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.     

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.     

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)     

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.     

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.     

Interpreting measures of frequency selectivity: is forward masking special?

In a previous article [Lutfi, J. Acoust. Soc. Am. 76, 1045-1050 (1984)], the following relation was used to predict measures of frequency selectivity obtained in forward masking from measures obtained in simultaneous masking: F(g) = G + H(g) - H(0), where, for a given masker level, F is the amount of forward masking (in dB) as a function of signal-masker frequency separation (g), H is the amount of simultaneous masking, and G is the amount of forward masking for g = 0. In the present study, the relation was tested for a wider range of signal and masker frequencies, masker levels, and signal delays. The relation described thresholds from all conditions well with the inclusion of one free parameter lambda corresponding to a constant frequency increment, F(g) = G + H(g + lambda) - H(lambda). The parameter lambda was required to account for observed shifts in the frequency of maximum forward masking. It is argued that a single tuning mechanism can account for commonly observed differences between simultaneous- and forward-masked measures of frequency selectivity.     

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.     

Psychophysical estimates of level-dependent best-frequency shifts in the apical region of the human basilar membrane

It is now undisputed that the best frequency (BF) of basal basilar-membrane (BM) sites shifts downwards as the stimulus level increases. The direction of the shift for apical sites is, by contrast, less well established. Auditory nerve studies suggest that the BF shifts in opposite directions for apical and basal BM sites with increasing stimulus level. This study attempts to determine if this is the case in humans. Psychophysical tuning curves (PTCs) were measured using forward masking for probe frequencies of 125, 250, 500, and 6000 Hz. The level of a masker tone required to just mask a fixed low-level probe tone was measured for different masker-probe time intervals. The duration of the intervals was adjusted as necessary to obtain PTCs for the widest possible range of masker levels. The BF was identified from function fits to the measured PTCs and it almost always decreased with increasing level. This result is inconsistent with most auditory-nerve observations obtained from other mammals. Several explanations are discussed, including that it may be erroneous to assume that low-frequency PTCs reflect the tuning of apical BM sites exclusively and that the inherent frequency response of the inner hair cell may account for the discrepancy.     

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.     

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.     

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.     

Comparisons of frequency selectivity in simultaneous and forward masking for subjects with unilateral cochlear impairments

Two experiments are described in which frequency selectivity was estimated, in simultaneous and forward masking, for each ear of subjects with moderate (25-60 dB HL) unilateral cochlear hearing losses. In both experiments, the signal level was fixed for a given ear and type of masking (simultaneous or forward), and the masker level was varied to determine threshold, using an adaptive, two-alternative forced-choice procedure. In experiment I, the masker was a noise with a spectral notch centered at the signal frequency (either 1.0 or 1.5 kHz); threshold was determined as a function of notch width. Signal levels were chosen so that the noise level required at threshold for a notch width of zero was similar for the normal and impaired ear of each subject in both simultaneous and forward masking. The function relating threshold to notch width had a steeper slope for the normal ear than for the impaired ear of each subject. For the normal ears, these functions were steeper in forward masking than in simultaneous masking. This difference was interpreted as resulting from suppression. For the impaired ears, significant differences in the same direction were observed for three of the five subjects, but the differences were smaller. In experiment II, psychophysical tuning curves (PTCs) were determined in the presence of a fixed notched noise centered at the signal frequency (1.0 kHz). For the normal ears, the PTCs were sharper in forward masking than in simultaneous masking. For the impaired ears, the PTCs were similar in simultaneous and forward masking, but those in forward masking tended to be sharper at masker frequencies far removed from the signal frequency. Overall, the results suggest that suppression is reduced, but not completely absent in cases of moderate cochlear hearing loss.     

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.     

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.     

Temporal effects in masking and their influence on psychophysical tuning curves

Psychophysical tuning curves (PTCs) were obtained in simultaneous and forward masking for a 20-ms, 1000-Hz signal presented at 10 dB SL. The signal was presented at the beginning of, at the temporal center of, at the end of, or immediately following a 400-ms masker. The first experiment was done in quiet; the second experiment was done in the presence of two bands of noise on either side of 1000 Hz. The results were similar in quiet and in noise. In simultaneous masking, the PTCs were broadest for the signal at masker onset, and generally sharpest for the signal at temporal center; the differences were largest on the high-frequency side. In most cases, there was virtually no difference in Q10 between the forward-masking PTC and the simultaneous-masking PTC with the signal temporally centered, although the high-frequency slope was always steeper in forward masking. These results indicate that, at least for brief signals, frequency selectivity measured with simultaneous-masking PTCs and the degree of sharpening revealed in forward-masking PTCs depend upon the temporal position of the signal within the simultaneous masker.     

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.     

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.     

Forward masking of amplitude modulation: basic characteristics

In this study we demonstrate an effect for amplitude modulation (AM) that is analogous to forward making of audio frequencies, i.e., the modulation threshold for detection of AM (signal) is raised by preceding AM (masker). In the study we focused on the basic characteristics of the forward-masking effect. Functions representing recovery from AM forward masking measured with a 150- ms 40- Hz masker AM and a 50- ms signal AM of the same rate imposed on the same broadband-noise carrier, showed an exponential decay of forward masking with increasing delay from masker offset. Thresholds remained elevated by more than 2 dB over an interval of at least 150 ms following the masker. Masked-threshold patterns, measured with a fixed signal rate (20, 40, and 80 Hz) and a variable masker rate, showed tuning of the AM forward-masking effect. The tuning was approximately constant across signal modulation rates used and consistent with the idea of modulation-rate selective channels. Combining two equally effective forward maskers of different frequencies did not lead to an increase in forward masking relative to that produced by either component alone. Overall, the results are consistent with modulation-rate selective neural channels that adapt and recover from the adaptation relatively quickly.