Intensity discrimination as a function of level and frequency and its relation to high-frequency hearing

This paper examines how intensity discrimination depends on the test frequency, the level, and the subjects's high-frequency hearing. Three experiments were performed. In the first experiment, intensity discrimination of pulsed tones was measured as a function of level at 1 and 14 kHz in five listeners. Results show less deviation from Weber's law at 14 kHz than at 1 kHz. In the second experiment, intensity discrimination was measured for a 1-kHz tone at 90-dB SPL as a function of the cutoff frequency of a high-pass masking noise in two listeners. Results show that the audibility of very high frequencies is important for frequency discrimination at 1 kHz. The DL increased by a factor between 1.5 and 2.0 as the cutoff frequency of the noise was lowered from 19 to 6 kHz. In the third experiment, thresholds from 6 to 20 kHz and intensity discrimination for a 1-kHz tone was measured in 12 listeners. Results show that the DLs at 80-dB SPL are correlated with the ability to hear very high frequencies. Results of all three experiments are consistent with the multiband version of the excitation-pattern model for intensity discrimination [Florentine and Buus, J. Acoust. Soc. Am. 70, 1646-1654 (1981)].     

Dead regions and pitch perception

The perception of pitch for pure tones with frequencies falling inside low- or high-frequency dead regions (DRs) was examined. Subjects adjusted a variable-frequency tone to match the pitch of a fixed tone. Matches within one ear were often erratic for tones falling in a DR, indicating unclear pitch percepts. Matches across ears of subjects with asymmetric hearing loss, and octave matches within ears, indicated that tones falling within a DR were perceived with an unclear pitch and/or a pitch different from "normal" whenever the tones fell more than 0.5 octave within a low- or high-frequency DR. One unilaterally impaired subject, with only a small surviving region between 3 and 4 kHz, matched a fixed 0.5-kHz tone in his impaired ear with, on average, a 3.75-kHz tone in his better ear. When asked to match the 0.5-kHz tone with an amplitude-modulated tone, he adjusted the carrier and modulation frequencies to about 3.8 and 0.5 kHz, respectively, suggesting that some temporal information was still available. Overall, the results indicate that the pitch of low-frequency tones is not conveyed solely by a temporal code. Possibly, there needs to be a correspondence between place and temporal information for a normal pitch to be perceived.     

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.     

Induced loudness reduction as a function of exposure time and signal frequency

Induced loudness reduction (ILR) is the decline in the loudness of a weaker tone induced by a preceding stronger tone. In this study we investigate how ILR depends on exposure time and signal frequency. For 12 listeners, successive magnitude estimation was used to measure the loudness of 70-dB-SPL test tones, presented with and without preceding 80-dB-SPL inducer tones at the same frequency. Experiment 1 measured the evolution of ILR over time at 0.5 kHz. The results suggest that ILR may begin after a single inducer presentation, and increases over at least 2 to 3 min as the inducer and test tones are repeated every few seconds. Following the cessation of the inducer, the recovery of loudness is slow and still incomplete after 1 min. Experiment 2 extended the measurements to additional signal frequencies. The results show that the amount of ILR and its evolution over time are approximately the same at frequencies from 0.5 to 8 kHz. Similarly, loudness matching showed no effect of frequency on ILR, which averaged 8.2 dB. These findings, together with previously noted similarities among ILR, ipsilaterally induced loudness adaptation, and temporary loudness shift, indicate that loudness reduction induced by stronger sounds is a very common phenomenon.     

Killer whale (Orcinus orca) hearing: auditory brainstem response and behavioral audiograms.

Killer whale (Orcinus orca) audiograms were measured using behavioral responses and auditory evoked potentials (AEPs) from two trained adult females. The mean auditory brainstem response (ABR) audiogram to tones between 1 and 100 kHz was 12 dB (re 1 mu Pa) less sensitive than behavioral audiograms from the same individuals (+/- 8 dB). The ABR and behavioral audiogram curves had shapes that were generally consistent and had the best threshold agreement (5 dB) in the most sensitive range 18-42 kHz, and the least (22 dB) at higher frequencies 60-100 kHz. The most sensitive frequency in the mean Orcinus audiogram was 20 kHz (36 dB), a frequency lower than many other odontocetes, but one that matches peak spectral energy reported for wild killer whale echolocation clicks. A previously reported audiogram of a male Orcinus had greatest sensitivity in this range (15 kHz, approximately 35 dB). Both whales reliably responded to 100-kHz tones (95 dB), and one whale to a 120-kHz tone, a variation from an earlier reported high-frequency limit of 32 kHz for a male Orcinus. Despite smaller amplitude ABRs than smaller delphinids, the results demonstrated that ABR audiometry can provide a useful suprathreshold estimate of hearing range in toothed whales.     

Comparing behavioral and physiological measures of combination tones: Sex and race differences

Both distortion-product otoacoustic emissions (DPOAEs) and performance in an auditory-masking task involving combination tones were measured in the same frequency region in the same ears. In the behavioral task, a signal of 3.6?kHz (duration 300?ms, rise/fall time 20?ms) was masked by a 3.0-kHz tone (62?dB SPL, continuously presented). These two frequencies can produce a combination tone at 2.4?kHz. When a narrowband noise (2.0-2.8?kHz, 17?dB spectrum level) was added as a second masker, detection of the 3.6-kHz signal worsened by 6-9?dB (the Greenwood effect), revealing that listeners had been using the combination tone at 2.4?kHz as a cue for detection at 3.6?kHz. Several outcomes differed markedly by sex and racial background. The Greenwood effect was substantially larger in females than in males, but only for the White group. When the magnitude of the Greenwood effect was compared with the magnitude of the DPOAE measured in the 2.4?kHz region, the correlations typically were modest, but were high for Non-White males. For many subjects, then, most of the DPOAE measured in the ear canal apparently is not related to the combination-tone cue that is masked by the narrowband noise.     

Effects of the use of personal music players on amplitude modulation detection and frequency discrimination

Measures of auditory performance were compared for an experimental group who listened regularly to music via personal music players (PMP) and a control group who did not. Absolute thresholds were similar for the two groups for frequencies up to 2 kHz, but the experimental group had slightly but significantly higher thresholds at higher frequencies. Thresholds for the frequency discrimination of pure tones were measured for a sensation level (SL) of 20 dB and center frequencies of 0.25, 0.5, 1, 2, 3, 4, 5, 6, and 8 kHz. Thresholds were significantly higher (worse) for the experimental than for the control group for frequencies from 3 to 8 kHz, but not for lower frequencies. Thresholds for detecting sinusoidal amplitude modulation (AM) were measured for SLs of 10 and 20 dB, using four carrier frequencies 0.5, 3, 4, and 6 kHz, and three modulation frequencies 4, 16, and 50 Hz. Thresholds were significantly lower (better) for the experimental than for the control group for the 4- and 6-kHz carriers, but not for the other carriers. It is concluded that listening to music via PMP can have subtle effects on frequency discrimination and AM detection.     

The perception of complex tones by a false killer whale (Pseudorca crassidens)

Complex tonal whistles are frequently produced by some odontocete species. However, no experimental evidence exists regarding the detection of complex tones or the discrimination of harmonic frequencies by a marine mammal. The objectives of this investigation were to examine the ability of a false killer whale to discriminate pure tones from complex tones and to determine the minimum intensity level of a harmonic tone required for the whale to make the discrimination. The study was conducted with a go/no-go modified staircase procedure. The different stimuli were complex tones with a fundamental frequency of 5 kHz with one to five harmonic frequencies. The results from this complex tone discrimination task demonstrated: (1) that the false killer whale was able to discriminate a 5 kHz pure tone from a complex tone with up to five harmonics, and (2) that discrimination thresholds or minimum intensity levels exist for each harmonic combination measured. These results indicate that both frequency level and harmonic content may have contributed to the false killer whale's discrimination of complex tones.     

The influence of middle-ear muscle contraction on auditory threshold for selected pure tones

The influence of middle-ear muscle (MEM) contraction on auditory threshold has been measured for pure tones of 0.25, 0.5, and 1.5 kHz. The reflex-activating signal was a 3-kHz pure tone. Signal paradigms were chosen to reduce or eliminate the effects of binaural loudness summation, contralateral direct masking, and contralateral remote and backward masking effects, and to maximize the influence of MEM contraction. Results indicate that under no condition was behavioral threshold affected by the MEM contraction induced using a pure-tone stimulus of 3 kHz, 105 dB SPL.     

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.     

Release from masking caused by envelope fluctuations

This paper examines how short-term energy fluctuations in a masker affect the thresholds for tones at frequencies above those of the masker. Two equally intense tones at 1060 and 1075 Hz produce up to 25 dB less masking than does a 1075-Hz tone set to the overall level of the two-tone complex. At wider frequency separations, two-tone complexes also produce less masking than the pure tone. These results indicate that envelope fluctuations in a masker, whose spectrum is confined to a single critical band, may result in release from masking. The release from masking probably is related to the comodulation masking release reported by Hall et al. [J. Acoust. Soc. Am. 76, 50-56 (1984b)] for modulated-noise maskers with bandwidths greater than one critical band. Further measurements with maskers, whose intensity level in the critical band around 1 kHz was 90 dB SPL, show similar masking by a pure tone and a 625- to 1075-Hz bandpass noise, but less masking by narrow-band noises. These results are inconsistent with a simple frequency selective energy-detector model and indicate that the auditory system can use periods of low masker energy as brief as a few ms to enhance detection of a tone. The results also imply that the upward spread of excitation is best represented by masking patterns for noises with bandwidths of several critical bands.     

Evidence for an analytic perception of multiharmonic sounds in the bat, Megaderma lyra, and its possible role for echo spectral analysis

For echolocation, the gleaning bat Megaderma lyra relies on short and broadband calls consisting of multiple harmonic components, each of which is downward frequency modulated. The harmonic components in M. lyra's calls have a relatively small frequency excursion and do not overlap spectrally. Broadband calls of other bat species, on the other hand, often consist of only a few harmonics which are modulated over broad and sometimes overlapping frequency ranges. A call consisting of narrow and nonoverlapping harmonic components may provide a less complete representation of target structure than a call which consists of broadly modulated components. However, a multiharmonic call may help the bats to perceive local spectral changes in the echo from shifts in the peak frequencies of single harmonics, and thereby to extract additional information about the target. To assess this hypothesis, the accuracy with which M. lyra can analyze frequency shifts of single partials in multiharmonic complex tones was investigated. A two-alternative, forced-choice behavioral task was used to measure M. lyra's frequency discrimination threshold for the third partial in complex tones whose spectral composition resembled that of the bat's sonar calls. The discrimination threshold for the third partial in a 21.5-kHz harmonic tone amounted to about 2% and was similar to the bat's pure-tone discrimination threshold at 64.5 kHz. Discrimination performance was essentially unaffected by random frequency changes of the other partials and by reducing stimulus duration from 50.5 to 1.5 ms. Both findings are in accordance with predictions made on the basis of the shape of M. Ivra's cochlear filters. The comparison between the observed frequency discrimination performance and a computational estimate of the expected frequency shift in the third harmonic of an echo reflected by a simple, two-front target showed that M. lyra's frequency resolution is sufficient for analyzing the target-specific information conveyed by shifts in the peak frequency of single echo components.     

Perception of across-frequency asynchrony and the role of cochlear delays

Cochlear filtering results in earlier responses to high than to low frequencies. This study examined potential perceptual correlates of cochlear delays by measuring the perception of relative timing between tones of different frequencies. A brief 250-Hz tone was combined with a brief 1-, 2-, 4-, or 6-kHz tone. Two experiments were performed, one involving subjective judgments of perceived synchrony, the other involving asynchrony detection and discrimination. The functions relating the proportion of "synchronous" responses to the delay between the tones were similar for all tone pairs. Perceived synchrony was maximal when the tones in a pair were gated synchronously. The perceived-synchrony function slopes were asymmetric, being steeper on the low-frequency-leading side. In the second experiment, asynchrony-detection thresholds were lower for low-frequency rather than for high-frequency leading pairs. In contrast with previous studies, but consistent with the first experiment, thresholds did not depend on frequency separation between the tones, perhaps because of the elimination of within-channel cues. The results of the two experiments were related quantitatively using a decision-theoretic model, and were found to be highly correlated. Overall the results suggest that frequency-dependent cochlear group delays are compensated for at higher processing stages, resulting in veridical perception of timing relationships across frequency.     

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.     

Assessing temporary threshold shift in a bottlenose dolphin (Tursiops truncatus) using multiple simultaneous auditory evoked potentials

Hearing sensitivity was measured in a bottlenose dolphin before and after exposure to an intense 20-kHz fatiguing tone in three different experiments. In each experiment, hearing was characterized using both the auditory steady-state response (ASSR) and behavioral methods. In experiments 1 and 2, ASSR stimuli consisted of seven frequency-modulated tones, each with a unique carrier and modulation frequency. The tones were simultaneously presented to the subject and the ASSR at each modulation rate measured to determine the effects of the sound exposure at the corresponding carrier frequency. In experiment 3 behavioral thresholds and ASSR input-output functions were measured at a single frequency before and after three exposures. Hearing loss was frequency-dependent, with the largest temporary threshold shifts occurring (in order) at 30, 40, and 20 kHz. ASSR threshold shifts reached 40-45 dB and were always larger than behavioral shifts (19-33 dB). The ASSR input-output functions were represented as the sum of two processes: a low threshold, saturating process and a higher threshold, linear process, that react and recover to fatigue at different rates. The loss of the near-threshold saturating process after exposure may explain the discrepancies between the ASSR and behavioral threshold shifts.     

Psychometric functions for level discrimination

To determine the form of psychometric functions for 2I,2AFC level discrimination (commonly called intensity discrimination), ten increment levels were presented in random order within blocks of 100 trials. Stimuli were chosen to encompass a wide range of conditions and difference limens: eight 10-ms tones had frequencies of 0.25, 1, 8, or 14 kHz and levels of 30, 60, or 90 dB SPL; two 500-ms stimuli also were tested: a 1-kHz tone at 90 dB SPL and broadband noise at 63 dB SPL. For each condition, at least 20 blocks were presented in mixed order. Results for five normal listeners show that the sensitivity, d', is nearly proportional to delta L (= 20 log [(p + delta p)/p], where p is sound pressure) over the entire range of difference limens. When d' is plotted against Weber fractions for sound pressure, delta p/p, or intensity, delta I/I, exponents of the best-fitting power functions decrease with increasing difference limens and are less than unity for large difference limens. The approximately proportional relation between d' and delta L agrees with modern multichannel models of level discrimination and with psychometric functions derived for single auditory-nerve fibers. The results also support the notion that the difference limen, expressed as delta LDL and plotted on a logarithmic scale, is an appropriate representation of performance in level-discrimination experiments.     

Subjective loudness level measurements and equal loudness contours in a bottlenose dolphin (Tursiops truncatus)

Loudness level measurements in human listeners are straightforward; however, it is difficult to convey the concepts of loudness matching or loudness comparison to (non-human) animals. For this reason, prior studies have relied upon objective measurements, such as response latency, to estimate equal loudness contours in animals. In this study, a bottlenose dolphin was trained to perform a loudness comparison test, where the listener indicates which of two sequential tones is louder. To enable reward of the dolphin, most trials featured tones with identical or similar frequencies, but relatively large sound pressure level differences, so that the loudness relationship was known. A relatively small percentage of trials were "probe" trials, with tone pairs whose loudness relationship was not known. Responses to the probe trials were used to construct psychometric functions describing the loudness relationship between a tone at a particular frequency and sound pressure level and that of a reference tone at 10 kHz with a sound pressure level of 90, 105, or 115 dB re 1 μPa. The loudness relationships were then used to construct equal loudness contours and auditory weighting functions that can be used to predict the frequency-dependent effects of noise on odontocetes.     

Spatial Spread of Visual Attention while Tracking a Moving Object

We conducted three experiments to investigate the spatial spread of visual attention. In Experiment 1, we measured the contrast sensitivities at various locations (spatial sensitivity function) relative to the moving target that the observer attended to track in an attentive tracking display. A probe was presented at a distance from the target at a location randomly chosen from within a certain range. The range of probe presentation location varied to examine whether the observer changes the area of attention to cope with this range. The results show that the probe range influenced the shape of spatial sensitivity function. The change in shape of this function suggests that the observer covers a wider area with attention for large probe ranges than small probe ranges. In the following experiments, we investigated the effect of the distance between the tracking target and a probe at a fixed location relative to the target (Experiment 2), or between the target and the center of a probe range of fixed size (Experiment 3). Since the relative probe location in a session was fixed in the experiments, the observer would pay attention to the target and probe locations independently of the relative distance if he/she could focus attention at multiple locations. Spatial sensitivity functions obtained in Experiments 2 and 3 showed that this was not the case. In both experiments the sensitivity to the probe decreased with increase in the relative distance as in Experiment 1, where the probe was presented at a location randomly chosen within each range. This indicates that attention cannot be divided among multiple locations, at least under the present experimental conditions. We will discuss a possible interpretation of the present results with a limited attentional resource and its spatial distribution.     

Simultaneous masking by gated and continuous sinusoidal maskers

Simultaneous masking of a 20-ms, 1-kHz signal was investigated using 50-ms gated and continuous sinusoidal maskers with frequencies below, at, and above 1 kHz. Gated maskers can produce considerably (5-20 dB) more masking than continuous maskers, and this difference does not appear to result from the spread of energy produced by gating either the masker or the signal. For masker frequencies below the signal frequency, this difference in masking is primarily due to the detection of the cubic difference tone in the continuous condition. For masker frequencies at and above the signal frequency, the difference appears to be an important property of masking. Implications of this frequency-dependent effect for measures of frequency selectivity are discussed.     

Contribution of spectral cues to human sound localization

The contribution of spectral cues to human sound localization was investigated by removing cues in 1/2-, 1- or 2-octave bands in the frequency range above 4 kHz. Localization responses were given by placing an acoustic pointer at the same apparent position as a virtual target. The pointer was generated by filtering a 100-ms harmonic complex with equalized head-related transfer functions (HRTFs). Listeners controlled the pointer via a hand-held stick that rotated about a fixed point. In the baseline condition, the target, a 200-ms noise burst, was filtered with the same HRTFs as the pointer. In other conditions, the spectral information within a certain frequency band was removed by replacing the directional transfer function within this band with the average transfer of this band. Analysis of the data showed that removing cues in 1/2-octave bands did not affect localization, whereas for the 2-octave band correct localization was virtually impossible. The results obtained for the 1-octave bands indicate that up-down cues are located mainly in the 6-12-kHz band, and front-back cues in the 8-16-kHz band. The interindividual spread in response patterns suggests that different listeners use different localization cues. The response patterns in the median plane can be predicted using a model based on spectral comparison of directional transfer functions for target and response directions.