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.     

Sound intensity processing by the goldfish

Capacities of the goldfish for intensity discrimination were studied using classical respiratory conditioning and a staircase psychophysical procedure. Physiological studies on single saccular (auditory) nerve fibers under similar stimulus conditions helped characterize the dimensions of neural activity used in intensity discrimination. Incremental intensity difference limens (IDLs in dB) for 160-ms increments in continuous noise, 500-ms noise bursts, and 500-ms, 800-Hz tone bursts are 2 to 3 dB, are independent of overall level, and vary with signal duration according to a power function with a slope averaging - 0.33. Noise decrements are relatively poorly detected and the silent gap detection threshold is about 35 ms. The IDLs for increments and decrements in an 800-Hz continuous tone are about 0.13 dB, are independent of duration, and are level dependent. Unlike mammalian auditory nerve fibers, some goldfish saccular fibers show variation in recovery time to tonal increments and decrements, and adaptation to a zero rate. Unit responses to tone increments and decrements show rate effects generally in accord with previous observations on intracellular epsp's in goldfish saccular fibers. Neurophysiological correlates of psychophysical intensity discrimination data suggest the following: (1) noise gap detection may be based on spike rate increments which follow gap offset; (2) detection of increments and decrements in continuous tones may be determined by steep low-pass filtering in peripheral neural channels which enhance the effects of spectral "splatter" toward the lower frequencies; (3) IDLs for pulsed signals of different duration can be predicted from the slopes of rate-intensity functions and spike rate variability in individual auditory nerve fibers; and (4) at different sound pressure levels, different populations of peripheral fibers provide the information used in intensity discrimination.     

A computer model of medial efferent suppression in the mammalian auditory system

Stimulation of the olivocochlear bundle reduces basilar membrane displacement, driven auditory nerve activity, and compound action potential (CAP) response to acoustic stimulation. These effects were simulated using a computer model of the auditory periphery. The model simulates the medial efferent activity by attenuating the basilar membrane response. The model was evaluated against three animal studies reporting measurements at three levels of the auditory system; basilar membrane, single auditory nerve fibers and whole auditory nerve CAP. The CAP data included conditions where tones were masked by noise and "unmasked" by stimulation of the olivocochlear bundle. The model was able to simulate the data both qualitatively and quantitatively. As a consequence, it may be a suitable platform for studying the contribution of the efferent system to auditory processing of more complex auditory sounds in distracting backgrounds.     

Rate-intensity functions and their modification by broadband noise for neurons in the guinea pig inferior colliculus

Rate-intensity functions (RIFs) were generated in response to characteristic frequency (CF) tones presented alone and in the presence of broadband noise for neurons in the central nucleus of the inferior colliculus (IC) of the anesthetized guinea pig. Seventy-six percent of the RIFs to CF tones were monotonic (some showing incomplete saturation), and 24% were nonmonotonic. The RIFs to continuous noise were more nonmonotonic than those to CF tones. In continuous or gated noise, the dynamic portion of the RIF to a tone was shifted to a higher tone level, with little change in the dynamic range. Above a threshold noise level, the shift was a linear function of noise level with slope 0.97. Little shift occurred when the noise was inversely gated with respect to the tone burst, suggesting that the underlying mechanism is suppression rather than adaptation. For 63% of units, the maximum discharge rate to a tone in low levels (less than 0-dB spectrum level) of noise (including inversely gated) was greater than to the tone alone. Although many of the effects of noise in the IC reflect peripheral mechanisms, they are supplemented by centrally based processes which enhance the detectability of tone intensity increments in the presence of noise.     

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.     

Asymmetry of masking between complex tones and noise: the role of temporal structure and peripheral compression

Thresholds for the detection of harmonic complex tones in noise were measured as a function of masker level. The rms level of the masker ranged from 40 to 70 dB SPL in 10-dB steps. The tones had a fundamental frequency (F0) of 62.5 or 250 Hz, and components were added in either cosine or random phase. The complex tones and the noise were bandpass filtered into the same frequency region, from the tenth harmonic up to 5 kHz. In a different condition, the roles of masker and signal were reversed, keeping all other parameters the same; subjects had to detect the noise in the presence of a harmonic tone masker. In both conditions, the masker was either gated synchronously with the 700-ms signal, or it started 400 ms before and stopped 200 ms after the signal. The results showed a large asymmetry in the effectiveness of masking between the tones and noise. Even though signal and masker had the same bandwidth, the noise was a more effective masker than the complex tone. The degree of asymmetry depended on F0, component phase, and the level of the masker. The maximum difference between masked thresholds for tone and noise was about 28 dB; this occurred when the F0 was 62.5 Hz, the components were in cosine phase, and the masker level was 70 dB SPL. In most conditions, the growth-of-masking functions had slopes close to 1 (on a dB versus dB scale). However, for the cosine-phase tone masker with an F0 of 62.5 Hz, a 10-dB increase in masker level led to an increase in masked threshold of the noise of only 3.7 dB, on average. We suggest that the results for this condition are strongly affected by the active mechanism in the cochlea.     

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.     

Decision rules in detection of simple and complex tones

Detection of simple and complex tones in the presence of a 64-dB SPL uniformly masking noise was examined in two experiments. In both experiments, the signals were either pure tones (220, 1100, or 3850 Hz) or an 18-tone complex consisting of equally intense components between 110 and 7260 Hz. In experiment 1, psychometric functions were obtained for detection in a 2I, 2AFC task. Results for eight normal listeners show that the psychometric functions are parallel for simple and complex tones. As expected, the masked thresholds for the pure tones are 43-44 dB SPL independent of frequency; the masked threshold for the complex tone is about 37 dB SPL per tone. These results indicate that the simultaneous presence of signal energy in many auditory channels aids detection. In experiment 2, psychometric functions were obtained with all four signals presented in random order within a block of trials. Results for four normal listeners show that the psychometric functions are parallel to one another and to those obtained in experiment 1. The thresholds are elevated to about 46 dB for the pure tones and to 40.5 dB for the complex tone. These results are nearly, but not quite, consistent with a multiband energy-detector model using an optimum decision rule; it appears that listeners may only make an unweighted sum of decision variables across an optimum selection of channels.     

Temporal resolution and temporal integration of short pulses at the auditory periphery of echolocating animals

To explain the temporal integration and temporal resolution abilities revealed in echolocating animals by behavioral and electrophysiological experiments, the peripheral coding of sounds in the high-frequency auditory system of these animals is modeled. The stimuli are paired pulses similar to the echolocating signals of the animals. Their duration is comparable with or smaller than the time constants of the following processes: formation of the firing rate of the basilar membrane, formation of the receptor potentials of internal hair cells, and recovery of the excitability of spiral ganglion neurons. The models of auditory nerve fibers differ in spontaneous firing rate, response thresholds, and abilities to reproduce small variations of the stimulus level. The formation of the response to the second pulse of a pair of pulses in the multitude of synchronously excited high-frequency auditory nerve fibers may occur in only two ways. The first way defined as the stochastic mechanism implies the formation of the response to the second pulse as a result of the responses of the fibers that did not respond to the first pulse. This mechanism is based on the stochastic nature of the responses of auditory nerve fibers associated with the spontaneous firing rate. The second way, defined as the repeatition mechanism, implies the appearance of repeated responses in fibers that already responded to the first pulse but suffered a decrease in their response threshold after the first spike generation. This mechanism is based on the deterministic nature of the responses of fibers associated with refractoriness. The temporal resolution of pairs of short pulses, which, according to the data of behavioral experiments, is about 0.1–0.2 ms, is explained by the formation of the response to the second pulse through the stochastic mechanism. A complete recovery of the response to the second pulse, which, according to the data of electrophysiological studies of short-latency evoked brainstem potentials in dolphins, occurs within 5 ms, is explained by the formation of the response to the second pulse through the repetition mechanism. The time constant of temporal integration, which, according to the behavioral experiments at threshold levels of pulses, is about 0.2–0.3 ms, is explained by the integrating properties of internal hair cells, etc. It is shown that, at the high-frequency auditory periphery, the temporal integration imposes no limitations on the temporal resolution, because both integration and resolution are different characteristics of the same multiple response of synchronously excited fibers.     

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

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

Masking patterns in the bullfrog (Rana catesbeiana). II: Physiological effects

Responses of individual eighth-nerve fibers in the bullfrog (Rana catesbeiana) were measured to tone bursts at best frequency against a background of continuous, broadband masking noise. These data were used to calculate critical masking ratios to describe the fibers' responses to tones embedded in noise. In the frequency response range of the amphibian papilla (100-1000 Hz), critical ratios increase with tone frequency. Critical ratios of basilar papilla fibers (1000-2000 Hz) are generally higher than those of amphibian papilla fibers. Critical ratios are also significantly related to fiber threshold such that fibers with high thresholds, regardless of their best frequencies, have higher critical ratios and are thus less selective to signals embedded in noise. Critical ratios based on neural responses show a somewhat different frequency-dependent trend than do critical ratios based on psychophysical data presented previously for this species [A. M. Simmons, J. Acoust. Soc. Am. 83, 1087-1092 (1988a)]. In addition, these neural critical ratios do not appear to be level independent, as are psychophysical critical ratios. The data suggest that frequency selectivity of hearing in the bullfrog as measured behaviorally is probably not mediated solely by spectral filtering in the auditory periphery.     

Louder sounds can produce less forward masking: effects of component phase in complex tones

The influence of the degree of envelope modulation and periodicity on the loudness and effectiveness of sounds as forward maskers was investigated. In the first experiment, listeners matched the loudness of complex tones and noise. The tones had a fundamental frequency (F0) of 62.5 or 250 Hz and were filtered into a frequency range from the 10th harmonic to 5000 Hz. The Gaussian noise was filtered in the same way. The components of the complex tones were added either in cosine phase (CPH), giving a large crest factor, or in random phase (RPH), giving a smaller crest factor. For each F0, subjects matched the loudness between all possible stimulus pairs. Six different levels of the fixed stimulus were used, ranging from about 30 dB SPL to about 80 dB SPL in 10-dB steps. Results showed that, at a given overall level, the CPH and the RPH tones were louder than the noise, and that the CPH tone was louder than the RPH tone. The difference in loudness was larger at medium than at low levels and was only slightly reduced by the addition of a noise intended to mask combination tones. The differences in loudness were slightly smaller for the higher than for the lower F0. In the second experiment, the stimuli with the lower F0s were used as forward maskers of a 20-ms sinusoid, presented at various frequencies within the spectral range of the maskers. Results showed that the CPH tone was the least effective forward masker, even though it was the loudest. The differences in effectiveness as forward maskers depended on masker level and signal frequency; in order to produce equal masking, the level of the CPH tone had to be up to 35 dB above that of the RPH tone and the noise. The implications of these results for models of loudness are discussed and a model is presented based on neural activity patterns in the auditory nerve; this predicts the general pattern of loudness matches. It is suggested that the effects observed in the experiments may have been influenced by two factors: cochlear compression and suppression.     

Physiological responses to the pulsation threshold paradigm. II: Representations of high-pass noise in average rate measures of auditory-nerve fiber discharge

In a companion article [L. I. Hellstrom, J. Acoust. Soc. Am. 85, 230-242 (1989)], it was shown that psychophysical pulsation threshold masking patterns (PTPs) for high-pass noise maskers are not a simple transformation of the profile of activity evoked in the auditory nerve by the masker. In this article, PTPs are compared with neural representations in which interactions of masker and probe are considered. It is hypothesized that, at pulsation threshold, some criterion value of rate change occurs when the stimulus switches from masker to probe. The iso-rate probe level, defined for single auditory-nerve fibers, is the probe level at which this rate change is zero. Iso-rate probe levels are lowest when probe frequency equals best frequency (BF) of the fiber. Profiles of iso-rate probe level versus BF (equal to probe frequency) are qualitatively similar to PTPs but differ quantitatively, e.g., in the rate of growth of probe level with masker level (1.2 dB/dB for PTPs, 0.54 dB/dB for iso-rate profiles). Quantitative differences can be further reduced by requiring a positive rate criterion. These results suggest that PTPs are not solely a reflection of the internal representation of the masker, but reflect responses to the probe tone as well.     

Auditory stream segregation in monkey auditory cortex: effects of frequency separation, presentation rate, and tone duration

Auditory stream segregation refers to the organization of sequential sounds into "perceptual streams" reflecting individual environmental sound sources. In the present study, sequences of alternating high and low tones, "...ABAB...," similar to those used in psychoacoustic experiments on stream segregation, were presented to awake monkeys while neural activity was recorded in primary auditory cortex (A1). Tone frequency separation (AF), tone presentation rate (PR), and tone duration (TD) were systematically varied to examine whether neural responses correlate with effects of these variables on perceptual stream segregation. "A" tones were fixed at the best frequency of the recording site, while "B" tones were displaced in frequency from "A" tones by an amount = delta F. As PR increased, "B" tone responses decreased in amplitude to a greater extent than "A" tone responses, yielding neural response patterns dominated by "A" tone responses occurring at half the alternation rate. Increasing TD facilitated the differential attenuation of "B" tone responses. These findings parallel psychoacoustic data and suggest a physiological model of stream segregation whereby increasing delta F, PR, or TD enhances spatial differentiation of "A" tone and "B" tone responses along the tonotopic map in A1.     

Effects of peripheral nonlinearity on psychometric functions for forward-masked tones

Psychometric functions (PFs) for forward-masked tones were obtained for conditions in which signal level was varied to estimate threshold at several masker levels (variable-signal condition), and in which masker level was varied to estimate threshold at several signal levels (variable-masker condition). The changes in PF slope across combinations of masker frequency, masker level, and signal delay were explored in three experiments. In experiment 1, a 2-kHz, 10-ms tone was masked by a 50, 70 or 90 dB SPL, 20-ms on-frequency forward masker, with signal delays of 2, 20, or 40 ms, in a variable-signal condition. PF slopes decreased in conditions where signal threshold was high. In experiments 2 and 3, the signal was a 4-kHz, 10-ms tone, and the masker was either a 4- or 2.4-kHz, 200-ms tone. In experiment 2, on-frequency maskers were presented at 30 to 90 dB SPL in 10-dB steps and off-frequency maskers were presented at 60 to 90 dB SPL in 10-dB steps, with signal delays of 0, 10, or 30 ms, in a variable-signal condition. PF slopes decreased as signal level increased, and this trend was similar for on- and off-frequency maskers. In experiment 3, variable-masker conditions with on- and off-frequency maskers and 0-ms signal delay were presented. In general, the results were consistent with the hypothesis that peripheral nonlinearity is reflected in the PF slopes. The data also indicate that masker level plays a role independent of signal level, an effect that could be accounted for by assuming greater internal noise at higher stimulus levels.     

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.     

Molecular analysis of the effect of relative tone level on multitone pattern discrimination

Molecular psychophysics attempts to model the observer's response to stimuli as they vary from trial to trial. The approach has gained popularity in multitone pattern discrimination studies as a means of estimating the relative reliance or decision weight listeners give to different tones in the pattern. Various factors affecting decision weights have been examined, but one largely ignored is the relative level of tones in the pattern. In the present study listeners detected a level-increment in a sequence of 5, 100-ms, 2.0-kHz tone bursts alternating in level between 40 and 80 dB SPL. The level increment was made largest on the 40-dB tones, yet despite this all four highly-practiced listeners gave near exclusive weight to the 80-dB tones. The effect was the same when the tones were replaced by bursts of broadband Gaussian noise alternating in level. It was reduced only when the level differences were made <10 dB, and it was entirely reversed only when the low-level tones alternated with louder bursts of Gaussian noise. The results are discussed in terms of the effects of both sensory and perceptual factors on estimates of decision weights.     

Signal transmission in noisy environments: auditory masking in the tympanic nerve of the bushcricket Metaballus litus (Orthoptera: Tettigoniinae)

Physiological responses of the auditory leg nerve were recorded in the tettigoniid Metaballus litus to suprathreshold tone pulses of 12.45 kHz, which is close to the carrier frequency of the male's call. This stimulus tone frequency was determined by characterizing the polar response of the foreleg. Physiological threshold of the receptors was calculated from intensity input/output curves, and the experimental stimulus was set at 40 dB above this threshold value. There was low variance in threshold values between preparations. Continuous octave filtered white noise centered on the stimulus frequency was presented at the same time as the tone pulse at increasing intensities. The summed action potentials (SAPs) of the whole leg nerve were averaged over 256 stimulus presentations and the magnitude of the response was calibrated to dB values. The range of noise levels was set between that inducing no decrease in the SAP response to the tone pulse stimulus, up to a masking intensity where the response to the tone pulse was only just observable. Decrement in SAP magnitude was linear, and complete masking occurred when the noise level was 20-25 dB above the initial level of zero masking. This final level was comparable in magnitude to the sound-pressure level of the tone pulse and within the natural range of the insect's auditory behavior. Following the cessation of the noise signal, the SAPs were monitored over intervals of 2 min until the SAP asymptoted to the preexperimental condition. The reduction in SAP magnitude during noise presentation was attributed to a loss in synchrony from the individual tympanic receptors.     

The attention filter for tones in noise has the same shape and effective bandwidth in the elderly as it has in young listeners

Listeners asked to detect tones masked by noise hear frequent signals but miss infrequent probes, suggesting that they attend to spectral regions where they expect the signals to occur. The narrow detection pattern centered on the frequent target approximates that obtained in notched noise, indicating that attention is focused on the auditory filter. We measured attention bands in young and elderly listeners (n=5, 4; 20-25 and 62-82 years of age) for targets (800 or 1200 Hz) and infrequent probe signals (target +/-25-100 Hz) masked in wideband noise. We anticipated that their width would increase with age, as has been reported for auditory filters. A yes-no single-interval procedure provided detection probabilities and detection response speeds. Both measures showed near-linear declines with decreasing signal level, and graded decay functions as probe frequency deviated from the target frequency. Probes deviating from the target by 25 to 50 Hz were equivalent to a 2-dB reduction in signal level for both measures. The equivalent rectangular bandwidth (ERB) for detection approximated 11% of the signal frequency for each age group. Confidence intervals (95%) showed that the elderly ERB could be at most only about 20% larger than that of younger listeners.     

Auditory thresholds in a sound-producing electric fish (Pollimyrus): behavioral measurements of sensitivity to tones and click trains

In this report we present the first behavioral measurements of auditory sensitivity for Pollimyrus adspersus. Pollimyrus is an electric fish (Mormyridae) that uses both electric and acoustic signals for communication. Tone detection was assessed from the fish's electric organ discharge rate. Suprathreshold tones usually evoked an accelerated rate in naive animals. This response (rate modulation > or =25%) was maintained in a classical conditioning paradigm by presenting a weak electric current near the offset of 3.5-s tone bursts. An adaptive staircase procedure was used to find detection thresholds at frequencies between 100 and 1700 Hz. The mean audiogram from six individuals revealed high sensitivity in the 200-900 Hz range, with the best thresholds near 500 Hz (66.5+/-4.2 SE dB re: 1 microPa). Sensitivity declined slowly (about 20 dB/octave) above and below this sensitivity maximum. Sensitivity fell off rapidly above 1 kHz (about 60 dB/octave) and no responses were observed at 5 kHz. This behavioral sensitivity matched closely the spectral content of the sounds that this species produced during courtship. Experiments with click trains showed that sensitivity (about 83-dB peak) was independent of inter-click-interval, within the 10-100 ms range.