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.     

Development of an electroacoustic analogue model of the middle ear and acoustic reflex

Experimental measurements of changes in the acoustic admittance of the eardrum caused by stapedius muscle contractions in human subjects are used to develop and electroacoustic analogue model of the middle ear. In this model the stapedius muscle is included as an explicit functional unit. The acoustical characteristics of the external ear canal are also included. The model is extensively evaluated by comparing its properties with the known characteristics of real ears of humans and other animals. Subsequently, the model is used to predict the effects of the acoustic reflex on middle ear sound transmission, which cannot easily be measured in humans. The model predicts attenuation of potentially hazardous high level sounds at frequencies below 1 kHz of up to about 10 dB, but very little effect at higher frequencies unless the reflex-eliciting stimulus is of sufficient intensity to cause partial disarticulation of the incudo-stapedial joint by stapedius muscle contraction. Overall attenuation for typical industrial noises is unlikely to be greater than approximately 3 dB(A) and will probably be even less in practice, resulting in little effective protection from the harmful effects of high intensity noise. It is considered that the model will be of benefit in the analysis of middle ear function, including the interpretation of audiological measurements of eardrum impedance and acoustic reflex response. This should lead to more versatile diagnosis of peripheral auditory dysfunction than has been possible hitherto.     

The effects of neural synchronization and peripheral compression on the acoustic-reflex threshold

This study investigates the acoustic reflex threshold (ART) dependency on stimulus phase utilizing low-level reflex audiometry [Neumann et al., Audiol. Neuro-Otol. 1, 359-369 (1996)]. The goal is to obtain optimal broadband stimuli for elicitation of the acoustic reflex and to obtain objective determinations of cochlear hearing loss. Three types of tone complexes with different phase characteristics were investigated: A stimulus that compensates for basilar-membrane dispersion, thus causing a large overall neural synchrony (basilar-membrane tone complex-BMTC), the temporally inversed stimulus (iBMTC), and random-phase tone complexes (rTC). The ARTs were measured in eight normal-hearing and six hearing-impaired subjects. Five different conditions of peak amplitude and stimulus repetition rate were used for each stimulus type. The results of the present study suggest that the ART is influenced by at least two different factors: (a) the degree of synchrony of neural activity across frequency, and (b) the fast-acting compression mechanism in the cochlea that is reduced in the case of a sensorineural hearing loss. The results allow a clear distinction of the two subjects groups based on the different ART for the utilized types and conditions of the stimuli. These differences might be useful for objective recruitment detection in clinical diagnostics.     

Fatigue and recovery of the human acoustic stapedius reflex in industrial noise.

The fatiguability of the acoustic stapedius muscle reflex in an actual noisy industrial environment was investigated in normal-hearing subjects. In a laboratory situation a small depression was found with a considerable individual variability. The stapedius reflex recovered slowly, approximately as a linear function of time. In a field study on an entire day of exposure in a ship-building yard the reflex depression was on the average 4 dB in response to a stimulation of 2000 Hz 10 min after the end of the workday. This corresponds to less than 8 dB immediately at the end of the exposure.     

The effects of external- and middle-ear filtering on auditory threshold and noise-induced hearing loss

A model of external- and middle-ear function is described that uses existing data to quantify the flow of sound power from the environment to the cochlea of humans, cats, and chinchillas. This model estimates the sound power produced at the entrance of the cochlea by an environmental sound stimulus, and can be used to predict the shape of the auditory threshold function and the relative potency of various traumatic acoustic stimuli. The shapes of the predicted and measured threshold functions in the three species are similar in best frequency, bandwidth, and low-frequency slope, and the model accurately predicts the hypersensitivity of the middle-frequency regions of the cochlea to acoustic trauma. The model assumes that the mechanics of the middle-ear system are linear even at high stimulus levels and does not include the effects of either middle-ear or cochlear efferent loops. The effects of these simplifications on the model are discussed as are the implications of the model results for hearing protection and damage risk criteria.     

Effects of frequency modulation on absolute, masked, and reflex thresholds

Behavioral and acoustic reflex thresholds were determined for five normal-hearing subjects in response to carrier signals of 500 and 2000 Hz which were unmodulated or modulated sinusoidally at rates of 2, 20, and 200 times per second with frequency deviations (delta f) of 30, 100, and 300 Hz. Behavioral (absolute and masked) thresholds were determined using an adaptive two-alternative forced-choice procedure. Acoustic reflex thresholds were determined by visual inspection of stored reflex waveforms. Frequency modulation was not found to exert a systematic effect at absolute threshold. Frequency modulation did affect threshold estimates systematically, but differentially, at masked threshold and acoustic reflex threshold. Increasing the frequency deviation of the modulation was associated with an increase in masked threshold and with a decrease in acoustic reflex threshold at both test frequencies. The findings are discussed in terms of critical band phenomena.     

Sensory threshold estimation from a continuously graded response produced by reflex modification audiometry

This article describes the use of reflex modification to determine sensory detection thresholds. The method is based upon the finding that low-intensity sensory stimuli presented shortly before a reflex eliciting stimulus are able to modify the amplitude of the reflex. The extent of such modification is related to the intensity of the initial low-intensity stimulus. In contrast to earlier reported procedures for threshold estimation, the method described in this article consists of fitting a smooth function to the relationship between startle response amplitude and the intensity of the inhibiting stimulus. The method entails fitting a cubic spline function to the medians of the square-root reflex amplitude at each prestimulus intensity. The resulting audiometric curves closely approximate audiometric data obtained from traditional operant methods both in sensitivity and shape. Parametric data are also presented that allow for optimizing stimulus presentation so as to obtain reliable thresholds using a minimal number of test trials. The procedures developed in this article may prove useful in other situations involving the estimation of a threshold effect from a continuously graded response.     

Repeated elicitation of the acoustic startle reflex leads to sensitisation in subsequent avoidance behaviour and induces fear conditioning

Background  

Autonomous reflexes enable animals to respond quickly to potential threats, prevent injury and mediate fight or flight responses. Intense acoustic stimuli with sudden onsets elicit a startle reflex while stimuli of similar intensity but with longer rise times only cause a cardiac defence response. In laboratory settings, habituation appears to affect all of these reflexes so that the response amplitude generally decreases with repeated exposure to the stimulus. The startle reflex has become a model system for the study of the neural basis of simple learning processes and emotional processing and is often used as a diagnostic tool in medical applications. However, previous studies did not allow animals to avoid the stimulus and the evolutionary function and long-term behavioural consequences of repeated startling remain speculative. In this study we investigate the follow-up behaviour associated with the startle reflex in wild-captured animals using an experimental setup that allows individuals to exhibit avoidance behaviour.     

Temporal integration of the contralateral acoustic-reflex threshold and its age-related changes

Although numerous studies have investigated temporal integration of the acoustic-reflex threshold (ART), research is lacking on the effect of age on temporal integration of the ART. Therefore the effect of age on temporal integration of the ART was investigated for a broad-band noise (BBN) activator. Subjects consisted of two groups of adults with normal-hearing sensitivity: one group of 20 young adults (ten males and ten females, ages 18-29 years, with a mean age of 24 years) and one group of 20 older adults (ten males and ten females, ages 59-75 years, with a mean age of 67.5 years). Activating stimulus durations were 12, 25, 50, 100, 200, 300, 500, and 1000 ms. Significant main effects for duration and age were obtained. That is, as the duration increased, the acoustic reflex threshold for BBN decreased. The interactions of duration x age group and duration x hearing level were not significant. The result of pair-wise analysis indicated statistically significant differences between the two age groups at durations of 20 ms and longer. The observed age effect on temporal integration of the ART for the BBN activator is interpreted in relation to senescent changes in the auditory system.     

The effects of aging on the stapedius reflex thresholds.

The acoustic reflex thresholds for broadband noise and 500-, 1000-, and 2000-Hz activating signals were measured in a group of young normal hearing adults and a group of elderly normal hearing subjects. The results indicated that the acoustic reflex thresholds for tonal activating signals in the young subjects were similar to those in the elderly subjects. However, the acoustic stapedius reflex thresholds for broadband noise activating signals is significantly higher in the elderly subjects than in the young. These differences were explained in light of Bredburg's findings [Acta Otolaryngol. Suppl. 236, 1--135 (1968)] regarding degeneration of outer hair cells as a function of aging.     

Detection of gaps in sinusoids and pulse trains by patients with cochlear implants

Gap detection thresholds were measured in patients with the Nucleus and Symbion cochlear implants as a function of several current waveform parameters. Detection of gaps in an electrical sinusoidal stimulus or in a train of biphasic pulses by implanted patients was similar to detection of gaps in comparable acoustic stimuli by normal listeners. Threshold gaps were 20-50 ms for low-level stimuli and improved with stimulus level to 2-5 ms for high-level stimuli. Gap detection performance was not affected by the electrode position in the cochlea or by the distance between stimulating electrodes. The data from most patients were well fitted by a trading relation between the duration of the gap and the square of stimulus intensity, indicating energy detection. The similarity of gap thresholds for normal subjects and implant patients suggests that many details of the peripheral neural activity are probably not important for this task, and that there is no retrocochlear loss of auditory temporal resolution with sensorineural hearing loss.     

The variability of acoustic reflex threshold measurements

Repeated measurements of the acoustic reflex threshold for various stimuli were obtained in normal subjects following a procedure similar to that commonly used in clinical audiological work, and also using a recording method with a well-defined criterion for determining the threshold value. The variability of measurements was estimated for each technique using a statistical model approach. The standard deviations of repeated measurements for a given stimulus were 2·8 and 2·5 dB, respectively, for the two methods. When the apparatus was refitted to the subject between measurements, the variability was increased. The small gain in repeatability resulting from the recording method is unlikely to be of practical significance in the clinical situation but may be of importance in research work where greater precision is desirable.     

Effects of background noise level on behavioral estimates of basilar-membrane compression

Hearing-impaired (HI) listeners often show poorer performance on psychoacoustic tasks than do normal-hearing (NH) listeners. Although some such deficits may reflect changes in suprathreshold sound processing, others may be due to stimulus audibility and the elevated absolute thresholds associated with hearing loss. Masking noise can be used to raise the thresholds of NH to equal the thresholds in quiet of HI listeners. However, such noise may have other effects, including changing peripheral response characteristics, such as the compressive input-output function of the basilar membrane in the normal cochlea. This study estimated compression behaviorally across a range of background noise levels in NH listeners at a 4 kHz signal frequency, using a growth of forward masking paradigm. For signals 5 dB or more above threshold in noise, no significant effect of broadband noise level was found on estimates of compression. This finding suggests that broadband noise does not significantly alter the compressive response of the basilar membrane to sounds that are presented well above their threshold in the noise. Similarities between the performance of HI listeners and NH listeners in threshold-equalizing noise are therefore unlikely to be due to a linearization of basilar-membrane responses to suprathreshold stimuli in the NH listeners.     

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.     

Monaural level discrimination under dichotic conditions

The ability to make judgments about the stimulus at one ear when a stimulus is simultaneously presented to the other ear was tested. Specifically, subjects discriminated the level of a 600 Hz target tone presented at the left ear while an identical-frequency distractor was simultaneously presented at the other ear. When there was no distractor, threshold was 0.7 dB. Threshold increased to 1.1 dB when a distractor with a fixed phase and level was introduced contra-aurally to the target. Further increases in threshold were observed when an across-presentation variability was introduced into the distractor phase (threshold of 1.6 dB) or level (threshold of 5.8 dB). When both the distractor level and phase varied, the largest threshold of 7.3 dB was obtained. These increases in threshold cannot be predicted by common binaural models, which assume that a target stimulus at one ear can be processed without interference from the stimulus at the nontarget ear. The measured thresholds are consistent with a model that utilizes two binaural dimensions that roughly correspond to the loudness and the position of a fused binaural image. The results show that, with binaurally fused tonal stimuli, subjects are unable to listen to one ear.     

Call recognition in the bullfrog, Rana catesbeiana: generalization along the duration continuum

Male bullfrogs emit multicroak, quasiharmonic advertisement calls that function in mate attraction and neighbor recognition. The degree of variability of acoustic features in these calls can influence perceptual decisions by conspecific receivers. Analysis of duration of individual croaks in spontaneous advertisement calls of a sample of males shows considerable intraindividual variability in this feature, even within short chorusing bouts. The influence of this intraindividual variability on behavior was examined in a series of evoked calling experiments. When presented with synthetic calls whose croak durations varied over the range of the natural variability in this feature, males responded similarly to intermediate and long duration croaks, but significantly less to short duration croaks. When presented with playbacks of calls with croak durations outside the natural range of variability, males again responded significantly less to shorter durations. The response gradient for duration is thus asymmetrical, with stimuli at the shorter end of the continuum evoking fewer responses than stimuli at the longer end. This asymmetry may be related to the biological demands of rejecting perception of heterospecific advertisement calls, and of mediating appropriate responses to conspecific aggressive calls. The shape of the response gradient for duration may reflect a process of stimulus generalization.     

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.     

Detection and discrimination of simulated motion of auditory targets in the horizontal plane

Three experiments investigated subjects' ability to detect and discriminate the simulated horizontal motion of auditory targets in an anechoic environment. "Moving" stimuli were produced by dynamic application of stereophonic balancing algorithms to a two-loudspeaker system with a 30 degree separation. All stimuli were 500-Hz tones. In experiment 1, subjects had to discriminate a left-to-right moving stimulus from a stationary stimulus pulsed for the same duration (300 or 600 ms). For both durations, minimum audible "movement" angles ("MAMA's") were on the order of 5 degrees for stimuli presented at 0 degrees azimuth (straight ahead), and increased to greater than 30 degrees for stimuli presented at +/- 90 degrees azimuth. Experiment 2 further investigated MAMA's at 0 degrees azimuth, employing two different procedures to track threshold: holding stimulus duration constant (at 100-600 ms) while varying velocity; or holding the velocity constant (at 22 degrees-360 degrees/s) while varying duration. Results from the two procedures agreed with each other and with the MAMA's determined by Perrott and Musicant for actually moving sound sources [J. Acoust. Soc. Am. 62, 1463-1466 (1977b)]: As stimulus duration decreased below 100-150 ms, the MAMA's increased sharply from 5 degrees-20 degrees or more, indicating that there is some minimum integration time required for subjects to perform optimally in an auditory spatial resolution task. Experiment 3 determined differential "velocity" thresholds employing simulated reference velocities of 0 degrees-150 degrees/s and stimulus durations of 150-600 ms. As with experiments 1 and 2, the data are more easily summarized by considering angular distance than velocity: For a given "extent of movement" of a reference target, about 4 degrees-10 degrees additional extent is required for threshold discrimination between two "moving" targets, more or less independently of stimulus duration or reference velocity. These data suggest that for the range of simulated velocities employed in these experiments, subjects respond to spatial changes--not velocity per se--when presented with a "motion" detection or discrimination task.     

Dolphin and sea lion auditory evoked potentials in response to single and multiple swept amplitude tones

Measurement of the auditory steady-state response (ASSR) is increasingly used to assess marine mammal hearing. These tests normally entail measuring the ASSR to a sequence of sinusoidally amplitude modulated tones, so that the ASSR amplitude function can be defined and the auditory threshold estimated. In this study, an alternative method was employed, where the ASSR was elicited by an amplitude modulated stimulus whose sound pressure level was slowly varied, or "swept," over a range of levels believed to bracket the threshold. The ASSR amplitude function was obtained by analyzing the resulting grand average evoked potential using a short-time Fourier transform. The suitability of this technique for hearing assessment of bottlenose dolphins and California sea lions was evaluated by comparing ASSR amplitude functions and thresholds obtained with swept amplitude and discrete, constant amplitude stimuli. When factors such as the number of simultaneous tones, the number of averages, and the frequency analysis window length were taken into account, the performance and time required for the swept-amplitude and discrete stimulus techniques were similar. The decision to use one technique over another depends on the relative importance of obtaining suprathreshold information versus the lowest possible thresholds.