Responses to amplitude-modulated tones in the auditory nerve of the cat.

Sinusoidally amplitude-modulated (AM) tones are frequently used in psychophysical and physiological studies, yet a comprehensive study on the coding of AM tones in the auditory nerve is lacking. AM responses of single auditory-nerve fibers of the cat are studied, systematically varying modulation depth, frequency, and sound level. Synchrony-level functions were nonmonotonic with maximum values that were inversely correlated with spontaneous rate (SR). In most fibers, envelope phase-locking showed a positive gain. Modulation transfer functions were uniformly low pass. Their corner frequency increased with characteristic frequency (CF), but changed little for CFs above 10 kHz. The highest modulation frequencies to which phase locking occurred were more than 0.8 oct lower than the highest frequencies to which phase locking to pure tones occurs. Cumulative, or unwrapped, phase increased linearly with modulation frequency: The slope was inversely related to CF, and slightly higher than group delays reported for pure tones. High SR, low CF fibers showed the poorest envelope phase locking. In some low CF fibers, phase locking increased at high levels, associated with "peak-splitting" phenomena. Changes in average rate due to modulation were small, and could be enhancement or suppression.     

Neuromagnetic steady-state responses to auditory stimuli

Steady-state magnetic responses to clicks presented at rates between 10 and 70 Hz have been recorded in healthy humans. The responses were highest in amplitude around 40 Hz. This amplitude enhancement is satisfactorily explained by summation of responses evoked by single clicks. The field maps suggest activation of the auditory cortex at all stimulus frequencies. Similar responses were obtained with gated noise bursts and by pauses in a series of clicks. The mean "apparent latency," determined from the phase lag at rates 30-70 Hz, was 54 ms. The physiological relevance of this quantity is shown to be questionable.     

Hybrid measurement of auditory steady-state responses and distortion product otoacoustic emissions using an amplitude-modulated primary tone

A maximum auditory steady-state response (ASSR) amplitude is yielded when the ASSR is elicited by an amplitude-modulated tone (f(c)) with a fixed modulation frequency (f(m) = 40 Hz), whereas the maximum distortion product otoacoustic emission (DPOAE) level is yielded when the DPOAE is elicited using a fixed frequency ratio of the primary tones (f2/f1 = 1.2). When eliciting the DPOAE and ASSR by the same tone pair, optimal stimulation is present for either DPOAE or ASSR and thus adequate simultaneous DPOAE/ASSR measurement is not possible across test frequency f2 or f(c), respectively. The purpose of the present study was to determine whether the ASSR and DPOAE can be measured simultaneously without notable restrictions using a DPOAE stimulus setting in which one primary tone is amplitude modulated. A DPOAE of frequency 2f1-f2 and ASSR of modulation frequency 41 Hz were measured in ten normal hearing subjects at a test frequency between 0.5 and 8 kHz (f2 = f(c)). The decrease in the DPOAE level and the loss in ASSR amplitude during hybrid mode stimulation amounted, on average, to only 2.60 dB [standard deviation (SD) = 1.38 dB] and 1.83 dB (SD = 2.38 dB), respectively. These findings suggest simultaneous DPOAE and ASSR measurements to be feasible across all test frequencies when using a DPOAE stimulus setting where the primary tone f2 is amplitude modulated.     

Across-critical-band processing of amplitude-modulated tones

Two experiments using two-tone sinusoidally amplitude-modulated stimuli were conducted to assess cross-channel effects in processing low-frequency amplitude modulation. In experiment I, listeners were asked to discriminate between two sets of two-tone amplitude-modulated complexes. In one set, the modulation phase of the lower frequency carrier tone was different from that of the upper frequency carrier tone. In the other stimulus set, both amplitude-modulated carriers had the same modulator phase. The amount of phase shift required to discriminate between the two stimulus sets was determined as a function of the separation between the two carriers, modulation depth, and modulation frequency. Listeners could discriminate a 50 degrees-60 degrees phase shift between the modulated envelopes for tones separated by more than a critical band. In experiment II, the modulation depth required to detect modulation of a probe carrier was measured in the presence of an amplitude-modulated masker. The threshold for detecting probe modulation was determined as a function of the separation between the masker and probe carriers, the phase difference between the masker and probe modulators, and masker modulation depth (in all conditions, the rate of probe and masker modulation was 10 Hz). The threshold for detecting probe modulation was raised substantially when the masker tone was also modulated. The results are consistent with theories suggesting that amplitude modulation helps form auditory objects from complex sound fields.     

Place specificity of multiple auditory steady-state responses

Auditory steady-state responses (ASSRs) were elicited by simultaneously presenting multiple AM (amplitude-modulated) tones with carrier frequencies of 500, 1000, 2000, and 4000 Hz and modulation frequencies of 77, 85, 93, and 102 Hz, respectively. Responses were also evoked by separately presenting single 500- or 2000-Hz AM tones. The objectives of this study were (i) to determine the cochlear place specificity of single and multiple ASSRs using high-pass noise masking and derived-band responses, and (ii) to determine if there were any differences between single- and multiple-stimulus conditions. For all carrier frequencies, derived-band ASSRs for 1-octave-wide derived bands ranging in center frequency from 0.25 to 8 kHz had maximum amplitudes within a 1/2 octave of the carrier frequency. For simultaneously presented AM tones of 500, 1000, 2000, and 4000 Hz, bandwidths for the function of derived-band ASSR amplitude by derived-band center frequency were 476, 737, 1177, and 3039 Hz, respectively. There were no significant differences when compared to bandwidths of 486 and 1371 for ASSRs to AM tones of 500 or 2000 Hz presented separately. Results indicate that ASSRs to moderately intense stimuli (60 dB SPL) reflect activation of reasonably narrow cochlear regions, regardless of presenting AM tones simultaneously or separately.     

Rate responses of auditory nerve fibers to tones in noise near masked threshold

The rate responses of auditory nerve fibers were measured for best frequency (BF) tone bursts in the presence of continuous background noise. Rate functions for BF tones were constructed over a 32-dB range of levels, centered on the behavioral masked thresholds of cats. The tone level at which noticeable rate changes are evoked by the tones corresponds closely to behavioral masked threshold at all noise levels used (-10- to 30-dB spectrum level). As the noise level increases, the response rate to the background noise approaches saturation, and the incremental rate response to tones decreases. At high noise levels, the rate responses to tones of low and medium spontaneous rate fibers are larger than those of high spontaneous rate fibers. Empirical statistics of auditory nerve fiber spike counts are reported; these differ from those expected of a Poisson process in that the variance is smaller than the mean. A new measure of discharge rate is described that allows rate changes to be expressed in units of a standard deviation. This measure allows tone-evoked responses to be interpreted in terms of their detectability in a signal detection task. Rate responses of low and medium spontaneous rate fibers are more detectable than those of high spontaneous rate fibers, especially at high noise levels. There appears to be sufficient information in the rate response of a small number of auditory nerve fibers to support behaviorally observed levels of detection performance.     

Bottlenose dolphin (Tursiops truncatus) steady-state evoked responses to multiple simultaneous sinusoidal amplitude modulated tones

Auditory steady-state evoked potentials were measured in a bottlenose dolphin (Tursiops truncatus) in response to single and multiple sinusoidal amplitude modulated (SAM) tones. Tests were conducted in air using a "jawphone" sound projector. Evoked potentials were recorded noninvasively using surface electrodes embedded in suction cups. Sound stimuli consisted of SAM tones with 1, 2, 3, or 4 carrier frequencies (10, 20, 30, 40 kHz), each with a unique modulation frequency. Stimulus sound pressure levels were varied in 5-dB steps from approximately 120 to 60-75 dB re 1 microPa, depending on frequency. Evoked potentials followed the temporal envelope of each stimulus, resulting in spectral components at each unique modulation frequency. Spectral analysis was used to evaluate the response amplitude for each carrier as a function of stimulus level. There were no significant differences between thresholds obtained with single and multiple stimuli at 10, 30, and 40 kHz. At 20 kHz, thresholds obtained with three components were higher than those obtained with four components, possibly revealing interactions between stimuli with less than one octave frequency separation. The use of multiple SAM stimuli may offer substantial advantages for studies of marine mammal hearing, where testing time and access to subjects are typically limited.     

Chinchilla auditory-nerve responses to low-frequency tones

Single unit activity was recorded in the auditory nerves of chinchillas. Period histograms were constructed for responses to tones with frequencies 30-1000 Hz. For low-frequency tones at near-threshold levels, peak period histogram phases for low- and medium-best-frequency (BF) neurons (less than or equal to kHz) ranged from synchronous with condensation at the eardrum to 90 degrees leading it. At near-threshold (but high absolute) levels, high-BF (greater than or equal to 8 kHz) neurons responded in phase with rarefaction. At even higher levels, period histograms for responses of high-BF neurons tended to become bimodal, with one of the modes lagging rarefaction by 90 degrees. Using cochlear microphonics as an indicator of basilar membrane (BM) displacement, at threshold levels, response phase of low- and medium-BF neurons fall within a range between displacement and velocity of the BM toward scala vestibuli. High-BF neurons respond, at threshold (but high) intensities, in phase with BM displacement toward scala tympani. The rates of growth of frequency sensitivity in responses of low-BF (+ 18 dB/oct) and high-BF (+ 12 dB/oct) neurons are consistent with preferred response phases corresponding to BM SV velocity and ST displacement, respectively. At supra-threshold levels high-BF neurons may fire preferentially to both scala tympani displacement and scala vestibuli velocity. These results support the notion that, for high-intensity, low-frequency stimuli, OHC hyperpolarization can induce excitation of the dendrites innervating IHCs.     

Classical conditioned responses to absent tones

Background  

Recent evidence for a tight coupling of sensorimotor processes in trained musicians led to the question of whether this coupling extends to preattentively mediated reflexes; particularly, whether a classically conditioned response in one of the domains (auditory) is generalized to another (tactile/motor) on the basis of a prior association in a second-order Pavlovian paradigm. An eyeblink conditioning procedure was performed in 17 pianists, serving as a model for overlearned audiomotor integration, and 14 non-musicians. Results: During the training session, subjects were conditioned to respond to auditory stimuli (piano tones). During a subsequent testing session, when subjects performed keystrokes on a silent piano, pianists showed significantly higher blink rates than non-musicians.     

Asymmetric lateral inhibitory neural activity in the auditory system: a magnetoencephalographic study

Background  

Decrements of auditory evoked responses elicited by repeatedly presented sounds with similar frequencies have been well investigated by means of electroencephalography and magnetoencephalography (MEG). However the possible inhibitory interactions between different neuronal populations remains poorly understood. In the present study, we investigated the effect of proceeding notch-filtered noises (NFNs) with different frequency spectra on a following test tone using MEG.     

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.     

Lateralization of sinusoidally amplitude-modulated tones: effects of spectral locus and temporal variation

It has long been recognized that listeners are sensitive to interaural temporal disparities (ITDs) of low-frequency (i.e., below 1600 Hz) stimuli. Within the last three decades, it has often been demonstrated that listeners are also sensitive to ITDs within the envelope of high-frequency, complex stimuli. Because these studies, for the most part, employed discrimination tasks, few data exist concerning the extent of laterality produced by ITDs as a function of the spectral locus of the stimulus. To this end, we employed an acoustic "pointing" task in which listeners varied the interaural intensity difference of a 500-Hz narrow-band noise (the pointer) so that it matched the intracranial position of a second, experimenter-controlled stimulus (the target). Targets were sinusoidally amplitude-modulated tones centered on 500 Hz, 1, 2, 3, or 4 kHz and modulated at rates ranging from 50 to 800 Hz. Targets were presented with either the entire waveform delayed or with only the envelope delayed. Our results suggest that: (1) for low-frequency targets, lateralization is influenced by ITDs in the envelope but is dominated by ITDs in the fine structure; (2) for high-frequency targets, envelope-based delays produce displacements of the acoustic images which are affected greatly by the rate of modulation; rather large extents of laterality could be produced with high rates of modulation; these data are consistent with those obtained previously in discrimination experiments; (3) for low rates of modulation (e.g., 100 Hz), delays of the entire waveform (both envelope and fine structure) produce much greater displacements of the acoustic image for low-frequency than for high-frequency targets (where fine-structure-based cues are not utilizable); (4) there appear to be no consistent relations among extent of laterality, rate of modulation, and the frequency of the carrier within and across listeners.     

Evoked magnetoencephalographic responses to omission of a tone in a musical scale

The musical scale is a basis for melodies and can be a simple melody by itself. The present study investigated magnetoencephalographic (MEG) responses to omissions of one tone out of the C major scale. The tone preceding the omitted "target" tone was either prolonged or repeated. In another series, the tone after the target tone was repeated. In "normal" oddball experiments, the complete C major scale was presented more frequently than an incomplete scale lacking one tone, and in "reverse" oddball experiments, the roles were exchanged. In the normal oddball experiments, omission of any tone produced a response significantly different in amplitude from the standard response in the group of non-musicians, although the responses differed depending on the types of omission. The leading tone (B in the C major scale) was shown to elicit a large response when omitted and also when its presence was emphasized. The Reverse oddball experiments showed that repeated presentation of an incomplete scale lacking one tone temporarily reduced the influence of the complete scale but could not even temporarily replace it working as "standard." In addition, an auxiliary study was done to see possible influence of rhythmic variations.     

Hearing threshold estimation using concurrent measurement of distortion product otoacoustic emissions and auditory steady-state responses

Both distortion product otoacoustic emissions (DPOAEs) and auditory steady-state responses (ASSRs) provide frequency-specific assessment of hearing. However, each method suffers from some restrictions. Hearing losses above 50 dB HL are not quantifiable using DPOAEs and their performance at frequencies below 1 kHz is limited, but their recording time is short. In contrast, ASSRs are a time-consuming method but have the ability to determine hearing thresholds in a wider range of frequencies and hearing losses. Thus, recording DPOAEs and ASSRs simultaneously at their adequate frequencies and levels could decrease the overall test time considerably. The goal of the present study was to develop a parameter-setting and test-protocol to measure DPOAEs and ASSRs binaurally and simultaneously at multiple frequencies. Ten normal-hearing and 23 hearing-impaired subjects participated in the study. The interaction of both responses when stimulated simultaneously at frequencies between 0.25 and 6 kHz was examined. Two limiting factors need to be kept. Frequency distance between ASSR carrier frequency f(c) and DPOAE primary tone f(2) needs to be at least 1.5 octaves, and DPOAEs may not be measured if the ASSR stimulus level is 70 dB SPL or above. There was a significant correlation between pure-tone and DPOAE/ASSR-thresholds in sensorineural hearing loss ears.     

Lateralization of bands of noise and sinusoidally amplitude-modulated tones: effects of spectral locus and bandwidth

Lateralization of narrow bands of noise was investigated while varying interaural temporal disparity (ITD), center frequency, and bandwidth, utilizing an acoustic pointing task. Stimuli were narrow bands of noise centered at octave intervals between 500 Hz and 4 kHz with bandwidths ranging from 50-400 Hz. In a second experiment, lateralization for bands of noise and sinusoidally amplitude-modulated (SAM) tones, whose spectral content was constrained to be no lower than 3.8 kHz, was assessed. Overall, relatively large extents of laterality were obtained from all four listeners for ITDs of low-frequency bands of noise. Increasing the bandwidth of these noises did not yield consistent changes in the extent of laterality across ITDs and listeners. Most targets centered at high frequencies were lateralized near the midline. However, three of the four listeners did exhibit rather large displacements of the intracranial image when the bandwidth of the high-frequency noises was 400 Hz or greater. Interestingly, ITDs within high-frequency SAM tones were relatively ineffective. Thus, it appears that ITDs of relatively wide-band, high-frequency stimuli can mediate rather substantial extents of laterality. However, these effects are highly listener-dependent.     

Neural responses to auditory temporal patterns

Sets of regularly repeating auditory stimuli elicit unique perceptions; listeners are able to identify specific temporal patterns. Some temporal patterns are unambiguous (only one pattern can be perceived), while others are ambiguous (numerous patterns can be detected). While the psychophysical properties of such percepts have been well studied, little is known about the underlying neurological bases of temporal pattern perception. In this experiment, the role of adaptation in temporal pattern perception is examined by studying neural responses in four cats to a temporal pattern that is perceptually unambiguous and one that is perceptually ambiguous. Measurements were made of the whole-nerve action potential, the auditory brainstem response, and potentials from the surface of the primary auditory cortex. The adaptation patterns corresponded with the perceptual organization of temporal patterns in humans at all levels of the nervous system studied.     

Listeners' sensitivity to "onset/offset" and "ongoing" interaural delays in high-frequency, sinusoidally amplitude-modulated tones

The relative potency of onset/offset and envelope-based ongoing interaural time delays (ITDs) was assessed using high-frequency stimuli. A two-cue, two-alternative, forced-choice adaptive task was employed to measure threshold ITDs with 100% sinusoidally amplitude-modulated tones centered at 4 kHz. Modulation rates of 125, 250, and 350 Hz were tested with durations of 32, 90, or 240 ms. In the first experiment, ITDs to be detected were imposed only at the onset/offset, only within the ongoing portion, or within both the onset/offset and ongoing portions of the stimuli. Results indicated that ongoing ITDs dominated onset/offset ITDs. The relative potency of ongoing ITDs was directly proportional to duration and inversely proportional to modulation rate. Quantitative analysis suggested that listeners effectively combine onset/offset and ongoing ITDs. Furthermore, the data could be largely accounted for by assuming that listeners attend to the interaural decorrelation of the stimulus resulting from onset/offset and/or ongoing ITDs. A second experiment showed that, (1) overall, an ongoing ITD of one-half period of the envelope had little impact on listeners' sensitivity to delays of the onset/offset and (2) sensitivity to delays within the onset/offset portion of the waveform was reduced by roving the delay within the ongoing portion of the waveform.     

Hemodynamic responses in human multisensory and auditory association cortex to purely visual stimulation

Background  

Recent findings of a tight coupling between visual and auditory association cortices during multisensory perception in monkeys and humans raise the question whether consistent paired presentation of simple visual and auditory stimuli prompts conditioned responses in unimodal auditory regions or multimodal association cortex once visual stimuli are presented in isolation in a post-conditioning run. To address this issue fifteen healthy participants partook in a "silent" sparse temporal event-related fMRI study. In the first (visual control) habituation phase they were presented with briefly red flashing visual stimuli. In the second (auditory control) habituation phase they heard brief telephone ringing. In the third (conditioning) phase we coincidently presented the visual stimulus (CS) paired with the auditory stimulus (UCS). In the fourth phase participants either viewed flashes paired with the auditory stimulus (maintenance, CS-) or viewed the visual stimulus in isolation (extinction, CS+) according to a 5:10 partial reinforcement schedule. The participants had no other task than attending to the stimuli and indicating the end of each trial by pressing a button.     

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.     

A population study of auditory-nerve fibers in unanesthetized decerebrate cats: response to pure tones

Population responses of single auditory-nerve (AN) fibers to 1-kHz pure tones were investigated in unanesthetized decerebrate cats. Driven rate (spikes/s), Fourier component amplitude (spikes/s), and d'e = (driven rate)/(standard deviation) were examined. These results extend the presently available information. First, the AN population responses, previously observed only in anesthetized animals, are now observed in unanesthetized animals. The anesthetized and unanesthetized animal responses are qualitatively similar. Second, a detectability measure d'e was applied, à la signal detection theory, to populations of AN fibers. A previously unknown behavior of the low-spontaneous-rate (SR) (less than 15 spikes/s) fibers was observed: the d'e increased to large values at moderate and high stimulus levels, giving rise to a large, sharply tuned peak in the d'e-place profile, even when the driven rate was saturated. This behavior was absent in the high-SR (greater than 15 spikes/s) fibers. This observation suggests that the d'e of the low-SR fibers is the best code for stimulus intensity among the different response measures examined, and that the d'e-place profile may be a more precise spatial code for stimulus frequency than the commonly considered rate-place code.