Two-tone suppression in the cricket, Eunemobius carolinus (Gryllidae, Nemobiinae)

Sounds with frequencies >15 kHz elicit an acoustic startle response (ASR) in flying crickets (Eunemobius carolinus). Although frequencies <15 kHz do not elicit the ASR when presented alone, when presented with ultrasound (40 kHz), low-frequency stimuli suppress the ultrasound-induced startle. Thus, using methods similar to those in masking experiments, we used two-tone suppression to assay sensitivity to frequencies in the audio band. Startle suppression was tuned to frequencies near 5 kHz, the frequency range of male calling songs. Similar to equal loudness contours measured in humans, however, equal suppression contours were not parallel, as the equivalent rectangular bandwidth of suppression tuning changed with increases in ultrasound intensity. Temporal integration of suppressor stimuli was measured using nonsimultaneous presentations of 5-ms pulses of 6 and 40 kHz. We found that no suppression occurs when the suppressing tone is >2 ms after and >5 ms before the ultrasound stimulus, suggesting that stimulus overlap is a requirement for suppression. When considered together with our finding that the intensity of low-frequency stimuli required for suppression is greater than that produced by singing males, the overlap requirement suggests that two-tone suppression functions to limit the ASR to sounds containing only ultrasound and not to broadband sounds that span the audio and ultrasound range.     

Variable-duration notched-noise experiments in a broadband noise context.

A variable-duration notched-noise experiment was conducted in a noise context. Broadband noise preceded and followed a tone and notched noise of similar duration. Thresholds were measured at four durations (10, 30, 100, and 300 ms), two center frequencies (0.6, 2.0 kHz), and five relative notch widths (0.0, 0.1, 0.2, 0.4, 0.8). At 0.6 kHz, 10-ms thresholds decrease 6 dB across notch widths, while 300-ms thresholds decrease over 35 dB. These trends are similar but less pronounced at 2 kHz. In a second experiment, the short-duration notched noise was replaced with a flat noise which provided an equivalent amount of simultaneous masking and thresholds dropped by as much as 20 dB. A simple combination of simultaneous and nonsimultaneous masking is unable to predict these results. Instead, it appears that the elevated thresholds at short durations are dependent on the spectral shape of the simultaneous masker.     

Temporal integration of trains of tone pulses by normal and by cochlearly impaired listeners

Two experiments investigated the temporal integration of trains of tone pulses by normal and by cochlearly impaired listeners. In the first experiment, thresholds were measured for a single 5-ms, 4-kHz tone pulse, and for ten such tone pulses as a function of interpulse interval (delta t). For normal listeners, temporal integration, defined as the threshold difference between one and ten pulses, was about 8 dB for delta t less than 20 ms, and about 5 dB at longer delta t's. For impaired listeners, temporal integration was only about 2-3 dB across the range of delta t's (5-160 ms) studied. A second experiment measured psychometric functions (log d' versus log signal power) for a single pulse and for ten pulses with delta t's of 5 ms and 80 ms. The normal listeners' functions had slopes close to unity in all three conditions, with a few exceptions. The impaired listeners' functions had slopes close to unity for ten pulses with delta t = 5 ms, but had slopes significantly greater than unity for delta t = 80 ms, and for a single pulse. At delta t = 80 ms, the increase in d' relative to the condition with a single tone was similar (a factor of square root of 10) for both impaired and normal listeners, but the threshold difference was smaller for the impaired listeners due to their steeper psychometric functions. For impaired listeners, then, temporal integration at delta t = 80 ms was normal in terms of a change in d' but abnormal when measured as a threshold difference.(ABSTRACT TRUNCATED AT 250 WORDS)     

Temporal integration in normal hearing, cochlear impairment, and impairment simulated by masking

To assess temporal integration in normal hearing, cochlear impairment, and impairment simulated by masking, absolute thresholds for tones were measured as a function of duration. Durations ranged from 500 ms down to 15 ms at 0.25 kHz, 8 ms at 1 kHz, and 2 ms at 4 and 14 kHz. An adaptive 2I, 2AFC procedure with feedback was used. On each trial, two 500-ms observation intervals, marked by lights, were presented with an interstimulus interval of 250 ms. The monaural signal was presented in the temporal center of one observation interval. The results for five normal and six impaired listeners show: (1) normal listeners' thresholds decrease by about 8 to 10 dB per decade of duration, as expected; (2) listeners with cochlear impairments generally show less temporal integration than normal listeners; and (3) listeners with impairments simulated using masking noise generally show the same amount of temporal integration as normal listeners tested in the quiet. The difference between real and simulated impairments indicates that the reduced temporal integration observed in impaired listeners probably is not due to splatter of energy to frequency regions where thresholds are low, but reflects reduced temporal integration per se.     

Stimulus parameters governing confusion effects in forward masking

In forward masking, performance may be affected by confusion, that is, by the difficulty of discriminating a suprathreshold signal from the preceding masker. This study investigated confusion effects for forward maskers composed of repeated bursts of a 100-Hz sinusoid followed by sinusoidal signals; such "pulsing" maskers produce confusion when the properties of the signal are identical to those of an individual masker "pulse." The level, frequency, and duration of the signal relative to an individual masker pulse, as well as offset-onset delay, were varied to determine the minimum change necessary to eliminate confusion. For maskers composed of 20-ms pulses, confusion was eliminated by changes in signal level of 5 dB or changes in signal frequency of 30 to 40 Hz. For maskers composed of 10-, 20-, or 40-ms pulses, confusion was eliminated by signal delays of 8 to 16 ms or by signal durations less than half or greater than twice the masker-pulse duration. Results with adaptive procedures designed to measure confusion-free or confusion-determined thresholds suggest that confusion effects can be minimized or avoided by extensive listener training with a procedure in which the signal and masker are not presented at similar intensities.     

Masking of short stimuli by noises with spiked spectra: II. Time summation and frequency selectivity of hearing in a narrow frequency range

The thresholds of masking of short high-frequency pulses with either different durations (1.25–25 ms) and similar central frequency or different central frequencies (3.6–4.4 kHz) but similar durations were measured to reveal manifestations of the properties of peripheral encoding in auditory perception. Noises with a spiked amplitude spectrum structure were used as maskers. The central frequency and the frequency band of a masker were 4 and 1 kHz, respectively. The central frequencies of a stimulus and a masker being equal, the noise the central frequency of which coincided with the frequency corresponding to a dip of an indented spectrum was called an off_(rip)-frequency masker. Owing to the off_(rip)-masker, stimuli-induced masking thresholds were formed taking into account excitation in a narrow region of a basila membrane and auditory nerve fibers with characteristic frequencies from a narrow range. High-frequency pulses with an envelope in the form of the Gaussian function and sinusoidal filling were used as stimuli. At masker levels of 30 dB above the auditory threshold, frequencies of off_(rip)-masker spectra spikes of 500–2000 Hz, and a central stimulus frequency of 4 kHz, the thresholds of tonal stimuli (25 ms in duration) masking in two out of three probationers were higher than the thresholds of masking of compact stimuli (1.25 ms in duration). In the third probationer, on the contrary, the thresholds of tonal stimuli masking were lower than the thresholds of compact stimuli masking. At masker levels of 50 dB, individual threshold differences disappeared. The obtained results were interpreted in the context of implementation of different methods of auditory encoding of the intensity. The methods were based on either the average frequency of auditory nerve pulsations or the number of fibers participating in the response. The interpretation was also carried out in the context of revealing manifestations of nonlinear properties of basila membrane displacements in auditory thresholds. The fact that the dependence of detection thresholds of compact stimuli on their central frequency in one of the two probationers did not reveal the minimum in case of coincidence of off_(rip)-masker and stimulus frequencies pointed to the presence of an auditory “problem zone” that was likely to be localized at the periphery of the auditory system.     

Age differences in backward masking

The present study was designed to assess the effects of age on the time course of backward masking. In experiment 1, thresholds for detecting a 10-ms, 500-Hz sinusoidal signal were measured as a function of the temporal separation between the signal and a 50-ms broadband masker. Subjects were younger (18-24) and older (over age 65) adults with normal hearing (thresholds less than 20 dB HL) for frequencies of 4 kHz and below. Younger subjects exhibited less overall masking and steeper recovery functions than did the older adults. Masked thresholds for younger participants approached unmasked thresholds for signal-masker delays greater than 6-8 ms. In contrast, older adults exhibited significant masking even at the longest delay tested (20 ms). In experiment 2, signal duration was decreased to 5 ms for a separate group of younger adults. Although overall thresholds were elevated for the shorter signal duration, the slope of the backward masking recovery function was not different from that observed for younger adults in experiment 1. The results suggest that age, independent of hearing loss, affects the temporal course of backward masking.     

Overshoot in normal-hearing and hearing-impaired subjects.

Overshoot was measured in both ears of four subjects with normal hearing and in five subjects with permanent, sensorineural hearing loss (two with a unilateral loss). The masker was a 400-ms broadband noise presented at a spectrum level of 20, 30, or 40 dB SPL. The signal was a 10-ms sinusoid presented 1 or 195 ms after the onset of the masker. Signal frequency was 1.0 or 4.0 kHz, which placed the signal in a region of normal (1.0 kHz) or impaired (4.0 kHz) absolute sensitivity for the impaired ears. For the normal-hearing subjects, the effects of signal frequency and masker level were similar to those published previously. In particular, overshoot was larger at 4.0 than at 1.0 kHz, and overshoot at 4.0 kHz tended to decrease with increasing masker level. At 4.0 kHz, overshoot values were significantly larger in the normal ears: Maximum values ranged from about 7-26 dB in the normal ears, but were always less than 5 dB in the impaired ears. The smaller overshoot values resulted from the fact that thresholds in the short-delay condition were considerably better in the hearing-impaired subjects than in the normal-hearing subjects. At 1.0 kHz, overshoot values for the two groups of subjects more or less overlapped. The results suggest that permanent, sensorineural hearing loss disrupts the mechanisms responsible for a large overshoot effect.     

Detection of intensity decrements by the chinchilla.

Three monaural chinchillas were trained to detect intensity decrements in broadband noise (20 kHz) using a shock-avoidance conditioning procedure. The intensity decrements were presented at one of nine different durations between 2 and 35 ms at noise levels of 25, 45, and 65 dB SPL. At each intensity-duration combination, the level of the decrement was varied to obtain a decrement threshold. The minimal detectable decrement decreased from approximately 20 dB at the shortest duration to an asymptote of roughly 4 dB at approximately 30 ms. The data were modeled by a low-pass filter with an 11-ms time constant. The decrement detection function of the chinchilla is similar to that of humans. However, long-duration decrement thresholds are larger in the chinchilla, as would be predicted from the large intensity difference limen of the chinchilla. In general, there was little change in the decrement function across background intensities except that 2-ms decrements were not detected at the 25-dB SPL background intensity.     

Gap detection by early-deafened cochlear-implant subjects

Two studies investigating gap-detection thresholds were conducted with cochlear-implant subjects whose onset of profound hearing loss was very early in life. The Cochlear Limited multiple-electrode prosthesis was used. The first study investigated the effects of pulse rate (200, 500, and 1000 pulses/s) and stimulus duration (500 and 1000 ms) on gap thresholds in 15 subjects. Average gap thresholds were 1.8 to 32.1 ms. There was essentially no effect of pulse rate and for almost all subjects, no effect of stimulus duration. For two subjects, performance was poorer for the 1000-ms stimulus duration. The second study investigated the relationships between gap thresholds, subject variables, and speech-perception scores. Data from the first study were combined with those from previous studies [Busby et al., Audiology 31, 95-111 (1992); Tong et al., J. Acoust. Soc. Am. 84, 951-962 (1988)], providing data from 27 subjects. A significant negative correlation was found between age at onset of deafness and gap thresholds and most variability in gap thresholds was for the congenitally deaf subjects. Significant negative correlations were found between gap thresholds and word scores for open-set Bamford-Kowal-Bench (BKB) sentences in the auditory-visual condition and lipreading enhancement scores for the same test.     

Auditory duration discrimination in Old World monkeys (Macaca, Cercopithecus) and humans

Auditory duration DLs at 2.0 kHz were measured in Old World monkeys (Macaca, Cercopithecus) and humans using a go, no-go repeating standard AX procedure and positive reinforcement operant conditioning techniques. For a 200-ms standard, monkey DLs were 45-125 ms, compared to 15-27 ms for humans. Weber fractions (delta T/T) for all species were smallest at standard durations of 200-400 ms and increased as standard duration decreased to 25 ms. Varying intensity from 30-70 dB SPL had only minor effects on DLs, except at the lowest levels tested, where DLs were elevated slightly. Monkeys had difficulty discriminating duration decrements, in contrast to humans. Results are discussed in relation to other comparative psychoacoustic data and primate vocal communication, including human speech.     

Evidence for a behavioral significance of saccular acoustic sensitivity in humans

In this article the results are reported of an experiment to provide direct evidence for a perceptual and behavioral significance of human saccular acoustic sensitivity. Ten human subjects were stimulated monaurally with 100-ms trains of 10-ms tone pulses with pulse repetition rate of 40 Hz, and were required to rate the pleasantness of the stimuli on a nine-point scale. The design included three within-subject factors: carrier frequency (two levels, 200 and 4,000 Hz), intensity [13 levels from 55 to 115 dB(A) in 5-dB steps] and ear (left and right). For intensities above 90 dB myogenic vestibular evoked potentials (MVEP) were also obtained from the ipsilateral sternocleidomastoid muscle from which it was possible to obtain thresholds by linear regression of MVEP amplitudes against intensity. A further between-subjects factor was added which assessed subjects' attitude to vestibular sensations. The results indicate that across subjects there is a general trend of decreasing pleasantness with increasing intensity, but for the 200-Hz condition there is a significant positive departure from monotonicity in pleasantness (p<0.05) above the mean saccular threshold. However, when split by the between-subjects factor, the positive departure was only evident for those subjects who have a positive attitude to vestibular sensations (p < 0.01). Implications of these results for human responses to loud sound and the possible evolutionary significance of saccular acoustic sensitivity are discussed.     

Measuring hearing in the harbor seal (Phoca vitulina): comparison of behavioral and auditory brainstem response techniques

Auditory brainstem response (ABR) and standard behavioral methods were compared by measuring in-air audiograms for an adult female harbor seal (Phoca vitulina). Behavioral audiograms were obtained using two techniques: the method of constant stimuli and the staircase method. Sensitivity was tested from 0.250 to 30 kHz. The seal showed good sensitivity from 6 to 12 kHz [best sensitivity 8.1 dB (re 20 microPa2 x s) RMS at 8 kHz]. The staircase method yielded thresholds that were lower by 10 dB on average than the method of constant stimuli. ABRs were recorded at 2, 4, 8, 16, and 22 kHz and showed a similar best range (8-16 kHz). ABR thresholds averaged 5.7 dB higher than behavioral thresholds at 2, 4, and 8 kHz. ABRs were at least 7 dB lower at 16 kHz, and approximately 3 dB higher at 22 kHz. The better sensitivity of ABRs at higher frequencies could have reflected differences in the seal's behavior during ABR testing and/or bandwidth characteristics of test stimuli. These results agree with comparisons of ABR and behavioral methods performed in other recent studies and indicate that ABR methods represent a good alternative for estimating hearing range and sensitivity in pinnipeds, particularly when time is a critical factor and animals are untrained.     

The effects of real and illusory glides on pure-tone frequency discrimination

Experiment 1 measured pure-tone frequency difference limens (DLs) at 1 and 4 kHz. The stimuli had two steady-state portions, which differed in frequency for the target. These portions were separated by a middle section of varying length, which consisted of a silent gap, a frequency glide, or a noise burst (conditions: gap, glide, and noise, respectively). The noise burst created an illusion of the tone continuing through the gap. In the first condition, the stimuli had an overall duration of 500 ms. In the second condition, stimuli had a fixed 50-ms middle section, and the overall duration was varied. DLs were lower for the glide than for the gap condition, consistent with the idea that the auditory system contains a mechanism specific for the detection of dynamic changes. DLs were generally lower for the noise than for the gap condition, suggesting that this mechanism extracts information from an illusory glide. In a second experiment, pure-tone frequency direction-discrimination thresholds were measured using similar stimuli as for the first experiment. For this task, the type of the middle section hardly affected the thresholds, suggesting that the frequency-change detection mechanism does not facilitate the identification of the direction of frequency changes.     

The behavioral response of mice to gaps in noise depends on its spectral components and its bandwidth

The purpose of these experiments was to determine whether detecting brief decrements in noise level ("gaps") varies with the spectral content and bandwidth of noise in mice as it does in humans. The behavioral effect of gaps was quantified by their inhibiting a subsequent acoustic startle reflex. Gap durations from 1 to 29 ms were presented in five adjacent 1-octave noise bands and one 5-octave band, their range being 2 kHz to 64 kHz. Gaps ended 60 ms before the startle stimulus (experiment 1) or at startle onset (experiment 2). Asymptotic inhibition was greater for higher-frequency 1-octave bands and highest for the 5-octave band in both experiments, but time constants were related to frequency only in experiment 1. For the lowest band (2-4 kHz) neither noise decrements (experiment 1 and 2) nor increments (experiment 3) had any behavioral consequence, but this band was effective when presented as a pulse in quiet (experiment 4). The lowest frequencies in the most effective 1-octave band were one octave above the spectral region where mice have their best absolute thresholds. These effects are similar to those obtained in humans, and reveal a special contribution of wide band, high-frequency stimulation to temporal acuity.     

Pitch of amplitude-modulated irregular-rate stimuli in acoustic and electric hearing

The pitch of stimuli was studied under conditions where place-of-excitation was held constant, and where pitch was therefore derived from "purely temporal" cues. In experiment 1, the acoustical and electrical pulse trains consisted of pulses whose amplitudes alternated between a high and a low value, and whose interpulse intervals alternated between 4 and 6 ms. The attenuated pulses occurred after the 4-ms intervals in condition A, and after the 6-ms intervals in condition B. For both normal-hearing subjects and cochlear implantees, the period of an isochronous pulse train equal in pitch to this "4-6" stimulus increased from near 6 ms at the smallest modulation depth to nearly 10 ms at the largest depth. Additionally, the modulated pulse trains in condition A were perceived as being lower in pitch than those in condition B. Data are interpreted in terms of increased refractoriness in condition A, where the larger pulses are more closely followed by the smaller ones than in condition B. Consistent with this conclusion, the A-B difference was reduced at longer interpulse intervals. These findings provide a measure of supra-threshold effects of refractoriness on pitch perception, and increase our understanding of coding of temporal information in cochlear implant speech processing schemes.     

Design of broadband RF pulses with polynomial-phase response

The achievable bandwidth of common linear-phase RF pulses is limited by the maximum feasible B1 amplitude of the MR system. It has been shown previously, that this limitation can be circumvented by overlaying a quadratic phase in the frequency domain, which spreads the power across the pulse duration. Quadratic-phase RF pulses are near optimal in terms of achieving minimal B1max. In this work, it is demonstrated that further B1max reduction can be achieved by combining quadratic with higher-order polynomial-phase functions. RF pulses with a phase response up to tenth order were designed using the Shinnar-Le Roux transformation, yielding considerable increases in bandwidth and selectivity as compared to pure quadratic-phase pulses. These benefits are studied for a range of pulse specifications and demonstrated experimentally. For B1max = 20 microT and a pulse duration of 2.1 ms, it was possible to increase the bandwidth from 3.1 kHz for linear and 3.8 kHz for a quadratic to 9.9 kHz for a polynomial-phase pulse.     

Forward masking as a mechanism of automatic gain control in odontocete biosonar: a psychophysical study

In a false killer whale Pseudorca crassidens, echo perception thresholds were measured using a go/no-go psychophysical paradigm and one-up-one-down staircase procedure. Computer controlled echoes were electronically synthesized pulses that were played back through a transducer and triggered by whale emitted biosonar pulses. The echo amplitudes were proportional to biosonar pulse amplitudes; echo levels were specified in terms of the attenuation of the echo sound pressure level near the animal's head relative to the source level of the biosonar pulses. With increasing echo delay, the thresholds (echo attenuation factor) decreased from -49.3 dB at 2 ms to -79.5 dB at 16 ms, with a regression slope of -9.5 dB per delay doubling (-31.5 dB per delay decade). At the longer delays, the threshold remained nearly constant around -80.4 dB. Levels of emitted pulses slightly increased with delay prolongation (threshold decrease), with a regression slope of 3.2 dB per delay doubling (10.7 dB per delay decade). The echo threshold dependence on delay is interpreted as a release from forward masking by the preceding emitted pulse. This release may compensate for the echo level decrease with distance, thus keeping the echo sensation level for the animal near constant within a certain distance range.     

Intensity discrimination of Gaussian-windowed tones: indications for the shape of the auditory frequency-time window.

The just-noticeable difference in intensity jnd(I) was measured for 1-kHz tones with a Gaussian-shaped envelope as a function of their spectro-temporal shape. The stimuli, with constant energy and a constant product of bandwidth and duration, ranged from a long-duration narrow-band "tone" to a short-duration broadband "click." The jnd(I) was measured in three normal-hearing listeners at sensation levels of 0, 10, 20, and 30 dB in 35 dB(A) SPL pink noise. At intermediate sensation levels, jnd(I) depends on the spectro-temporal shape: at the extreme shapes (tones and clicks), intensity discrimination performance is best, whereas at intermediate shapes the jnd(I) is larger. Similar results are observed at a higher overall sound level, and at a higher carrier frequency. The maximum jnd(I) is observed for stimuli with an effective bandwidth of about 1/3 octave and an effective duration of 4 ms at 1 kHz (1 ms at 4 kHz). A generalized multiple-window model is proposed that assumes that the spectro-temporal domain is partitioned into "internal" auditory frequency-time windows. The model predicts that intensity discrimination thresholds depend upon the number of windows excited by a signal: jnd(I) is largest for stimuli covering one window.     

The effect of diotic and dichotic level-randomization on the binaural masking-level difference

Detection thresholds for tones in narrow-band noise were measured for two binaural configurations: N(o)S(o) and N(o)S(pi). The 30-Hz noise band had a mean overall level of 65 dB SPL and was centered on 250, 500, or 5000 Hz. Signals and noise were simultaneously gated for 500, 110, or 20 ms. Three conditions of level randomization were tested: (1) no randomization; (2) diotic randomization--the stimulus level (common to both ears) was randomly chosen from an uniformly distributed 40-dB range every presentation interval; and (3) dichotic randomization--the stimulus levels for each ear were each independently and randomly chosen from the 40-dB range. Regardless of binaural configuration, level randomization had small effects on thresholds at 500 and 110 ms, implying that binaural masking-level differences (BMLDs) do not depend on interaural level differences for individual stimuli. For 20-ms stimuli, both diotic and dichotic randomization led to markedly poorer performance than at 500- and 110-ms durations; BMLDs diminished with no randomization and dichotic randomization but not with diotic randomization. The loss of BMLDs at 20 ms, with degrees-of-freedom (2WT) approximately 1, implies that changes in intracranial parameters occurring during the course of the observation interval are necessary for BMLDs when mean-level and mean-intracranial-position cues have been made unhelpful.