Detection of silent intervals between noises activating different perceptual channels: some properties of "central" auditory gap detection

This article describes four experiments on gap detection by normal listeners, with the general goal being to examine the consequences of using noises in different perceptual channels to delimit a silent temporal gap to be detected. In experiment 1, subjects were presented with pairs of narrow-band noise sequences. The leading element in each pair had a center frequency of 2 kHz and the trailing element's center frequency was parametrically varied. Gap detection thresholds became increasingly poor, sometimes by up to an order of magnitude, as the spectral disparity was increased between the noise bursts that marked the gap. These data suggested that gap-detection performance is impoverished when the underlying perceptual timing operation requires a comparison of activity in different perceptual channels rather than a discontinuity detection within a given channel. In experiment 2, we assessed the effect of leading-element duration in within-channel and between-channel gap detection tasks. Gap detection thresholds rose when the duration of the leading element was less than about 30 ms, but only in the between-channel case. In experiment 3, the gap-detection stimulus was redesigned so that we could probe the perceptual mechanisms that might be involved in stop consonant discrimination. The leading element was a wideband noise burst, and the trailing element was a 300-ms bandpassed noise centered on 1.0 kHz. The independent variable was the duration of the leading element, and the dependent variable was the smallest detectable gap between the elements. When the leading element was short in duration (5-10 ms), gap thresholds were close to 30 ms, which is close to the voice onset time that parses some voiced from unvoiced stop consonants. In experiment 4, the generality of the leading-element duration effect in between-channel gap detection was examined. Spectrally identical noises defining the leading and trailing edges of the gap were presented to the same or to different ears. There was a leading-element duration effect only for the between channel case. The mean gap threshold was again close to 30 ms for short leading-element durations. Taken together, the data suggest that gap detection requiring a temporal correlation of activity in different perceptual channels is a fundamentally different task to the discontinuity detection used to execute gap detection performance in the traditional, within-channel paradigm.     

Diotic and dichotic detection using multiplied-noise maskers

Detection thresholds were measured with a multiplied-noise masker that was in phase in both ears and a sinusoidal signal which was either in phase or out of phase (NoSo and NoS pi conditions). The masker was generated by multiplying a low-pass noise with a sinusoidal carrier. The signal was a sinusoid with the same frequency as the carrier and a constant phase offset, theta, with respect to the carrier. By adjusting the phase offset, the stimulus properties were varied in such a way that only interaural time delays (theta = pi/2) or interaural intensity differences (theta = 0) were present within the NoS pi stimulus. Thresholds were measured at a center frequency of 4 kHz as a function of bandwidth for theta = pi/2 and for theta = 0. In a second experiment thresholds were measured for a bandwidth of 25 Hz as a function of the center frequency. The results show that narrow-band BMLDs at 4 kHz can amount to 30 dB for the theta = 0 condition. For this condition, narrow-band BMLDs are also reasonably constant across frequency, in contrast to results obtained with standard Gaussian-noise maskers. For theta = pi/2, BMLDs are restricted to the frequency region below 2 kHz provided that the masker is narrow band, but BMLDs of up to 15 dB are found at 4 kHz if the masker is 50 Hz or wider. The frequency dependence of the binaural thresholds seems to be best explained by assuming that the stimulus waveforms are compressed before binaural interaction.     

Hepatic sinusoidal responses to intraportal injections of phenylephrine and isoprenaline in the rat

1. Specific alpha- and beta-adrenoreceptor agonists, phenylephrine and isoprenaline, were injected intraportally into the intact rat liver under direct microscopic observation by an in vivo transillumination technique. 2. The diameter of a hepatic sinusoid and the intra-sinusoidal erythrocyte velocity were quantitatively measured, and the sinusoidal volume flow was calculated from these two parameters. 3. Results show that phenylephrine causes a sinusoidal constriction and an increased sinusoidal blood flow, whereas isoprenaline causes the opposite effects on the sinusoids. 4. All the sinusoidal responses to phenylephrine and isoprenaline were dose-dependent and were possibly related to the direct effect of these drugs on the sinusoids.     

Behavioral auditory thresholds for sinusoidal electrical stimuli in the cat

Behavioral thresholds were measured in four cats by training them to respond behaviorally to acoustic auditory stimuli using food as a reward in an operant reinforcement paradigm. Following training, the subjects were implanted unilaterally with either a scaled-UCSF electrode containing four contacts or an electrode array containing eight intracochlear contacts and one extracochlear contact under temporalis muscle. Behavioral thresholds for electrical stimuli were obtained as a function of sinusoidal frequency, duration, and electrode configuration. Threshold functions for both electrode types and all animals had minima between 48 and 125 Hz and, in general, were relatively flat below this minima; functions increased at 3-6 dB/octave from 96 Hz to 1 kHz. Threshold varied predictably as a function of electrode configuration, with thresholds decreasing as much as 20 dB as electrode spacing was widened from a radial bipolar (200-microns separation) to a monopolar configuration (apical stimulating and temporalis return). With long-duration stimuli, increasing the electrode separation systematically increased the slope of the threshold-versus-frequency contours in all animals. Irrespective of electrode type or configuration, charge/phase thresholds for single-cycle sinusoids were relatively flat for stimulus periods up to 1-5 ms, approximating a constant charge/phase determination of threshold. At phase durations greater than 5 ms, charge thresholds increased at a rate slightly above 6 dB/octave (constant peak current), which was suggestive of neural accommodation. Thresholds for the cat share many features with those reported for implanted humans and monkeys.     

Effects of modulator phase for comodulation masking release and modulation detection interference

In an effort to evaluate the importance of across-frequency comparisons of envelope patterns in comodulation masking release (CMR) experiments and to compare joint effects of target-masker frequency separation for both CMR and modulation detection interference (MDI) tasks, thresholds were measured for three tasks. These tasks were: (a) the detection of sinusoidal amplitude modulation (SAM) of a tone, (b) the detection of a reduction in the modulation depth of a fully modulated SAM tone, and (c) the detection of a tone added to a narrow band of noise. Thresholds were obtained for the target alone and for the target presented with two maskers. For the detection of SAM, thresholds did not depend on whether the modulation patterns of the target and masker elements were the same or random. For the latter two tasks, modulator phase effects were apparent for target-masker frequency separations less than 1-2 oct. In contrast, past work has shown that observers can compare modulator envelope phases across frequency separations larger than 1-2 oct [Strickland et al., J. Acoust. Soc. Am. 86, 2160-2166 (1989); Yost and Sheft, J. Acoust. Soc. Am. 85, 848-857 (1989)]. In a second experiment, thresholds for the detection of SAM were obtained after prolonged exposure to a fully modulated SAM tone. For four of the five observers, modulation-rate specific adaptation was obtained for test/adapting carrier-frequency separations approaching 2 oct below and 1 oct above the adaptor.     

COAE thresholds: 1. Effects of equal-amplitude versus subtraction methods

Although research has demonstrated that click-evoked otoacoustic emissions (COAEs) elicited by high-level stimuli are useful for identifying hearing loss, the ability of COAEs to predict behavioral thresholds has not been adequately tested. Results of studies comparing COAE thresholds and behavioral thresholds have been equivocal, perhaps due to the need for a more rigorous approach to COAE threshold estimation. The present study was designed to address several methodological concerns in COAE threshold testing, particularly the effects of two methods of stimulus presentation on COAE testing and threshold calculation. In an attempt to make COAE threshold estimation consistent across participants, COAE threshold calculations were based on mean noise floor levels across participants. COAE and noise floor levels were measured in 15 participants using both equal-amplitude clicks and a subtraction method. Broadband COAEs were analyzed into 1/3 octave bands, so that input/output functions could be examined and COAE thresholds could be calculated for each 1/3 octave band. Comparison of the two stimulus methods indicated several differences. Mean noise floor levels for the equal-amplitude method were approximately 6 dB lower than those measured for the subtraction method across frequency. In many cases COAEs evoked using the equal-amplitude method were higher in amplitude than those evoked using the subtraction method. COAE thresholds measured using the equal-amplitude click stimuli were significantly lower than those measured using the subtraction method. The significantly higher thresholds obtained using the subtraction method may be attributed in part to the reduction of COAE amplitude by the subtraction procedure, and not merely to the higher noise level. Slopes of the input/output functions were not significantly different between the two stimulus methods. These results suggest that the equal-amplitude method is preferable for COAE threshold testing because lower noise floor and larger amplitude COAEs may be obtained in the same test time.     

Gap encoding by inferior collicular neurons is altered by minimal changes in signal envelope

Neural correlates of temporal resolution in the central auditory system are currently under intense investigation. The gap detection paradigm offers a simple, yet important, test of temporal acuity because changes in behavioral gap thresholds have been correlated with deficits in complex stimulus processing, such as speech perception. In gap detection studies, silent gaps are typically shaped by rapid (< 1.0 ms) rise/fall (R/F) times, i.e., rapid decreases and increases in sound intensity. However, in nature, the envelopes surrounding silent periods can vary significantly in R/F time. Therefore, we investigated whether changes in the R/F time surrounding the silent gap affect neural processing by inferior collicular (IC) neurons. Gap R/F times were varied between 0.5 and 16 ms and the discharge pattern, response rate, and first spike latency of IC neurons were measured for gap widths up to 100 ms. Neurons were classified into phasic or tonic discharge patterns based on peri-stimulus time histograms elicited to 100 ms noise carriers. The results indicate that (1) minimal gap thresholds increased with R/F time regardless of response type, (2) first spike latency variance increased systematically with R/F time for units which had small first spike standard deviations at short R/F times, and (3) the response rate of some units (called 'gap-tuned') changed as a function of both R/F time and gap width. Gap-tuned units responded strongly to a particular gap width only when the envelope of the gap was shaped by a particular R/F time. For gap-tuned units, increases in R/F time shifted the tuning to larger gap widths and also broadened the response profile. These results show that temporal acuity of neurons in the IC, as measured by the gap detection paradigm, is sensitive to the envelope surrounding gaps embedded in noise carriers.     

Auditory masking-level differences in the cat.

Trained 4 cats to avoid shock by responding to the intermittent occurrence of 1-kHz tone pulses at one ear, while a continual train of noise pulses was simultaneously presented either to the signal ear alone or to both ears. Using the masked threshold levels determined with monaural noise as a reference, the amount of unmasking produced by the addition of noise to the nonsignal ear was measured. Significantly lower tonal detection thresholds were observed when noise equal in intensity to that at the signal ear was added to the nonsignal ear. Additional unmasking occurred when the intensity of the noise at the latter ear was raised to a level 10 db higher than that at the signal ear. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Effects of pulse separation on detection thresholds for electrical stimulation of the human cochlea

Effects of pulse separation on detection of electrical stimulation of the cochlea were studied in 12 profoundly deaf human subjects with Nucleus 22 cochlear implants. Biphasic symmetric pulses were used. Pulse separation is the time from offset of one biphasic pulse to the onset of the next biphasic pulse in the train. Effects of pulse separation were studied in the context of different covariables in four stages of the experiment. Effects of pulse separation seen in the different stages were similar, despite the different covariables. Both pulse separation and the total number of pulses per stimulus seem to be important variables affecting stimulus detection. For 0.5 ms/phase pulses, thresholds were lowest at the shortest pulse separations tested (0.2-1.1 ms) and increased as a function of pulse separation. For 2 ms/phase pulses, detection thresholds were lowest at pulse separations around 7.5 ms, in most cases, and higher at both longer and shorter pulse separations. These results suggest that interactions among adjacent pulses can either hinder or facilitate detection of the signal depending on the magnitudes of pulse separation and phase duration. Pulse separations at which thresholds measured for 2 ms/phase pulses were minimum were fairly consistent across subjects and did not correlate well with speech recognition scores. However, significant variation in this measure across species has been seen.     

Auditory spatial responses of young guinea pigs (Cavia porcellus) during and after ear blocking.

Tested 30 newborn guinea pigs to determine their ability to approach an auditory stimulus early in development. Observations of the behavior of 1–4 day old Ss in a circular 8-choice maze revealed a pronounced tendency to orient toward and approach a tape-recorded signal of guinea pig vocalizations. The occurrence of approach responses was reduced to chance in Ss tested with one ear occluded by wax ear plugs which attenuated but did not totally eliminate sound. The effect of monaural ear blocks was more severe than binaural blocks, which reflects the importance of binaural cues in the maintenance of approach responses to sound. In a 2nd study with 40 Ss the ability of older animals, 11–31 days of age, was examined. Directional approach responses to sound were also evident at this age, and ear plugs disrupted performance only under monaural conditions. Furthermore, in Ss raised from birth with monaural ear blocks but tested without ear plugs, there was a subsequent disruption of performance for at least 21 days. Results indicate the importance of binaural cues in the development of early auditory spatial reponses and suggest the need for appropriate binaural experience for subsequent localization of sounds. (52 ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

The effects of attenuation of frequency segments on binaural localization of sound

Perceived location of tonal stimuli d and narrow noise bands presented in two-dimensional space varies in an orderly manner with changes in stimulus frequency. Hence, frequency has a referent in space that is most apparent during monaural listening. The assumption underlying the present study is that maximum sound pressure level measured at the ear canal entrance for the various frequencies serves as a prominent spectral cue for their spatial referents. Even in binaural localization, location judgments in the vertical plane are strongly influenced by spatial referents. We measured sound pressure levels at the left ear canal entrance for 1.0-kHz-wide noise bands, centered from 4.0 kHz through 10.0 kHz, presented at locations from 60 degrees through -45 degrees in the vertical plane; the horizontal plane coordinate was fixed at -90 degrees. On the basis of these measurements, we fabricated three different bandstop stimuli in which differently centered 2.0-kHz-wide frequency segments were filtered from a broadband noise. Unfiltered broadband noise served as the remaining stimulus. Localization accuracy differed significantly among stimulus conditions (p < .01). Where in the vertical plane most errors were made depended on which frequency segment was filtered from the broadband noise.     

Age-related alteration in processing of temporal sound features in the auditory midbrain of the CBA mouse

The perception of complex sounds, such as speech and animal vocalizations, requires the central auditory system to analyze rapid, ongoing fluctuations in sound frequency and intensity. A decline in temporal acuity has been identified as one component of age-related hearing loss. The detection of short, silent gaps is thought to reflect an important fundamental dimension of temporal resolution. In this study we compared the neural response elicited by silent gaps imbedded in noise of single neurons in the inferior colliculus (IC) of young and old CBA mice. IC neurons were classified by their temporal discharge patterns. Phasic units, which accounted for the majority of response types encountered, tended to have the shortest minimal gap thresholds (MGTs), regardless of age. We report three age-related changes in neural processing of silent gaps. First, although the shortest MGTs (1-2 msec) were observed in phasic units from both young and old animals, the number of neurons exhibiting the shortest MGTs was much lower in old mice, regardless of the presentation level. Second, in the majority of phasic units, recovery of response to the stimulus after the silent gap was of a lower magnitude and much slower in units from old mice. Finally, the neuronal map representing response latency versus best frequency was found to be altered in the old IC. These results demonstrate a central auditory system correlate for age-related decline in temporal processing at the level of the auditory midbrain.     

Temporal integration at 6 kHz as a function of masker bandwidth

Thresholds were measured for a 6-kHz sinusoidal signal presented within a 500-ms masker. The masker was either a bandpass Gaussian noise of varying bandwidth, or a sinusoid of the same frequency as the signal. The spectrum level of the noise masker was kept constant at 20 dB SPL, and the level of the sinusoidal masker was 40 dB SPL. Thresholds for signal durations between 2 and 300 ms were measured for masker bandwidths ranging from 60 to 12,000 Hz. The masker was spectrally centered around 6 kHz. For masker bandwidths less than 600 Hz, the slope of the temporal integration function decreased with decreasing masker bandwidth. The results are not consistent with current models of temporal integration or temporal resolution. It is suggested that the results at narrow bandwidths can be understood in terms of changes in the power spectrum of the stimulus envelope or modulation spectrum. According to this view, the onset and offset ramps of the signal introduce detectable high-frequency components into the modulation spectrum, which provide a salient cue in narrowband maskers. For broadband maskers, these high-frequency components are masked by the inherent rapid fluctuations in the masker envelope. Additionally, for signal durations between 7 and 80 ms, signal thresholds decreased by up to 5 dB as the masker bandwidth increased from 1200 to 12,000 Hz. The mechanisms underlying this effect are not yet fully understood.     

Modulation detection interference in cochlear implant subjects

The aim of this study was to determine whether detection thresholds for amplitude modulated signals on a single electrode were influenced by a masking modulation on a second electrode in cochlear implant users. Data were collected from four post-linguistically deafened subjects using the Cochlear Limited prosthesis. Investigated were the effects of the spatial separation between test and masker electrodes, 0 to 5 electrodes (0 to 3.75 mm), and the amount of masking modulation: 24%, 48%, 72%, and 96% above detection thresholds. Initially, modulation detection thresholds for stimulation on a single electrode without masking modulation were obtained for a set of six electrodes in the middle of the array. Modulation detection thresholds on a fixed test electrode were then obtained with unmodulated and modulated masking on a second electrode, which was one of the six electrodes in the initial study. In both studies, thresholds were measured for modulated pulse duration at the modulation frequencies of 10-200 Hz. In the first study, the shape of the detection thresholds as a function of modulation frequency, the temporal modulation transfer function, generally resembled a low-pass filter for two subjects. For the other two subjects, the functions were relatively flat across modulation frequencies. In the second study, unmodulated masking resulted in a small elevation in detection thresholds across electrodes. Modulation detection interference (MDI), the difference between thresholds for the modulated maskers and the unmodulated masker, was greater for larger amounts of masking modulation than for smaller amounts of masking modulation. For three of the four subjects, MDI was higher for smaller spatial separations between the two electrodes than for larger spatial separations suggesting that a portion of MDI may be due to overlap of neural excitation distributions produced by stimulation on two electrodes in close proximity on the array.     

Spike train encoding by regular-spiking cells of the visual cortex

1. To study the encoding of input currents into output spike trains by regular-spiking cells, we recorded intracellularly from slices of the guinea pig visual cortex while injecting step, sinusoidal, and broadband noise currents. 2. When measured with sinusoidal currents, the frequency tuning of the spike responses was markedly band-pass. The preferred frequency was between 8 and 30 Hz, and grew with stimulus amplitude and mean intensity. 3. Stimulation with broadband noise currents dramatically enhanced the gain of the spike responses at low and high frequencies, yielding an essentially flat frequency tuning between 0.1 and 130 Hz. 4. The averaged spike responses to sinusoidal currents exhibited two nonlinearities: rectification and spike synchronization. By contrast, no nonlinearity was evident in the averaged responses to broadband noise stimuli. 5. These properties of the spike responses were not present in the membrane potential responses. The latter were roughly linear, and their frequency tuning was low-pass and well fit by a single-compartment passive model of the cell membrane composed of a resistance and a capacitance in parallel (RC circuit). 6. To account for the spike responses, we used a "sandwich model" consisting of a low-pass linear filter (the RC circuit), a rectification nonlinearity, and a high-pass linear filter. The model is described by six parameters and predicts analog firing rates rather than discrete spikes. It provided satisfactory fits to the firing rate responses to steps, sinusoids, and broadband noise currents. 7. The properties of spike encoding are consistent with temporal nonlinearities of the visual responses in V1, such as the dependence of response frequency tuning and latency on stimulus contrast and bandwidth. We speculate that one of the roles of the high-frequency membrane potential fluctuations observed in vivo could be to amplify and linearize the responses to lower, stimulus-related frequencies.     

Acoustic distortion as a measure of frequency selectivity: relation to psychophysical equivalent rectangular bandwidth

The magnitude of cubic intermodulation distortion generated when two tones are progressively separated in frequency reaches a broad maximum when the distortion frequency falls just over half an octave below the high-frequency stimulus (f2), when this distortion is measured with a microphone in the ear canal. For the component 2f1-f2, this peak occurs at an f2/f1 ratio of approximately 1.2. The tuning, magnitude, and mean group delay of this distortion peak was measured for a fixed f2 of 4 kHz at 40 dB SPL and a varied f1 at 55 dB SPL in eight human subjects with normal hearing. The distortion peak measures were compared with the frequency selectivity at 4 kHz of the same eight subjects derived using a forward-masking notched-noise paradigm. In the six subjects from whom good, repeatable levels of distortion were measured, a significant negative correlation was found between the tuning of the distortion peak and the psychophysical bandwidth at f2. It is concluded that the tuning of the distortion peak may provide an objective measure of frequency selectivity in the human cochlea.     

Free-field binaural unmasking in ferrets.

Free-field detection by normal and monaural ferrets (N?=?4) of a 500-Hz tone presented over 1 laterally placed loudspeaker and partially masked by narrowband noise from 2 sources was studied at 2 angular separations of the noise sources (0° and 180°). Monaural listening was achieved either by plugging 1 ear canal or removing 1 cochlea. Normal ferrets showed an improvement in detectability of the tone when there was a 180° separation between the noise sources. This unmasking of the tone was abolished in both groups of monaural ferrets, suggesting that the unmasking was due to binaural processing. The development of an animal model demonstrating free-field binaural unmasking, in a species other than humans, will allow investigation into the functional consequences of experimental hearing loss. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Middle latency responses to electrical stimulation of the auditory nerve in unanaesthetized guinea pigs

Middle latency responses (MLR) to sinusoidal and pulsatile electrical stimulation (ES) of the cochlea and to acoustical stimulation (AS) were evaluated in awake guinea pigs with chronically implanted electrodes. The ear, which was later electrically stimulated, was deafened by local intracochlear application of gentamicin, the opposite ear was left intact. Waveforms and P1-P2 interpeak intervals of the electrically evoked MLR (ES-MLR) were similar to those evoked by acoustical stimulation of the intact ear (AS-MLR) and the latencies of the ES-MLR were shorter by about 1-3 ms. Thresholds of ES-MLR in the frequency range 0.5-32 kHz increased with increasing ES frequency (slope 3.2 dB/octave), thresholds were 3.5-9.5 dB lower for intracochlear than for extracochlear ES. Dynamic ranges for ES-MLR varied between 6-20 dB. MLR amplitude-intensity functions for ES were steeper (slope 2-12 microV/dB) than those for AS (slope 0.2-2 microV/dB). Maximal ES-MLR amplitudes exceeded usually 1.5-3 times the amplitudes of the acoustically evoked MLR. Both types of stimulations evoked larger MLR amplitudes to contralateral stimulation than to ipsilateral stimulation (average ratio = 4.1 +/- 2.2 for AS and 3.3 +/- 2.2 for ES). Because of the relatively long latency and therefore insensitivity to electrical artifact, the ES-MLR can be used for the evaluation of different strategies of the electrical stimulation of the cochlea in awake guinea pig.     

Discrimination of interaural envelope delays: the effect of randomizing component starting phase

The goal of this study was to examine the nature of envelope extraction in the discrimination of high-frequency waveforms on the basis of envelope delay. Threshold interaural envelope delays were measured for complexes consisting of three or five components for which the starting phases of all sinusoids were either sine phase or randomized between intervals of a two-alternative forced-choice (2-AFC) task. The center frequency was 4 kHz and the frequency separation was varied from 25 to 500 Hz. The results showed that thresholds were greater for the phase-randomized conditions than the sine-phase conditions. The phase effect tended to diminish with increasing frequency separation for three-component complexes but not for the five-component complexes. Sensitivity to envelope delay was better for five-component complexes than for three-component complexes at most frequency separations. In general, the results showed superior lateralization performance for conditions in which the envelope fluctuations were greater, a finding that is consistent with models of high-frequency binaural processing that include envelope extraction prior to binaural comparison.     

Neurophysiological correlate of the auditory after-image ('Zwicker tone')

The 'Zwicker tone' (ZT) is an auditory after-image that can be evoked most effectively when a band-suppressed noise (relative width of gap 1/3 octave) presented for a certain period of time has been switched off. The sensation of this purely monaural phenomenon is that of a pure tone with a frequency corresponding to the center frequency of the gap and an equivalent level of 10-15 dB above auditory threshold. The sensation decays gradually; it may last as long as 10 s depending on how long the evoking noise was presented. The search for a physiological correlate has been futile so far, probably because the search was confined to more peripheral levels of the auditory system (inferior colliculus). A neuromagnetic study was performed in normal-hearing subjects in order to look for a neurophysiological correlate of the ZT in the auditory cortex. With a stimulation paradigm especially designed for this study, we have been able to isolate poststimulus activity which appears to be related to the ZT and which originates in the supratemporal auditory cortex. It is a sustained neuromagnetic activity that shows a clear-cut dipolar field distribution, and it appears that this activity has certain similarities with the tone-evoked auditory sustained response. The hypothesis is put forward that during the sensation of the ZT a process takes place in the auditory cortex which is similar to that underlying the sustained response, and which gives rise to the sensation of the ZT. In contrast to the sustained response, however, which is due to neural activity evoked by an external acoustic stimulus, the sustained activity associated with the ZT is due to a temporary absolute or relative reduction of neural activity originating from those regions in which the ZT exciting stimulus caused an adaptation. These differences in neural activity cannot be distinguished by the auditory system from a corresponding external acoustic signal. Preliminary studies in patients suffering from tonal tinnitus yielded results which exhibit a certain similarity with those obtained in the ZT experiment.