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)].     

Learning to perceive pitch differences

This paper reports two experiments concerning the stimulus specificity of pitch discrimination learning. In experiment 1, listeners were initially trained, during ten sessions (about 11,000 trials), to discriminate a monaural pure tone of 3000 Hz from ipsilateral pure tones with slightly different frequencies. The resulting perceptual learning (improvement in discrimination thresholds) appeared to be frequency-specific since, in subsequent sessions, new learning was observed when the 3000-Hz standard tone was replaced by a standard tone of 1200 Hz, or 6500 Hz. By contrast, a subsequent presentation of the initial tones to the contralateral ear showed that the initial learning was not, or was only weakly, ear-specific. In experiment 2, training in pitch discrimination was initially provided using complex tones that consisted of harmonics 3-7 of a missing fundamental (near 100 Hz for some listeners, 500 Hz for others). Subsequently, the standard complex was replaced by a standard pure tone with a frequency which could be either equal to the standard complex's missing fundamental or remote from it. In the former case, the two standard stimuli were matched in pitch. However, this perceptual relationship did not appear to favor the transfer of learning. Therefore, the results indicated that pitch discrimination learning is, at least to some extent, timbre-specific, and cannot be viewed as a reduction of an internal noise which would affect directly the output of a neural device extracting pitch from both pure tones and complex tones including low-rank harmonics.     

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.     

Frequency discrimination of complex tones with overlapping and non-overlapping harmonics

These experiments address the following issues. (1) When two complex tones contain different harmonics, do the differences in timbre between them impair the ability to discriminate the pitches of the tones? (2) When two complex tones have only a single component in common, and that component is the most discriminable component in each tone, is the frequency discrimination of the component affected by differences in residue pitch between the two tones? (3) How good is the pitch discrimination of complex tones with no common components when each tone contains multiple harmonics, so as to avoid ambiguity of pitch? (4) Is the pitch discrimination of complex tones with common harmonics impaired by shifting the component frequencies to nonharmonic values? In all experiments, frequency difference limens (DLCs) were measured for multiple-component complex tones, using an adaptive two-interval, two-alternative, forced-choice task. Three highly trained subjects were used. The main conclusions are as follows. (1) When two tones have the first six harmonics in common, DLCs are larger when the upper harmonics are different than when the upper harmonics are in common or are absent. It appears that differences in timbre impair DLCs. (2) Discrimination of the frequency of a single common partial in two complex tones is worse when the two tones have different residue pitches than when they have the same residue pitch. (3) DLCs for complex tones with no common harmonics are generally larger than those for complex tones with common harmonics. For the former, large individual differences occur, probably because subjects are affected differently by differences in timbre. (4) DLCs for harmonic complex tones are smaller than DLCs for complex tones in which the components are mistuned from harmonic values. This can probably be attributed to the less distinct residue pitch of the inharmonic complexes, rather than to reduced discriminability of partials. Overall, the results support the idea that DLCs for complex tones with common harmonics depend on residue pitch comparisons, rather than on comparisons of the pitches of partials.     

Single-lobed frequency-dependent beam shape in an echolocating false killer whale (Pseudorca crassidens)

Recent studies indicate some odontocetes may produce echolocation beams with a dual-lobed vertical structure. The shape of the odontocete echolocation beam was further investigated in a false killer whale performing an echolocation discrimination task. Clicks were recorded with an array of 16 hydrophones and frequency-dependent amplitude plots were constructed to assess beam shape. The majority of the echolocation clicks were single-lobed in structure with most energy located between 20 and 80 kHz. These data indicate the false killer whale does not produce a dual-lobed structure, as has been shown in bottlenose dolphins, which may be a function of lowered frequencies in the emitted signal due to hearing loss.     

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.     

Frequency and intensity difference limens for harmonics within complex tones

A two-interval, two-alternative forced choice task was used to estimate frequency difference limens (DLs) for individual harmonics within complex tones, and DLs for the periodicity (i.e., number of periods per s) of the whole complexes. For complex tones with equal-amplitude harmonics, the DLs for the lowest harmonics were small (less than one percent). The DLs increased rather abruptly around the fifth to seventh harmonic. The highest harmonic in each complex was also well discriminated, and the discriminability of a single high harmonic was markedly improved by increasing its level relative to the other components. The DL for a complex tone was generally smaller than the frequency DL of its most discriminable component. The DL for a complex was found to be predictable from the DLs of the harmonics comprising the complex, using a formula derived by Goldstein [J. Acoust. Soc. Am. 54, 1496-1516 (1973)] from his optimum processor theory for the formation of the pitch of complex tones. The DL for a complex is sometimes primarily determined by high harmonics, such as the highest harmonic, or a harmonic whose level exceeds that of adjacent harmonics. We also measured intensity DLs for individual harmonics within complex tones. The intensity DLs were smallest for low harmonic numbers, and for the highest harmonic in a complex. An excitation-pattern model was used to determine whether the frequency DLs of harmonics within complex tones could be explained in terms of place mechanisms, i.e., in terms of changes in the amount of excitation at appropriate frequency places. We conclude that place mechanisms are not adequate, and that information about the frequencies of individual harmonics is probably carried in the time patterning of neural impulses.     

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.     

Pitch shift of pure and complex tones induced by masking noise

Psychoacoustic experiments were performed to measure the pitch-shift effects of pure and complex tones resulting from the addition of a masking noise to the tonal stimuli. Harmonic residue tones with either two or three harmonics and a fundamental frequency of 200 Hz were chosen as test tones. The pitch shifts of virtual and spectral pitches of the residue tones were measured as a function of the intensity of a low-pass noise with 600-Hz cutoff frequency. The SPL of this noise varied between 30 and 70 dB. In another experiment, the pitch shifts of single pure tones corresponding to the frequencies and SPLs of the harmonics of the residue tones were measured using the same masking noise. The results from five subjects for the harmonic residue tones show only a weak dependence of pitch shift on masking noise intensity. This dependence exists for both spectral and virtual pitches. In the case of single pure tones, pitch shift depends more distinctly on noise intensity. Pitch shifts of up to 5% were found in the range of noise intensity investigated. The magnitude of pitch shift shows pronounced interindividual differences, but the direction of the shift effect is always the same. In all cases pitch increases with higher masking noise levels.     

The effect of hearing loss on the resolution of partials and fundamental frequency discrimination

The relationship between the ability to hear out partials in complex tones, discrimination of the fundamental frequency (F0) of complex tones, and frequency selectivity was examined for subjects with mild-to-moderate cochlear hearing loss. The ability to hear out partials was measured using a two-interval task. Each interval included a sinusoid followed by a complex tone; one complex contained a partial with the same frequency as the sinusoid, whereas in the other complex that partial was missing. Subjects had to indicate the interval in which the partial was present in the complex. The components in the complex were uniformly spaced on the ERB(N)-number scale. Performance was generally good for the two "edge" partials, but poorer for the inner partials. Performance for the latter improved with increasing spacing. F0 discrimination was measured for a bandpass-filtered complex tone containing low harmonics. The equivalent rectangular bandwidth (ERB) of the auditory filter was estimated using the notched-noise method for center frequencies of 0.5, 1, and 2 kHz. Significant correlations were found between the ability to hear out inner partials, F0 discrimination, and the ERB. The results support the idea that F0 discrimination of tones with low harmonics depends on the ability to resolve the harmonics.     

Differential sensitivity to vowel continua in Old World monkeys (Macaca) and humans

Previous studies indicate that monkey pure tone frequency discrimination is quantitatively and qualitatively very different from that of humans: Monkey DLs at 1.0 and 2.0 kHz are up to 20 times larger than human DLs, and monkeys DLs increase as sensation level increases, in contrast to human DLs [Sinnott et al., J. Acoust. Soc. Am. 78, 1977-1985 (1985); Sinnott et al., J. Comp. Psychol. 101, 126-131 (1987)]. These results led to an hypothesis that monkey frequency discrimination is more dependent upon "rate" coding than is that of humans. The present study compared monkey and human DLs for formant frequency changes along three synthetic vowel continua /I-i/, /ae-epsilon/, and /a-v/. Here, monkey DLs for formants near 1.0 and 2.0 kHz (32-48 Hz) were only about two to three times larger than human DLs (11-21 Hz), and both monkeys and humans exhibited relatively similar, flat sensation level functions. Taken together, these data indicate that monkey and human frequency discrimination is more similar in the case of a complex vowel stimulus than in the case of a simple pure tone stimulus. Results are discussed in relation to "rate" versus "temporal" coding of tones and vowels in the auditory system.     

Cats exhibit the Franssen Effect illusion

The Franssen Effect (FE) is a striking auditory illusion previously demonstrated only in humans. To elicit the FE, subjects are presented with two spatially-separated sounds; one a transient tone with an abrupt onset and immediate ramped offset and the other a sustained tone of the same frequency with a ramped onset which remains on for several hundred ms. The FE illusion occurs when listeners localize the tones at the location of the transient signal, even though that sound has ended and the sustained one is still present. The FE illusion occurs most readily in reverberant environments and with pure tones of approximately 1-2.5 kHz in humans, conditions where sound localization is difficult in humans. Here, we demonstrate this illusion in domestic cats using, for the first time, localization procedures. Previous studies in humans employed discrimination procedures, making it difficult to link the FE to sound localization mechanisms. The frequencies for eliciting the FE in cats were higher than in humans, corresponding to frequencies where cats have difficulty localizing pure tones. These findings strengthen the hypothesis that difficulty in accurately localizing sounds is the basis for the FE.     

Perception and cortical neural coding of harmonic fusion in ferrets

This study examined the perception and cortical representation of harmonic complex tones, from the perspective of the spectral fusion evoked by such sounds. Experiment 1 tested whether ferrets spontaneously distinguish harmonic from inharmonic tones. In baseline sessions, ferrets detected a pure tone terminating a sequence of inharmonic tones. After they reached proficiency, a small fraction of the inharmonic tones were replaced with harmonic tones. Some of the animals confused the harmonic tones with the pure tones at twice the false-alarm rate. Experiment 2 sought correlates of harmonic fusion in single neurons of primary auditory cortex and anterior auditory field, by comparing responses to harmonic tones with those to inharmonic tones in the awake alert ferret. The effects of spectro-temporal filtering were accounted for by using the measured spectrotemporal receptive field to predict responses and by seeking correlates of fusion in the predictability of responses. Only 12% of units sampled distinguished harmonic tones from inharmonic tones, a small percentage that is consistent with the relatively weak ability of the ferrets to spontaneously discriminate harmonic tones from inharmonic tones in Experiment 1.     

Target detection, shape discrimination, and signal characteristics of an echolocating false killer whale (Pseudorca crassidens).

This study demonstrated the ability of a false killer whale (Pseudorca crassidens) to discriminate between two targets and investigated the parameters of the whale's emitted signals for changes related to test conditions. Target detection performance comparable to the bottlenose dolphin's (Tursiops truncatus) has previously been reported for echolocating false killer whales. No other echolocation capabilities have been reported. A false killer whale, naive to conditioned echolocation tasks, was initially trained to detect a cylinder in a "go/no-go" procedure over ranges of 3 to 8 m. The transition from a detection task to a discrimination task was readily achieved by introducing a spherical comparison target. Finally, the cylinder was successfully compared to spheres of two different sizes and target strengths. Multivariate analyses were used to evaluate the parameters of emitted signals. Duncan's multiple range tests showed significant decreases (df = 185, p less than 0.05) in both source level and bandwidth in the transition from detection to discrimination. Analysis of variance revealed a significant decrease in the number of clicks over test conditions [F(5.26) = 5.23, p less than 0.0001]. These data suggest that the whale relied on cues relevant to target shape as well as target strength, that changes in source level and bandwidth were task-related, that the decrease in clicks was associated with learning experience, and that Pseudorca's ability to discriminate shapes using echolocation may be comparable to that of Tursiops truncatus.     

Detection and discrimination of frequency modulation of complex signals

Detection and discrimination of frequency modulation were studied for harmonic signals with triangular spectral envelopes. The center frequency of the stimuli was near 2 kHz; the fundamental frequency was near 100 Hz. To prevent the possibility that the discrimination was based on differences of initial or final frequencies, these frequencies were equal within and across modulations in each individual experiment. Differences between modulations consisted of differences in the trajectories between the initial and final frequencies. Performance worsened as the slopes of the spectral envelopes decreased. Addition of noise also impaired modulation discrimination. The dependence on the signal-to-noise ratio was similar to what is found for stationary stimuli: Discrimination of frequency modulation deteriorated more rapidly with decreasing signal-to-noise ratio when stimuli had shallow spectral slopes than when they had steep spectral slopes. In spite of the precautions taken (i.e., initial and final frequency the same), the discrimination of these stimuli was more likely based on quasistationary frequency discrimination than on discrimination of modulation rate. This conclusion is consistent with previous findings for pure tones presented in quiet that frequency discrimination is more acute than modulation-rate discrimination.     

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.     

Effect of level on the discrimination of harmonic and frequency-shifted complex tones at high frequencies

Moore and Se?k [J. Acoust. Soc. Am. 125, 3186-3193 (2009)] measured discrimination of a harmonic complex tone and a tone in which all harmonics were shifted upwards by the same amount in Hertz. Both tones were passed through a fixed bandpass filter and a background noise was used to mask combination tones. Performance was well above chance when the fundamental frequency was 800 Hz, and all audible components were above 8000 Hz. Moore and Se?k argued that this suggested the use of temporal fine structure information at high frequencies. However, the task may have been performed using excitation-pattern cues. To test this idea, performance on a similar task was measured as a function of level. The auditory filters broaden with increasing level, so performance based on excitation-pattern cues would be expected to worsen as level increases. The results did not show such an effect, suggesting that the task was not performed using excitation-pattern cues.     

Characteristics of whistles from the acoustic repertoire of resident killer whales (Orcinus orca) off Vancouver Island, British Columbia

The acoustic repertoire of killer whales (Orcinus orca) consists of pulsed calls and tonal sounds, called whistles. Although previous studies gave information on whistle parameters, no study has presented a detailed quantitative characterization of whistles from wild killer whales. Thus an interpretation of possible functions of whistles in killer whale underwater communication has been impossible so far. In this study acoustic parameters of whistles from groups of individually known killer whales were measured. Observations in the field indicate that whistles are close-range signals. The majority of whistles (90%) were tones with several harmonics with the main energy concentrated in the fundamental. The remainder were tones with enhanced second or higher harmonics and tones without harmonics. Whistles had an average bandwidth of 4.5 kHz, an average dominant frequency of 8.3 kHz, and an average duration of 1.8 s. The number of frequency modulations per whistle ranged between 0 and 71. The study indicates that whistles in wild killer whales serve a different function than whistles of other delphinids. Their structure makes whistles of killer whales suitable to function as close-range motivational sounds.     

The relationship between frequency selectivity and pitch discrimination: effects of stimulus level

Three experiments tested the hypothesis that fundamental frequency (fo) discrimination depends on the resolvability of harmonics within a tone complex. Fundamental frequency difference limens (fo DLs) were measured for random-phase harmonic complexes with eight fo's between 75 and 400 Hz, bandpass filtered between 1.5 and 3.5 kHz, and presented at 12.5-dB/component average sensation level in threshold equalizing noise with levels of 10, 40, and 65 dB SPL per equivalent rectangular auditory filter bandwidth. With increasing level, the transition from large (poor) to small (good) fo DLs shifted to a higher fo. This shift corresponded to a decrease in harmonic resolvability, as estimated in the same listeners with excitation patterns derived from measures of auditory filter shape and with a more direct measure that involved hearing out individual harmonics. The results are consistent with the idea that resolved harmonics are necessary for good fo discrimination. Additionally, fo DLs for high fo's increased with stimulus level in the same way as pure-tone frequency DLs, suggesting that for this frequency range, the frequencies of harmonics are more poorly encoded at higher levels, even when harmonics are well resolved.     

Thresholds for hearing mistuned partials as separate tones in harmonic complexes

When a low harmonic in a harmonic complex tone is mistuned from its harmonic value by a sufficient amount it is heard as a separate tone, standing out from the complex as a whole. This experiment estimated the degree of mistuning required for this phenomenon to occur, for complex tones with 10 or 12 equal-amplitude components (60 dB SPL per component). On each trial the subject was presented with a complex tone which either had all its partials at harmonic frequencies or had one partial mistuned from its harmonic frequency. The subject had to indicate whether he heard a single complex tone with one pitch or a complex tone plus a pure tone which did not "belong" to the complex. An adaptive procedure was used to track the degree of mistuning required to achieve a d' value of 1. Threshold was determined for each ot the first six harmonics of each complex tone. In one set of conditions stimulus duration was held constant at 410 ms, and the fundamental frequency was either 100, 200, or 400 Hz. For most conditions the thresholds fell between 1% and 3% of the harmonic frequency, depending on the subject. However, thresholds tended to be greater for the first two harmonics of the 100-Hz fundamental and, for some subjects, thresholds increased for the fifth and sixth harmonics. In a second set of conditions fundamental frequency was held constant at 200 Hz, and the duration was either 50, 110, 410, or 1610 ms. Thresholds increased by a factor of 3-5 as duration was decreased from 1610 ms to 50 ms. The results are discussed in terms of a hypothetical harmonic sieve and mechanisms for the formation of perceptual streams.