New data on the physiology of the olivo-cochlear efferent systems

The medial part of the olivocochlear efferent tract seems to be involved in two distinct phenomena. The crossed part is involved in the masking function and the uncrossed one in the control, through a tonic action, of the cochlear micromechanics.     

诱发性耳声发射的掩蔽

３４例听觉正常受试者（共４８耳）进行疏波短声诱发性耳声发射（ＥＯＡＥ）掩蔽实验,项目包括同侧同时掩蔽、同侧后掩蔽和对倒后掩蔽。同时掩蔽的掩蔽声是稳态白噪声,后掩蔽的掩蔽声是宽带噪声。同侧同时掩蔽强度达３０ｄＢＳＬ时,未观察到对ＥＯＡＥ的掩蔽效应,但对主观听觉感受有掩蔽作用,表明ＥＯＡＥ的客观属性反映听觉行为有其局限性、同侧及对侧后掩蔽出现掩蔽效应时的掩蔽强度分别为３０和５０ｄＢＳＬ,掩蔽阈约分别为５９和６８ｄＢＳＬ。耳蜗的机械特性－非线性或耳蜗内存在的某种功能性的反馈调节系统可能是同侧后掩蔽的作用机理。下行的对侧橄榄耳蜗内侧束对外毛细胞主动收缩的抑制性作用,可有效解释对倒后掩蔽的ＥＯＡＥ变化。     

Effect of forward masking on evoked potentials of auditory cortex in guinea pigs

Thresholds of the event-related potentials (ERPs) appearance were measured for one stationary and four moving auditory images presented in silence or under forward masking conditions. The difference between thresholds in silence and after noise masker was considered as masking level. Under the forward masking, the amplitude of the ERP to the first click in the test series decreased in guinea pig auditory cortex. Masking level decreased with the time lag between signal and masker and didn't depend on the fused auditory image localization that corresponded to the first click in different test signals. This fact can support the hypothesis that for the long test signals the initial part can be masked more than the final one. The ERPs amplitude to next clicks in test series depended on interaction of two factors: forward masking in the "masker-signal" system and interaction of separate ERPs in the series evoked by the test signal.     

听神经复合动作电位适应性的定量研究

本文研究了短声串作为刺激时,听神经复合动作电位(CAP)中发生的掩蔽作用随短声串的各项参数变化的关系.结果表明:①双短声诱发的CAP中掩蔽作用随两个短声间距的增大而减小;②掩蔽声越强,或被掩蔽声越弱,掩蔽作用越明显.恢复所需的时间也越长;③掩蔽作用随掩蔽声个数的增多而增大,当个数增大到一定数目时,掩蔽作用趋于饱和.     

A neuro-synaptic model of the auditory masking and unmasking process

A model of bilateral information processing in the auditory system was presented on the basis of the interaction of postsynaptic potentials intra- and internuclei in order to analyze the mechanism of binaural unmasking as well as monaural masking. The system was composed of a bilateral pair of auditory nuclei, which were organized in two parallel afferent systems as well as an efferent system. In the model, bilateral inputs were processed in three stages, i.e., the detection of interaural differences by the first afferent system, the equalization of relative neural timing by the efferent system, and the cancellation of specific spectral components by the second afferent system. Assuming the masking process to be forward and backward inhibitions on the auditory memory of signal by the ones of noise, the unmasking process could be explained as a result of disinhibition by the cancellation of bilateral masker inputs.This study was supported in part by a Grant-in-Aid for Scientific Research (No. 59870082) from the Japanese Ministry of Education, Science, and Culture     

Hearing Sensitivity to Shifts of Rippled-Spectrum Sound Signals in Masking Noise

The goal of the study was to enlarge knowledge of discrimination of complex sound signals by the auditory system in masking noise. For that, influence of masking noise on detection of shift of rippled spectrum was studied in normal listeners. The signal was a shift of ripple phase within a 0.5-oct wide rippled spectrum centered at 2 kHz. The ripples were frequency-proportional (throughout the band, ripple spacing was a constant proportion of the ripple center frequency). Simultaneous masker was a 0.5-oct noise below-, on-, or above the signal band. Both the low-frequency (center frequency 1 kHz) and on-frequency (the same center frequency as for the signal) maskers increased the thresholds for detecting ripple phase shift. However, the threshold dependence on the masker level was different for these two maskers. For the on-frequency masker, the masking effect primarily depended on the masker/signal ratio: the threshold steeply increased at a ratio of 5 dB, and no shift was detectable at a ratio of 10 dB. For the low-frequency masker, the masking effect primarily depended on the masker level: the threshold increased at a masker level of 80 dB SPL, and no shift was detectable at a masker level of 90 dB (for a signal level of 50 dB) or 100 dB (for a signal level of 80 dB). The high-frequency masker had little effect. The data were successfully simulated using an excitation-pattern model. In this model, the effect of the on-frequency masker appeared to be primarily due to a decrease of ripple depth. The effect of the low-frequency masker appeared due to widening of the auditory filters at high sound levels.     

Evidence that adaptation of suppression cannot account for auditory enhancement or enhanced forward masking.

Delaying the onset of a signal relative to the onset of a simultaneous notched masker often improves the ability of listeners to 'hear out' the signal at both threshold and suprathreshold levels. Viemeister & Bacon (J. acoust. Soc. Am., 71, 1502-1507 (1982)) suggested that such auditory enhancement effects could be accounted for if the suppression produced by the masker on the signal frequency adapted, thereby releasing the signal from suppression. In support of their hypothesis, Viemeister & Bacon reported that a masker preceded by an enhancer having no component at the signal frequency produced more forward masking than did the masker by itself. Here evidence is provided from five new experiments showing that adaptation of psychophysical two-tone suppression is inadequate to account either for auditory enhancement effects or for the enhanced forward masking demonstrated by Viemeister & Bacon.     

后掩蔽效应对大棕蝠下丘神经元声反应的影响

以回声定位蝙蝠为模式动物,采用在体动物细胞外单位记录法,研究了后掩蔽效应对下丘神经元声反应的影响。结果显示,部分神经元(38％,12／31)对测试声刺激的反应明显受到掩蔽声的抑制,其后掩蔽效应强弱与掩蔽声和测试声的相对强度差(inter-stimulus level difference,SLD),以及测试声与掩蔽声之间的间隔时间(inter-stimulus onset asynchrony,SOA)有关：当掩蔽声强度升高或测试声强度降低时,后掩蔽效应增强;而SOA的缩短,亦可见后掩蔽效应增强。另外,相当数量的神经元(52%,16／31)对测试声刺激的反应并不受掩蔽声的影响,其中有的神经元只有在特定SLD和SOA时,才表现出后掩蔽效应。而少数下丘神经元(10％,3／31)在特定SLD和SOA时,掩蔽声对测试声反应有易化作用。上述结果表明,部分下丘神经元参与了声认知活动中的后掩蔽形成过程,推测下丘神经元在定型声反应特性中,对掩蔽声诱导的兴奋前抑制性输入与测试声诱导的兴奋性输入之间的时相性动态整合起关键作用。     

Contralateral Masking in Bilateral Cochlear Implant Patients: A Model of Medial Olivocochlear Function Loss

Contralateral masking is the phenomenon where a masker presented to one ear affects the ability to detect a signal in the opposite ear. For normal hearing listeners, contralateral masking results in masking patterns that are both sharper and dramatically smaller in magnitude than ipsilateral masking. The goal of this study was to investigate whether medial olivocochlear (MOC) efferents are needed for the sharpness and relatively small magnitude of the contralateral masking function. To do this, bilateral cochlear implant patients were tested because, by directly stimulating the auditory nerve, cochlear implants circumvent the effects of the MOC efferents. The results indicated that, as with normal hearing listeners, the contralateral masking function was sharper than the ipsilateral masking function. However, although there was a reduction in the magnitude of the contralateral masking function compared to the ipsilateral masking function, it was relatively modest. This is in sharp contrast to the results of normal hearing listeners where the magnitude of the contralateral masking function is greatly reduced. These results suggest that MOC function may not play a large role in the sharpness of the contralateral masking function but may play a considerable role in the magnitude of the contralateral masking function.     

The psychoacoustic phenomenon of masking level differences in the human subject studied by auditory evoked potentials

Auditory brainstem responses (ABRs), middle latency responses (MLRs), and slow cortical potentials (SCPs) were recorded in normal-hearing adults to trains of low-frequency acoustic signals delivered binaurally against a background of a continuous masking noise. Two stimulus conditions, labelled as binaural homophasic and binaural antiphasic paradigms, respectively, were systematically compared. In the homophasic paradigm both the signals and the masker were in-phase at two ears. In the antiphasic paradigm the signals were 180 degrees out-of-phase at two ears, while the masker was in-phase. The psychoacoustic release from masking in the antiphasic vs. the homophasic paradigm was regularly accompanied by an increase in amplitudes and a shortening in peak latencies of the SCPs. In contrast, no differences were evidenced between the homophasic and the antiphasic paradigms with respect to the ABRs and the MLRs. Considering the generation loci of the studied electric responses, it is concluded that the binaural psychoacoustic phenomenon, referred to as the masking level difference, is operated primarily at the cortical level.     

Binaural masking and sensitivity to interaural delay in the inferior colliculus.

The binaural masking level difference (BMLD) is a psychophysical effect whereby signals masked by a noise at one ear become unmasked by sounds reaching the other. BMLD effects are largest at low frequencies where they depend on signal phase, suggesting that part of the physiological mechanism responsible for the BMLD resides in cells that are sensitive to interaural time disparities. We have investigated a physiological basis for unmasking in the responses of delay-sensitive cells in the central nucleus of the inferior colliculus in anaesthetized guinea pigs. The masking effects of a binaurally presented noise, as a function of the masker delay, were quantified by measuring the number of discharges synchronized to the signal, and by measuring the masked threshold. The noise level for masking was lowest at the best delay for the noise. The mean magnitude of the unmasking across our neural population was similar to the human psychophysical BMLD under the same signal and masker conditions.     

Effect of harmonicity on the detection of a signal in a complex masker and on spatial release from masking

The amount of masking of sounds from one source (signals) by sounds from a competing source (maskers) heavily depends on the sound characteristics of the masker and the signal and on their relative spatial location. Numerous studies investigated the ability to detect a signal in a speech or a noise masker or the effect of spatial separation of signal and masker on the amount of masking, but there is a lack of studies investigating the combined effects of many cues on the masking as is typical for natural listening situations. The current study using free-field listening systematically evaluates the combined effects of harmonicity and inharmonicity cues in multi-tone maskers and cues resulting from spatial separation of target signal and masker on the detection of a pure tone in a multi-tone or a noise masker. A linear binaural processing model was implemented to predict the masked thresholds in order to estimate whether the observed thresholds can be accounted for by energetic masking in the auditory periphery or whether other effects are involved. Thresholds were determined for combinations of two target frequencies (1 and 8 kHz), two spatial configurations (masker and target either co-located or spatially separated by 90 degrees azimuth), and five different masker types (four complex multi-tone stimuli, one noise masker). A spatial separation of target and masker resulted in a release from masking for all masker types. The amount of masking significantly depended on the masker type and frequency range. The various harmonic and inharmonic relations between target and masker or between components of the masker resulted in a complex pattern of increased or decreased masked thresholds in comparison to the predicted energetic masking. The results indicate that harmonicity cues affect the detectability of a tonal target in a complex masker.     

Detection of sonar signals in the presence of pulses of masking noise by the echolocating bat,Eptesicus fuscus

Summary Two big brown bats (Eptesicus fuscus) were trained to report the presence or absence of a virtual sonar target. The bats' sensitivity to transient masking was investigated by adding 5 ms pulses of white noise delayed from 0 to 16 ms relative to the target echo. When signal and masker occurred simultaneously, the bats required a signal energy to noise spectrum level ratio of 35 dB for 50% probability of detection. When the masker was delayed by 2 ms or more there was no significant masking and echo energy could be reduced by 30 dB for the same probability of detection. The average duration of the most energetic sonar signal of each trial was measured to be 1.7 ms and 2.4 ms for the two bats, but a simple relation between detection performance and pulse duration was not found.In a different experiment the masking noise pulses coincided with the echo, and the duration of the masker was varied from 2 to 37.5 ms. The duration of the masker had little or no effect on the probability of detection.The findings are consistent with an aural integration time constant of about 2 ms, which is comparable to the duration of the cries. This is an order of magnitude less than found in backward masking experiments with humans and may be an adaptation to the special constraints of echolocation. The short time of sensitivity to masking may indicate that the broad band clicks of arctiid moths produced as a countermeasure to bat predation are unlikely to function by masking the echo of the moth.Abbreviations SPL sound pressure level - SD standard deviation - SE standard error - BW bandwidth     

Forward masking in the evoked responses of the guinea pig auditory cortex: effects of variation of the interaural time and phase differences

In anaesthesized guinea pigs the evoked potentials of the auditory cortex were studied in a forward masking paradigm. In-phase and out-of-phase binaurally presented clicks with interaural time delay (ITD) were used as masker, in-phase click with ITD = 0 served as probe signal. Addition of the masking stimulus suppressed the probe-evoked response that followed the masker. The magnitude of the suppression correlated with the amount of the masker-evoked response: an increase in masker-evoked excitation caused a greater reduction in probe response magnitude. Amplitude of masker-evoked response was seen to be a monotonic or non-monotonic function of ITD. The non-monotonic response exhibited a sensitivity to the interaural phase differences when in-phase and out-of-phase maskers were presented, and showed the tendency to be periodic function of ITD in the expanded range of ITD values. Phase-sensitive responses differed in recovery time following the in-phase and out-of-phase masking stimuli. At near-threshold levels of a forward masker an enhancement of the probe-evoked response was observed.     

Compressive nonlinearity of human hearing in sound spectra discrimination

In the psychophysical experiments reported here, cochlear compression function was derived by comparing on-frequency and off-frequency masking. The signal was rippled spectrum noise. The ripple density discrimination threshold was measured in the ripple phase reversion test. An increase in masker intensity led to a decrease in a resolvable ripple density threshold. The on-frequency masker level at threshold increased proportionally to the signal intensity. The off-frequency masker level at threshold also increased proportionally to the signal at signal intensity levels below 50 dB, whereas at signal levels above 60 dB SPL, the ratio of the masker level at threshold gradient to signal level gradient was 1 : 5 dB/dB, revealing cochlear compression.     

Compressive Nonlinearity in the Auditory System: Manifestation in the Action of Complex Sound Signals

In psychophysical experiments on humans, a cochlear compression was revealed by comparing onfrequency and low-frequency masking in the conditions of discrimination of complex spectra of sound signals. It has been established that, with an increase in the level of the masker, the limit of the discriminable density of the spectrum ripples decreases. It was noted that when the signal level increased, the on-frequency masker level at threshold grew proportionally to the level of the signal, whereas the low-frequency masker level grew much more slowly, indicating the compression of the signal, which was not affected by lateral suppression and off-frequency listening.     

Spatial features of vibrotactile masking effects on airpuff-elicited sensations in the human hand

It was recently shown that the cutaneous sensitivity to airpuffs is decreased by a low-frequency vibrotactile masker in the hairy skin, and by a low-frequency but especially by a high-frequency masker in the glabrous skin. In the current study, the spatial features of this masking effect were determined in four healthy human subjects, using a reaction time paradigm. The masking effect decreased monotonically with increasing interstimulus distance, and identically in longitudinal and transverse (i.e., lateral) directions in the palm or dorsal surface of the hand. The masking effect was stronger in the glabrous than in the hairy skin, especially in the fingers. In the glabrous skin, the spread of masking effect produced by a high-frequency masker was more extensive than that produced by a low-frequency masker. The mechanical spread of high-frequency vibration was less extensive than that of low-frequency vibration in the skin. In the glabrous skin, a masker applied to the tip of the finger produced a stronger masking effect on sensations in the base of the finger than when the masker was located at the base and the test stimulus was located at the tip. It is concluded that mechanical spread of vibration in the skin is of minor importance in explaining the masking effects. Different peripheral neural mechanisms underlie the airpuff-elicited sensations in the hairy and glabrous skin. The afferent inhibitory mechanisms are stronger for signals coming from the glabrous skin of the fingers than for signals coming from the hairy skin. Furthermore, the peripheral innervation density and size of the cortical representational areas may be of importance in determining the magnitude of the masking effect.     

Spatial Features of Vibrotactile Masking Effects on Airpuff-Elicited Sensations in the Human Hand

It was recently shown that the cutaneous sensitivity to airpuffs is decreased by a low-frequency vibrotactile masker in the hairy skin, and by a low-frequency but especially by a high-frequency masker in the glabrous skin. In the current study, the spatial features of this masking effect were determined in four healthy human subjects, using a reaction time paradigm. The masking effect decreased monotonically with increasing interstimulus distance, and identically in longitudinal and transverse (i.e., lateral) directions in the palm or dorsal surface of the hand. The masking effect was stronger in the glabrous than in the hairy skin, especially in the fingers. In the glabrous skin, the spread of masking effect produced by a high-frequency masker was more extensive than that produced by a low-frequency masker. The mechanical spread of high-frequency vibration was less extensive than that of low-frequency vibration in the skin. In the glabrous skin, a masker applied to the tip of the finger produced a stronger masking effect on sensations in the base of the finger than when the masker was located at the base and the test stimulus was located at the tip. It is concluded that mechanical spread of vibration in the skin is of minor importance in explaining the masking effects. Different peripheral neural mechanisms underlie the airpuff-elicited sensations in the hairy and glabrous skin. The afferent inhibitory mechanisms are stronger for signals coming from the glabrous skin of the fingers than for signals coming from the hairy skin. Furthermore, the peripheral innervation density and size of the cortical representational areas may be of importance in determining the magnitude of the masking effect.     

Change in Auditory Image Movement Trajectories under Conditions of Direct Nonsimultaneous Masking

The authors studied fused auditory image (FAI) movement trajectories under conditions of direct nonsimultaneous masking. This movement was created by a gradual change in a dichotically presented series of clicks with interaural differences in stimulation from 0 to ±700 s or from ±700 to 0 s. Binaurally presented transmissions of wide-band noise served as maskers. The location and length of the trajectories were evaluated without a masker and with five values of the time lag between the signal beginning and masker end. When the test signal duration was 200 ms, the length of the trajectories was 33–44° without a masker. In the first test group, this trajectory lay close to the median line of the head without a masker (irrespective of the movement direction) and moved away from it under masking conditions. When the FAI moved from the median line towards the right or left ear, the initial part of the trajectory was masked; when the movement direction was opposite, the final part was masked. In the second group, the trajectories were located near the ears when the FAI moved from either ear and shifted towards the median line as a result of masking. When the movement direction was opposite, they were close to the median line and shifted towards the ear under masking conditions. When the FAI moved along all trajectories, their initial parts were masked.     

Detection of Tastes in Mixture with Other Tastes: Issues of Masking and Aging

When one taste (masker) is strong enough, it can completelymask another taste (target) of different quality. How strongthe masker must be to do this depends on how strong the targetis. As the target concentration is increased, the masking concentrationmust be increased, too, but in ever-increasing proportion. Toquantify the conditions for such complete masking, the target'sdetection threshold was measured as a function of the masker'sconcentration, from zero to strong. This was done for 12 binarycombinations of sucrose, sodium chloride, citric acid and quininehydrochloride. The 12 functions generated show that some tastantsmask each other much more efficiently than others. Masking givesnew insight into the role of aging in taste: older (66–90years) subjects' thresholds, regardless of masking concentration,always measured a constant factor higher than younger (18–29years) subjects' thresholds (about two to seven times higher,depending on target tastant). Thus, with increasing level ofthe masker, the thresholds of young and elderly go up in parallel.Thresholds of tastants in water alone are false predictors ofelderly persons' ability to perceive ingredients like salt andsugar condiments in foods, where, because of masking, theirthresholds can be several times higher than in water. Age manifesteditself relatively mildly in sucrose and citric acid, moderatelyin sodium chloride, and strongly in quinine hydrochloride. Chem.Senses 21: 211–221, 1996.