 88 or 250 Hz. It was presented either alone or with an interferer, filtered between 1375 and 1875 Hz with an F0 corresponding to the nominal target F0. PDI was larger in the presence of the resolved (250 Hz F0) than in the presence of the unresolved (88 Hz F0) interferer, presumably because the pitch of the former was more salient than that of the latter. Experiments 4A and 4B showed that PDI was reduced but not eliminated when the interferer was gated on 200 ms before and off 200 ms after the target, and that some PDI was observed with a continuous interferer. The current findings provide an alternative interpretation of a study supposedly providing strong evidence for the existence of two pitch mechanisms.     

Louder sounds can produce less forward masking: effects of component phase in complex tones

The influence of the degree of envelope modulation and periodicity on the loudness and effectiveness of sounds as forward maskers was investigated. In the first experiment, listeners matched the loudness of complex tones and noise. The tones had a fundamental frequency (F0) of 62.5 or 250 Hz and were filtered into a frequency range from the 10th harmonic to 5000 Hz. The Gaussian noise was filtered in the same way. The components of the complex tones were added either in cosine phase (CPH), giving a large crest factor, or in random phase (RPH), giving a smaller crest factor. For each F0, subjects matched the loudness between all possible stimulus pairs. Six different levels of the fixed stimulus were used, ranging from about 30 dB SPL to about 80 dB SPL in 10-dB steps. Results showed that, at a given overall level, the CPH and the RPH tones were louder than the noise, and that the CPH tone was louder than the RPH tone. The difference in loudness was larger at medium than at low levels and was only slightly reduced by the addition of a noise intended to mask combination tones. The differences in loudness were slightly smaller for the higher than for the lower F0. In the second experiment, the stimuli with the lower F0s were used as forward maskers of a 20-ms sinusoid, presented at various frequencies within the spectral range of the maskers. Results showed that the CPH tone was the least effective forward masker, even though it was the loudest. The differences in effectiveness as forward maskers depended on masker level and signal frequency; in order to produce equal masking, the level of the CPH tone had to be up to 35 dB above that of the RPH tone and the noise. The implications of these results for models of loudness are discussed and a model is presented based on neural activity patterns in the auditory nerve; this predicts the general pattern of loudness matches. It is suggested that the effects observed in the experiments may have been influenced by two factors: cochlear compression and suppression.     

Intensity discrimination and increment detection in cochlear-implant users

Intensity difference limens (DLs) were measured in users of the Nucleus 22 and Clarion v1.2 cochlear implants and in normal-hearing listeners to better understand mechanisms of intensity discrimination in electric and acoustic hearing and to evaluate the possible role of neural adaptation. Intensity DLs were measured for three modes of presentation: gated (intensity increments gated synchronously with the pedestal), fringe (intensity increments delayed 250 or 650 ms relative to the onset of the pedestal), and continuous (intensity increments occur in the presence of a pedestal that is played throughout the experimental run). Stimuli for cochlear-implant listeners were trains of biphasic pulses; stimuli for normal-hearing listeners were a 1-kHz tone and a wideband noise. Clarion cochlear-implant listeners showed level-dependent effects of presentation mode. At low pedestal levels, gated thresholds were generally similar to thresholds obtained in the fringe and continuous conditions. At higher pedestal levels, however, the fringe and continuous conditions produced smaller intensity DLs than the gated condition, similar to the gated-continuous difference in intensity DLs observed in acoustic hearing. Nucleus cochlear-implant listeners did not show consistent threshold differences for the gated and fringe conditions, and were not tested in the continuous condition. It is not clear why a difference between gated and fringe thresholds occurred for the Clarion but not the Nucleus subjects. Normal-hearing listeners showed improved thresholds for the continuous condition relative to the gated condition, but the effect was larger for the 1-kHz tonal carrier than for the noise carrier. Findings suggest that adaptation occurring central to the inner hair cell synapse mediates the gated-continuous difference observed in Clarion cochlear-implant listeners and may also contribute to the gated-continuous difference in acoustic hearing.     

Effects of signal envelope on the pitch of short sinusoidal tones

The pitch of short sinusoidal tones with exponentially rising or decaying envelopes is judged higher than the pitch of a gated tone of the same frequency, duration, and energy. The upward pitch shift depends on the rise or decay rate, the intensity, and the frequency. The effect, which requires a nonlinearity in the auditory system, cannot be adequately explained by existing models of hearing. Control experiments on pitch matching for short tones of varying duration and varying intensity are described. These suggest that envelope-induced pitch effects are linked to changes in average intensity, so that they are essentially the same as intensity-induced pitch changes. A model based on these considerations is proposed.     

Comparison of behavioral and auditory brainstem response measures of threshold shift in rats exposed to loud sound

The purpose of this study was to determine how closely the auditory brainstem response (ABR) can estimate sensorineural threshold shifts in rats exposed to loud sound. Behavioral and ABR thresholds were obtained for tones or noise before and after exposure to loud sound. The results showed that the ABR threshold shift obtained with tone pips estimated the initial pure-tone threshold shifts to within +/-5 dB 11% of the time and the permanent pure-tone threshold shifts 55% of the time, both with large errors. Determining behavioral thresholds for the same tone pips used for the ABR did not improve the agreement between the measures. In contrast, the ABR obtained with octave noise estimated the initial threshold shifts for that noise to within +/-5 dB 25% of the time and the permanent threshold shifts 89% of the time, with much smaller errors. Thus, it appears that the noise-evoked ABR is more accurate in estimating threshold shift than the tone-evoked ABR.     

Masking patterns in the bullfrog (Rana catesbeiana). II: Physiological effects

Responses of individual eighth-nerve fibers in the bullfrog (Rana catesbeiana) were measured to tone bursts at best frequency against a background of continuous, broadband masking noise. These data were used to calculate critical masking ratios to describe the fibers' responses to tones embedded in noise. In the frequency response range of the amphibian papilla (100-1000 Hz), critical ratios increase with tone frequency. Critical ratios of basilar papilla fibers (1000-2000 Hz) are generally higher than those of amphibian papilla fibers. Critical ratios are also significantly related to fiber threshold such that fibers with high thresholds, regardless of their best frequencies, have higher critical ratios and are thus less selective to signals embedded in noise. Critical ratios based on neural responses show a somewhat different frequency-dependent trend than do critical ratios based on psychophysical data presented previously for this species [A. M. Simmons, J. Acoust. Soc. Am. 83, 1087-1092 (1988a)]. In addition, these neural critical ratios do not appear to be level independent, as are psychophysical critical ratios. The data suggest that frequency selectivity of hearing in the bullfrog as measured behaviorally is probably not mediated solely by spectral filtering in the auditory periphery.     

The effect of amplitude envelope on the pitch of sine wave tones

Psychophysical experiments show that the pitch of a short sine wave tone depends upon the amplitude envelope of the tone. Subjects find that the pitch of an exponentially decaying tone (1dB/ms) is higher than the pitch of a (20-ms) rectangularly gated tone of equal frequency. The percentage difference in frequency required to produce equal pitches with the two envelopes depends upon frequency fo: 2.6% at fo = 412 Hz, 1.4% at fo = 825 Hz, 1% at fo = 1650 Hz, and 0.7% at fo = 3300 Hz. The pitch change is insensitive to the relative intensities of the two tones. The spectra of tones with the two different envelopes suggest no obvious explanation for the pitch change. However, the weighted time-varying spectra for tones with two different envelopes evolve differently with time. Alternatively the pitch change can be derived from a modified version of the auditory phase theory of Huggins.     

Monaural and interaural intensity discrimination: level effects and the "binaural advantage"

This study examined whether the level effects seen in monaural intensity discrimination (Weber's law and the "near miss") in a two-interval task are also observed in discrimination of interaural intensity differences (IIDs) in a single-interval task. Both tasks were performed for various standard levels of 4-kHz pure tones and broadband noise. The Weber functions (10 log deltaI/I versus I in dB) in the monaural and binaural conditions were parallel. For noise, the Weber functions had slopes close to zero (Weber's law) while the Weber functions for the tones had a mean slope of -0.089 (near miss). The near miss for the monaural and binaural tasks with tones was eliminated when a high-pass masker was gated with the listening intervals. The near-miss was also observed for 250- and 1000-Hz tones in the binaural task despite overall decreased sensitivity to changes in IID at 1000 Hz. The binaural thresholds showed a small (about 2-dB) advantage over monaural thresholds only in the broadband noise conditions. More important, however, is the fact that the level effects seen monaurally are also seen binaurally. This suggests that the basic mechanisms responsible for Weber's law and the near miss are common to monaural and binaural processing.     

Interactions between test- and inducer-tone durations in induced loudness reduction

A tone usually declines in loudness when preceded by a more intense inducer tone. This phenomenon is called "loudness recalibration" or "induced loudness reduction" (ILR). The present study investigates how ILR depends on level, loudness, and duration. A 2AFC procedure was used to obtain loudness matches between 2500-Hz comparison tones and 500-Hz test tones at 60 and 70 dB SPL, presented with and without preceding 500-Hz inducer tones. For 200-ms test and comparison tones, the amount of ILR did not depend on inducer level (set at 80 dB SPL and above), but ILR was greater with 200- than with 5-ms inducers, even when both were equally loud. For 5-ms tones, ILR was as great with 5- as with 200-ms inducers and about as great as when test and inducer tones both lasted 200 ms. These results suggest that (1) neither the loudness nor the SPL of the inducer alone governs ILR, and (2) inducer duration must equal or exceed test-tone duration to yield maximal amounts of ILR. Further analysis indicates that the efferent system may be partly responsible for ILR of 200-ms test tones, but is unlikely to account for ILR of 5-ms tones.     

The influence of linguistic and musical experience on Cantonese word learning

Adult non-native speech perception is subject to influence from multiple factors, including linguistic and extralinguistic experience such as musical training. The present research examines how linguistic and musical factors influence non-native word identification and lexical tone perception. Groups of native tone language (Thai) and non-tone language listeners (English), each subdivided into musician and non-musician groups, engaged in Cantonese tone word training. Participants learned to identify words minimally distinguished by five Cantonese tones during training, also completing musical aptitude and phonemic tone identification tasks. First, the findings suggest that either musical experience or a tone language background leads to significantly better non-native word learning proficiency, as compared to those with neither musical training nor tone language experience. Moreover, the combination of tone language and musical experience did not provide an additional advantage for Thai musicians above and beyond either experience alone. Musicianship was found to be more advantageous than a tone language background for tone identification. Finally, tone identification and musical aptitude scores were significantly correlated with word learning success for English but not Thai listeners. These findings point to a dynamic influence of musical and linguistic experience, both at the tone dentification level and at the word learning stage.