Growth and recovery of temporary threshold shift at 3 kHz in bottlenose dolphins: experimental data and mathematical models

Measurements of temporary threshold shift (TTS) in marine mammals have become important components in developing safe exposure guidelines for animals exposed to intense human-generated underwater noise; however, existing marine mammal TTS data are somewhat limited in that they have typically induced small amounts of TTS. This paper presents experimental data for the growth and recovery of larger amounts of TTS (up to 23 dB) in two bottlenose dolphins (Tursiops truncatus). Exposures consisted of 3-kHz tones with durations from 4 to 128 s and sound pressure levels from 100 to 200 dB re 1 μPa. The resulting TTS data were combined with existing data from two additional dolphins to develop mathematical models for the growth and recovery of TTS. TTS growth was modeled as the product of functions of exposure duration and sound pressure level. TTS recovery was modeled using a double exponential function of the TTS at 4-min post-exposure and the recovery time.     

Temporary threshold shift in bottlenose dolphins (Tursiops truncatus) exposed to mid-frequency tones

A behavioral response paradigm was used to measure hearing thresholds in bottlenose dolphins before and after exposure to 3 kHz tones with sound exposure levels (SELs) from 100 to 203 dB re 1 microPa2 s. Experiments were conducted in a relatively quiet pool with ambient noise levels below 55 dB re 1 microPa2/Hz at frequencies above 1 kHz. Experiments 1 and 2 featured 1-s exposures with hearing tested at 4.5 and 3 kHz, respectively. Experiment 3 featured 2-, 4-, and 8-s exposures with hearing tested at 4.5 kHz. For experiment 2, there were no significant differences between control and exposure sessions. For experiments 1 and 3, exposures with SEL=197 dB re 1 microPa2 s and SEL > or = 195 dB re 1 microPa2 s, respectively, resulted in significantly higher TTS4 than control sessions. For experiment 3 at SEL= 195 dB re 1 microPa2 s, the mean TTS4 was 2.8 dB. These data are consistent with prior studies of TTS in dolphins exposed to pure tones and octave band noise and suggest that a SEL of 195 dB re 1 microPa2 s is a reasonable threshold for the onset of TTS in dolphins and white whales exposed to midfrequency tones.     

Squirrel monkey temporary threshold shift from 48-h exposures to low-frequency noise.

Five squirrel monkeys were exposed for 1, 2, 4, 8, 16, 24, and 48 h to a 375--750-Hz band noise at an overall SPL of 95 dB. The TTS4.5 growth pattern for the 750-Hz test frequency was biphasic and did not reach an asymptote after 48 h of exposure. For all exposures, the mean thresholds of the five monkeys returned to within 5 dB of the preexposure mean 20 h after exposure. Recovery curves from all exposures at the 750-Hz test frequency appeared biphasic. Increasing SPL from 95 to 105 dB increased TTS4.5 by 4 dB at 750 Hz for a 1-h exposure. Recovery from the 105-dB exposure followed the same pattern as recovery from the 95-DB exposure. When compared with data collected from human subjects under similar conditions, these experiments indicate that the growth and recovery of TTS in squirrel monkeys are sufficiently similar to growth and recovery in man to justify further comparative investigation.     

Temporary threshold shift caused by combined steady-state and impulse noises

Temporary threshold shift (TTS) measurements on 11 subjects, resulting from exposures to steady-state noise, impulse noise, and combinations of both types of noise are reported. Twenty minute exposures to wide-band steady-state noise at levels of 78, 84, 90 and 96 dBA, and impulse noise at levels of 96, 102, 108, 114, 120, 126 and 132 dB(peak), and repetition rate of 3·2 pulses/s, were used. When a hazardous level of steady-state noise was combined with various levels of impulse noise, there was a significant reduction in the measured TTS at 4 and 6 kHz. This reduction was greatest when the peak level of the pulse exceeded the r.m.s. level of the steady-state noise by 6–18 dB. When a hazardous level of the pulse was combined with several levels of steady-state noise, no significant reduction in TTS was observed. These findings are interpreted as a result of acoustic reflex stimulation; the pulses superimposed on a hazardous steady-state noise continually re-activated the reflex and prevented fatigue. The converse did not apply, that is, the non-hazardous (but high-level) steady-state noise did not appear to counteract fatigue resulting from hazardous impulse noise.     

Pitch shift of pure and complex tones induced by masking noise

Psychoacoustic experiments were performed to measure the pitch-shift effects of pure and complex tones resulting from the addition of a masking noise to the tonal stimuli. Harmonic residue tones with either two or three harmonics and a fundamental frequency of 200 Hz were chosen as test tones. The pitch shifts of virtual and spectral pitches of the residue tones were measured as a function of the intensity of a low-pass noise with 600-Hz cutoff frequency. The SPL of this noise varied between 30 and 70 dB. In another experiment, the pitch shifts of single pure tones corresponding to the frequencies and SPLs of the harmonics of the residue tones were measured using the same masking noise. The results from five subjects for the harmonic residue tones show only a weak dependence of pitch shift on masking noise intensity. This dependence exists for both spectral and virtual pitches. In the case of single pure tones, pitch shift depends more distinctly on noise intensity. Pitch shifts of up to 5% were found in the range of noise intensity investigated. The magnitude of pitch shift shows pronounced interindividual differences, but the direction of the shift effect is always the same. In all cases pitch increases with higher masking noise levels.     

Overexposure effects of a 1-kHz tone on the distortion product otoacoustic emission in humans

The effects of overexposure on the properties of distortion product otoacoustic emissions (DPOAEs) are investigated. In total, 39 normal-hearing humans were monaurally exposed to a 1-kHz tone lasting for 3 min at an equivalent threshold sound-pressure level of 105.5 dB. The effects of overexposure were studied in two experiments (1) on the broadband DPOAE and (2) on the DPOAE fine structure, measured using a higher frequency resolution in a narrower frequency range. The obtained DPOAE shifts were compared to temporary threshold shift (TTS) obtained after a similar exposure. Similarities between DPOAE shifts and TTS were found in the affected frequency range and the time course of recovery. The amount of TTS was higher in the early recovery time (1-4-min postexposure), but similar to the DPOAE shift (even in absolute terms) at later recovery times (5-20-min postexposure). The DPOAE fine structure was not systematically changed after the exposure.     

Repeated TTS exposures in monkeys: alterations in hearing, cochlear structure, and single-unit thresholds

The findings of a number of studies investigating the effects of excessive sound on hearing have indicated that the correspondence between behavioral, physiological, and histological measures of noise-induced hearing loss may be markedly dependent upon the sensitivity of the particular measure. Recent studies demonstrating significant changes in the responses of single auditory neurons following brief exposures to pure tones suggest that single-unit activity may be a sensitive indicator of physiological insult to the organ of Corti's sensory cells. In addition, the long-lasting nature of the changes in neural responsiveness suggests that each temporary threshold shift (TTS) episode may produce an increment of damage to the ear that eventually contributes to a measurable permanent threshold shift (PTS). A logical extension of this implication is the proposal that repeated episodes of TTS would first affect single-unit thresholds, and that such damage would eventually manifest itself as PTS. A test of this notion was performed by repeatedly exposing monkeys to short-lasting TTS sounds for many months. Behavioral thresholds were monitored using a reaction-time task before and after each inducement of TTS. Two subjects participated in exposure sessions for 18 months, while the remaining monkey was exposed to identical stimuli for 6 months. At the end of behavioral testing, the monkeys were prepared for chronic recording from single cells of the cochlear nucleus. Following the recording period, cochleas were prepared for examination as plastic-embedded whole mounts. Flat preparations of the cochlear duct were made and the position and extent of damage to the organ of Corti and myelinated nerve fibers were determined. No elevations in behavioral threshold were noted for the monkey receiving 6 months of sound-exposure experience, while for both subjects exposed for 18 months, a significant high-frequency hearing loss became apparent during the final months of exposure. For damaged ears, the thresholds of ipsilateral cochlear nucleus units were elevated for characteristic frequencies (CFs) corresponding to the frequency regions where behavioral thresholds were shifted. Thresholds for units with high-frequency CFs in the animal exposed for 6 months also demonstrated a loss in sensitivity. Histological examination of the cochleas of monkeys with permanent hearing losses revealed corresponding damage to the high-frequency region of the organ of Corti. The monkey exposed for 6 months, which demonstrated only elevated unit thresholds, also had high-frequency lesions.(ABSTRACT TRUNCATED AT 400 WORDS)     

Onset, growth, and recovery of in-air temporary threshold shift in a California sea lion (Zalophus californianus)

A California sea lion (Zalophus californianus) was tested in a behavioral procedure to assess noise-induced temporary threshold shift (TTS) in air. Octave band fatiguing noise was varied in both duration (1.5-50 min) and level (94-133 dB re 20 muPa) to generate a variety of equal sound exposure level conditions. Hearing thresholds were measured at the center frequency of the noise (2500 Hz) before, immediately after, and 24 h following exposure. Threshold shifts generated from 192 exposures ranged up to 30 dB. Estimates of TTS onset [159 dB re (20 muPa)(2) s] and growth (2.5 dB of TTS per dB of noise increase) were determined using an exponential function. Recovery for threshold shifts greater than 20 dB followed an 8.8 dB per log(min) linear function. Repeated testing indicated possible permanent threshold shift at the test frequency, but a later audiogram revealed no shift at this frequency or higher. Sea lions appear to be equally susceptible to noise in air and in water, provided that the noise exposure levels are referenced to absolute sound detection thresholds in both media. These data provide a framework within which to consider effects arising from more intense and/or sustained exposures.     

Human temporary threshold shift (TTS) and damage risk

Information regarding the relation of human temporary threshold shift (TTS) to properties of steady-state and intermittent noise published since the 1966 appearance of the CHABA damage risk contours is reviewed. The review focuses on results from four investigative areas relevant to potential revision of the CHABA contours including effects of long-duration exposure and asymptotic threshold shifts (ATS); equivalent quiet and/or safe noise levels; effects of intermittency; and use of noise-induced temporary threshold shift (NITTS) to predict susceptibility to noise-induced permanent threshold shift (NIPTS). These data indicate that two of three major postulates on which the original contours were based are not valid. First, recovery from TTS is not independent of the conditions that produced the TTS as was assumed. Second, the assumption that all exposures that produce equal TTS2 are equally hazardous is not substantiated. The third postulate was that NIPTS produced by 10 years of daily exposure is approximately equal to the TTS2 produced by the same noise after an 8-h exposure. Based upon several TTS experiments showing that TTS reaches an asymptote after about 8 h of exposure, the third CHABA postulate can be reworded to state the hypothesis that ATS produced by sound of fixed level and spectrum represents an upper bound on PTS produced by that sound regardless of the exposure duration or the number of times exposed. This hypothesis has a strong, logical foundation if ATS represents a true asymptote for TTS, not a temporary plateau, and if threshold shifts do not increase after the noise exposure ceases.     

Towards a better understanding of temporary threshold shift of hearing

It was thought that temporary threshold shift of hearing due to exposure to noise might be more easily understood if the shifts were considered in terms of the rms pressure rather than in decibels. Therefore, the forms to be expected if the rate of shift of the pressure threshold were proportional to the difference between itself and the ultimate threshold were calculated and compared with a limited selection of published data. Good agreement with data for individuals during growth of TTS, and for one example of intermittent exposure to noise, was found and moderately good agreement with recovery. Agreement with averaged data, particularly for intermittent exposures, was poor, possibly because averaging the widely disparate figures obtained for individuals masks the true effects. It is also shown that the maximum ultimate TTS due to exposure to noise may be simply related to the mean square pressure of that noise.Further consideration of the mass of published work is needed, but this study suggests that at least some facets of TTS can be simply described in terms of exponential pressure shifts.     

Vowel amplitude variation associated with the heart cycle

Modulation of the acoustic amplitude of a sustained vowel across the cardiac (ECG) cycle was examined by signal-averaging techniques. Ten normal men prolonged [a] at a comfortable Fo maintained within three SPL ranges: 60-68, 70-78, and 80-88 dB. Peak-to-peak amplitude variation associated with the heart cycle averaged 8.5% (s.d. = 5.4) re: mean, varying from about 14% at low SPLs to approximately 3% at high SPLs. The amplitude modulation was estimated to account for 11.8% of the measured short-term amplitude perturbation (shimmer), ranging from about 5% to almost 22% for individual samples. The mean deterministic shimmer (Sd) was 0.036 dB (s.d. = 0.019), with a trend toward decreasing Sd with increasing SPL. Additionally, fundamental frequency variation across the heart cycle within these phonations was comparable to that observed by Orlikoff and Baken [J. Acoust. Soc. Am. 85, 888-893 (1989)], and was shown to be uninfluenced by vocal SPL, although deterministic jitter (Jd) did decrease with vocal intensity. The results are discussed in terms of how the phonovascular relationship may affect the reliability and interpretation of acoustic shimmer measures.     

Sonar inter-ping noise field characterization during cetacean behavioral response studies off Southern California

The potential negative effects of sound, particularly active sonar, on marine mammals has received considerable attention in the past decade. Numerous behavioral response studies are ongoing around the world to examine such direct exposures. However, detailed aspects of the acoustic field (beyond simply exposure level) in the vicinity of sonar operations both during real operations and experimental exposures have not been regularly measured. For instance, while exposures are typically repeated and intermittent, there is likely a gradual decay of the intense sonar ping due to reverberation that has not been well described. However, it is expected that the sound field between successive sonar pings would exceed natural ambient noise within the sonar frequency band if there were no sonar activity. Such elevated sound field between the pings may provide cues to nearby marine mammals on source distances, thus influencing potential behavioral response. Therefore, a good understanding of the noise field in these contexts is important to address marine mammal behavioral response to MFAS exposure. Here we investigate characteristics of the sound field during a behavioral response study off California using drifting acoustic recording buoys. Acoustic data were collected before, during, and after playbacks of simulated mid-frequency active sonar (MFAS). An incremental computational method was developed to quantify the inter-ping sound field during MFAS transmissions. Additionally, comparisons were made between inter-ping sound field and natural background in three distinctive frequency bands: low-frequency (<3 kHz), MFA-frequency (3–4.5 kHz), and high-frequency (>4.5 kHz) bands. Results indicate significantly elevated sound pressure levels (SPLs) in the inter-ping interval of the MFA-frequency band compared to natural background levels before and after playbacks. No difference was observed between inter-ping SPLs and natural background levels in the low- and high-frequency bands. In addition, the duration of elevated inter-ping sound field depends on the MFAS source distance. At a distance of 900–1300 m from the source, inter-ping sound field at the exposure frequency is observed to remain 5 dB above natural background levels for approximately 15 s, or 65%, of the entire inter-ping interval. However, at a distance of 2000 m, the 5 dB elevation of the inter-ping SPLs lasted for just 7 s, or 30% of the inter-ping interval. The prolonged elevation of sound field beyond the brief sonar ping at such large distances is most likely due to volume reverberation of the marine environment, although multipath propagation may also contribute to this.     

Temporary threshold shift in human subject exposed to noise

Using an audiometer,the effect of the noise level upon temporarythreshold shift(TTS)for five trained normal subjects(left ear only)was studied.The measurements were carried out after 6 min exposure(in third octave band)for different sound pressure levels ranging between 75-105 dB at three test fre-quencies 2,3,and 4 kHz.The results indicated that at exposure to noise of soundpressure level(SPL)above 85 dB,TTS increases linearly with ths SPL for all thetest frequencies.The work had extended to study the recovery curves for the sameears.The results indicated that the reduction in TTS on doubling the recoverytimes,for the two sound pressure levels 95 dB and 105 dB,occurs at a rate of near-ly 3 dB.The comparison of the recovery curve at 3 kHz with that calculated usingWard's general equation for recovery was made.Finally,to study the values ofTTS produced by exposure to certain noise at different test frequencies,distribu-tion curves for two recovery times were plotted representing TTS values,for anexposure     

Hazard from intense low-frequency acoustic impulses

It was predicted that because the ear is spectrally tuned, it should be most affected by intense impulses with spectral peaks near the frequency where it is tuned best (3.0 kHz for the human ear) and progressively less affected by impulses at lower frequencies [G.R. Price, Scand. Audiol. Suppl. 16, 111-121 (1982)]. This prediction is counter to all the DRCs for impulse noise; therefore an adequate test is essential. In order to augment the data on hearing loss to low-spectral-frequency impulses, three groups of cats (eight, nine, and ten animals) were exposed on one occasion to 50 impulses from a 105-mm howitzer at peak SPLs of 153, 159, and 166 dB. Threshold shifts were measured electrophysiologically on the day of exposure (CTS) and following a 2-month recovery period (PTS). Maximum PTSs appeared at 4 kHz (even though the spectral peak of the impulse had been at about 100 Hz), and CTSs recovered into PTSs about half as large. Furthermore, for group data, even small CTSs tended to have a permanent component. These data raise the question as to whether or not any threshold shift persisting an hour or two after exposure to high levels should be considered tolerable. When compared with data from rifle fire exposures, the data confirmed the earlier prediction that as the spectral frequency drops, hazard declines at the rate of a little more than 3 dB/oct, contrary to the rating by existing DRCs.     

One source for distortion product otoacoustic emissions generated by low- and high-level primaries

Distortion product otoacoustic emissions (DPOAE) elicited by tones below 60-70 dB sound pressure level (SPL) are significantly more sensitive to cochlear insults. The vulnerable, low-level DPOAE have been associated with the postulated active cochlear process, whereas the relatively robust high-level DPOAE component has been attributed to the passive, nonlinear macromechanical properties of the cochlea. However, it is proposed that the differences in the vulnerability of DPOAEs to high and low SPLs is a natural consequence of the way the cochlea responds to high and low SPLs. An active process boosts the basilar membrane (BM) vibrations, which are attenuated when the active process is impaired. However, at high SPLs the contribution of the active process to BM vibration is small compared with the dominating passive mechanical properties of the BM. Consequently, reduction of active cochlear amplification will have greatest effect on BM vibrations and DPOAEs at low SPLs. To distinguish between the "two sources" and the "single source" hypotheses we analyzed the level dependence of the notch and corresponding phase discontinuity in plots of DPOAE magnitude and phase as functions of the level of the primaries. In experiments where furosemide was used to reduce cochlear amplification, an upward shift of the notch supports the conclusion that both the low- and high-level DPOAEs are generated by a single source, namely a nonlinear amplifier with saturating I/O characteristic.     

Intense sounds may alter the mechanical properties of the cochlear partition

Morphological and structural changes have been observed in various cochlear elements following exposure to intense sounds. Whether these changes are sufficient to locally alter the mechanical properties of the cochlear partition is unknown. Here a psychophysical test for mechanical changes in the partition is developed and applied. Monaural exposures to an intense 1700-Hz tone were preceded and followed by a binaural task in which the subject adjusted the interaural time difference of a 250-Hz tone in order to perceive it as centered inside his head. If the local gradients of mass, stiffness, and/or coupling were altered by the exposure, then the post-exposure settings should contain a time lead toward one ear or the other, depending upon the direction of the mechanical changes. Following exposure to an intense tone, the maximum temporary threshold shift (TTS) is often displaced upward in frequency from the exposure stimulus--an effect widely known as the half-octave shift in TTS. Time leads toward the nonexposed ear in the post-exposure centering task would be in accord with a mechanical-change explanation of the half-octave shift. Settings in this direction were observed with the most intense and longest durations of exposure used, but with less intense or shorter exposures, the post-exposure settings were initially toward the exposed ear. Thus, the psychophysical evidence reported here supports the idea that exposure to intense sounds does produce temporary, local changes in cochlear mechanics, but it fails to provide a simple confirmation of the possibility that these mechanical changes underlie the half-octave shift in TTS.     

Underwater temporary threshold shift in pinnipeds: effects of noise level and duration

Behavioral psychophysical techniques were used to evaluate the residual effects of underwater noise on the hearing sensitivity of three pinnipeds: a California sea lion (Zalophus californianus), a harbor seal (Phoca vitulina), and a northern elephant seal (Mirounga angustirostris). Temporary threshold shift (TTS), defined as the difference between auditory thresholds obtained before and after noise exposure, was assessed. The subjects were exposed to octave-band noise centered at 2500 Hz at two sound pressure levels: 80 and 95 dB SL (re: auditory threshold at 2500 Hz). Noise exposure durations were 22, 25, and 50 min. Threshold shifts were assessed at 2500 and 3530 Hz. Mean threshold shifts ranged from 2.9-12.2 dB. Full recovery of auditory sensitivity occurred within 24 h of noise exposure. Control sequences, comprising sham noise exposures, did not result in significant mean threshold shifts for any subject. Threshold shift magnitudes increased with increasing noise sound exposure level (SEL) for two of the three subjects. The results underscore the importance of including sound exposure metrics (incorporating sound pressure level and exposure duration) in order to fully assess the effects of noise on marine mammal hearing.     

Temporal integration of loudness in listeners with hearing losses of primarily cochlear origin.

To investigate how hearing loss of primarily cochlear origin affects the loudness of brief tones, loudness matches between 5- and 200-ms tones were obtained as a function of level for 15 listeners with cochlear impairments and for seven age-matched controls. Three frequencies, usually 0.5, 1, and 4 kHz, were tested in each listener using a two-interval, two--alternative forced--choice (2I, 2AFC) paradigm with a roving-level, up-down adaptive procedure. Results for the normal listeners generally were consistent with published data [e.g., Florentine et al., J. Acoust Soc. Am. 99, 1633-1644 (1996)]. The amount of temporal integration--defined as the level difference between equally loud short and long tones--varied nonmonotonically with level and was largest at moderate levels. No consistent effect of frequency was apparent. The impaired listeners varied widely, but most showed a clear effect of level on the amount of temporal integration. Overall, their results appear consistent with expectations based on knowledge of the general properties of their loudness-growth functions and the equal-loudness-ratio hypothesis, which states that the loudness ratio between equal-SPL long and brief tones is the same at all SPLs. The impaired listeners' amounts of temporal integration at high SPLs often were larger than normal, although it was reduced near threshold. When evaluated at equal SLs, the amount of temporal integration well above threshold usually was in the low end of the normal range. Two listeners with abrupt high-frequency hearing losses (slopes > 50 dB/octave) showed larger-than-normal maximal amounts of temporal integration (40 to 50 dB). This finding is consistent with the shallow loudness functions predicted by our excitation-pattern model for impaired listeners [Florentine et al., in Modeling Sensorineural Hearing Loss, edited by W. Jesteadt (Erlbaum, Mahwah, NJ, 1997), pp. 187-198]. Loudness functions derived from impaired listeners' temporal-integration functions indicate that restoration of loudness in listeners with cochlear hearing loss usually will require the same gain whether the sound is short or long.     

Impact noise: the importance of level, duration, and repetition rate

The applicability of the equal energy hypothesis (EEH) to impact noise exposures was studied using chinchillas. Hearing thresholds were estimated by recording the evoked potentials from a chronic electrode implanted in the inferior colliculus. The animals were exposed to broadband impacts of 200-ms duration. The study was carried out in two parts. In experiment I, six exposure levels (107, 113, 119, 125, 131, and 137 dB SPL) and three repetition rates (4/s, 1/s and 1/4s) were employed. In the second experiment, the total duration of the exposure as well as the total energy were kept constant by trading level and rate. Results indicate that hearing loss resulting from exposure to impact noise does not conform to the predictions of the EEH. The permanent threshold shift as well as the hair cell loss are more or less equal across the lower peak exposure levels. However, both the hearing loss and the hair cell damage increase for exposures with higher peak levels. Furthermore, hearing loss and cochlear damage are dependent upon the rate of exposure. Thus the amount of hearing loss and hair cell damage appears to depend on the interaction of several factors including peak level, rate, and the susceptibility of the animal.     

Cortical responses to the 2f1-f2 combination tone measured indirectly using magnetoencephalography

The simultaneous presentation of two tones with frequencies f(1) and f(2) causes the perception of several combination tones in addition to the original tones. The most prominent of these are at frequencies f(2)-f(1) and 2f(1)-f(2). This study measured human physiological responses to the 2f(1)-f(2) combination tone at 500 Hz caused by tones of 750 and 1000 Hz with intensities of 65 and 55 dB SPL, respectively. Responses were measured from the cochlea using the distortion product otoacoustic emission (DPOAE), and from the auditory cortex using the 40-Hz steady-state magnetoencephalographic (MEG) response. The perceptual response was assessed by having the participant adjust a probe tone to cause maximal beating ("best-beats") with the perceived combination tone. The cortical response to the combination tone was evaluated in two ways: first by presenting a probe tone with a frequency of 460 Hz at the perceptual best-beats level, resulting in a 40-Hz response because of interaction with the combination tone at 500 Hz, and second by simultaneously presenting two f(1) and f(2) pairs that caused combination tones that would themselves beat at 40 Hz. The 2f(1)-f(2) DPOAE in the external auditory canal had a level of 2.6 (s.d. 12.1) dB SPL. The 40-Hz MEG response in the contralateral cortex had a magnitude of 0.39 (s.d. 0.1) nA m. The perceived level of the combination tone was 44.8 (s.d. 11.3) dB SPL. There were no significant correlations between these measurements. These results indicate that physiological responses to the 2f(1)-f(2) combination tone occur in the human auditory system all the way from the cochlea to the primary auditory cortex. The perceived magnitude of the combination tone is not determined by the measured physiological response at either the cochlea or the cortex.