Auditory brain stem responses from human adults and infants: wave V tuning curves

Decrement in ABR wave V amplitude was measured in the presence of simultaneous tonal maskers. Probe stimuli were 1.0, 4.0, and 8.0-kHz third-octave-filtered clicks. Adults and 3-month-old infants served as subjects. The resultant amplitude-decrement functions for each tonal masker were fit with regression lines. The sound pressure level (SPL) required to reduce wave V to 50% of the unmasked probe amplitude was plotted for each masker to develop tuning curves. The tuning curves were quantified by calculations of tip-to-tail difference, Q 10, and SPL at maximum masker frequency (MMF). Tuning curves for adult and infant subjects were similar for the 1.0-kHz probe. For the high-frequency probes (4.0 and 8.0 kHz), smaller tip-to-tail differences and lower Q 10 values were observed for the infant subjects. Ranges of MMF level were similar across adult and infant subjects. For the 8.0-kHz probe, tuning curves from infant subjects consistently showed maximum masker frequencies which were lower than the probe.     

Intermodulation distortion in the cochlea as shown by offset action potential (AP) masking curves

In their recent article "Offset AP masker tuning curve and the FFT of the stimulus" [J. Acoust. Soc. Am. 84, 1354-1362 (1988)], Henry and Lewis demonstrated that the tuning curve obtained by the simultaneous masking of the whole nerve action potential (AP) could have two tips when the AP is generated at the offset of the envelope of a high-level probe. The primary tip falls below the probe frequency, whereas the secondary tip falls above the probe frequency. Curves obtained for the onset response with either forward or simultaneous masking did not show the secondary peak, nor did curves obtained for the offset response with forward masking. Henry and Lewis discussed various reasons for the secondary tip, but came to no conclusion as to the underlying mechanisms. Here, it is reasoned that the secondary tip of the offset curve can be simply explained by the generation within the cochlea of intermodulation distortion (IMD), which acts as a forward masker to the offset response. The IMD is dominated by the cubic component (2f1-f2) and arises from the interaction of the probe tone and the simultaneous masker. Finally, it is reasoned that the lower sideband of the frequency splatter present at probe offset is the primary stimulus for the evoked neural response under probe offset conditions. Thus the offset curve will always have a primary tip that is lower in frequency than that of the respective onset curve. These hypotheses are supported by single-fiber data.     

Psychophysical tuning curves measured in simultaneous and forward masking

The level of a masker necessary to mask a probe fixed in frequency and level was determined as a function of masker frequency using a two-interval forced-choice technique. Both simultaneous- and forward- masking techniques were used. Parameters investigated include the level of the probe tone and the frequency of the probe tone. The general form of the psychophysical tuning curves obtained in this way is quite similar to that of single-neurone tuning curves, when low-level probe tones are used. However, the curves obtained to forward masking generally show sharper tips and steeper slopes than those found in simultaneous masking, and they are also generally sharper than neurophysiological tuning curves. For frequencies of the masker close to that of the probe a simultaneous masker was sometimes less effective than a forward masker. The results are discussed in relation to possible lateral suppression effects in simultaneous masking, and in relation to the observer's use of pitch cues in forward masking. It is concluded that neither the simultaneous-masking curves nor the forward-masking curves are likely to give an accurate representation of human neural tuning curves.     

The effects of cochlear hypothermia on compound action potential tuning

The effects of lowered cochlear temperature on eighth-nerve tuning were assessed by using forward masking of whole nerve action potential (AP) responses to generate AP tuning curves (APTCs) at cochlear temperatures ranging from 38.5 degrees to 30 degrees C for probe frequencies from 8 to 36 kHz. The data indicate that subnormal cochlear temperatures result in: broadened APTCs for probe frequencies above 10 kHz which are interpreted as resulting from reduced hair-cell frequency selectivity, lowered or more sensitive APTC tips where tone-burst thresholds are unchanged, and raised or less sensitive tips where thresholds to tone bursts were elevated. Increased tip sensitivity is explained in terms of enhanced eighth-nerve adaptation which occurred during hypothermia. Experiments directly addressing adaptation were performed, in which the masker-probe interval (delta t) was systematically lengthened. The normalized AP decrement versus delta t functions indicate an enhancement of both the amount and duration of adaptation during hypothermia. Functions relating the growth of response to the masker (AP decrement versus masker intensity functions) were reduced at low temperatures.     

Frequency and level dependent masking of the multiple auditory steady-state response in the bottlenose dolphin (Tursiops truncatus)

The potential for interactions between steady-state evoked responses to simultaneous auditory stimuli was investigated in two bottlenose dolphins (Tursiops truncatus). Three experiments were conducted using either a probe stimulus (probe condition) or a probe in the presence of a masker (probe-plus-masker condition). In the first experiment, the probe and masker were sinusoidal amplitude-modulated (SAM) tones. Probe and masker frequencies and masker level were manipulated to provide variable masking conditions. Probe frequencies were 31.7, 63.5, 100.8, and 127.0 kHz. The second experiment was identical to the first except only the 63.5 kHz probe was used and maskers were pure tones. For the third experiment, thresholds were measured for the probe and probe-plus-masker conditions using two techniques, one based on the lowest detectable response and the other based on a regression analysis. Results demonstrated localized masking effects where lower frequency maskers suppressed higher frequency probes and higher amplitude maskers produced a greater masking effect. The pattern of pure tone masking was nearly identical to SAM tone masking. The two threshold estimates were similar in low masking conditions, but in high masking conditions the lowest detectable response tended to overestimate thresholds while the regression-based analysis tended to underestimate thresholds.     

Offset AP masker tuning curve and the FFT of the stimulus

In previous experiments, it was noted that a cochlear compound action potential (CAP) can be produced by the offset of a tone, provided that the amplitude of the tone is modulated by a trapezoid with slopes that are typically much steeper than required to produce onset responses. Subsequently, such trapezoidal tone bursts with steep slopes were used as probe stimuli in simultaneous and forward masking experiments that were designed to evaluate the tuning characteristics of these offset CAPs. Masker tuning curves (MTCs) were generated by plotting the masker frequency necessary to reduce the amplitude of the offset CAP by 50%. Simultaneous masking of the offset CAP generated a W-shaped MTC, with two sharply tuned tips and one sharply tuned peak. Forward masking generated a sharply tuned V-shaped offset MTC. By contrast, for onset CAPs, both simultaneous and forward masking generated V-shaped MTCs. The very steep stimulus slopes required to produce an offset CAP are likely to generate much more acoustic splatter than the more gradual slopes required to produce an onset CAP, and this may be related to the different shapes of the onset and offset simultaneous MTCs. To explore this possibility, the relationship of the spectral characteristics (determined by fast Fourier transform, or FFT) to the shape of the MTC was studied.(ABSTRACT TRUNCATED AT 250 WORDS)     

Compound action potential thresholds and tuning curves in the alligator lizard

Compound action potential (AP) thresholds and tuning curves were measured in the alligator lizard using techniques similar to those used with the mammal. For tone burst stimuli, the AP was a major component of the gross response, the electrical response to sound recorded by a wire electrode near the cochlea. The AP thresholds agreed well with threshold estimates based on single-fiber data. A forward masking paradigm produced AP tuning curves which resemble single-fiber tuning curves. A simultaneous masking paradigm produced curves which match the boundaries of single-fiber two-tone rate suppression. The AP thresholds and tuning curves provide useful information about the peripheral physiology of the ear of the alligator lizard.     

The temporal evolution of masking and frequency selectivity in the goldfish (Carassius auratus)

The temporal evolution of masking and frequency selectivity was studied in the goldfish using classical respiratory conditioning and a tracking psychophysical procedure. The temporal position of a brief tonal signal within a longer duration, tonal masker has little or no effect on signal detectability when the frequency of the masker is less than or equal to that of the signal. For masker frequencies above that of the signal, signal detectability improves as the signal onset is delayed relative to that of the masker. These patterns of tone-on-tone masking are quite similar to those observed for humans. These temporal masking patterns are qualitatively similar in shape to the peristimulus-time histogram profiles of the low-frequency saccular fibers thought to be used in this task. Frequency- and time-dependent changes in signal detectability result in specific changes in the sharpness of psychophysical tuning curves (PTC). In general, PTCs determined for signals occurring at masker onset are the most broadly tuned, and PTCs determined in forward masking are the most sharply tuned. The PTCs for signals temporally centered in the masker are intermediate. These results suggest that temporal tone-on-tone masking patterns and the temporal evolution of psychophysical tuning curves result from the response properties of peripheral auditory-nerve fibers.     

Neural correlates of psychophysical release from masking

Responses of chinchilla auditory-nerve fibers were measured for stimulus conditions analogous to those in which psychophysical release from masking has been observed in humans. The maskers were two equal power, narrow-band noise stimuli with different amplitude envelopes. The neurons in the sample fell into three groups that resolved the maskers' envelopes with varying degrees of accuracy. The boundaries of these groups were not sharply delineated by characteristic frequency (CF) but were dependent on the relationship between the masker level and the neurons' thresholds at the masker frequency. For the neurons that best preserved the maskers' envelope fluctuations, a neural release from masking was observed; rate-based neural masked thresholds were higher for the masker with the least fluctuating envelope. The results suggest that neural and psychophysical release from masking arises because the probe evokes larger rate changes, relative to the background response to the masker, during periods of low masker energy. Between two otherwise equivalent maskers, the one with the periods of lowest energy will produce the lower masked thresholds because rate changes are larger and more detectable.     

Relationship between psychophysical tuning curves and "suppression"

This study examined two-tone unmasking and auditory frequency selectivity about 3 kHz for the purpose of demonstrating a qualitative relationship between the two. An adaptive 2IFC forward-masking procedure was used to collect psychophysical tuning curves (PTC's) and two-tone masking data under a quiet and noise condition for the same normal-hearing listeners. In the noise condition, a narrowband noise masker, centered one decade down from the probe, was gated on with the tonal masker(s). Kiang and Moxon [J. Acoust. Soc. Am. 55, 620-630 (1974)] have found that low-frequency narrowband noise serves to decrease the sharpness of electrophysiological tuning curves by affecting only the tip segments. The data for four highly practiced listeners indicate that the gated-noise masker was effective in broadening the PTC's and in lessening the magnitude of two-tone unmasking. The mutually reflected changes in tuning curves and in two-tone unmasking indicate a close relationship between frequency selectivity and unmasking: the greater the magnitude of unmasking above the center frequency of the PTC, the sharper the tuning of the PTC.     

A reexamination of forward masking in the auditory nerve

Forward masking, as measured behaviorally, is defined as an increase in a signal's detection threshold resulting from a preceding masker. Previously, forward masking in the auditory nerve has been measured as a reduction in the neural response to a signal when preceded by a masker. However, detection threshold depends on both the magnitude of the response to the signal and the variance of the response. Thus changes in detectability cannot be inferred from response reduction alone. Relkin and Pelli (1987) have described a two-interval forced-choice procedure that may be used to measure the threshold for the detection of a probe signal in recordings of spike counts in single auditory neurons. These methods have been used to study the forward masking of characteristic frequency probe tones by characteristic frequency maskers as masker intensity was varied. Although the masker does reduce the detectability of the probe tone, it was found that the threshold shifts are much less than those observed behaviorally, particularly for intense maskers. In part, the small threshold shifts can be attributed to the reduction in response variance following the masker, which is the result of the adaptation of spontaneous activity. These results imply that behavioral forward masking must result from suboptimal processing of spike counts from auditory neurons at a location central to the auditory nerve.     

Detection cues in forward masking and their relationship to off-frequency listening

Conventional and restricted-listening psychophysical tuning curve paradigms were used to evaluate the masking ability of forward maskers of different envelope characteristics, and the relationship between off-frequency listening effects and the envelope characteristics of both the variable masker and stationary masker used to restrict listening to a narrow region surrounding the probe. Maskers were selected so as to differ widely in the degree of fluctuation in their envelopes. The masker with the largest amplitude fluctuations exhibited greater forward-masking ability than other stimuli; this effect was observed on the high-frequency branch and within the tip region of the tuning curve. It is suggested that differences in masking ability may reflect the use of different signal detection cues. The results are also consistent with previous reports [D. Johnson-Davies and R.D. Patterson, J. Acoust. Soc. Am. 65, 756-770 (1979); B.J. O'Loughlin and B.C.J. Moore, J. Acoust. Soc. Am. 69, 1119-1125 (1981a)] which suggest that off-frequency listening is a factor contributing to the sharpness of psychophysical tuning curves. This effect is largely dependent, however, on the envelope characteristics of both variable and stationary maskers.     

Growth of masking as a measure of response growth in hearing-impaired listeners

Growth-of-masking functions were obtained from 19 normal and 5 hearing-impaired listeners using a simultaneous-masking paradigm. When masker and probe frequency are identical, the slope of masking approximates 1.0 for both normal-hearing and impaired listeners. For masker frequencies less than or greater than probe frequency, the slopes for impaired listeners are shallower than those of normals. These findings are consistent with previously reported physiological data (single-fiber rate versus level and AP masking functions) for animals with induced cochlear lesions. Results are discussed in terms of a potential masking technique to estimate the growth of response in normal and impaired ears.     

Effects of stimulus level on forward-masked psychophysical tuning curves in quiet and in noise

Forward-masked psychophysical tuning curves were obtained from normal-hearing listeners at different probe levels in quiet and in a broadband background noise. In quiet, tuning-curve shape changed with probe level. For six listeners, tuning curves became broader with increasing probe level, primarily due to a decrease in the low-frequency slopes. For one listener, tuning curves became narrower with increasing probe level. The addition of a background noise, which was presented continuously at a level 10 dB below the noise level required to mask the probe tone, reduced the masker levels required to mask the probe tone. The reduction was greater near the tip of the tuning curve than on the tail, so that tuning curves in background noise were narrower than those obtained in quiet. Tuning curves with comparable masker levels near the tip of the tuning curve (Lmtip) were similar in shape, regardless of probe level or whether tuning curves were obtained in quiet or noise. Comparisons of tuning-curve characteristics derived by fitting tuning curves with least-squares procedures, indicated that low-frequency slopes decreased with Lmtip. As a consequence, Q10 dB values decreased with Lmtip. These results are consistent with the interpretation that tuning-curve shapes are determined by the intensities of the maskers required to mask the probe tone. The addition of a background noise restricted (partially masked) the excitation pattern of the probe so that lower masker intensities were required to "forward mask" the probe tone, and narrower tuning curves resulted from less intense markers. The results are well described by a two-process model of auditory excitation patterns.     

A new procedure for measuring peripheral compression in normal-hearing and hearing-impaired listeners.

Forward-masking growth functions for on-frequency (6-kHz) and off-frequency (3-kHz) sinusoidal maskers were measured in quiet and in a high-pass noise just above the 6-kHz probe frequency. The data show that estimates of response-growth rates obtained from those functions in quiet, which have been used to infer cochlear compression, are strongly dependent on the spread of probe excitation toward higher frequency regions. Therefore, an alternative procedure for measuring response-growth rates was proposed, one that employs a fixed low-level probe and avoids level-dependent spread of probe excitation. Fixed-probe-level temporal masking curves (TMCs) were obtained from normal-hearing listeners at a test frequency of 1 kHz, where the short 1-kHz probe was fixed in level at about 10 dB SL. The level of the preceding forward masker was adjusted to obtain masked threshold as a function of the time delay between masker and probe. The TMCs were obtained for an on-frequency masker (1 kHz) and for other maskers with frequencies both below and above the probe frequency. From these measurements, input/output response-growth curves were derived for individual ears. Response-growth slopes varied from >1.0 at low masker levels to <0.2 at mid masker levels. In three subjects, response growth increased again at high masker levels (>80 dB SPL). For the fixed-level probe, the TMC slopes changed very little in the presence of a high-pass noise masking upward spread of probe excitation. A greater effect on the TMCs was observed when a high-frequency cueing tone was used with the masking tone. In both cases, however, the net effects on the estimated rate of response growth were minimal.     

Temporal factors associated with cochlear nerve tuning to dual and single tones: a qualitative study.

The simultaneous presentation of a 10- and 10.86-kHz tone produces an 860-Hz cochlear nerve difference tone (DT) response in the gerbil which persists for the duration of the stimulus. Forward masking shows this response is generated by neurons sharply tuned to the stimulus frequencies. When compared with the DT response, the cochlear nerve compound action potential (CAP) to a single tone is smaller in amplitude, has a higher nonmasked threshold, and produces a less sensitive tuning curve (TC). Forward maskers can also produce amplitude enhancement of the CAP, but this was not observed for the onset portion of the DT response. The CAP TC is as sharply tuned as the TC of either the DT onset response or the entire DT response. A comparison was made of tuning of the DT response to the onset, the first half and second half of the 23-ms duration probe stimulus, using either a 5- or 15-ms masker-probe interval. An increase of the tip threshold of the TC to all three portions of the stimulus occurred as the interval was increased between the end of the masker and the midpoint of the portion of the stimulus under question. The 15-ms masker-probe interval produced sharper TCs.     

Sinusoidal and noise maskers in simultaneous and forward masking

We compare psychophysical tuning curves obtained with sinusoidal and narrow-band (50-Hz wide) noise maskers in both simultaneous and forward masking. In one experiment, we examine the effects of different combinations of duration and intensity of the 1-kHz sinusoidal signal. In a second experiment, we compare tuning curves obtained with a sinusoidal signal to those obtained with a noise signal. In both experiments, a narrow-band noise is a more effective simultaneous masker than a sinusoid for masker frequencies near the signal frequency. We argue that this is probably due to the use of different detection cues in the presence of sinusoidal and noise maskers, and that the greater masking produced by the noise is not simply due to its greater variability. As observed in other studies, tuning curves are narrower in forward masking than in simultaneous masking.     

Modulation masking produced by beating modulators.

This study examined whether "modulation masking" could be produced by temporal similarity of the probe and masker envelopes, even when the masker envelope did not contain a spectral component close to the probe frequency. Both masker and probe amplitude modulation were applied to a single 4-kHz sinusoidal or narrow-band noise carrier with a level of 70 dB SPL. The threshold for detecting 5-Hz probe modulation was affected by the presence of a pair of masker modulators beating at a 5-Hz rate (40 and 45 Hz, 50 and 55 Hz, or 60 and 65 Hz). The threshold was dependent on the phase of the probe modulation relative to the beat cycle of the masker modulators; the threshold elevation was greatest (12-15 dB for the sinusoidal carrier and 9-11 dB for the noise carrier, expressed as 20 log m) when the peak amplitude of the probe modulation coincided with a peak in the beat cycle. The maximum threshold elevation of the 5-Hz probe produced by the beating masker modulators was 7-12 dB greater than that produced by the individual components of the masker modulators. The threshold elevation produced by the beating masker modulators was 2-10 dB greater for 5-Hz probe modulation than for 3- or 7-Hz probe modulation. These results cannot be explained in terms of the spectra of the envelopes of the stimuli, as the beating masker modulators did not produce a 5-Hz component in the spectra of the envelopes. The threshold for detecting 5-Hz probe modulation in the presence of 5-Hz masker modulation varied with the relative phase of the probe and masker modulation. The pattern of results was similar to that found with the beating two-component modulators, except that thresholds were highest when the masker and probe were 180 degrees out of phase. The results are consistent with the idea that nonlinearities within the auditory system introduce distortion in the internal representation of the envelopes of the stimuli. In the case of two-component beating modulators, a weak component is introduced at the beat rate, and it has an amplitude minimum when the beat cycle is at its maximum. The results could be fitted well using two models, one based on the concept of a sliding temporal integrator and one based on the concept of a modulation filter bank.     

Auditory brain stem responses from human adults and infants: restriction of frequency contribution by notched-noise masking

The frequency contribution to the click-evoked ABR wave V was examined in adults and 3-month-old infants through the use of notch-filtered broadband noise. Notch center frequencies were set at 1.0, 4.0, and 8.0 kHz. Responses were obtained at 20, 40, and 60 dBnHL during the simultaneous presentation of each notched-noise masker as well as in an unmasked condition. The ABR wave V was analyzed for absolute latency and amplitude, as well as latency and amplitude changes resulting from the introduction of masking. Analyses showed wave V latency and amplitude values to be similar for adults and infants within the 1.0-kHz notch. Differences between adult and infant groups were observed as the notch was shifted to the high frequencies. Further, latency and amplitude shifts resulting from the introduction of masking noise produced differential effects on infant responses when compared to adults.     

Frequency specificity of chirp-evoked auditory brainstem responses

This study examines the usefulness of the upward chirp stimulus developed by Dau et al. [J. Acoust. Soc. Am. 107, 1530-1540 (2000)] for retrieving frequency-specific information. The chirp was designed to produce simultaneous displacement maxima along the cochlear partition by compensating for frequency-dependent traveling-time differences. In the first experiment, auditory brainstem responses (ABR) elicited by the click and the broadband chirp were obtained in the presence of high-pass masking noise, with cutoff frequencies of 0.5, 1, 2, 4, and 8 kHz. Results revealed a larger wave-V amplitude for chirp than for click stimulation in all masking conditions. Wave-V amplitude for the chirp increased continuously with increasing high-pass cutoff frequency while it remains nearly constant for the click for cutoff frequencies greater than 1 kHz. The same two stimuli were tested in the presence of a notched-noise masker with one-octave wide spectral notches corresponding to the cutoff frequencies used in the first experiment. The recordings were compared with derived responses, calculated offline, from the high-pass masking conditions. No significant difference in response amplitude between click and chirp stimulation was found for the notched-noise responses as well as for the derived responses. In the second experiment, responses were obtained using narrow-band stimuli. A low-frequency chirp and a 250-Hz tone pulse with comparable duration and magnitude spectrum were used as stimuli. The narrow-band chirp elicited a larger response amplitude than the tone pulse at low and medium stimulation levels. Overall, the results of the present study further demonstrate the importance of considering peripheral processing for the formation of ABR. The chirp might be of particular interest for assessing low-frequency information.