Latency of auditory brain-stem responses and otoacoustic emissions using tone-burst stimuli

A comparison of the latency of auditory brain-stem responses (ABR) and evoked otoacoustic emissions (EOAE) has led to an interpretation for the travel of transients in the peripheral auditory system that is consistent with both sets of data. The "cochlear echo" theory for the origin of the EOAE indicates that the latency of a particular frequency component back to the ear canal should be twice the forward latency of its characteristic place in the cochlea. The latency of wave V of the ABR to tone-burst stimuli can be described as the sum of two components: (1) a component that varies with intensity and frequency in an orderly and predictable manner and (2) a component that is independent of both intensity and frequency. Because the EOAE data can be predicted by taking twice the value of component (1) of the ABR latency, this component is interpreted to be due to mechanical travel through the cochlea. A consequence of this interpretation is that the remaining neural component of the ABR latency must be relatively independent of frequency and intensity.     

Adaptation and recovery from adaptation in single fiber responses of the cat auditory nerve.

This study examined the time course of adaptation and recovery from adaptation of single auditory-nerve fiber responses. The conditions studied were: (1) adaptation response using low level, 800 Hz or characteristic frequency (CF) stimuli; and (2) onset recovery and whole tone response recovery of a probe tone following a masker of equal frequency with variable silent intervals between the masker offset and probe onset. Single unit responses to 290 ms long, 800 Hz or CF tones presented at 10-30 dB SL were recorded from the auditory nerve of the cat. Adaptation properties were determined and fit to the equation: A(tp) = Yre(-tp/tau Rr) + Yse(-tp/tau Rs) + Ass. Recovery from adaptation was determined by recording the response of a probe tone following a 100-ms masker tone equal in frequency to the probe, and with amplitudes ranging from 20- to 30-dB relative to the probe amplitude. Both the onset recovery and the whole tone recovery were determined for the single unit responses. The onset data were analyzed and fit to either the equation: A (delta xt,tp) = Ass - Yre(-tp/tau Rr) - Yse(- delta t/tau Rs) or A (delta t,tp) = Ass - Yre(- delta t/tau R). The whole tone response showed two distinctive time patterns that could be fit to either an adaptation equation or to the two-time-constant recovery equation, depending on the relative amplitude of the masker and the length of the silent interval between masker offset and probe onset. The results of this study indicate that single fiber time constants are comparable to those measured in previous studies using the auditory-nerve neurophonic (ANN). Likewise, the pattern of recovery of the whole tone response for single fiber responses is comparable to the ANN. Possible sites and mechanisms for adaptation and recovery from adaptation taking into account recent data from electrical stimulation studies and receptor channel morphology and kinetics are discussed.     

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)     

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.     

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.     

An auditory-periphery model of the effects of acoustic trauma on auditory nerve responses

Acoustic trauma degrades the auditory nerve's tonotopic representation of acoustic stimuli. Recent physiological studies have quantified the degradation in responses to the vowel /E/ and have investigated amplification schemes designed to restore a more correct tonotopic representation than is achieved with conventional hearing aids. However, it is difficult from the data to quantify how much different aspects of the cochlear pathology contribute to the impaired responses. Furthermore, extensive experimental testing of potential hearing aids is infeasible. Here, both of these concerns are addressed by developing models of the normal and impaired auditory peripheries that are tested against a wide range of physiological data. The effects of both outer and inner hair cell status on model predictions of the vowel data were investigated. The modeling results indicate that impairment of both outer and inner hair cells contribute to degradation in the tonotopic representation of the formant frequencies in the auditory nerve. Additionally, the model is able to predict the effects of frequency-shaping amplification on auditory nerve responses, indicating the model's potential suitability for more rapid development and testing of hearing aid schemes.     

Human brain-stem auditory evoked responses obtained by cross correlation to trains of clicks, noise bursts, and tone bursts

Brain-stem auditory evoked responses (BAERs) were obtained in eight normal-hearing young adults. Stimuli included clicks, noise bursts, and tone bursts. Tone bursts included carrier frequencies of 1, 2, 4, and 8 kHz. All stimuli were presented at 60 dB nHL. BAERs were obtained by presenting stimuli in pseudorandom trains, called maximum length sequences (MLSs). BAERs were recovered by cross correlating the responses with a recovery sequence. MLS-BAERs were obtained with minimum pulse intervals (MPIs) of 6, 4, and 2 ms. Conventional BAERs were also obtained for stimuli presented at a rate of 30 Hz. BAERs were obtained for all stimuli, for both the conventional averaging technique and for the cross-correlation technique. BAERs were observed for MPIs as short as 2 ms for all stimuli. Wave V was the only peak consistently identifiable for these stimuli. For all stimuli, wave V latency increased and wave V amplitude decreased with decreasing MPI. This is the first demonstration of the use of maximum length sequences combined with cross correlation to obtain BAERs to noise burst and tone burst stimuli.     

Suppression of auditory nerve responses. II. Suppression threshold and growth, iso-suppression contours

Two-tone "synchrony suppression" was studied in responses of single auditory nerve fibers recorded from anesthetized cats. Suppression thresholds for suppressor tones set to a fiber's characteristic frequency (CF) were approximately equal to discharge rate thresholds for CF tones. Suppression thresholds above and below CF were usually lower than the corresponding discharge rate thresholds. However, at all frequencies studied (including CF), suppression thresholds were higher than the corresponding thresholds for discharge synchronization. Across fibers, rates of suppression growth for suppressors at CF were greatest in low-CF fibers and least in high-CF fibers, and there was a systematic decrease in suppression growth rate at CF as CF increased. Within fibers, rates of suppression growth above CF were typically less than at CF, and slopes were monotonically decreasing functions of frequency. Within-fiber rates of suppression growth below CF were variable, but they usually were greater than rates of growth at CF. Iso-suppression contours (frequencies and intensities producing criterion amounts of suppression) indicated that tones near CF are the most potent suppressors at near-threshold intensities, and that the frequency producing the most suppression usually shifts downward as the amount of suppression increases. These data support the notion that synchrony suppression arises primarily as a passive consequence of hair cell activation.     

Stimulus dependencies of the gerbil brain-stem auditory-evoked response (BAER). III: Additivity of click level and rate with noise level

Two experiments were performed that evaluated the effects of ipsilateral-direct broadband noise maskers on the gerbil brain-stem auditory-evoked response (BAER) to click stimuli. In experiment 1, clicks were presented at 27 Hz at levels including 70, 80, 90, and 100 dB pSPL. Noise conditions included a no-noise control, and included noise levels varying in 10-dB increments from 20 dB SPL to a maximum noise level of 50, 60, 70, and 80 dB SPL for click levels of 70, 80, 90, and 100 dB pSPL, respectively. Gerbil BAER peaks were labeled with small roman numerals to distinguish them from human BAER peaks. The dependent variables included waves i and v latencies and amplitudes. Peak latencies increased and peak amplitudes decreased with decreasing click level and increasing noise level. To a first approximation, peak latencies and amplitudes showed changes with increasing noise level that were similar across click level. With increasing click level, there was little or no effect on the i-v interval. There was an increase in the i-v interval with increasing noise level. In experiment 2, click level was held constant at 90 dB pSPL, and click rates included 15, 40, 65, and 90 Hz. For each click rate, noise conditions included a no-noise control, and noise levels included 20, 30, 40, 50, 60, and 70 dB SPL. With increasing click rate and noise level, there was an increase in peak latencies, an increase in the i-v interval, and a decrease in peak amplitudes. The magnitude of peak latency and amplitude shifts with increasing click rate was dependent on noise level. Specifically, the magnitude of rate-dependent changes decreased with increasing level of broadband noise. These data are compared to human BAER experiments, and are found to be in fundamental agreement.     

Stimulus dependencies of the gerbil brain-stem auditory-evoked response (BAER). I: Effects of click level, rate, and polarity

Three experiments evaluating the effects of various stimulus manipulations on the click-evoked gerbil brain-stem auditory-evoked response (BAER) are reported. In experiment 1, click polarity and level were covaried. With increasing click level, there is a parallel decrease in the latency of the first five BAER peaks (i-v) and an increase in BAER peak amplitudes. Mean wave i amplitude was greater for rarefaction than condensation clicks at high click levels; mean wave v amplitude was greater for condensation clicks at higher click levels. Experiment 2 covaried click rate and polarity. The latency of the BAER peaks increased with increasing click repetition rate. This rate-dependent latency increase was greater for the later BAER peaks, resulting in an increase in the i-v interval with increasing click rate. As rate increased, the amplitudes of waves i and v decreased monotonically, whereas the amplitudes of waves ii-iv were largely uninfluenced by click rate. As in experiment 1, mean wave i amplitude was greater for rarefaction clicks, whereas mean wave v amplitude was greater for condensation clicks. The magnitude of these polarity dependencies on waves i and v amplitude decreased with increasing click rate. Experiment 3 evaluated the effects of click polarity on BAERs to high-intensity (100 dB pSPL) clicks presented at a rate of 10 Hz. In eight of ten gerbils evaluated, wave i amplitude was greater to rarefaction clicks, and, in all ten animals, wave v amplitude was greater to condensation clicks. The effects of click level and rate on BAER peak amplitudes, latencies, and interwave intervals are reminiscent of stimulus dependencies reported for the human BAER. The effects of click polarity on the amplitudes of waves i and v of the gerbil BAER have also been reported for the human BAER.     

Representation of the vowel /epsilon/ in normal and impaired auditory nerve fibers: model predictions of responses in cats

The temporal response of auditory-nerve (AN) fibers to a steady-state vowel is investigated using a computational auditory-periphery model. The model predictions are validated against a wide range of physiological data for both normal and impaired fibers in cats. The model incorporates two parallel filter paths, component 1 (C1) and component 2 (C2), which correspond to the active and passive modes of basilar membrane vibration, respectively, in the cochlea. The outputs of the two filters are subsequently transduced by two separate functions, added together, and then low-pass filtered by the inner hair cell (IHC) membrane, which is followed by the IHC-AN synapse and discharge generator. The C1 response dominates at low and moderate levels and is responsible for synchrony capture and multiformant responses seen in the vowel responses. The C2 response dominates at high levels and contributes to the loss of synchrony capture observed in normal and impaired fibers. The interaction between C1 and C2 responses explains the behavior of AN fibers in the transition region, which is characterized by two important observations in the vowel responses: First, all components of the vowel undergo the C1/C2 transition simultaneously, and second, the responses to the nonformant components of the vowel become substantial.     

Stochastic properties of cat auditory nerve responses to electric and acoustic stimuli and application to intensity discrimination

Statistical properties of electrically stimulated (ES) and acoustically stimulated (AS) auditory nerve fiber responses were assessed in undeafened and short-term deafened cats, and a detection theory approach was used to determine fibers' abilities to signal intensity changes. ES responses differed from AS responses in several ways. Rate-level functions were an order of magnitude steeper, and discharge rate normally saturated at the stimulus pulse rate. Dynamic ranges were typically 1-4 dB for 200 pps signals, as compared with 15-30 dB for AS signals at CF, and they increased with pulse rate without improving threshold or changing absolute rate-level function slopes. For both ES and AS responses, variability of spike counts elicited by repeated trials increased with level in accord with Poisson-process predictions until the discharge rate exceeded 20-40 spikes/s. AS variability continued increasing monotonically at higher discharge rates, but more slowly. In contrast, maximum ES variability was usually attained at 100 spikes/s, and at higher discharge rates variability reached a plateau that was either maintained or decreased slightly until discharge rate approached the stimulus pulse rate. Variability then decreased to zero as each pulse elicited a spike. Increasing pulse rate did not substantially affect variability for rates up to 800 pps; rather, higher pulse rates simply extended the plateau region. Spike count variability was unusually high for some ES fibers. This was traced to response nonstationarities that stemmed from two sources, namely level-dependent fluctuations in excitability that occurred at 1-3 s intervals and, for responses to high-rate, high-intensity signals, fatigue that arose when fibers discharged at their maximum possible rates. Intensity discrimination performance was assessed using spike count as the decision variable in a simulated 2IFC task. Neurometric functions (percent correct versus intensity difference) were obtained at several levels of the standard (I), and the intensity difference (delta I) necessary for 70% correct responses was estimated. AS Weber fractions (10 log delta I/I) averaged +0.2 dB (delta IdB = 3.1 dB) for 50 ms tones at CF. ES Weber fractions averaged -12.8 dB (delta IdB = 0.23 dB) for 50 ms, 200 pps signals, and performance was approximately constant between 100 and 1000 pps. Intensity discrimination by single cells in ES conditions paralleled human psychophysical performance for similar signals. High ES sensitivity to intensity changes arose primarily from steeper rate-level functions and secondarily from reduced spike count variability.     

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.     

Stimulus dependencies of the gerbil brain-stem auditory-evoked response (BAER). II: Effects of broadband noise level and high-pass masker cutoff frequency across click polarity

Two experiments concerning the effects of masking noise on the gerbil brain-stem auditory-evoked response (BAER) are reported. Experiment 1 evaluated the effects of broadband masking noise on the BAER obtained to condensation and rarefaction clicks. With increasing noise level, there was an increase in BAER peak latencies, an increase in the i-v interval, and a decrease in peak amplitudes. Experiment 2 evaluated the effects of high-pass masking noise on the BAER obtained to condensation and rarefaction clicks. Both high-pass responses and derived-band responses were evaluated. For high-pass responses, with decreasing masker cutoff frequency, there was an increase in BAER peak latencies, a decrease in the i-v interval, and a decrease in peak amplitudes. For derived-band responses, with decreasing derived-band frequency, there was an increase in peak latencies and a decrease in the i-v interval. A comparison of wave i and wave v amplitudes across derived-band frequency demonstrates a greater contribution of high-frequency cochlear regions to wave i than wave v. Small, insignificant, effects of click polarity on BAER peak amplitudes were observed. These trends were in the direction seen in a companion paper [R. Burkard and H. F. Voigt, J. Acoust. Soc. Am. 85, 2514-2525 (1989)] and were, in general, reduced by the presence of broadband or high-pass maskers.     

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.     

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.     

Encoding of steady-state vowels in the auditory nerve: representation in terms of discharge rate

Responses of large populations of auditory-nerve fibers to synthesized steady-state vowels were recorded in anesthetized cats. Driven discharge rate to vowels, normalized by dividing by saturation rate (estimated from the driven rate to CF tones 50 dB above threshold), was plotted versus fiber CF for a number of vowel levels. For the vowels /I/ and /e/, such rate profiles showed a peak in the region of the first formant and another in the region of the second and third formants, for sound levels below about 70 dB SPL. For /a/ at levels below about 40 dB SPL there are peaks in the region of the first and second formants. At higher levels these peaks disappear for all the vowels because of a combination of rate saturation and two-tone suppression. This must be qualified by saying that rate profiles plotted separately for units with spontaneous rates less than one spike per second may retain peaks at higher levels. Rate versus level functions for units with CFs above the first formant can saturate at rates less than the saturation rate to CF to-es or they can be nonmonotonic; these effects are most likely produced by the same mechanism as that involved in two-tone suppression.     

Auditory brainstem responses in the Eastern Screech Owl: an estimate of auditory thresholds

The auditory brainstem response (ABR), a measure of neural synchrony, was used to estimate auditory sensitivity in the eastern screech owl (Megascops asio). The typical screech owl ABR waveform showed two to three prominent peaks occurring within 5 ms of stimulus onset. As sound pressure levels increased, the ABR peak amplitude increased and latency decreased. With an increasing stimulus presentation rate, ABR peak amplitude decreased and latency increased. Generally, changes in the ABR waveform to stimulus intensity and repetition rate are consistent with the pattern found in several avian families. The ABR audiogram shows that screech owls hear best between 1.5 and 6.4 kHz with the most acute sensitivity between 4-5.7 kHz. The shape of the average screech owl ABR audiogram is similar to the shape of the behaviorally measured audiogram of the barn owl, except at the highest frequencies. Our data also show differences in overall auditory sensitivity between the color morphs of screech owls.     

Cochlear microphonics and the initiation of spikes in the auditory nerve: correlation of single-unit data with neural and receptor potentials recorded from the round window

On the basis of comparisons of responses of guinea pig ganglion cells and inner hair cells to intense low-frequency tones, Sellick et al. [Hear. Res. 7, 199-221 (1982)] have proposed that basal inner hair cells can be depolarized (and thus, VIII-N. spikes generated) by the extracellular microphonic generated during hyperpolarization of outer hair cells. VIII-N. data for the chinchilla have been presented that, to a first approximation, support such a hypothesis [Ruggero and Rich, J. Acoust. Soc. Am. 73, 2096-2108 (1983)]. However, an apparent discrepancy exists in our results, vis à vis Sellick et al.'s hypothesis, in that basal fiber near-threshold responses precede maximal negativity of the round window microphonic (i.e., maximal hyperpolarization of outer hair cells) by up to 90 degrees (but generally less than 45 degrees), depending on frequency. It is shown here that the discrepancy is resolved if certain nonlinear phase changes and overall distortion of the microphonic waveshapes, both of which occur at intense stimulus levels, are taken into account. It is also shown that compound action potentials (AP's), superimposed on the round window microphonics, can be identified at multiple times within each stimulus cycle, closely matching the near-threshold response phases of single-unit excitation. AP1 is nearly synchronous with the negative-to-positive transition of round window microphonics and with the excitation of fibers innervating apical-to-middle cochlear regions. AP2 is synchronous with the positive-to-negative transition of the microphonics and with the excitation of basal fibers. One or two other AP's probably reflect "peak splitting" in the responses of both basal and apical fibers.