Diversity of adaptation patterns in responses of eighth nerve fibers in the bullfrog, Rana catesbeiana

The firing patterns of eighth nerve fibers in the bullfrog, Rana catesbeiana, were analyzed for responses to long duration tone bursts at best excitatory frequency ( BEF ) and at frequencies along the upper and lower boundaries of the excitatory tuning curve of each fiber. These firing patterns were used as an index of the degree of short-term adaptation of each fiber. Amphibian papilla fibers (with BEFs 100-1000 Hz) exhibited marked diversity in their firing patterns to BEF tones, ranging from very flat or tonic (sustained responses throughout the duration of the stimulus) to very peaked or phasic (responding primarily or exclusively to stimulus onset). Moreover, the degree of short-term adaptation shown by an individual fiber varied with stimulating frequency. The firing patterns of amphibian papilla fibers tended to become more tonic as stimulus frequency was lowered below BEF ; conversely, as stimulus frequency was increased above BEF , firing patterns either showed little change from that at BEF , or became more phasic. A similar frequency dependence of adaptation has not been reported in responses of mammalian eighth nerve fibers with comparable BEFs . The firing patterns of basilar papilla fibers ( BEFs greater than 1000 Hz) remained similar in response to both BEF and non- BEF tones. These data reveal that the firing patterns and degrees of short-term adaptation of amphibian papilla fibers vary considerably across the tuning curve, whereas those of basilar papilla fibers remain relatively more constant with changes in stimulating frequency.(ABSTRACT TRUNCATED AT 250 WORDS)     

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.     

Frequency selectivity of single cochlear-nerve fibers based on the temporal response pattern to two-tone signals

The physiological basis of auditory frequency selectivity was investigated by recording the temporal response patterns of single cochlear-nerve fibers in the cat. The characteristic frequency and sharpness of tuning was determined for low-frequency cochlear-nerve fibers with two-tone signals whose frequency components were of equal amplitude and starting phase. The measures were compared with those obtained with sinusoidal signals. The two-tone characteristic frequency (2TCF) is defined as the arithmetic-center frequency at which the fiber is synchronized to both signal frequencies in equal measure. The 2TCF closely corresponds to the characteristic frequency as determined by the frequency threshold curve. Moreover, the 2TCF changes relatively little (2%-12%) over a 60-dB intensity range. The 2TCF generally shifts upward with increasing intensity for cochlear-nerve fibers tuned to frequencies below 1 kHz and shifts downward as a function of intensity for units with characteristic frequencies (CF's) above 1 kHz. The shifts in the 2TCF are considerably smaller than those observed with sinusoidal signals. Filter functions were derived from the synchronization pattern to the two-tone signal by varying the frequency of one of the components over the fiber's response area while maintaining the other component at the 2TCF. The frequency selectivity of the two-tone filter function was determined by dividing the vector strength to the variable frequency signal by the vector strength to the CF tone. The filter function was measured 10 dB down from the peak (2T Q 10 dB) and compared with the Q 10 dB of the frequency threshold curve. The correlation between the two measures of frequency selectivity was 0.72. The 2T Q 10 dB does change as a function of intensity. The magnitude and direction of the change is dependent on the sharpness of tuning at low and moderate sound-pressure levels (SPL's). The selectivity of the more sharply tuned fibers (2T Q 10 dB greater than 3) diminishes at intensities above 60 dB SPL. However, the broadening of selectivity is relatively small in comparison to discharge rate-based measures of selectivity. The selectivity of the more broadly tuned units remains unchanged or improves slightly at similar intensity levels. The present data indicate that the frequency selectivity and tuning of low-frequency cochlear-nerve fibers are relatively stable over a 60-dB range of SPL's when measured in terms of their temporal discharge properties.     

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.     

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.     

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.     

Effects of frequency modulation on absolute, masked, and reflex thresholds

Behavioral and acoustic reflex thresholds were determined for five normal-hearing subjects in response to carrier signals of 500 and 2000 Hz which were unmodulated or modulated sinusoidally at rates of 2, 20, and 200 times per second with frequency deviations (delta f) of 30, 100, and 300 Hz. Behavioral (absolute and masked) thresholds were determined using an adaptive two-alternative forced-choice procedure. Acoustic reflex thresholds were determined by visual inspection of stored reflex waveforms. Frequency modulation was not found to exert a systematic effect at absolute threshold. Frequency modulation did affect threshold estimates systematically, but differentially, at masked threshold and acoustic reflex threshold. Increasing the frequency deviation of the modulation was associated with an increase in masked threshold and with a decrease in acoustic reflex threshold at both test frequencies. The findings are discussed in terms of critical band phenomena.     

Forward masking of amplitude modulation: basic characteristics

In this study we demonstrate an effect for amplitude modulation (AM) that is analogous to forward making of audio frequencies, i.e., the modulation threshold for detection of AM (signal) is raised by preceding AM (masker). In the study we focused on the basic characteristics of the forward-masking effect. Functions representing recovery from AM forward masking measured with a 150- ms 40- Hz masker AM and a 50- ms signal AM of the same rate imposed on the same broadband-noise carrier, showed an exponential decay of forward masking with increasing delay from masker offset. Thresholds remained elevated by more than 2 dB over an interval of at least 150 ms following the masker. Masked-threshold patterns, measured with a fixed signal rate (20, 40, and 80 Hz) and a variable masker rate, showed tuning of the AM forward-masking effect. The tuning was approximately constant across signal modulation rates used and consistent with the idea of modulation-rate selective channels. Combining two equally effective forward maskers of different frequencies did not lead to an increase in forward masking relative to that produced by either component alone. Overall, the results are consistent with modulation-rate selective neural channels that adapt and recover from the adaptation relatively quickly.     

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.     

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.     

Comparison of level discrimination, increment detection, and comodulation masking release in the audio- and envelope-frequency domains

In general, the temporal structure of stimuli must be considered to account for certain observations made in detection and masking experiments in the audio-frequency domain. Two such phenomena are (1) a heightened sensitivity to amplitude increments with a temporal fringe compared to gated level discrimination performance and (2) lower tone-in-noise detection thresholds using a modulated masker compared to those using an unmodulated masker. In the current study, translations of these two experiments were carried out to test the hypothesis that analogous cues might be used in the envelope-frequency domain. Pure-tone carrier amplitude-modulation (AM) depth-discrimination thresholds were found to be similar using both traditional gated stimuli and using a temporally modulated fringe for a fixed standard depth (ms = 0.25) and a range of AM frequencies (4-64 Hz). In a second experiment, masked sinusoidal AM detection thresholds were compared in conditions with and without slow and regular fluctuations imposed on the instantaneous masker AM depth. Release from masking was obtained only for very slow masker fluctuations (less than 2 Hz). A physiologically motivated model that effectively acts as a first-order envelope change detector accounted for several, but not all, of the key aspects of the data.     

The influence of spread of excitation on the detection of amplitude modulation imposed on sinusoidal carriers at high levels

The improvement in amplitude modulation (AM) detection thresholds with increasing level of a sinusoidal carrier has been attributed to listening on the high-frequency side of the excitation pattern, where the growth of excitation is more linear, or to an increase in the number of "channels" via spread of excitation. In the present study, AM detection thresholds were measured using a 1000-Hz sinusoidal carrier. Thresholds for modulation frequencies of 4-64 Hz improved by about 10-20 dB as the carrier level increased from 10 dB SL (14.5 dB SPL on average) to 80 dB SPL. To minimize the use of spread of excitation with an 80-dB carrier, tonal "restrictors" with frequencies of 501, 801, 1210, and 1510 Hz were used alone and in combination. High-frequency restrictors elevated AM detection thresholds, whereas low-frequency restrictors did not, indicating that excitation on the high side is more important for detecting AM. Results of modeling suggest that the improvement in AM detection thresholds at high levels is likely due to the use of a relatively linear growth of response on the high-frequency side of the excitation pattern.     

Spectral modulation masking patterns reveal tuning to spectral envelope frequency

Auditory processing appears to include a series of domain-specific filtering operations that include tuning in the audio-frequency domain, followed by tuning in the temporal modulation domain, and perhaps tuning in the spectral modulation domain. To explore the possibility of tuning in the spectral modulation domain, a masking experiment was designed to measure masking patterns in the spectral modulation domain. Spectral modulation transfer functions (SMTFs) were measured for modulation frequencies from 0.25 to 14 cycles/octave superimposed on noise carriers either one octave (800-1600 Hz, 6400-12,800 Hz) or six octaves wide (200-12,800 Hz). The resulting SMTFs showed maximum sensitivity to modulation between 1 and 3 cycles/octave with reduced sensitivity above and below this region. Masked spectral modulation detection thresholds were measured for masker modulation frequencies of 1, 3, and 5 cycles/octave with a fixed modulation depth of 15 dB. The masking patterns obtained for each masker frequency and carrier band revealed tuning (maximum masking) near the masker frequency, which is consistent with the theory that spectral envelope perception is governed by a series of spectral modulation channels tuned to different spectral modulation frequencies.     

Digital EPR with an arbitrary waveform generator and direct detection at the carrier frequency

A digital EPR spectrometer was constructed by replacing the traditional bridge with an arbitrary waveform generator (AWG) to produce excitation patterns and a high-speed digitizer for direct detection of the spin system response at the carrier frequency. Digital down-conversion produced baseband signals in quadrature with very precise orthogonality. Real-time resonator tuning was performed by monitoring the Fourier transforms of signals reflected from the resonator during frequency sweeps generated by the AWG. The capabilities of the system were demonstrated by rapid magnetic field scans at 256 MHz carrier frequency, and FID and spin echo experiments at 1 and 10 GHz carrier frequencies. For the rapid scan experiments the leakage through a cross-loop resonator was compensated by adjusting the amplitude and phase of a sinusoid at the carrier frequency that was generated with another AWG channel.     

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.     

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

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

Dynamic all-optical tuning of transverse resonant cavity modes in photonic bandgap fibers

Photonic bandgap fibers for transverse illumination containing half-wavelength microcavities have recently been designed and fabricated. We report on the fabrication and characterization of an all-optical tunable microcavity fiber. The fiber is made by incorporating a photorefractive material inside a Fabry-Perot cavity structure with a quality factor Q >200 operating at 1.5 microm. Under short-wavelength transverse external illumination, a 2 nm reversible shift of the cavity resonant mode is achieved. Dynamic all-optical tuning is reported at frequencies up to 400 Hz. Experimental results are compared with simulations based on the amplitude and kinetics of the transient photodarkening effect measured in situ in thin films.     

Inner hair cell response patterns: implications for low-frequency hearing.

Inner hair cell (IHC) responses to tone-burst stimuli were measured from three locations in the apical half of the guinea pig cochlea. In addition to the measurement of ac receptor potentials, average intracellular voltages, reflecting both ac and dc components of the receptor potential, were computed and compared to determine how bandwidth changes with level. Companion phase measures were also obtained and evaluated. Data collected from turn 2, where best frequency (BF) is approximately 4000 Hz, indicate that frequency response functions are asymmetrical with steeper slopes above the best frequency of the cell. However, in turn 4, where BF is around 250 Hz, the opposite behavior is observed and the steepest slopes are measured below BF. The data imply that cochlear filters are generally asymmetrical with steeper slopes above BF. High-pass filtering by the middle ear serves to reduce this asymmetry in turn 3 and to reverse it in turn 4. Apical response patterns are used to assess the degree to which the middle ear transfer function, the IHC's velocity dependence and the shunting effect of the helicotrema influence low-frequency hearing in guinea pigs. Implications for low-frequency hearing in man are also discussed.     

20 Hz~10 kHz光纤水听器相移灵敏度校准

利用相位生成载波解调技术精确测量光纤水听器的光相移量,在20 Hz~10 kHz频率范围实现了光纤水听器探头相移灵敏度的校准.20 Hz~1.25 kHz频段采用驻波管比较法进行校准,1.25 kHz~10 kHz频段采用自由场脉冲比较法进行校准.利用本文建立的校准系统,对TMD 35#光纤水听器的相移灵敏度进行校准,校准结果表明,两种方法测得的相移灵敏度具有很好的一致性,在1.25 kHz频率点的相移灵敏度值偏差为0.8 dB.不确定度分析表明,该校准系统的扩展不确定度(k=2)为0.9 dB.