Relationship between psychophysical tuning curves and "suppression"

This study examined two-tone unmasking and auditory frequency selectivity about 3 kHz for the purpose of demonstrating a qualitative relationship between the two. An adaptive 2IFC forward-masking procedure was used to collect psychophysical tuning curves (PTC's) and two-tone masking data under a quiet and noise condition for the same normal-hearing listeners. In the noise condition, a narrowband noise masker, centered one decade down from the probe, was gated on with the tonal masker(s). Kiang and Moxon [J. Acoust. Soc. Am. 55, 620-630 (1974)] have found that low-frequency narrowband noise serves to decrease the sharpness of electrophysiological tuning curves by affecting only the tip segments. The data for four highly practiced listeners indicate that the gated-noise masker was effective in broadening the PTC's and in lessening the magnitude of two-tone unmasking. The mutually reflected changes in tuning curves and in two-tone unmasking indicate a close relationship between frequency selectivity and unmasking: the greater the magnitude of unmasking above the center frequency of the PTC, the sharper the tuning of the PTC.     

Acoustic response and tuning in saccular nerve fibers of the goldfish (Carassius auratus)

The acoustic frequency selectivity of over 500 saccular nerve fibers of the goldfish was studied using automated threshold tracking based on spike rate increments defined statistically. Saccular fibers of the goldfish show great variation in (1) best sensitivity (-26 to + 35 dB re: 1 dyn/cm2), (2) best frequency (below 100 to 1770 Hz), (3) spontaneous rate (0 to over 200 spikes/s), (4) spontaneous type (silent, regular, irregular, burst), and (5) degree of tuning (Q 10 dB from less than 0.1 to 2). Saccular fibers may be grouped into four nonoverlapping categories based on tuning and best frequency: (1) untuned (less than 10-dB variation in sensitivity between 100 and 1000 Hz), (2) low frequency (BF from below 120 to 290 Hz), (3) midfrequency (BF between 330 and 670 Hz), and (4) high frequency (BF between 790 and 1770 Hz). Within each category, all spontaneous rates and types, and all degrees of tuning can be observed. The least sensitive fibers within each group have zero spontaneous rates. The goldfish is like all other vertebrates studied in that the peripheral auditory system is adapted for frequency selectivity throughout the animal's entire frequency range of hearing. Peripheral tuning most likely accounts for behavioral determinations of the "auditory filter" and for the detectability of signals masked by noise. The signal-to-noise ratio enhancement provided by these peripheral filters is likely to be of primary biological significance. A "place principle" of sound quality analysis based on lines "labeled" according to best frequency in the brain cannot be ruled out on the basis of the peripheral physiology.     

Basilar membrane mechanics at the base of the chinchilla cochlea. I. Input-output functions, tuning curves, and response phases

Basilar membrane (BM) velocity was measured at a site 3.5 mm from the basal end of the chinchilla cochlea using the M?ssbauer technique. The threshold of the compound action potential recorded at the round window in response to tone bursts was used as an indicator of the physiological state of the cochlea. The BM input-output functions display a compressive nonlinearity for frequencies around the characteristic frequency (CF, 8 to 8.75 kHz), but are linear for frequencies below 7 and above 10.5 kHz. In preparations with little surgical damage, isovelocity tuning curves at 0.1 mm/s are sharply tuned, have Q10's of about 6, minima as low as 13 dB SPL, tip-to-tail ratios (at 1 kHz) of 56 to 76 dB, and high-frequency slopes of about 300 dB/oct. These mechanical responses are as sharply tuned as frequency-threshold curves of chinchilla auditory nerve fibers with corresponding CF. There is a progressive loss of sensitivity of the mechanical response with time for the frequencies around CF, but not for frequencies on the tail of the tuning curve. In some experiments the nonlinearity was maintained for several hours, in spite of a considerable loss of sensitivity of the BM response. High-frequency plateaus were observed in both isovelocity tuning curves and phase-frequency curves.     

Response dynamics of goldfish saccular fibers: effects of stimulus frequency and intensity on fibers with different tuning, sensitivity, and spontaneous activity

The effects of stimulus frequency and intensity on response patterns (PST histograms) to tone burst stimulation were examined in differently tuned saccular fibers of the goldfish. In addition, the sensitivity of these fibers to amplitude-modulated (AM) signals of different carrier frequencies was measured. The response patterns evoked by unmodulated signals were a complex function of tuning, spontaneous activity and sensitivity of the fiber, and the frequency and intensity of the signal. Frequency-dependent response patterns were found in low-frequency fibers with best frequencies (BF) below 200 Hz. Responses in these fibers ranged from tonic to phasic in nonspontaneous fibers and included more complex patterns in spontaneously active fibers, such as suppression of evoked activity below spontaneous levels. Midfrequency fibers (BF = 500-600 Hz) showed responses similar to those in low-frequency fibers, but with less dependence on frequency. In contrast, both high-frequency (BF = 800-1000 Hz) and wideband, untuned fibers showed frequency-invariant patterns of adaptation. High-frequency fibers were equally sensitive to AM signals at all frequencies tested. The sensitivity of low-frequency fibers to AM, however, increased as a function of carrier frequency and corresponded to the degree of adaptation in response to unmodulated tones. In general, the AM sensitivity of a fiber could be predicted more by its pattern of response to unmodulated signals than by its tuning characteristics.     

The temporal evolution of masking and frequency selectivity in the goldfish (Carassius auratus)

The temporal evolution of masking and frequency selectivity was studied in the goldfish using classical respiratory conditioning and a tracking psychophysical procedure. The temporal position of a brief tonal signal within a longer duration, tonal masker has little or no effect on signal detectability when the frequency of the masker is less than or equal to that of the signal. For masker frequencies above that of the signal, signal detectability improves as the signal onset is delayed relative to that of the masker. These patterns of tone-on-tone masking are quite similar to those observed for humans. These temporal masking patterns are qualitatively similar in shape to the peristimulus-time histogram profiles of the low-frequency saccular fibers thought to be used in this task. Frequency- and time-dependent changes in signal detectability result in specific changes in the sharpness of psychophysical tuning curves (PTC). In general, PTCs determined for signals occurring at masker onset are the most broadly tuned, and PTCs determined in forward masking are the most sharply tuned. The PTCs for signals temporally centered in the masker are intermediate. These results suggest that temporal tone-on-tone masking patterns and the temporal evolution of psychophysical tuning curves result from the response properties of peripheral auditory-nerve fibers.     

A spatial template for the shape of tuning curves in the mammalian cochlea

The shape of the tuning curve of primary auditory neurons of four mammals is characterized using a simple exponential model. The regression analysis formalizes a distinction between the characteristic frequency of a neuron and its "nominal" characteristic frequency in cases of temporary threshold loss in high-frequency neurons. Second, the model offers a stronger quality test for sharpness of tuning than the Q10dB since it takes into account the threshold of the neuron at its characteristic frequency and its "characteristic place" of origin along the cochlear partition. Third, the model reveals that the low-frequency side of the tip segment of the tuning curve is bounded by a constraint or template which is most simply expressed in spatial terms. The template describes the basal-side boundary of an "excitatory region" whose length along the cochlear partition is proportional to the square root of the sound pressure. Tuning curve variability arises because biological dependencies influence the basic template. A "spatial-filter" hypothesis is developed and its generality is discussed, particularly in regard to the case of the acoustic "fovea" of the horseshoe bat. Finally, the possibility is discussed that the template possesses a simple physiological correlate in the form of a spatially localized region marked by a "dc" shift of the mean position of the basilar membrane which sets the sensitivity of the tuning mechanism [E.L. LePage, J. Acoust. Soc. Am. 82, 139-154 (1987)].     

Distortion-product otoacoustic emission suppression tuning curves in humans

Distortion-product otoacoustic emission (DPOAE) suppression data as a function of suppressor level (L(3)) for f(2) frequencies from 0.5 to 8 kHz and L(2) levels from 10 to 60 dB sensation level were used to construct suppression tuning curves (STCs). DPOAE levels in the presence of suppressors were converted into decrement versus L(3) functions, and the L(3) levels resulting in 3 dB decrements were derived by transformed linear regression. These L(3) levels were plotted as a function of f(3) to construct STCs. When f(3) is represented on an octave scale, STCs were similar in shape across f(2) frequency. These STCs were analyzed to provide estimates of gain (tip-to-tail difference) and tuning (Q(ERB)). Both gain and tuning decreased as L(2) increased, regardless of f(2), but the trend with f(2) was not monotonic. A roughly linear relation was observed between gain and tuning at each frequency, such that gain increased by 4-16 dB (mean ≈ 5 dB) for every unit increase in Q(ERB), although the pattern varied with frequency. These findings suggest consistent nonlinear processing across a wide frequency range in humans, although the nonlinear operation range is frequency dependent.     

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.     

Two-tone suppression in the cricket, Eunemobius carolinus (Gryllidae, Nemobiinae)

Sounds with frequencies >15 kHz elicit an acoustic startle response (ASR) in flying crickets (Eunemobius carolinus). Although frequencies <15 kHz do not elicit the ASR when presented alone, when presented with ultrasound (40 kHz), low-frequency stimuli suppress the ultrasound-induced startle. Thus, using methods similar to those in masking experiments, we used two-tone suppression to assay sensitivity to frequencies in the audio band. Startle suppression was tuned to frequencies near 5 kHz, the frequency range of male calling songs. Similar to equal loudness contours measured in humans, however, equal suppression contours were not parallel, as the equivalent rectangular bandwidth of suppression tuning changed with increases in ultrasound intensity. Temporal integration of suppressor stimuli was measured using nonsimultaneous presentations of 5-ms pulses of 6 and 40 kHz. We found that no suppression occurs when the suppressing tone is >2 ms after and >5 ms before the ultrasound stimulus, suggesting that stimulus overlap is a requirement for suppression. When considered together with our finding that the intensity of low-frequency stimuli required for suppression is greater than that produced by singing males, the overlap requirement suggests that two-tone suppression functions to limit the ASR to sounds containing only ultrasound and not to broadband sounds that span the audio and ultrasound range.     

Two-tone suppression in auditory nerve of the cat: rate-intensity and temporal analyses

Responses to two-tone stimuli were recorded from auditory-nerve fibers in anesthetized cats. One tone, the suppressor, was set at a frequency above characteristic frequency and was fixed in intensity. A second tone was set at an excitatory frequency and was varied in intensity. The suppressor tone, when set at a sufficient level, always reduced the response to the excitatory tone by an amount equivalent to a fixed number of decibels, regardless of the excitatory tone's intensity. Estimates of suppression magnitude were derived from shifts in rate-intensity function obtained when the suppressor tone was present relative to the functions obtained for the excitatory tone alone. When suppressor-tone intensity was increased, suppression magnitude likewise increased. When the two tones were increasingly separated in frequency, either by varying the excitor or by varying the suppressor, suppression magnitude decreased monotonically. Suppression behaved in the same manner regardless of whether suppresor tone was excitatory or nonexcitatory. When frequency separation was small enough and when both tones were above the neuron's characteristic frequency, responses synchronized to low-order combination tones could be elicited. These responses usually possessed different rate-intensity characteristics and resulted in estimates of suppression magnitude which were spuriously low. When frequency separation is normalized with regard to position of traveling wave maxima within the cochlear duct, the magnitude of two-tone suppression for a given suppressor-tone intensity is seen to be frequency independent.     

Inner hair cell responses to the 2f1-f2 intermodulation distortion product

Recordings of dc and ac receptor potentials from pigmented guinea pig inner hair cells indicate strong responses to the 2f1-f2 intermodulation tone when f1 and f2 are greater than the hair cell characteristic frequency and do not cause a response when given individually. The effective magnitude of this cubic distortion product (CDP) was about 25-30 dB below equal sound level primaries over a 20-30-dB range of their sound levels. The relative strength of the CDP declined at a rate greater than 180-dB/oct separation of the primaries. When magnitude of f1 or f2 was held constant, the growth of CDP was nonmonotonic, exhibiting a distinct maximum. With a constant level of f1 or f2, optimal CDP was produced when the level of f2 was 10-15 dB greater than f1. Strong two-tone suppression from the primaries has a role in shaping the CDP growth. The ac receptor potentials of the CDP show a 150 degrees-200 degrees phase shift when the primaries are increased over a 50-dB range. These results support the hypothesis of a propagated CDP in the cochlea and are consistent with the major features of related studies of human psychoacoustic experiments, afferent nerve neural rate functions, and ear canal distortion products.     

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.     

The detection of temporal gaps as a function of frequency region and absolute noise bandwidth.

Temporal gap detection was measured as a function of absolute signal bandwidth at a low-, a mid-, and a high-frequency region in six listeners with normal hearing sensitivity. Gap detection threshold decreased monotonically with increasing stimulus bandwidth at each of the three frequency regions. Given conditions of equivalent absolute bandwidth, gap detection thresholds were not significantly different for upper cutoff frequencies ranging from 600 to 4400 Hz. A second experiment investigated gap detection thresholds at two pressure-spectrum levels, conditions typically resulting in substantially different estimates of frequency selectivity. Estimates of frequency selectivity were collected at the two levels using a notched-noise masker technique. The gap threshold-signal bandwidth functions were almost identical at pressure-spectrum levels of 70 dB and 40 dB for the two subjects in experiment II, while estimates of frequency selectivity showed poorer frequency selectivity at the 70-dB level than at 40 dB. Data from both experiments indicated that gap detection in bandlimited noise was inversely related to signal bandwidth and that gap detection did not vary significantly with changes in signal frequency over the range of 600 to 4400 Hz. Over the range of frequencies investigated, the results indicated no clear relation between gap detection for noise stimuli and peripheral auditory filtering.     

Nonlinear phenomena as observed in the ear canal and at the auditory nerve

We report here several measures of nonlinear effects in the mammalian ear made in the external auditory meatus and in single neurons of the auditory nerve. We have measured the 2f1-f2 and the f2-f1 distortion products and we have found that the neural distortion product threshold curve for 2f1-f2 mirrors the low-frequency side of the frequency threshold curve, when the neural distortion product threshold curve of 2f1-f2 is plotted versus log(f2/f1) its slope is about 50 dB/oct and its intercept is 10-20 dB above the frequency threshold at the characteristic frequency CF, substantial 2f1-f2 distortion was seen in all animals studied while the f2-f1 distortion product was only rarely found at substantial levels, and the distortion product pressure observed in the ear canal was at a level equal to that detected at threshold by the neural units under study. We have also made measurements of two-tone rate suppression thresholds using two new and consistent threshold paradigms. We find that for high and intermediate characteristic frequency neural units the suppression threshold is independent of frequency and at a level of about 70 dB SPL, the suppression above CF is much less than below CF, and the tip of the frequency tuning curve can be suppressed by up to 40 dB by a low-frequency suppressor.     

Influence of rate of change of frequency on the overall pitch of frequency-modulated tones

The mechanism(s) determining pitch may assign less weight to portions of a sound where the frequency is changing rapidly. The present experiments explored the possible effect of this on the overall pitch of frequency-modulated sounds. Pitch matches were obtained between an adjustable unmodulated sinusoid and a sinusoidal carrier that was frequency modulated using a highly asymmetric function with the form of a repeating U or inverted U shaped function. The amplitude was constant during the 400-ms presentation time of each stimulus, except for 10-ms raised-cosine onset and offset ramps. In experiment 1, the carrier level was 50 dB SPL and the geometric mean of the instantaneous frequency of the modulated carrier, fc, was either 0.5, 1, 2, or 8 kHz. The modulation rate (fm) was 5, 10, or 20 Hz. The overall depth (maximum to minimum) of the FM was 8% of fc. For all carrier frequencies, the matched frequency was shifted away from the mean carrier frequency, downwards for the U shaped function stimuli and upwards for the repeated inverted U shaped function stimuli. The shift was typically slightly greater than 1% of fc, and did not vary markedly with fc. The effect of fm was small, but there was a trend for the shifts to decrease with increasing fm for fc = 0.5 kHz and to increase with increasing fm for fc = 2 kHz. In experiment 2, the carrier level was reduced to 20 dB SL and matches were obtained only for fc = 2 kHz. Shifts in matched frequency of about 1% were still observed, but the trend for the shifts to increase with increasing fm no longer occurred. In experiment 3, matches were obtained for a 4-kHz carrier at 50 dB SPL. Shifts of about 1% again occurred, which did not vary markedly with fm. The shifts in matched frequency observed in all three experiments are not predicted by models based on the amplitude- or intensity-weighted average of instantaneous frequency (EWAIF or IWAIF). The shifts (and the pitch shifts observed earlier for two-tone complexes and for stimuli with simultaneous AM and FM) are consistent with a model based on the assumption that the overall pitch of a frequency-modulated sound is determined from a weighted average of period estimates, with the weight attached to a given estimate being inversely related to the short-term rate of change of period and directly related to a compressive function of the amplitude.     

Acoustic distortion in the ear canal. I. Cubic difference tones: effects of acute noise injury

Acoustic emissions in the form of cubic difference tones (CDT's), 2f1-f2, were measured in the ear canals of gerbils and cats. The state of the cochlea was manipulated by means of acute exposure to noise and was monitored with the aid of the whole-nerve response to tone pips. The resulting shifts in the levels of emissions generated by pairs of primary tones of equal intensity were then compared to the corresponding threshold shifts of the whole-nerve response across frequency. Data obtained from normal ears before injury indicate that the absolute thresholds of the whole-nerve responses across frequency are not necessarily good predictors of the absolute levels of CDT emissions generated by 70- and 80-dB SPL primaries. While high emission levels were often linked to low whole-nerve thresholds in pre-exposed ears, instances of animals with sensitive whole-nerve thresholds coupled with very weak emissions were also found. Conversely, animals with poor whole-nerve thresholds (shifted by up to 30 dB) could occasionally have high levels of emissions. After acute noise injury, however, the shifts of emission levels as a function of the center frequency of the primary-tone pair largely corresponded to the threshold shifts seen in the whole-nerve response. In other words, the temporary level shift of an acoustic emission largely reflected the acute change to a specific cochlear region associated with the primary frequencies.     

Vibrotactile forward masking: evidence for channel independence

Threshold shifts for the detection of vibrotactile test stimuli were determined as a function of the intensity of a masker. A 50-ms sinusoidal test stimulus was applied to the thenar eminence of the hand 25 ms after the termination of a 700-ms sinusoidal masker applied to the same site. The intensity of the masker was varied over a range of 0-44 dB SL. The frequency of the masker was either 15 or 250 Hz and the frequency of the test stimulus was either 15, 25, 100, or 250 Hz. The results support the hypothesis that the detection of vibrotactile stimuli is mediated by at least two receptor systems which do not mask each other.     

Intensity discrimination of pulsed tones by the goldfish (Carassius auratus)

Intensity discrimination thresholds for 500-ms pure-tone bursts were measured as a function of frequency in the goldfish (Carassius auratus) using classical respiratory conditioning. At 55-dB sensation level (SL), thresholds range from 1.44-2.2 dB between 100 and 1600 Hz. There is not important effect of frequency on intensity discrimination. Thresholds at 35-dB SL average 0.7 dB higher than at 55-dB SL. This is a small difference in the context of the threshold variability. In intensity discrimination acuity, the goldfish is quantitatively similar to other vertebrates, including birds and mammals.     

Photonic generation of high quality frequency-tunable millimeter wave and terahertz wave

A scheme for the photonic generation of frequency-tunable millimeter wave and terahertz wave signals based on a highly flat optical frequency comb is proposed and demonstrated experimentally.The frequency comb is generated using two cascaded phase modulators(PMs)and an electro-absorption modulator(EAM).The frequency comb covers a 440-GHz frequency range,with 40-GHz comb spacing and less than 2-dB amplitude variation.By filtering out two of the comb lines with 50 dB out of the band suppression ratio,high frequency-purity and low phase noise millimeter wave or terahertz wave signals are successfully generated,with frequencies ranging from 40 to 440 GHz.