Short-term poststimulatory response characteristics of the human acoustic stapedius reflex: monotic and dichotic stimulation.

Two experiments were performed to study short-term poststimulatory response characteristics of the human acoustic stapedius reflex in the time and intensity domains. In experiment 1, monotic magnitude-intensity functions (MIFs) were obtained for a 20-ms test stimulus preceded by a conditioning stimulus varying in duration (20, 50, 100, 500 ms) and level (-10, 0, +10 dB re: stapedius-reflex threshold) and temporally separated from the test stimulus by various interstimulus intervals (ISIs) (0, 20, 50, 100, 500 ms). Experiment 2 was similar in design except that conditioner and test stimuli were presented dichotically and fewer ISIs were used. Both experiments demonstrated that a prior conditioning stimulus produced significant increases in test-stimulus response magnitude. These poststimulatory effects were characterized by complex interactions among stimulus variables (conditioner duration, conditioner level, and interstimulus interval) with similar interactions occurring for both monotic and dichotic stimuli. A simple superposition effect of the responses to the conditioner and test stimulus does not account for the effect of prior stimulation since responses often exceeded the sum of the responses to the conditioner and the test stimulus alone.     

Frequency specificity of the human auditory brainstem and middle latency responses using notched noise masking

This study investigated the frequency specificity of the auditory brainstem and middle latency responses to 80 and 90 dB ppe SPL 500-Hz and 90 dB ppe SPL 2000-Hz tonebursts. The stimuli were brief (2-1-2 cycle) linear-gated tonebursts. ABR/MLRs were recorded using two electrode montages: (1) Cz-nape of neck and (2) Cz-ipsilateral earlobe. Cochlear contributions to ABR wave V-Na and MLR waves Na-Pa and Pa-Nb were assessed by plotting notched noise tuning curves which showed amplitudes and latencies as a function of center frequency of the noise masker [Abdala and Folsom, J. Acoust. Soc. Am. 97, 2394 (1995); ibid. 98, 921 (1995)]. Maxima in the response amplitude profiles for the ABR and MLR to 80 dB ppe SPL tonebursts occurred within one-half octave of the nominal stimulus frequency, with minimal contributions to the responses from frequencies greater than one octave away. At 90 dB ppe SPL, contributions came from a slightly broader frequency region for both stimulus frequencies. Thus, the ABR/MLR to 80 dB ppe SPL tonebursts shows good frequency specificity which decreases at 90 dB ppe SPL. No significant differences exist in frequency specificity of: (1) ABR wave V-Na versus MLR waves Na-Pa and Pa-Nb at either stimulus frequency or intensity; and (2) ABR/MLRs recorded using the two electrode montages.     

Frequency specificity of human auditory brainstem responses as revealed by pure-tone masking profiles

The frequency specificity of the auditory brainstem response (ABR) was examined by means of pure-tone masking profiles using click, 4000-Hz, and 1000-Hz filtered-click stimuli. Simultaneous pure-tone maskers were presented at one-half octave intervals around stimulus center frequency. Masking profiles at two intensities (60 and 40 dB SL) were obtained by measuring both latency and amplitude shifts in wave V as a result of the discrete-frequency maskers. Both latency and amplitude analyses showed masking profiles at 40 dB SL that were narrow and centered around stimulus frequency, whereas profiles at 60 dB SL showed high-frequency spread of the cochlear excitation area.     

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.     

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.     

Nonlinear phenomena as observed in the ear canal and at the auditory nerve

We report here several measures of nonlinear effects in the mammalian ear made in the external auditory meatus and in single neurons of the auditory nerve. We have measured the 2f1-f2 and the f2-f1 distortion products and we have found that the neural distortion product threshold curve for 2f1-f2 mirrors the low-frequency side of the frequency threshold curve, when the neural distortion product threshold curve of 2f1-f2 is plotted versus log(f2/f1) its slope is about 50 dB/oct and its intercept is 10-20 dB above the frequency threshold at the characteristic frequency CF, substantial 2f1-f2 distortion was seen in all animals studied while the f2-f1 distortion product was only rarely found at substantial levels, and the distortion product pressure observed in the ear canal was at a level equal to that detected at threshold by the neural units under study. We have also made measurements of two-tone rate suppression thresholds using two new and consistent threshold paradigms. We find that for high and intermediate characteristic frequency neural units the suppression threshold is independent of frequency and at a level of about 70 dB SPL, the suppression above CF is much less than below CF, and the tip of the frequency tuning curve can be suppressed by up to 40 dB by a low-frequency suppressor.     

Vibrotactile forward masking: evidence for channel independence

Threshold shifts for the detection of vibrotactile test stimuli were determined as a function of the intensity of a masker. A 50-ms sinusoidal test stimulus was applied to the thenar eminence of the hand 25 ms after the termination of a 700-ms sinusoidal masker applied to the same site. The intensity of the masker was varied over a range of 0-44 dB SL. The frequency of the masker was either 15 or 250 Hz and the frequency of the test stimulus was either 15, 25, 100, or 250 Hz. The results support the hypothesis that the detection of vibrotactile stimuli is mediated by at least two receptor systems which do not mask each other.     

Signal transmission in noisy environments: auditory masking in the tympanic nerve of the bushcricket Metaballus litus (Orthoptera: Tettigoniinae)

Physiological responses of the auditory leg nerve were recorded in the tettigoniid Metaballus litus to suprathreshold tone pulses of 12.45 kHz, which is close to the carrier frequency of the male's call. This stimulus tone frequency was determined by characterizing the polar response of the foreleg. Physiological threshold of the receptors was calculated from intensity input/output curves, and the experimental stimulus was set at 40 dB above this threshold value. There was low variance in threshold values between preparations. Continuous octave filtered white noise centered on the stimulus frequency was presented at the same time as the tone pulse at increasing intensities. The summed action potentials (SAPs) of the whole leg nerve were averaged over 256 stimulus presentations and the magnitude of the response was calibrated to dB values. The range of noise levels was set between that inducing no decrease in the SAP response to the tone pulse stimulus, up to a masking intensity where the response to the tone pulse was only just observable. Decrement in SAP magnitude was linear, and complete masking occurred when the noise level was 20-25 dB above the initial level of zero masking. This final level was comparable in magnitude to the sound-pressure level of the tone pulse and within the natural range of the insect's auditory behavior. Following the cessation of the noise signal, the SAPs were monitored over intervals of 2 min until the SAP asymptoted to the preexperimental condition. The reduction in SAP magnitude during noise presentation was attributed to a loss in synchrony from the individual tympanic receptors.     

Near-threshold equal-loudness contours for harbor seals (Phoca vitulina) derived from reaction times during underwater audiometry: a preliminary study

Equal-loudness functions describe relationships between the frequencies of sounds and their perceived loudness. This pilot study investigated the possibility of deriving equal-loudness contours based on the assumption that sounds of equal perceived loudness elicit equal reaction times (RTs). During a psychoacoustic underwater hearing study, the responses of two young female harbor seals to tonal signals between 0.125 and 100 kHz were filmed. Frame-by-frame analysis was used to quantify RT (the time between the onset of the sound stimulus and the onset of movement of the seal away from the listening station). Near-threshold equal-latency contours, as surrogates for equal-loudness contours, were estimated from RT-level functions fitted to mean RT data. The closer the received sound pressure level was to the 50% detection hearing threshold, the more slowly the animals reacted to the signal (RT range: 188-982 ms). Equal-latency contours were calculated relative to the RTs shown by each seal at sound levels of 0, 10, and 20 dB above the detection threshold at 1 kHz. Fifty percent detection thresholds are obtained with well-trained subjects actively listening for faint familiar sounds. When calculating audibility ranges of sounds for harbor seals in nature, it may be appropriate to consider levels 20 dB above this threshold.     

Basilar membrane mechanics at the base of the chinchilla cochlea. II. Responses to low-frequency tones and relationship to microphonics and spike initiation in the VIII nerve

Low-frequency stimuli (40- to 1000-Hz tones) have been used to correlate the motion of the 8-to 9-kHz place of the chinchilla basilar membrane with the cochlear microphonics recorded at the round window and with the responses of auditory nerve fibers with appropriate characteristic frequency. At the lowest stimulus frequencies, maximum displacement of the basilar membrane toward scala tympani occurs in near synchrony with maximum rarefaction at the eardrum and maximum negativity at the round window; at higher frequencies, the mechanical and microphonic response phases progressively lag rarefaction, reaching - 240 deg at 1000 Hz. At most frequencies (40-1000 Hz) near-threshold neural responses, once corrected for neural travel-time and synaptic delays, somewhat lead (by some 40 deg) maximal scala tympani displacement and maximal negativity of the round window microphonics. The variation of sensitivity with frequency is similar for basilar membrane displacement and microphonic responses: Under open-bulla conditions, sensitivity is constant for frequencies between 100 and 1000 Hz; below 100 Hz, sensitivity decreases at rates close to 12 dB/oct toward lower frequencies. Neural response sensitivity matches BM displacement more closely than BM velocity.     

Noise-induced temporary threshold shift and recovery in Yangtze finless porpoises Neophocaena phocaenoides asiaeorientalis

In Yangtze finless porpoises Neophocaena phocaenoides asiaeorientalis, the effects of fatiguing noise on hearing thresholds at frequencies of 32, 45, 64, and 128 kHz were investigated. The noise parameters were: 0.5-oct bandwidth, -1 to +0.5 oct relative to the test frequency, 150 dB re 1 μPa (140-160 dB re 1 μPa in one measurement series), with 1-30 min exposure time. Thresholds were evaluated using the evoked-potential technique allowing the tracing of threshold variations with a temporal resolution better than 1 min. The most effective fatiguing noise was centered at 0.5 octave below the test frequency. The temporary threshold shift (TTS) depended on the frequencies of the fatiguing noise and test signal: The lower the frequencies, the bigger the noise effect. The time-to-level trade of the noise effect was incomplete: the change of noise level by 20 dB resulted in a change of TTS level by nearly 20 dB, whereas the tenfold change of noise duration resulted in a TTS increase by 3.8-5.8 dB.     

Group delay measurement from spiral ganglion cells in the basal turn of the guinea pig cochlea

Measurements of group delay were made extracellularly from spiral ganglion cells in the 3.7 to 5.0-mm region of the guinea pig cochlea, using sinusoidally amplitude modulated tones with constant modulating frequency (100 Hz) and depth of modulation (0.19). Threshold cochlear tuning was accompanied by frequency-dependent group delays. The group delay on the low-frequency tail was independent of carrier frequency; the interunit variation was 0.28-1.28 ms. The difference in group delay between CF and the low-frequency tail decreased as the CF threshold increased (-0.09 +/- 0.02 ms per 10 dB, beginning at 0.62 +/- 0.07 ms at 0 dB SPL). The group delay decreased above CF; at the units' maximum frequency it was less than the low-frequency tail value, and was sometimes negative. Following arterial injections of furosemide the CF threshold increased and the group delay peak decreased; the low-frequency tail was unaffected. The group delay decreased with increasing intensity; the reduction near and above CF was not only larger than that on the low-frequency tail, but also the change at 5-10 dB above threshold was far greater than expected from the Q10dB of the suprathreshold iso-rate tuning curves. A minimum-phase analysis suggested that the group delay response above CF, together with its nonlinear behavior, can be accounted for by a high-frequency, level-independent, amplitude plateau, in combination with the single unit, amplitude nonlinearity which is known to exist above CF.     

Multicomponent stimulus interactions observed in basilar-membrane vibration in the basal region of the chinchilla cochlea.

Multicomponent stimuli consisting of two to seven tones were used to study suppression of basilar-membrane vibration at the 3-4-mm region of the chinchilla cochlea with a characteristic frequency between 6.5 and 8.5 kHz. Three-component stimuli were amplitude-modulated sinusoids (AM) with modulation depth varied between 0.25 and 2 and modulation frequency varied between 100 and 2000 Hz. For five-component stimuli of equal amplitude, frequency separation between adjacent components was the same as that used for AM stimuli. An additional manipulation was to position either the first, third, or fifth component at the characteristic frequency (CF). This allowed the study of the basilar-membrane response to off-CF stimuli. CF suppression was as high as 35 dB for two-tone combinations, while for equal-amplitude stimulus components CF suppression never exceeded 20 dB. This latter case occurred for both two-tone stimuli where the suppressor was below CF and for multitone stimuli with the third component=CF. Suppression was least for the AM stimuli, including when the three AM components were equal. Maximum suppression was both level- and frequency dependent, and occurred for component frequency separations of 500 to 600 Hz. Suppression decreased for multicomponent stimuli with component frequency spacing greater than 600 Hz. Mutual suppression occurred whenever stimulus components were within the compressive region of the basilar membrane.     

Differential responses to acoustic damage and furosemide in auditory brainstem and otoacoustic emission measures

Characteristics of distortion product otoacoustic emissions (DPOAEs) and auditory brainstem responses (ABRs) were measured in Mongolian gerbil before and after the introduction of two different auditory dysfunctions: (1) acoustic damage with a high-intensity tone, or (2) furosemide intoxication. The goal was to find emission parameters and measures that best differentiated between the two dysfunctions, e.g., at a given ABR threshold elevation. Emission input-output or "growth" functions were used (frequencies f1 and f2, f2/f1 = 1.21) with equal levels, L1 = L2, and unequal levels, with L1 = L2 + 20 dB. The best parametric choice was found to be unequal stimulus levels, and the best measure was found to be the change in the emission threshold level, delta x. The emission threshold was defined as the stimulus level required to reach a criterion emission amplitude, in this case -10 dB SPL. (The next best measure was the change in emission amplitude at high stimulus levels, specifically that measured at L1 x L2 = 90 x 70 dB SPL.) For an ABR threshold shift of 20 dB or more, there was essentially no overlap in the emission threshold measures for the two conditions, sound damage or furosemide. The dividing line between the two distributions increased slowly with the change in ABR threshold, delta ABR, and was given by delta x(t) = 0.6 delta ABR + 8 dB. For a given delta ABR, if the shift in emission threshold was more than the calculated dividing line value, delta x(t), the auditory dysfunction was due to acoustic damage, if less, it was due to furosemide.     

Thresholds for the detection of inharmonicity in complex tones

Thresholds were measured for the detection of inharmonicity in complex tones. Subjects were required to distinguish a complex tone whose partials were all at exact harmonic frequencies from a similar complex tone with one of the partials slightly mistuned. The mistuning which allowed 71% correct identification in a two-alternative forced-choice task was estimated for each partial in turn. In experiment I the fundamental frequency was either 100, 200, or 400 Hz, and the complex tones contained the first 12 harmonics at equal levels of 60 dB SPL per component. The stimulus duration was 410 ms. For each fundamental the thresholds were roughly constant when expressed in Hz, having a mean value of about 4 Hz (range 2.4-7.3 Hz). In experiment II the fundamental frequency was fixed at 200 Hz, and thresholds for inharmonicity were measured for stimulus durations of 50, 110, 410, and 1610 ms. For harmonics above the fifth the thresholds increased from less than 1 Hz to about 40 Hz as duration was decreased from 1610-50 ms. For the lower harmonics (up to the fourth) threshold changed much less with duration, and for the three shorter durations thresholds for each duration were roughly a constant proportion of the harmonic frequency. The results suggest that inharmonicity is detected in different ways for high and low harmonics. For low harmonics the inharmonic partial appears to "stand out" from the complex tone as a whole. For high harmonics the mistuning is detected as a kind of "beat" or "roughness," presumably reflecting a sensitivity to the changing relative phase of the mistuned harmonic relative to the other harmonics.(ABSTRACT TRUNCATED AT 250 WORDS)     

Vibrotactile masking: effects of stimulus onset asynchrony and stimulus frequency

Vibrotactile thresholds for the detection of a 50-ms vibratory stimulus on the thenar eminence of the hand were measured in the presence of and in the absence of a 700-ms suprathreshold vibratory masking stimulus. When thresholds were measured in the presence of the masking stimulus, stimulus onset asynchrony (SOA) was varied so that backward, simultaneous, and forward masking could be measured. The amount of masking, expressed as threshold shift, was greatest when the test stimulus was presented near the onset or offset of the masking stimulus. For both backward and forward masking, the amount of masking decreased as a function of increasing stimulus onset asynchrony. Comparisons were made of the amounts of masking measured when the test and masking stimuli were both sinusoids, and when the test stimulus was a sinusoid and the masking stimulus was noise. In all conditions, the masked threshold decreased approximately 4.0 dB when SOA was increased from 100 to 650 ms with reference to the onset of the 700-ms masking stimulus. More simultaneous masking was observed when sinusoidal test stimuli were detected in the presence of noise than when they were detected in the presence of sinusoidal maskers of the same frequency. The functions were essentially identical for detection of a low-frequency (20 Hz) test stimulus mediated by a non-Pacinian channel and detection of a high-frequency (250 Hz) test stimulus mediated by the Pacinian channel.     

The time required to focus on a cued signal frequency

How quickly can a listener focus on a single tonal cue that indicates the frequency of an upcoming signal? Initial measurements were made with frequency uncertainty (signal frequency varies randomly from trial to trial) and with certainty (same frequency on all trials). Measured by a yes-no procedure, thresholds for 40- and 20-ms signals presented in continuous broadband noise at 50 dB SPL were higher in uncertainty than in certainty; the difference decreased monotonically from 5 dB at frequencies below 500 Hz to under 3 dB above about 2500 Hz. This decrease in the detrimental effect from uncertainty, which comes about with increasing signal frequency, may result from preferential attention to higher frequencies. In a second experiment, frequency again varied randomly, but each trial now began with a cue at the signal frequency. The critical variable was the delay from cue onset to signal onset. A delay of 352 ms eliminated the detrimental effect of frequency uncertainty at all frequencies. At the shortest delays of 52 and 82 ms the detrimental effect was reduced primarily at lower frequencies. Our analysis suggests that shifting focus to a cued frequency region, under optimal stimulus conditions, requires less than 52 ms.     

Fine structure of hearing threshold and loudness perception

Hearing thresholds measured with high-frequency resolution show a quasiperiodic change in level called threshold fine structure (or microstructure). The effect of this fine structure on loudness perception over a range of stimulus levels was investigated in 12 subjects. Three different approaches were used. Individual hearing thresholds and equal loudness contours were measured in eight subjects using loudness-matching paradigms. In addition, the loudness growth of sinusoids was observed at frequencies associated with individual minima or maxima in the hearing threshold from five subjects using a loudness-matching paradigm. At low levels, loudness growth depended on the position of the test- or reference-tone frequency within the threshold fine structure. The slope of loudness growth differs by 0.2 dB/dB when an identical test tone is compared with two different reference tones, i.e., a difference in loudness growth of 2 dB per 10-dB change in stimulus. Finally, loudness growth was measured for the same five subjects using categorical loudness scaling as a direct-scaling technique with no reference tone instead of the loudness-matching procedures. Overall, an influence of hearing-threshold fine structure on loudness perception of sinusoids was observable for stimulus levels up to 40 dB SPL--independent of the procedure used. Possible implications of fine structure for loudness measurements and other psychoacoustic experiments, such as different compression within threshold minima and maxima, are discussed.     

Level discrimination as a function of level for tones from 0.25 to 16 kHz

Difference limens for level (delta L in dB = 20 log [(p + delta p)/p], where p is pressure) were measured as a function of level for tones at 0.25, 0.5, 1, 2, 4, 8, 10, 12, 14, and 16 kHz. At each frequency, test levels encompassed the range from near threshold to 95 dB SPL in steps of 10 dB or smaller. The stimulus duration was 500 ms and the interstimulus interval was 250 ms. An adaptive two-alternative forced-choice procedure with feedback was used. Results for six normal listeners show individual differences among listeners, but the general trends seen in the average data clearly are present in the individual data and show the following. First, the delta Ls at all but the highest frequencies are generally smaller at high levels than at low levels. Second, the delta Ls at equal SPLs are largely independent of frequency up to about 4 kHz, but increase with frequency above 4 kHz. Third, at 8 and 10 kHz, the delta Ls are clearly nonmonotonic functions of level, showing consistent deterioration in the mid-level delta Ls relative to the low- and high-level delta Ls. The present data are discussed qualitatively in terms of current models of level discrimination.     

Excess masking among listeners with a sensorineural hearing loss

Three experiments were conducted to determine whether listeners with a sensorineural hearing loss exhibited greater than normal amounts of masking at frequencies above the frequency of the masker. Excess masking was defined as the difference (in dB) between the masked thresholds actually obtained from a hearing-impaired listener and the expected thresholds calculated for the same individual. The expected thresholds were the power sum of the listener's thresholds in quiet and the average masked thresholds obtained from a group of normal-hearing subjects at the test frequency. Hearing-impaired listeners, with thresholds in quiet ranging from approximately 35-70 dB SPL (at test frequencies between 500-3000 Hz), displayed approximately 12-15 dB of maximum excess masking. The maximum amount of excess masking occurred in the region where the threshold in quiet of the hearing-impaired listener and the average normal masked threshold were equal. These findings indicate that listeners with a sensorineural hearing loss display one form of reduced frequency selectivity (i.e., abnormal upward spread of masking) even when their thresholds in quiet are taken into account.