Coding of spectral fine structure in the auditory nerve. I. Fourier analysis of period and interspike interval histograms

The temporal fine structure of discharge patterns of single auditory-nerve fibers in adult cats was analyzed in response to signals consisting of a variable number of equal-intensity, in-phase harmonics of a common low-frequency fundamental. Two analytic methods were employed. The first method considered Fourier spectra of period histograms based on the period of the fundamental, and the second method considered Fourier spectra of interspike interval histograms (ISIH's). Both analyses provide information about fiber tuning properties, but Fourier spectra of ISIH's also allow estimates to be made of the degree of resolution of individual stimulus components. At low intensities (within 20-40 dB of threshold), indices of synchronization to individual components of complex tones were similar to those obtained for pure tones. This was true even when fibers were capable of responding to several signal components simultaneously. Response spectra obtained at low intensities resembled fibers' tuning curves, and fibers with low spontaneous discharge rates tended to provide better resolution of stimulus components than fibers with high spontaneous rates. Strongly nonlinear behavior existed at higher stimulus intensities. In this, information was transmitted about progressively fewer signal components and about frequencies not present in the acoustic stimulus, and the component eliciting the largest response shifted away from the fiber's characteristic frequency and toward the edges of the stimulus spectrum. This high-intensity "edge enhancement" can result from the combined effects of a compressive input-output nonlinearity, suppression, and the fortuitous addition of internally generated combination tones. The data indicate that sufficient information exists for the auditory system to determine the frequencies of narrowly spaced stimulus components from the temporal fine structure of nerve fiber's responses.     

Coding of spectral fine structure in the auditory nerve. II: Level-dependent nonlinear responses

Phase-locked discharge patterns of single cat auditory-nerve fibers were analyzed in response to complex tones centered at fiber characteristic frequency (CF). Signals were octave-bandwidth harmonic complexes defined by a center frequency F and an intercomponent spacing factor N, such that F/N was the fundamental frequency. Parameters that were manipulated included the phase spectrum, the number of components, and the intensity of the center component. Analyses employed Fourier transforms of period histograms to assess the degree to which responses were synchronized to the frequencies present in the acoustic stimulus. Several nonlinearities were observed in the response as intensity was varied between threshold and 80-90 dB SPL. Response nonlinearities were strong for all signals except those with random phase spectra. The most commonly observed nonlinearity was an emphasis of one or more stimulus components in the response. The degree of nonlinearity usually increased with intensity and signal complexity and decreased with fiber frequency selectivity. Half-wave rectification introduced synchronization to the missing fundamental. The strength of the response at the fundamental was related to stimulus crest factor. Signals with low center frequencies and high crest factors often elicited instantaneous discharge rates at the theoretical maximum of pi CF. This suggests that the probability of spike generation approaches one during high-amplitude waveform segments. Response nonlinearity was interpreted as arising from three sources, namely, cochlear mechanics, compression of instantaneous discharge rate, and saturation of average discharge rate. At near-threshold intensities, fibers with high spontaneous rates exhibited responses that were linear functions of stimulus waveshape, whereas fibers with low spontaneous spike rates produced responses that were best described in terms of an expansive nonlinearity.     

Representation of steady-state vowels in the temporal aspects of the discharge patterns of populations of auditory-nerve fibers.

This paper is concerned with the representation of the spectra of synthesized steady-state vowels in the temporal aspects of the discharges of auditory-nerve fibers. The results are based on a study of the responses of large numbers of single auditory-nerve fibers in anesthetized cats. By presenting the same set of stimuli to all the fibers encountered in each cat, we can directly estimate the population response to those stimuli. Period histograms of the responses of each unit to the vowels were constructed. The temporal response of a fiber to each harmonic component of the stimulus is taken to be the amplitude of the corresponding component in the Fourier transform of the unit's period histogram. At low sound levels, the temporal response to each stimulus component is maximal among units with CFs near the frequency of the component (i.e., near its place). Responses to formant components are larger than responses to other stimulus components. As sound level is increased, the responses to the formants, particularly the first formant, increase near their places and spread to adjacent regions, particularly toward higher CFs. Responses to nonformant components, exept for harmonics and intermodulation products of the formants (2F1,2F2,F1 + F2, etc), are suppressed; at the highest sound levels used (approximately 80 dB SPL), temporal responses occur almost exclusively at the first two or three formants and their harmonics and intermodulation products. We describe a simple calculation which combines rate, place, and temporal information to provide a good representation of the vowels' spectra, including a clear indication of at least the first two formant frequencies. This representation is stable with changes in sound level at least up to 80 dB SPL; its stability is in sharp contrast to the behavior of the representation of the vowels' spectra in terms of discharge rate which degenerates at stimulus levels within the conversational range.     

Auditory nerve representation of a complex communication sound in background noise.

A population study of auditory nerve responses in the bullfrog, Rana catesbeiana, analyzed the relative contributions of spectral and temporal coding in representing a complex, species-specific communication signal at different stimulus intensities and in the presence of background noise. At stimulus levels of 70 and 80 dB SPL, levels which approximate that received during communication in the natural environment, average rate profiles plotted over fiber characteristic frequency do not reflect the detailed spectral fine structure of the synthetic call. Rate profiles do not change significantly in the presence of background noise. In ambient (no noise) and low noise conditions, both amphibian papilla and basilar papilla fibers phase lock strongly to the waveform periodicity (fundamental frequency) of the synthetic advertisement call. The higher harmonic spectral fine structure of the synthetic call is not accurately reflected in the timing of fiber firing, because firing is "captured" by the fundamental frequency. Only a small number of fibers synchronize preferentially to any harmonic in the call other than the first, and none synchronize to any higher than the third, even when fiber characteristic frequency is close to one of these higher harmonics. Background noise affects fiber temporal responses in two ways: It can reduce synchronization to the fundamental frequency, until fiber responses are masked; or it can shift synchronization from the fundamental to the second or third harmonic of the call. This second effect results in a preservation of temporal coding at high noise levels. These data suggest that bullfrog eighth nerve fibers extract the waveform periodicity of multiple-harmonic stimuli primarily by a temporal code.     

Neuronal responses to amplitude-modulated and pure-tone stimuli in the guinea pig inferior colliculus, and their modification by broadband noise

Neuronal responses were recorded to pure and to sinusoidally amplitude-modulated (AM) tones at the characteristic frequency (CF) in the central nucleus of the inferior colliculus of anesthetized guinea pigs. Temporal (synchronized) and mean-rate measures were derived from period histograms locked to the stimulus modulation waveform to characterize the modulation response. For stimuli presented in quiet, the modulation gain at low frequencies of modulation (approx less than 50 Hz) was inversely proportional to the neuron's mean firing rate in response to both the modulated stimulus and to a pure tone at an equivalent level. In 43% of units the mean discharge rates in response to the AM stimuli were greatest for those modulation frequencies that generated the largest temporal responses. These discharge-rate maxima occurred at signal intensities corresponding to the steeply sloping part of the neuron's pure-tone rate-intensity function (RIF). The change in mean-rate response to modulated stimuli, as a function of intensity, was qualitatively similar to the pure-tone RIF. Adding broadband noise to the modulated stimulus increased the neuron's temporal response to low modulation frequencies. This increase in modulation gain was correlated with mean firing rate in response to the modulation but did not bear a simple relationship to the noise-induced shift in the RIF measured for a pure tone.     

A temporal analysis of auditory-nerve fiber responses to spoken stop consonant-vowel syllables

Auditory-nerve fiber spike trains were recorded in response to spoken English stop consonant-vowel syllables, both voiced (/b,d,g/) and unvoiced (/p,t,k/), in the initial position of syllables with the vowels /i,a,u/. Temporal properties of the neural responses and stimulus spectra are displayed in a spectrographic format. The responses were categorized in terms of the fibers' characteristic frequencies (CF) and spontaneous rates (SR). High-CF, high-SR fibers generally synchronize to formants throughout the syllables. High-CF, low/medium-SR fibers may also synchronize to formants; however, during the voicing, there may be sufficient low-frequency energy present to suppress a fiber's synchronized response to a formant near its CF. Low-CF fibers, from both SR groups, synchronize to energy associated with voicing. Several proposed acoustic correlates to perceptual features of stop consonant-vowel syllables, including the initial spectrum, formant transitions, and voice-onset time, are represented in the temporal properties of auditory-nerve fiber responses. Nonlinear suppression affects the temporal features of the responses, particularly those of low/medium-spontaneous-rate fibers.     

A high-precision magnetoencephalographic study of human auditory steady-state responses to amplitude-modulated tones

The cerebral magnetic field of the auditory steady-state response (SSR) to sinusoidal amplitude-modulated (SAM) tones was recorded in healthy humans. The waveforms of underlying cortical source activity were calculated at multiples of the modulation frequency using the method of source space projection, which improved the signal-to-noise ratio (SNR) by a factor of 2 to 4. Since the complex amplitudes of the cortical source activity were independent of the sensor position in relation to the subject's head, a comparison of the results across experimental sessions was possible. The effect of modulation frequency on the amplitude and phase of the SSR was investigated at 30 different values between 10 and 98 Hz. At modulation frequencies between 10 and 20 Hz the SNR of harmonics near 40 Hz were predominant over the fundamental SSR. Above 30 Hz the SSR showed an almost sinusoidal waveform with an amplitude maximum at 40 Hz. The amplitude decreased with increasing modulation frequency but was significantly different from the magnetoencephalographic (MEG) background activity up to 98 Hz. Phase response at the fundamental and first harmonic decreased monotonically with increasing modulation frequency. The group delay (apparent latency) showed peaks of 72 ms at 20 Hz, 48 ms at 40 Hz, and 26 ms at 80 Hz. The effects of stimulus intensity, modulation depth, and carrier frequency on amplitude and phase of the SSR were also investigated. The SSR amplitude decreased linearly when stimulus intensity or the modulation depth were decreased in logarithmic steps. SSR amplitude decreased by a factor of 3 when carrier frequency increased from 250 to 4000 Hz. From the phase characteristics, time delays were found in the range of 0 to 6 ms for stimulus intensity, modulation depth, and carrier frequency, which were maximal at low frequencies, low intensities, or maximal modulation depth.     

Speech processing in the auditory system. II: Lateral inhibition and the central processing of speech evoked activity in the auditory nerve

A biologically realistic model of a uniform lateral inhibitory network (LIN) is shown capable of extracting from the complex spatio-temporal firing patterns of the cat's auditory nerve the formants and low-order harmonics of synthetic voiced speech stimuli. The model provides a realistic mechanism to utilize the temporal aspects of the firing and thus supports the hypothesis that the neural coding of complex sounds in terms of average rates can be supplemented by the information coded in the synchronous firing. At low levels of intensity the LIN can sharpen the average rate profiles. At moderate and high levels the LIN uses the cues available in the distribution of phases of the synchronous activity which exhibit rapid relative phase shifts at specific characteristic frequency (CF) locations (corresponding to the frequencies of the low-order harmonics in the stimulus). These temporal phase shifts manifest themselves at the input of the LIN as steep and localized spatial discontinuities in the instantaneous pattern of activity across the fiber array. The LIN enhances its output from these spatially steep input regions while suppressing its output from spatially smooth input regions (where little phase shifts occur). In this manner the LIN recreates from the response patterns a representation of the stimulus spectrum using the temporal cues as spatial markers of the stimulus components rather than as absolute measures of their frequencies. Similar results are obtained with various lateral inhibitory topologies, e.g., recurrent versus nonrecurrent, single versus double layer, and linear versus nonlinear.     

Comparing stimulus-frequency otoacoustic emissions measured by compression, suppression, and spectral smoothing

Stimulus-frequency otoacoustic emissions (SFOAEs) have been measured in several different ways, including (1) nonlinear compression, (2) two-tone suppression, and (3) spectral smoothing. Each of the three methods exploits a different cochlear phenomenon or signal-processing technique to extract the emission. The compression method makes use of the compressive growth of emission amplitude relative to the linear growth of the stimulus. The emission is defined as the complex difference between ear-canal pressure measured at one intensity and the rescaled pressure measured at a higher intensity for which the emission is presumed negligible. The suppression method defines the SFOAE as the complex difference between the ear-canal pressure measured with and without a suppressor tone at a nearby frequency. The suppressor tone is presumed to substantially reduce or eliminate the emission. The spectral smoothing method involves convolving the complex ear-canal pressure spectrum with a smoothing function. The analysis exploits the differing latencies of stimulus and emission and is equivalent to windowing in the corresponding latency domain. Although the three methods are generally assumed to yield identical emissions, no equivalence has ever been established. This paper compares human SFOAEs measured with the three methods using procedures that control for temporal drifts, contamination of the calibration by evoked emissions, and other potential confounds. At low stimulus intensities, SFOAEs measured using all three methods are nearly identical. At higher intensities, limitations of the procedures contribute to small differences, although the general spectral shape and phase of the three SFOAEs remain similar. The near equivalence of SFOAEs measured by compression, suppression, and spectral smoothing indicates that SFOAE characteristics are not mere artifacts of measurement methodology.     

Spectral characteristics and synchrony in primary auditory-nerve fibers in response to pure-tone acoustic stimuli

Under pure-tone stimulation, the spectrum of the period histogram recorded from primary auditory-nerve fibers at low and medium frequencies contains components at DC, at the applied tone frequency (the fundamental), and at a small number of harmonics of the tone frequency. The magnitudes and phases of these spectral components are examined. The spectral magnitudes of the fundamental and various harmonic components generally bear a fixed proportionality to each other over a broad range of signal conditions and nerve-fiber characteristics. This implies that the shape of the underlying rectified wave remains essentially unchanged over a broad range of stimulus intensities. For high-frequency stimuli, the fundamental and harmonic components are substantially attenuated. We provide a theoretical basis for the decrease of the spectralcomponent magnitudes with increasing harmonic number. For low-frequency pure-tone signals, the decrease is caused principally by the uncertainty in the position of neural-event occurrences within the half-wave-rectified period histogram. The lower the stimulus frequency, the greater this time uncertainty and therefore the lower the frequency at which the spectral components begin to diminish. For high-frequency pure-tone signals, on the other hand, the decrease is caused principally by the frequency rolloff associated with nervespike time jitter (it is then called loss of phase locking or loss of synchrony). Since some of this jitter arises from noise in the auditory nerve, it can be minimized by using peak detection rather than level detection. Using a specially designed microcomputer that measures the times at which the peaks of the action potentials occur, we have demonstrated the presence of phase locking to tone frequencies as high as 18 kHz. The traditional view that phase locking is always lost above 6 kHz is clearly not valid. This indicates that the placeversus-periodicity dichotomy in auditory theory requires reexaraination.     

Quantitative aes: the problems of the energy dependent phase shift and modulation amplitude and of the non-ideal behaviour of the channel electron multiplier

In pursuit of the objective to obtain identical Auger electron spectra from different types of instruments, the Varian 10 keV CMA system has been studied. On one particular unit it is shown that the high frequency of 17 kHz, chosen for modulating the spectrometer to produce differential spectra, causes two major errors, each of which affects the relative intensities of peaks in the spectrum. Firstly, it is shown that the phase of the AC signal changes by 56° through the spectrum to 2000 eV, and secondly the amplitude of the AC modulation varies by 100% over the same range. The latter causes low energy peaks to be reduced to half of their true intensity, but the former can cause intensities at either high or low energy to be reduced even more strongly. It is recommended that instruments should be tested and operated, if necessary, at a lower frequency such as 1.7 kHz at which the problems should disappear. A further factor affecting intensities in both the direct and differentiated spectra is the operation of the bias control for the cone of the channel electron multiplier. Field penetration of the cone causes an unwanted behaviour. It is recommended that the cone be fitted with a transparent wire mesh to remove this field penetration and then the bias may be operated at the required potential of at least 200V.     

Conservation of adapting components in auditory-nerve responses

The responses of single auditory-nerve fibers of Mongolian gerbil were studied using tonal stimuli. The peristimulatory adaptation of firing rate in response to tone bursts presented in quiet and during a background stimulus is described quantitatively. The total transient response which can be produced to the onset of a tone burst, whether presented in quiet or as an intensity increment, is limited and appears to demonstrate a form of conservation. Specifically, the total numbers of spikes produced by the rapidly adapting component, and the slower short-term adaptation component, are proportional at all intensities, and are limited for each fiber. Furthermore, when an incremental stimulus is presented on a background, the total transient response to the background and to the increment is limited and depends upon the final intensity, not the background intensity. When the presumed underlying synaptic drive is derived by removing the effects of refractoriness from the spike train, the same conservation of the transient response components, and proportionality between rapid and short-term components, are observed.     

Strategies for the representation of a tone in background noise in the temporal aspects of the discharge patterns of auditory-nerve fibers

The responses of populations of auditory-nerve fibers to both a 1.0-kHz tone, and 1.0-kHz tone in broadband noise, have been measured. Period histograms were generated from fiber spike trains and discrete Fourier transforms (DFTs) with a resolution of 125 Hz were computed from each histogram. Sample mean and sample variance statistics were generated for period histograms of response and for temporal response measures derived from discrete Fourier transforms. It is demonstrated how the statistical properties of auditory-nerve fiber response determine the strategy for the estimation and discrimination of particular stimulus components. When the tone is presented alone, the entire population of auditory-nerve fibers provides statistically reliable estimates of the 1.0-kHz tone. Upon addition of the broadband noise stimulus only those units with characteristic frequencies which are close in frequency to the 1.0-kHz stimulus provide spectral estimates which have high signal-to-noise ratios (mean-squared-to-variance ratios). Estimates of the 1.0-kHz-tone stimulus derived from auditory-nerve fibers with characteristic frequencies which are far from the 1.0-kHz stimulus are statistically unreliable. Based on the responses of the population of auditory-nerve fibers, the strategy for estimating the 1.0-kHz-tone stimulus is to derive estimates of the 1.0-kHz stimulus from the subpopulation of neurons with characteristic frequencies close to the 1.0-kHz stimulus. It is concluded that neurons which are tuned close to 1.0-kHz provide the central nervous system (CNS) with the most salient information about the 1.0-kHz stimulus in the presence of the broadband background.     

Two phase effects in roughness perception.

The respective influences of spectral and temporal aspects of sound in roughness perception are examined by way of phase manipulations. In a first experiment, the phase of the central component of three-component signals is shown to modify perceived roughness, for a given amplitude spectrum, regardless of whether it modifies the waveform envelope. A second experiment shows that the shape of the waveform envelope, for a given amplitude spectrum and a given modulation depth, also influences perceived roughness. We interpret both of these results by considering the envelope of an internal representation that is deduced from the physical signal by taking into account peripheral auditory processing. The results indicate that the modulation depth of such an internal representation is not the only determinant of roughness, but that an effect of temporal asymmetry is also to be taken into account.     

Speech coding in the auditory nerve: IV. Sounds with consonant-like dynamic characteristics

Discharge patterns of auditory-nerve fibers in anesthetized cats were obtained for two stimulus levels in response to synthetic stimuli with dynamic characteristics appropriate for selected consonants. A set of stimuli was constructed by preceding a signal that was identified as /da/by another sound that was systematically manipulated so that the entire complex would sound like either /da/, /ada/, /na/, /sa/, /sa/, or others. Discharge rates of auditory-nerve fibers in response to the common /da/-like formant transitions depended on the preceding context. Average discharge rates during these transitions decreased most for fibers whose CFs were in frequency regions where the context had considerable energy. Some effect of the preceding context on fine time patterns of response to the transitions was also found, but the identity of the largest response components (which often corresponded to the formant frequencies) was in general unaffected. Thus the response patterns during the formant transitions contain cues about both the nature of the transitions and the preceding context. A second set of stimuli sounding like /s/ and /c/ was obtained by varying the duration of the rise in amplitude at the onset of a filtered noise burst. At both 45 and 60 dB SPL, there were fibers which showed a more prominent peak in discharge rate at stimulus onset for /c/ than for /s/, but the CF regions that reflected the clearest distinctions depended on stimulus level. The peaks in discharge rate that occur in response to rapid changes in amplitude or spectrum might be used by the central processor as pointers to portions of speech signals that are rich in phonetic information.     

Speech processing in the auditory system. I: The representation of speech sounds in the responses of the auditory nerve

In a previous paper the speech evoked spatio-temporal response patterns recorded in large populations of auditory-nerve fibers in the cat were examined [M.I. Miller and M.B. Sachs, J. Acoust. Soc. Am. 74, 502-517 (1983)]. The distribution of the relative phases of synchronized activity emerges as an important response feature reflecting the stimulus spectral parameters. Specifically, each strong low-order harmonic of the stimulus (less than or equal to 1.5-2 kHz) dominates the synchrony of a relatively broad segment of fibers near its corresponding characteristic frequency (CF) location in a pattern which mirrors the underlying traveling wave component. Each such fiber segment can be roughly subdivided into two regions: (1) a region basal to the point of resonance of the harmonic where the fiber PST histograms accumulate only small delays (or phase shifts) relative to each other reflecting the fast speed of propagation of the traveling wave, and (2) a region at or very near the point of resonance where the responses exhibit drastic relative phase shifts owing to the sudden slow down of the traveling wave and the consequent rapid accumulation of phase shifts. These rapid phase shifts thus manifest themselves as steep and localized spatial discontinuities in an otherwise relatively uniform instantaneous pattern of activity across the fiber array, all occurring at the CF locations corresponding to the low-order harmonics of the stimulus.     

Effect of blowing snow and ground blizzards on millimeter wave scintillation spectra

We describe the effect of ground blizzards and blowing snow on millimeter wave scintillation spectra at 116 and 230 GHz. During snowstorms, multipath propagation can occur for various reasons. Whatever the reason for the multipath, it will affect the scintillation spectrum. Enhancement of the scintillation spectrum is observed at both low and high temporal frequencies. In some cases, two additional corner frequencies are evident at the high-frequency end of the scintillation spectrum. The method of obtaining the cross-path wind component fails when the wind is nearly parallel to the propagation path.     

基于时频波谱对易原理分析莫尔条纹谱

数值解析信号波形时频域对易演变过程,研究任意信号波形与频率组分的内在联系, 并将该对易原理应用于莫尔条纹相位分析,提取相位信息。采用矩形窗模拟冲激波形和直流波形的变换过程,通过控制矩形窗函数窗宽,获得各种宽度矩形脉冲,窗宽趋于零情况下获得冲激波形,趋于∞获得直流波形。自开发快速傅里叶变换(fast Fourier transform, FFT)系统,对矩形脉冲实施离散傅里叶变换,方便快捷获得相应频谱数值解析波形,分析波形与频谱对易关系。结果发现,矩形窗函数频谱是Sa函数,窗宽变化导致Sa函数波形变化。窗宽减小时,Sa函数波形展宽,振动舒缓,趋于零极限时,变成直流波形。窗宽增大时,Sa函数波形紧缩,振动加剧,趋于∞的极限时,演变成δ冲激波形,信号波形时频域是对易的。根据时频域波形与频谱对易关系,在分析莫尔条纹时,将莫尔条纹的一级谱滤出并归一,由波谱对易原理,时域信号将体现Sa函数,使条纹对比分明,便于提取相位信息。     

Speech coding in the auditory nerve: II. Processing schemes for vowel-like sounds

Several processing schemes by which phonetically important information for vowels can be extracted from responses of auditory-nerve fibers are analyzed. The schemes are based on power spectra of period histograms obtained in response to a set of nine two-formant, steady-state, vowel-like stimuli presented at 60 and 75 dB SPL. One class of "local filtering" schemes, which was originally proposed by Young and Sachs [J. Acoust. Soc. Am. 66, 1381-1403 (1979)], consists of analyzing response patterns by filters centered at the characteristic frequencies (CF) of the fibers, so that a tonotopically arranged measure of synchronized response can be obtained. Various schemes in this class differ in the characteristics of the filter. For a wide range of filter bandwidths, formant frequencies correspond approximately to the CFs for which the response measure is maximal. If in addition, the bandwidths of the analyzing filters are made compatible with psychophysical measures of frequency selectivity, low-frequency harmonics of the stimulus fundamental are resolved in the output profile, so that fundamental frequency can also be estimated. In a second class of processing schemes, a dominant response component is defined for each fiber from a 1/6 octave spectral representation of the response pattern, and the formant frequencies are estimated from the most frequent values of the dominant component in the ensemble of auditory-nerve fibers. The local filtering schemes and the dominant component schemes can be related to "place" and "periodicity" models of auditory processing, respectively.     

The role of the envelope in processing iterated rippled noise.

Iterated rippled noise (IRN) is generated by a cascade of delay and add (the gain after the delay is 1.0) or delay and subtract (the gain is -1.0) operations. The delay and add/subtract operations impart a spectral ripple and a temporal regularity to the noise. The waveform fine structure is different in these two conditions, but the envelope can be extremely similar. Four experiments were used to determine conditions in which the processing of IRN stimuli might be mediated by the waveform fine structure or by the envelope. In experiments 1 and 3 listeners discriminated among three stimuli in a single-interval task: IRN stimuli generated with the delay and add operations (g = 1.0), IRN stimuli generated using the delay and subtract operations (g = -1.0), and a flat-spectrum noise stimulus. In experiment 2 the listeners were presented two IRN stimuli that differed in delay (4 vs 6 ms) and a flat-spectrum noise stimulus that was not an IRN stimulus. In experiments 1 and 2 both the envelope and waveform fine structure contained the spectral ripple and temporal regularity. In experiment 3 only the envelope had this spectral and temporal structure. In all experiments discrimination was determined as a function of high-pass filtering the stimuli, and listeners could discriminate between the two IRN stimuli up to frequency regions as high as 4000-6000 Hz. Listeners could discriminate the IRN stimuli from the flat-spectrum noise stimulus at even higher frequencies (as high as 8000 Hz), but these discriminations did not appear to depend on the pitch of the IRN stimuli. A control experiment (fourth experiment) suggests that IRN discriminations in high-frequency regions are probably not due entirely to low-frequency nonlinear distortion products. The results of the paper imply that pitch processing of IRN stimuli is based on the waveform fine structure.