Influence of benzodiazepines on auditory perception.

The aim of this study was to test for an influence of benzodiazepine (BZD) on various perceptual and/or cognitive auditory processes. Loudness, auditory selective attention, and the ability of subjects to form perceptual streams out of alternating tone sequences were tested. Nine subjects were tested before, 1, 3, 7, and 24 h after a single-dose oxazepam vs placebo administration in a crossover design. A sample of blood allows us to measure plasma oxazepam concentration. The results revealed a significant reduction in stream segregation expressed as d' scores 1 h after oxazepam intake in the test subjects. No significant change occurred across time in the same subjects when they were administrated a placebo in another session. Furthermore, oxazepam had no substantial and systematic influence either on auditory selective attention or on loudness perception. Altogether, these results suggest that the perceptual organization of sound sequences involves inhibitory neural mechanisms, which can be affected by BZDs. This outcome is consistent with existing models of auditory stream segregation and may be paralleled with earlier findings on the effect of BZDs on perceptual binding in the visual modality.

Medial olivocochlear bundle activation and perceived auditory intensity in humans.

In order to test the hypothesis of a role of cochlear efferent activity in intensity perception in humans, loudness functions, loudness integration, and loudness summation were measured in the absence and in the presence of contralateral noise in normal-hearing subjects. Additionally, relationships with the effect of the noise on evoked otoacoustic emissions (EOAEs) were tested, and comparisons with vestibular neurotomy patients were performed. Overall, the results failed to demonstrate significant effects of contralateral noise stimulation on loudness functions and loudness integration. Furthermore, no significant differences were found in vestibular neurotomy patients. A significant effect of contralateral noise on loudness summation was noted, but was not related to the effect on otoacoustic emissions. The present results fail to support the notion that efferent influences onto the cochlear compression have a significant perceptual effect.

Perceptual correlates of neural plasticity related to spontaneous otoacoustic emissions?

The inner ear contains receptor cells that oscillate spontaneously, generating waves that propagate backward in the cochlea, ultimately causing sound to be radiated into the ear canal--the spontaneous otoacoustic emissions (SOAEs). Except in rare conditions, these internally generated signals appear to go unheard. The intensity of SOAEs admittedly hovers near the threshold of detection, but they are essentially continuous and perhaps last a lifetime. The hypothesis is tested that the frequency difference limen (DL(F)) is affected by SOAEs. The results show that the DL(F) systematically improves near SOAE frequencies, determined ipsi- or contralaterally to the SOAEs, arguing for a central effect. The results are discussed in the context of central plasticity.

Transfer of learning across durations and ears in auditory frequency discrimination.

Frequency-discrimination thresholds (FDTs) for 1-kHz tone pips with durations of 40, 100, and 200 msec were measured in the left and right ears of 10 normal-hearing listeners, before and after six 2-h frequency-discrimination training sessions involving, exclusively, the 200-msec duration and the right ear. In the trained ear, highly significant improvements in FDTs were observed at all durations. Further inspection of the data suggested complete generalization between 200 and 100 msec, but not at 40 msec. Posttraining FDTs were not found to differ between the two ears for the two untrained durations, but proved significantly smaller in the right (trained) than in the left (untrained) ear at the trained (200-msec) duration only. A control experiment involving 10 additional subjects allowed us to establish the absence of intrinsic differences in pretraining FDTs between the right and left ears. Overall, these findings indicate that frequency-discrimination learning generalizes widely across stimulus durations and across ears, but that part of the improvement is specific to the range of durations and to the ear used in training.

Perceptual auditory stream segregation of sequences of complex sounds in subjects with normal and impaired hearing.

The influence of hearing loss and aging on the perceptual organization of sound sequences was investigated by comparing the ability of young normal-hearing subjects and elderly subjects having either impaired or normal hearing for their age to form perceptual auditory streams from sequences of harmonic complex tones as a function of differences in fundamental frequency (F0). The sequences consisted of repeating triplets of harmonic complex tones separated by a silence (ABA-). In conditions in which the F0s of the A and B tone were so low that the harmonics could not be individually resolved by the peripheral auditory system even in the young normal-hearing subjects, those subjects showed similar stream segregation performance to the elderly hearing-impaired subjects. In contrast, when the F0s of the tones were high enough for the harmonics to be largely resolved at the auditory periphery in normal-hearing subjects, but presumably unresolved in the elderly subjects, the former showed significantly more stream segregation than the latter. These results, which cannot be consistently explained in terms of age differences, suggest that auditory stream segregation is adversely affected by reduced peripheral frequency selectivity of elderly individuals. This finding has implications for the understanding of the listening difficulties experienced by elderly individuals in cocktail-party situations.

Influence of focused auditory attention on cochlear activity in humans.

The mammalian auditory system contains descending pathways that originate in the cortex and relay at various intermediate levels before reaching the peripheral sensory organ of Corti. The last link in this chain consists of the olivocochlear bundle. The activity of this bundle can be measured through otoacoustic emissions, which are acoustic signatures of the cochlear biomechanical activity. In the present study, it was hypothesized that frequency-specific activation of the olivocochlear bundle in the contralateral ear would show up as frequency-specific variations in otoacoustic emission amplitude in the ipsilateral ear. Two groups of young adult subjects participated in this experiment. Evoked otoacoustic emissions were recorded in the ipsilateral ear at two test frequencies (1 and 2 kHz). Subjects had to detect probe tones at a given frequency in background noise in the contralateral ear. Larger efferent activation was measured at test frequencies on which attention is focused. This result provides evidence for an influence of attention on the auditory periphery via descending projections.

Effects of supine body position and low radial accelerations on the visually perceived apparent zenith.

The visually perceived eye level (VPEL) has been shown to shift toward the lower part of the body in upright subjects facing toward the axis of rotation on a centrifuge. This shift occurs in the same direction as the shift in the gravito-inertial forces (Gis) produced by very low radial acceleration (centrifugation) combined with gravity. The purpose of this study was to determine whether the same phenomenon affects the visually perceived apparent zenith (VPAZ) in subjects in a supine position. Twelve supine subjects were instructed to set a luminous target to the VPAZ, either while they were in total darkness and motionless or while undergoing very low centrifugation. Data showed that Gis induced a VPAZ shift similar to that observed for the VPEL. Thus, as is the case for the VPEL, the corresponding logarithmic psychophysical function of the VPAZ may be considered to be a type of oculogravic illusion phenomenon with differences in the subjects' that differs from subject to subject, depending on the subject's sensitivity to low radial accelerations. Data on VPEL and VPAZ support the notion that the subjective perception of eye level in total darkness takes into account changes--even if extremely slight-in the direction of the gravito-inertial forces produced by the combination of gravity and low radial accelerations, although subjects are unaware of the Gi shift. However, depending on the intensity of the radial acceleration and the angular deviation of Gi relative to G, the shift of the VPEL and the VPAZ can be either amplified or attenuated. Moreover, differences between VPEL and VPAZ responses suggest two explanatory assumptions--namely, that this is (1) a peripheral phenomenon dependent on the neurophysiological anisotropy of the otolithic system or (2) a central phenomenon dependent on the relevance assigned to the peripheral information by the integrative sensory functions and the associative processes.

Delay and temporal integration in medial olivocochlear bundle activation in humans.

OBJECTIVE: Contralateral suppression of the transient-evoked otoacoustic emissions (TEOAEs) provides a means of studying auditory efferent function, but the temporal dynamics of the reflex are not fully understood. The most fundamental parameter is the time-course of activation of contralateral suppression. The stimulus parameters are likely to be important; this may include temporal dynamics of the suppressor itself. This investigation thus was devoted to the further study of 1) delay of contralateral suppression of TEOAEs-effect of delay of the ipsilateral probe-and 2) temporal variation of the suppressor-effect of amplitude modulation of the contralateral noise stimulus. DESIGN: Measurements were made in three samples of normal-hearing subjects (N(total) = 71), employing well-established methods of TEOAE assessment. RESULTS: Statistically significant contralateral suppression occurred some 60 msec after onset of the contralateral noise; thereafter, the effect was essentially constant (i.e., to >180 msec). The results for click delays less than 60 msec, nevertheless, were systematic and readily fitted by a sloping straight line (dB suppression versus time) reminiscent of the concept of threshold power integration. The onset of suppression may thus be characterized by a time constant. The delay of suppression also was found to be reduced by contralateral amplitude-modulated noise. CONCLUSIONS: These findings reinforce a growing consensus in the literature that, despite initiation perhaps some milliseconds after onset of the contralateral stimulus, there is a substantial delay, i.e., in the tens of milliseconds, before maximal suppression is achieved. The exact time constant of this effect appears to depend upon the combination of probe and suppressor levels, including the temporality of the suppressor. These factors are likely to delimit the role/influence of this reflex in real-world function, favoring perhaps more-or-less sustained suppression that is activated in a time-varying sound environment.

An auditory negative after-image as a human model of tinnitus.

The Zwicker tone (ZT) is an auditory after-image, i.e. a tonal sensation that occurs following the presentation of notched noise. In the present study, the hypothesis that neural lateral inhibition is involved in the generation of this auditory illusion was investigated in humans through differences in perceptual detection thresholds measured following broadband noise, notched noise, and low-pass noise stimulation. The detection thresholds were measured using probe tones at several frequencies, within as well as outside the suppressed frequency range of the notched noise, and below as well as above the corner frequency of the low-pass noise. Thresholds measured after broadband noise using a sequence of four 130-ms probe tones (with a 130-ms inter-burst interval) proved to be significantly smaller that those measured using the same probe tones after notched noise at frequencies falling within the notch, but larger for frequencies on the outer edges of the noise. Thresholds measured following low-pass noise using the same sequence of probe tones were found to be smaller at frequencies slightly above the corner, but larger at lower, neighboring frequencies. This pattern of results is consistent with the hypothesis that the changes in auditory sensitivity induced by stimuli containing sharp spectral contrasts reflect lateral inhibition processes in the auditory system. The potential implications of these findings for the understanding of the mechanisms underlying the generation of auditory illusions like the ZT or tinnitus are discussed.

Influence of peripheral resolvability on the perceptual segregation of harmonic complex tones differing in fundamental frequency.

Two experiments investigated the influence of resolvability on the perceptual organization of sequential harmonic complexes differing in fundamental frequency (F0). Using a constant-stimuli method, streaming scores for ABA-... sequences of harmonic complexes were measured as a function of the F0 difference between the A and B tones. In the first experiment, streaming scores were measured for harmonic complexes having two different nominal F0s (88 and 250 Hz) and filtered in three frequency regions (a LOW, a MID, and a HIGH region with corner frequencies of 125-625 Hz, 1375-1875 Hz, and 3900-5400 Hz, respectively). Some streaming was observed in the HIGH region (in which the harmonics were always unresolved) but streaming scores remained generally lower than in the LOW and MID regions. The second experiment verified that the streaming observed in the HIGH region was not due to the use of distortion products. Overall, the results indicated that although streaming can occur in the absence of spectral cues, the degree of resolvability of the harmonics has a significant influence.

The effect of modulation rate on the detection of frequency modulation and mistuning of complex tones.

Experiment 1 measured frequency modulation detection thresholds (FMTs) for harmonic complex tones as a function of modulation rate. Six complexes were used, with fundamental frequencies (F0s) of either 88 or 250 Hz, bandpass filtered into a LOW (125-625 Hz), MID (1375-1875 Hz) or HIGH (3900-5400 Hz) frequency region. The FMTs were about an order of magnitude greater for the three complexes whose harmonics were unresolved by the peripheral auditory system (F0 = 88 Hz in the MID region and both F0s in the HIGH region) than for the other three complexes, which contained some resolved harmonics. Thresholds increased with increases in FM rate above 2 Hz for all conditions. The increase was larger when the F0 was 88 Hz than when it was 250 Hz, and was also larger in the LOW than in the MID and HIGH regions. Experiment 2 measured thresholds for detecting mistuning produced by modulating the F0s of two simultaneously presented complexes out of phase by 180 degrees. The size of the resulting mistuning oscillates at a rate equal to the rate of FM applied to the two carriers. At low FM rates, thresholds were lowest when the harmonics were either resolved for both complexes or unresolved for both complexes, and highest when resolvability differed across complexes. For pairs of complexes with resolved harmonics, mistuning thresholds increased dramatically as the FM rate was increased above 2-5 Hz, in a way which could not be accounted for by the effect of modulation rate on the FMTs for the individual complexes. A third experiment, in which listeners detected constant ("static") mistuning between pairs of frequency-modulated complexes, provided evidence that this deterioration was due the harmonics in one of the two "resolved" complexes becoming unresolved at high FM rates, when analyzed over some finite time window. It is concluded that the detection of time-varying mistuning between groups of harmonics is limited by factors that are not apparent in FM detection data.

Tones disappear faster in the right ear than in the left.

In order to gain further information on the characteristics and physiological correlates of tone decay in humans, the tone decay test was administered to 58 normal-hearing subjects, successively in the left and right ears and in absence and presence of a contralateral noise. The results revealed that tone decay was greater in the right than in the left ear and was increased by contralateral noise. The contralateral effect of this noise on cochlear biomechanisms was then estimated by measuring contralaterally induced variations in the amplitude of click-evoked otoacoustic emissions in the same subjects. In the right ear, the increase in tone decay and the decrease in otoacoustic emission amplitude--both induced by contralateral noise--were positively correlated (r = .315, p = .016). Furthermore, the contralateral changes in otoacoustic emission amplitude were found to be on average larger in the right than in the left ear, this asymmetry being correlated with that observed for the tone decay. These findings are discussed in relation to previous results on simple and induced loudness adaptation in the vicinity of threshold, on contralateral attenuation of otoacoustic emissions and on the influence of the auditory efferents on cochlear biomechanisms.

Activation of medial olivocochlear efferent system in humans: influence of stimulus bandwidth.

The activity of the medial olivocochlear bundle (MOCB) can be studied in humans through variations in the level of evoked otoacoustic emissions (EOAEs) elicited by contralateral acoustic stimuli (CAS). The present study sought to investigate how the activity of the MOC system at a given frequency, as measured through the contralateral suppression of tone-pip EOAEs, depends on the bandwidth of the contralateral stimulus. EOAEs were recorded in 155 normal-hearing subjects, successively with and without contralateral stimuli whose bandwidth, center frequency and level were systematically varied. We showed a clear dependence of contralateral EOAE suppression on bandwidth demonstrating increased suppression with increased bandwidth over about two octaves around the center frequency of the noise. This effect was obtained irrespective of whether contralateral noise energy was kept constant independently of bandwidth or not, which indicates a role of bandwidth per se in contralateral EOAE suppression. Results are interpreted in terms of a simple model of MOCB activation mechanisms including peripheral bandpass filtering, within-channel compression and across-channel spatial summation by the afferent paths. Complementary experiments suggested a greater effectiveness of increases in bandwidth on the upper than on the lower side and of frequency components akin to or remote from the test frequency than of intermediate bands. Finally, these results were complemented by detailed spectrum analyses of the EOAE level variations induced by the different noises, which revealed that whilst noise components close to or remote from the center frequency generally attenuated EOAE level, intermediate components could in some cases lead to a relative increase in EOAE level. These results can further be explained by assuming different positive and negative weights on the inputs to the spatial summation process depending on their position relative to the center frequency.

Effect of frequency content on categorical loudness normalization.

In this study, increases in loudness with increases in bandwidth, termed loudness summation, were derived from loudness growth functions estimated using a loudness-scaling procedure. The results revealed that at equal loudness category, categorical loudness summation was generally larger in normal-hearing than in hearing-impaired subjects; furthermore, the increase in loudness summation at intermediate loudness levels observed in the former, was absent in the latter. These results, in broad agreement with recent data from the literature, can be explained in the light of physiological data on cochlear compression. One implication of these results regarding hearing aid fitting was that channel-by-channel loudness normalization was effective only when the incoming sound was closed in bandwidth to one of the test stimuli.

Informational masking in normal-hearing and hearing-impaired listeners.

The present study aimed to test whether central, across-channel, informational auditory processing abilities are altered by hearing loss. The informational masking effect exerted on a 1 kHz tone-pip by a simultaneous four-tone masker, whose spectral content changed within as well as across trials, was measured in the left and right ears of normal-hearing subjects and hearing-impaired subjects with either symmetrical or asymmetrical hearing loss between the two ears. In the subjects with normal-hearing or symmetrical hearing loss, the level of the masker was set to 40 dB SL in each ear, in the subjects with asymmetrical hearing loss, the masker was set to 40 dB SL in the best ear and loudness-balanced in the other ear. The results failed to reveal significant differences in informational masking between normal-hearing and hearing-impaired subjects. However, in subjects with asymmetric hearing loss, less informational masking was observed in the ear with the more elevated absolute thresholds than in the opposite ear. Since the latter finding can be explained in terms of across-ear differences in loudness recruitment, it is suggested that central, across-channel, informational processing abilities are not substantially different in hearing-impaired than in normal-hearing ears.

Effect of signal duration on categorical loudness scaling in normal and in hearing-impaired listeners.

The present study sought to determine whether the duration of white-noise bursts affects their loudness category rating in the same way for hearing-impaired as for normally-hearing subjects. Twelve normally-hearing and 12 hearing-impaired subjects took part. Categorical loudness growth functions were obtained for 16.25 ms, 32.5 ms, 75 ms, 150 ms and 300 ms white noise bursts. Temporal integration of loudness was defined as the intensity difference needed for stimuli of different durations to result in identical category ratings. In normally-hearing subjects, temporal integration of loudness occurred mainly with the short-duration (16.25 ms and 32.5 ms) stimuli, whereas it was found with almost every stimulus duration in hearing-impaired subjects. In other words, temporal integration of loudness between 16.25 ms and 300 ms stimulus duration was greater in hearing-impaired listeners and there was a difference between normal and hearing-impaired subjects regarding change in loudness perception with stimulus duration. Consequently, the use of fixed-duration stimuli hinders loudness normalization.

Temporal loudness integration and spectral loudness summation in normal-hearing and hearing-impaired listeners.

The aim of this study was to test for differences between normal-hearing and hearing-impaired listeners regarding two fundamental aspects of intensity perception: loudness integration and loudness summation. Loudness functions for three different stimuli were measured using categorical loudness scaling in 8 normal-hearing and 12 hearing-impaired subjects. The results indicated that temporal loudness integration, defined as the difference in SPL between 16.25-ms and 300-ms noise bursts of equal loudness, was larger in the hearing-impaired than in the normal-hearing listeners. Loudness summation, defined as the difference in SPL between a 300-ms, 1,600-Hz tone pip and a white noise burst of the same duration and loudness, did not differ between the two groups. Implications of these results for hearing aid fitting strategies based on loudness normalization are discussed.

Loudness growth functions and EABR characteristics in Digisonic cochlear implantees.

Electrically evoked auditory brainstem responses (EABRs) and loudness functions were measured in 14 subjects equipped with an MXM Digisonic cochlear implant. EABRs were evoked by 75-Hz pulse trains presented on the apical electrode. Loudness functions at the same rate and at a rate more conventional for psychoacoustic measurements (300 Hz) were measured using a categorical loudness-scaling procedure. The results revealed a significant difference in the loudness functions measured at 75 and 300 Hz, loudness increasing more steeply with stimulus intensity for the latter rate. Significant correlations between EABR wave V thresholds and perceptual thresholds measured at both 75 and 300 Hz were observed. Furthermore, in 8 out of the 14 patients, EABR wave V saturated at a stimulus level corresponding precisely to the loudest bearable, i.e. "Too loud" level for the 300-Hz stimulation rate; this same level corresponded to the "Comfortable" loudness level for the 75-Hz stimulation rate. On average, an almost linear relationship was observed over the first half of the loudness range between the stimulus intensity, expressed as a pulse duration in log units, and wave V amplitude in dB. Although further investigation is required before maximum comfort levels can be predicted reliably from EABR measures in individual subjects, these results indicate new directions regarding the estimation of perceptual dynamic range limits on the basis of EABR measures in cochlear implantees.

Sinusoidal amplitude modulation alters contralateral noise suppression of evoked otoacoustic emissions in humans.

It is well established that low-level broad band noise can elicit an amplitude decrease in evoked otoacoustic emissions recorded in the opposite ear. However, the influence of the temporal characteristics of the contralateral stimulus on this effect remains largely unknown. In the present study, otoacoustic emissions evoked by 60 dB SPL clicks were recorded in 19 normal-hearing subjects using the Otodynamics IL088, successively in absence and presence of a contralateral noise that was either steady or modulated sinusoidally in amplitude at different depths (from 25% to 100% in 25 point steps) and rates (from 50 Hz to 800 Hz in half-octave steps). The energy was kept constant whatever the modulation depth. The results showed that the evoked otoacoustic-emission attenuation effect induced by contralateral stimulation varied depending on the modulation depth and frequency of the contralateral amplitude-modulated noise. The largest suppression effect was observed at the 100 Hz modulation frequency and the 100% modulation depth. The 50 Hz modulation resulted in less suppression than with unmodulated noise. An interpretation of these results in terms of the influence of temporal amplitude fluctuations falling within a certain range on medial olivocochlear bundle activity is discussed.

Stronger bilateral efferent influences on cochlear biomechanical activity in musicians than in non-musicians.

The auditory sensory end organ is under the control of the central nervous system via efferent projections. Contralateral suppression of otoacoustic emissions (acoustic signatures of the cochlear biomechanical activity) provides a non-invasive approach to assess olivocochlear efferent activity in humans. Using this approach, the present study compared professional musicians with musically-inexperienced subjects. The results revealed stronger bilateral cochlear suppression, suggesting larger efferent influences in both ears, in musicians. Furthermore, in indicating no difference in left/right asymmetry of efferent-mediated suppression between the two groups, the present findings suggest that the observed differences in olivocochlear activity reflect bilaterally-enhanced activity of the cortical auditory structures in musicians rather than differences in cerebral hemispheric asymmetry between the two groups.