Stimulus dependencies of the gerbil brain-stem auditory-evoked response (BAER). I: Effects of click level, rate, and polarity

Three experiments evaluating the effects of various stimulus manipulations on the click-evoked gerbil brain-stem auditory-evoked response (BAER) are reported. In experiment 1, click polarity and level were covaried. With increasing click level, there is a parallel decrease in the latency of the first five BAER peaks (i-v) and an increase in BAER peak amplitudes. Mean wave i amplitude was greater for rarefaction than condensation clicks at high click levels; mean wave v amplitude was greater for condensation clicks at higher click levels. Experiment 2 covaried click rate and polarity. The latency of the BAER peaks increased with increasing click repetition rate. This rate-dependent latency increase was greater for the later BAER peaks, resulting in an increase in the i-v interval with increasing click rate. As rate increased, the amplitudes of waves i and v decreased monotonically, whereas the amplitudes of waves ii-iv were largely uninfluenced by click rate. As in experiment 1, mean wave i amplitude was greater for rarefaction clicks, whereas mean wave v amplitude was greater for condensation clicks. The magnitude of these polarity dependencies on waves i and v amplitude decreased with increasing click rate. Experiment 3 evaluated the effects of click polarity on BAERs to high-intensity (100 dB pSPL) clicks presented at a rate of 10 Hz. In eight of ten gerbils evaluated, wave i amplitude was greater to rarefaction clicks, and, in all ten animals, wave v amplitude was greater to condensation clicks. The effects of click level and rate on BAER peak amplitudes, latencies, and interwave intervals are reminiscent of stimulus dependencies reported for the human BAER. The effects of click polarity on the amplitudes of waves i and v of the gerbil BAER have also been reported for the human BAER.     

A comparison of steady-state evoked potentials to modulated tones in awake and sleeping humans.

Steady-state evoked potential responses were measured to binaural amplitude-modulated (AM) and combined amplitude- and frequency-modulated (AM/FM) tones. For awake subjects, AM/FM tones produced larger amplitude responses than did AM tones. Awake and sleeping responses to 30-dB HL AM/FM tones were compared. Response amplitudes were lower during sleep and the extent to which they differed from awake amplitudes was dependent on both carrier and modulation frequencies. Background EEG noise at the stimulus modulation frequency was also reduced during sleep and varied with modulation frequency. A detection efficiency function was used to indicate the modulation frequencies likely to be most suitable for electrical estimation of behavioral threshold. In awake subjects, for all carrier frequencies tested, detection efficiency was highest at a modulation frequency of 45 Hz. In sleeping subjects, the modulation frequency regions of highest efficiency varied with carrier frequency. For carrier frequencies of 250 Hz, 500 Hz, and 1 kHz, the highest efficiencies were found in two modulation frequency regions centered on 45 and 90 Hz. For 2 and 4 kHz, the highest efficiencies were at modulation frequencies above 70 Hz. Sleep stage affected both response amplitude and background EEG noise in a manner that depended on modulation frequency. The results of this study suggest that, for sleeping subjects, modulation frequencies above 70 Hz may be best when using steady-state potentials for hearing threshold estimation.     

Stimulus dependencies of the gerbil brain-stem auditory-evoked response (BAER). III: Additivity of click level and rate with noise level

Two experiments were performed that evaluated the effects of ipsilateral-direct broadband noise maskers on the gerbil brain-stem auditory-evoked response (BAER) to click stimuli. In experiment 1, clicks were presented at 27 Hz at levels including 70, 80, 90, and 100 dB pSPL. Noise conditions included a no-noise control, and included noise levels varying in 10-dB increments from 20 dB SPL to a maximum noise level of 50, 60, 70, and 80 dB SPL for click levels of 70, 80, 90, and 100 dB pSPL, respectively. Gerbil BAER peaks were labeled with small roman numerals to distinguish them from human BAER peaks. The dependent variables included waves i and v latencies and amplitudes. Peak latencies increased and peak amplitudes decreased with decreasing click level and increasing noise level. To a first approximation, peak latencies and amplitudes showed changes with increasing noise level that were similar across click level. With increasing click level, there was little or no effect on the i-v interval. There was an increase in the i-v interval with increasing noise level. In experiment 2, click level was held constant at 90 dB pSPL, and click rates included 15, 40, 65, and 90 Hz. For each click rate, noise conditions included a no-noise control, and noise levels included 20, 30, 40, 50, 60, and 70 dB SPL. With increasing click rate and noise level, there was an increase in peak latencies, an increase in the i-v interval, and a decrease in peak amplitudes. The magnitude of peak latency and amplitude shifts with increasing click rate was dependent on noise level. Specifically, the magnitude of rate-dependent changes decreased with increasing level of broadband noise. These data are compared to human BAER experiments, and are found to be in fundamental agreement.     

Potentials evoked by the sinusoidal modulation of the amplitude or frequency of a tone

Steady state responses to the sinusoidal modulation of the amplitude or frequency of a tone were recorded from the human scalp. For both amplitude modulation (AM) and frequency modulation (FM), the responses were most consistent at modulation frequencies between 30 and 50 Hz. However, reliable responses could also be recorded at lower frequencies, particularly at 2-5 Hz for AM and at 3-7 Hz for FM. With increasing modulation depth at 40 Hz, both the AM and FM response increased in amplitude, but the AM response tended to saturate at large modulation depths. Neither response showed any significant change in phase with changes in modulation depth. Both responses increased in amplitude and decreased in phase delay with increasing intensity of the carrier tone, the FM response showing some saturation of amplitude at high intensities. Both responses could be recorded at modulation depths close to the subjective threshold for detecting the modulation and at intensities close to the subjective threshold for hearing the stimulus. The responses were variable but did not consistently adapt over periods of 10 min. The 40-Hz AM and FM responses appear to originate in the same generator, this generator being activated by separate auditory systems that detect changes in either amplitude or frequency.     

Physiological studies of central masking in man. II: Tonepip SSRs and the masking level difference.

The auditory steady-state response (SSR), an evoked response generated in the auditory cortex, was initiated by monaural trains of 500-Hz tonepips repeated at rates near 40 Hz while wideband noise was being delivered to the same or opposite ear. Contralateral noise reduced SSR amplitudes in an intensity-dependent manner, whereas ipsilateral noise enhanced the SSR amplitudes at low levels and depressed them at high levels. Systematic phase changes accompanied the amplitude changes. These results, obtained with tonepips, closely resemble those previously reported for clicks. A third experiment, a masking level difference (MLD) experiment, examined changes in the SSR measures during four successive tonepip-plus-noise conditions: (1) monaural tonepips alone; (2) adding ipsilateral noise; (3) then adding contralateral noise; (4) finally, adding contralateral tonepips. The SSR amplitude changes measured in the experiment did not always correspond with the changes in perception reported by the subject.     

Human brain-stem auditory evoked responses obtained by cross correlation to trains of clicks, noise bursts, and tone bursts

Brain-stem auditory evoked responses (BAERs) were obtained in eight normal-hearing young adults. Stimuli included clicks, noise bursts, and tone bursts. Tone bursts included carrier frequencies of 1, 2, 4, and 8 kHz. All stimuli were presented at 60 dB nHL. BAERs were obtained by presenting stimuli in pseudorandom trains, called maximum length sequences (MLSs). BAERs were recovered by cross correlating the responses with a recovery sequence. MLS-BAERs were obtained with minimum pulse intervals (MPIs) of 6, 4, and 2 ms. Conventional BAERs were also obtained for stimuli presented at a rate of 30 Hz. BAERs were obtained for all stimuli, for both the conventional averaging technique and for the cross-correlation technique. BAERs were observed for MPIs as short as 2 ms for all stimuli. Wave V was the only peak consistently identifiable for these stimuli. For all stimuli, wave V latency increased and wave V amplitude decreased with decreasing MPI. This is the first demonstration of the use of maximum length sequences combined with cross correlation to obtain BAERs to noise burst and tone burst stimuli.     

Near-field responses from the round window, inferior colliculus, and auditory cortex of the unanesthetized chinchilla: manipulations of noiseburst level and rate.

Few studies have compared the response properties of near-field potentials from multiple levels of the auditory nervous system of unanesthetized animals. The purpose of this study was to investigate the effects of brief-duration noisebursts on neural responses recorded from electrodes chronically implanted at the round window, inferior colliculus and auditory cortex of chinchillas. Responses were obtained from seven unanesthetized chinchillas to a noiseburst-level and noiseburst-rate series. For the noiseburst-rate series, a 70 dB pSPL noiseburst was varied in rate from 10 to 100 Hz using conventional averaging procedures, and from 100 to 500 Hz using pseudorandom pulse trains called maximum length sequences (MLSs). Response thresholds were similar for the compound action potential (CAP), inferior colliculus potential (ICP) and auditory cortex potential (ACP). With decreasing noiseburst level, there were decreases in the amplitudes and increases in the latencies of the CAP, ICP and ACP. The shapes of the mean normalized amplitude input/output (I/O) functions were similar for the ICP and ACP, while the normalized I/O functions for the first positive peak (P1) and first negative peak (N1) of the CAP differed from each other and from the ICP and ACP. The slopes of the latency/intensity functions were shallowest for the CAP, intermediate for the ICP, and steepest for the ACP. With increasing rate, the latency shift was least for the CAP, intermediate for the ICP and greatest for the ACP. The amplitude of P1 of the CAP varied little with rate. All other potentials showed a pronounced decrease in amplitude at high stimulation rates. Excluding CAP P1, proportional amplitude decrease with rate was greatest for the ACP, intermediate for N1 of the CAP and least for the ICP. Responses were present in most animals at all recording sites, even for the highest rate (500 Hz) used in this study. For all potentials, the MLS procedure allowed the collection of a response at rates well above those where sequential responses would have overlapped using conventional averaging procedures.     

Hearing sensitivity during target presence and absence while a whale echolocates

Hearing sensitivity was measured in a false killer whale during echolocation. Sensitivity was measured using probe stimuli as sinusoidally amplitude modulated signals with a 22.5-kHz carrier frequency and recording auditory evoked potentials as envelope-following responses. The probes were presented and responses were recorded during short 2-s periods when the animal echolocated to detect the presence or absence of a target in a go/no-go paradigm. In the target-absent trials, a hearing threshold of 90.4 dB re 1 muPa was found; in the target-present trials, the threshold was 109.8 dB. Thus, a 19.4-dB difference was found between thresholds in the target-present and target-absent trials. To check the possibility that this difference was the result of different masking degree of the probe by the emitted sonar clicks, click statistics were investigated in similar trials. No indication was found that the energy of the emitted clicks was higher in the target-present than in target-absent trials; on the contrary, mean click level, mean number of clicks per train, and overall train energy was slightly higher in the target-absent trials. Thus the data indicate that the hearing sensitivity of the whale varied depending on target presence or absence.     

The effect of broadband noise on the human brainstem auditory evoked response. II. Frequency specificity

A series of experiments evaluated the effects of broadband noise (ipsilateral) on wave V of the brainstem auditory evoked response (BAER) elicited by tone bursts or clicks in the presence of high-pass masking noise. Experiment 1 used 1000- and 4000-Hz, 60-dB nHL tone bursts in the presence of broadband noise. With increasing noise level, wave V latency shift was greater for the 1000-Hz tone bursts, while amplitude decrements were similar for both tone-burst frequencies. Experiment 2 varied high-pass masker cutoff frequency and the level of subtotal masking in the presence of 50-dB nHL clicks. The effects of subtotal masking on wave V (increase in latency and decrease in amplitude) increased with increasing derived-band frequency. Experiment 3 covaried high-pass masker cutoff frequency and subtotal masking level for 1000- and 4000-Hz tone-burst stimuli. The effect of subtotal masking on wave V latency was reduced for both tone-burst frequencies when the response-generating region of the cochlear partition was limited by high-pass maskers. The results of these three experiments suggest that most of the wave V latency shift associated with increasing levels of broadband noise is mediated by a place mechanism when the stimulus is a moderate intensity (60 dB nHL), low-frequency (1000 Hz) tone burst. However, the interpretation of the latency shifts produced by broadband noise for 4000-Hz tone-burst stimuli is made more complex by multiple technical factors discussed herein.     

The role of auditory feedback in sustaining vocal vibrato

Vocal vibrato and tremor are characterized by oscillations in voice fundamental frequency (F0). These oscillations may be sustained by a control loop within the auditory system. One component of the control loop is the pitch-shift reflex (PSR). The PSR is a closed loop negative feedback reflex that is triggered in response to discrepancies between intended and perceived pitch with a latency of approximately 100 ms. Consecutive compensatory reflexive responses lead to oscillations in pitch every approximately 200 ms, resulting in approximately 5-Hz modulation of F0. Pitch-shift reflexes were elicited experimentally in six subjects while they sustained /u/ vowels at a comfortable pitch and loudness. Auditory feedback was sinusoidally modulated at discrete integer frequencies (1 to 10 Hz) with +/- 25 cents amplitude. Modulated auditory feedback induced oscillations in voice F0 output of all subjects at rates consistent with vocal vibrato and tremor. Transfer functions revealed peak gains at 4 to 7 Hz in all subjects, with an average peak gain at 5 Hz. These gains occurred in the modulation frequency region where the voice output and auditory feedback signals were in phase. A control loop in the auditory system may sustain vocal vibrato and tremorlike oscillations in voice F0.     

The influence of carrier level and frequency on modulation and beat-detection thresholds for sinusoidal carriers

This paper is concerned with modulation and beat detection for sinusoidal carriers. In the first experiment, temporal modulation transfer functions (TMTFs) were measured for carrier frequencies between 1 and 10 kHz. Modulation rates covered the range from 10 Hz to about the rate equaling the critical bandwidth at the carrier frequency. In experiment 2, TMTFs for three carrier frequencies were obtained as a function of the carrier level. In the final experiment, thresholds for the detection of either the lower or the upper modulation sideband (beat detection) were measured for "carrier" frequencies of 5 and 10 kHz, using the same range of modulation rates as in experiment 1. The TMTFs for carrier frequencies of 2 kHz and higher remained flat up to a modulation rate of about 100-130 Hz and had similar values across carrier frequencies. For higher rates, modulation thresholds initially increased and then decreased rapidly, reflecting the subjects' ability to resolve the sidebands spectrally. Detection thresholds generally improved with increasing carrier level, but large variations in the exact level dependence were observed, across subjects as well as across carrier frequencies. For beat rates up to about 70 Hz (at 5 kHz) and 100 Hz (at 10 kHz), beat detection thresholds were the same for the upper and the lower sidebands and were about 6 dB higher than the level per sideband at the modulation-detection threshold. At higher rates the threshold for both sidebands increased, but the increase was larger for the lower sideband. This reflects an asymmetry in masking with more masking towards lower frequencies. Only at rates well beyond the maximum of the TMTF did detection for the lower sideband start to be better than that for the upper sideband. The asymmetry at intermediate frequency separations can be explained by assuming that detection always takes place in filters centered above the stimulus spectrum. The shape of the TMTF and the beat-detection data reflects a limitation in resolving fast amplitude variations, which must occur central to the inner-ear filtering. Its characteristic resembles that of a first-order low-pass filter with a cutoff frequency of about 150 Hz.     

Acoustically dependent latency shifts of BSER (wave V) in man

The latencies of wave V in Brain Stem Evoked Responses (BSER) elicited by a set of acoustic transients were measured. The stimuli were produced by delivering pulses to two filters, arranged in series. The filters were set so that the maximum acoustic energy in the transients, i.e., filtered clicks, occurred at 0.5, 1, 2, 4, or 8 kHz. The filtered clicks were presented via earphones at a rate of 30/s at 20, 40, or 60 dB HL to ten subjects with normal hearing. The latencies of wave V varied systematically with center frequency of the filtered clicks when they were each at the same HL. Stimuli presented at 40 dB HL produced the greatest opportunity for relating stimulus frequency to latency. The latencies for a smaller set of responses to stimuli presented at 10/s were the same as those for the principal data taken at 30/s. The changes in latency of wave V due to frequency are similar to those observed by other investigators in whole-nerve responses recorded in man.     

Intrinsic envelope fluctuations and modulation-detection thresholds for narrow-band noise carriers

A model is presented which calculates the intrinsic envelope power of a bandpass noise carrier within the passband of a hypothetical modulation filter tuned to a specific modulation frequency. Model predictions are compared to experimentally obtained amplitude modulation (AM) detection thresholds. In experiment 1, thresholds for modulation rates of 5, 25, and 100 Hz imposed on a bandpass Gaussian noise carrier with a fixed upper cutoff frequency of 6 kHz and a bandwidth in the range from 1 to 6000 Hz were obtained. In experiment 2, three noises with different spectra of the intrinsic fluctuations served as the carrier: Gaussian noise, multiplied noise, and low-noise noise. In each case, the carrier was spectrally centered at 5 kHz and had a bandwidth of 50 Hz. The AM detection thresholds were obtained for modulation frequencies of 10, 20, 30, 50, 70, and 100 Hz. The intrinsic envelope power of the carrier at the output of the modulation filter tuned to the signal modulation frequency appears to provide a good estimate for AM detection threshold. The results are compared with predictions on the basis of the more complex auditory processing model by Dau et al.     

A perceptual learning investigation of the pitch elicited by amplitude-modulated noise

Noise that is amplitude modulated at rates ranging from 40 to 850 Hz can elicit a sensation of pitch. Here, the processing of this temporally based pitch was investigated using a perceptual-learning paradigm. Nine listeners were trained (1 hour per day for 6-8 days) to discriminate a standard rate of sinusoidal amplitude modulation (SAM) from a faster rate in a single condition (150 Hz SAM rate, 5 kHz low-pass carrier). All trained listeners improved significantly on that condition. These trained listeners subsequently showed no more improvement than nine untrained controls on pure-tone and rippled-noise discrimination with the same pitch, and on SAM-rate discrimination with a 30 Hz rate, although they did show some improvement with a 300 Hz rate. In addition, most trained, but not control, listeners were worse at detecting SAM at 150 Hz after, compared to before training. These results indicate that listeners can learn to improve their ability to discriminate SAM rate with multiple-hour training and that the mechanism that is modified by learning encodes (1) the pitch of SAM noise but not that of pure tones and rippled noise, (2) different SAM rates separately, and (3) differences in SAM rate more effectively than cues for SAM detection.     

Auditory brainstem responses to chirps delivered by different insert earphones

The frequency response and sensitivity of the ER-3A and ER-2 insert earphones are measured in the occluded-ear simulator using three ear canal extensions. Compared to the other two extensions, the DB 0370 (Bru?el & Kj?r), which is recommended by the international standards, introduces a significant resonance peak around 4500 Hz. The ER-3A has an amplitude response like a band-pass filter (1400 Hz, 6 dB/octave -4000 Hz, -36 dB/octave), and a group delay with "ripples" of up to ±0.5 ms, while the ER-2 has an amplitude response, and a group delay which are flat and smooth up to above 10000 Hz. Both earphones are used to record auditory brainstem responses, ABRs, from 22 normal-hearing ears in response to two chirps and a click at levels from 20 to 80 dB nHL. While the click-ABRs are slightly larger for ER-2 than for ER-3A, the chirp-ABRs are much larger for ER-2 than for ER-3A at levels below 60 dB nHL. With a simulated amplitude response of the ER-3A and the smooth group delay of the ER-2 it is shown that the increased chirp-ABR amplitude with the ER-2 is caused by its broader amplitude response and not by its smoother group delay.     

Modeling auditory evoked brainstem responses to transient stimuli

A quantitative model is presented that describes the formation of auditory brainstem responses (ABRs) to tone pulses, clicks, and rising chirps as a function of stimulation level. The model computes the convolution of the instantaneous discharge rates using the "humanized" nonlinear auditory-nerve model of Zilany and Bruce [J. Acoust. Soc. Am. 122, 402-417 (2007)] and an empirically derived unitary response function which is assumed to reflect contributions from different cell populations within the auditory brainstem, recorded at a given pair of electrodes on the scalp. It is shown that the model accounts for the decrease of tone-pulse evoked wave-V latency with frequency but underestimates the level dependency of the tone-pulse as well as click-evoked latency values. Furthermore, the model correctly predicts the nonlinear wave-V amplitude behavior in response to the chirp stimulation both as a function of chirp sweeping rate and level. Overall, the results support the hypothesis that the pattern of ABR generation is strongly affected by the nonlinear and dispersive processes in the cochlea.     

Subcomponent cues in binaural unmasking

The addition of a signal in the N0Sπ binaural configuration gives rise to fluctuations in interaural phase and amplitude. Sensitivity to these individual cues was measured by applying sinusoidal amplitude modulation (AM) or quasi-frequency modulation (QFM) to a band of noise. Discrimination between interaurally in-phase and out-of-phase modulation was measured using an adaptive task for narrow bands of noise at center frequencies from 250 to 1500 Hz, for modulation rates of 2-40 Hz, and with or without flanking bands of diotic noise. Discrimination thresholds increased steeply for QFM with increasing center frequency, but increased only modestly for AM, and mainly for modulation rates below 10 Hz. Flanking bands of noise increased thresholds for AM, but had no consistent effect for QFM. The results suggest that two underlying mechanisms may support binaural unmasking: one most sensitive to interaural amplitude modulations that is susceptible to across-frequency interference, and a second, most sensitive to interaural phase modulations that is immune to such effects.     

Click lateralization is related to the beta component of the dichotic brainstem auditory evoked potentials of human subjects

The changes in perception and in the binaural difference waveform (BD) for dichotic clicks with interaural time and level differences (ITDs and ILDs) are compared. Only beta, the first major peak of the BD, correlated with the perceptual measurements. Whenever beta is clearly present, the clicks are perceived as a unitary fused image. Whenever the clicks are perceived as not fused, beta is undetectable by our methods. The amplitude of beta remains nearly constant as the ITD is increased to about 1 ms, while the click's position is perceived as moving from midline toward the leading ear. Over about the next 0.2 ms, beta becomes undetectable, as the image stops moving and loses its fused quality. As the ILD is increased, beta amplitude decreases gradually, while the image remains unitary and moves toward the unattenuated earphone. Thus beta becomes undetectable for ILDs of 30 to 35 dB, and the dichotic clicks become indistinguishable from monotic clicks for ILDs of 44 to 53 dB. The ITD and ILD matching curve for beta latency is similar to the ITD and ILD psychophysical matching curve for lateralization. These results suggest that beta is a physiological correlate of the categorical percept, binaural fusion, and is generated by a brainstem structure essential for the preception of click lateralization.     

Sperm whales (Physeter catodon L. 1758) do not react to sounds from detonators

A number of observations show that sperm whales (Physeter catodon L. 1758) react to various man-made pulses with moderate source levels. The behavioral responses are described to vary from silence to fear. Click rates of five submerged male sperm whales were measured during the discharge of eight detonators off Andenes, northern Norway. In addition, the behavioral response of a surfaced specimen was observed. Click rates of the submerged whales and the behavior of the surfaced specimen did not change during the discharges with received sound levels of some 180 dB re 1 microPa peRMS. The apparent lack of response to the discharges could be due to similarity between sperm whale clicks and detonations. Accordingly, it can be speculated that the discharges may have been perceived as isolated clicks from conspecifics.     

Auditory evoked potentials in two short-finned pilot whales (Globicephala macrorhynchus)

The hearing sensitivities of two short-finned pilot whales (Globicephala macrorhynchus) were investigated by measuring auditory evoked potentials generated in response to clicks and sinusoidal amplitude modulated (SAM) tones. The first whale tested, an adult female, was a long-time resident at SeaWorld San Diego with a known health history. Click-evoked responses in this animal were similar to those measured in other echolocating odontocetes. Auditory thresholds were comparable to dolphins of similar age determined with similar evoked potential methods. The region of best sensitivity was near 40 kHz and the upper limit of functional hearing was between 80 and 100 kHz. The second whale tested, a juvenile male, was recently stranded and deemed non-releasable. Click-evoked potentials were not detected in this animal and testing with SAM tones suggested severe hearing loss above 10 kHz.