Top-Down Influences of the Medial Olivocochlear Efferent System in Speech Perception in Noise

One of the putative functions of the medial olivocochlear (MOC) system is to enhance signal detection in noise. The objective of this study was to elucidate the role of the MOC system in speech perception in noise. In normal-hearing human listeners, we examined (1) the association between magnitude of MOC inhibition and speech-in-noise performance, and (2) the association between MOC inhibition and the amount of contralateral acoustic stimulation (CAS)-induced shift in speech-in-noise acuity. MOC reflex measurements in this study considered critical measurement issues overlooked in past work by: recording relatively low-level, linear click-evoked otoacoustic emissions (CEOAEs), adopting 6 dB signal-to-noise ratio (SNR) criteria, and computing normalized CEOAE differences. We found normalized index to be a stable measure of MOC inhibition (mean = 17.21%). MOC inhibition was not related to speech-in-noise performance measured without CAS. However, CAS in a speech-in-noise task caused an SNR_SP enhancement (mean = 2.45 dB), and this improvement in speech-in-noise acuity was directly related to their MOC reflex assayed by CEOAEs. Individuals do not necessarily use the available MOC-unmasking characteristic while listening to speech in noise, or do not utilize unmasking to the extent that can be shown by artificial MOC activation. It may be the case that the MOC is not actually used under natural listening conditions and the higher auditory centers recruit MOC-mediated mechanisms only in specific listening conditions–those conditions remain to be investigated.     

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。     

Cochlear Delay and Medial Olivocochlear Functioning in Children with Suspected Auditory Processing Disorder

Behavioral manifestations of processing deficits associated with auditory processing disorder (APD) have been well documented. However, little is known about their anatomical underpinnings, especially cochlear processing. Cochlear delays, a proxy for cochlear tuning, measured using stimulus frequency otoacoustic emission (SFOAE) group delay, and the influence of the medial olivocochlear (MOC) system activation at the auditory periphery was studied in 23 children suspected with APD (sAPD) and 22 typically developing (TD) children. Results suggest that children suspected with APD have longer SFOAE group delays (possibly due to sharper cochlear tuning) and reduced MOC function compared to TD children. Other differences between the groups include correlation between MOC function and SFOAE delay in quiet in the TD group, and lack thereof in the sAPD group. MOC-mediated changes in SFOAE delay were in opposite directions between groups: increase in delay in TD vs. reduction in delay in the sAPD group. Longer SFOAE group delays in the sAPD group may lead to longer cochlear filter ringing, and potential increase in forward masking. These results indicate differences in cochlear and MOC function between sAPD and TD groups. Further studies are warranted to explore the possibility of cochlea as a potential site for processing deficits in APD.     

听神经元的频率调谐

频率作为声音的一个重要参数,在听敏感神经元对声音进行分析和编码过程中扮演重要角色。一般用频率调谐曲线来表示听敏感神经元的频率调谐特性,并用Qn(10,30,50)值表达频率调谐曲线的尖锐程度,Qn值越大,频率调谐曲线也越尖锐,神经元的频率调谐能力越好,对频率的分辨能力越高。从听觉外周到中枢,听敏感神经元的频率调谐逐级锐化,而这种锐化主要是由听中枢的多种抑制性神经递质的作用而产生的,其中起主要作用的是GABA能和甘氨酸能神经递质。此外,离皮层调控,双侧下丘间的联合投射以及弱噪声前掩蔽等因素也会影响听敏感神经元的频率调谐特性。     

Interaural Level Differences Do Not Suffice for Restoring Spatial Release from Masking in Simulated Cochlear Implant Listening

Spatial release from masking refers to a benefit for speech understanding. It occurs when a target talker and a masker talker are spatially separated. In those cases, speech intelligibility for target speech is typically higher than when both talkers are at the same location. In cochlear implant listeners, spatial release from masking is much reduced or absent compared with normal hearing listeners. Perhaps this reduced spatial release occurs because cochlear implant listeners cannot effectively attend to spatial cues. Three experiments examined factors that may interfere with deploying spatial attention to a target talker masked by another talker. To simulate cochlear implant listening, stimuli were vocoded with two unique features. First, we used 50-Hz low-pass filtered speech envelopes and noise carriers, strongly reducing the possibility of temporal pitch cues; second, co-modulation was imposed on target and masker utterances to enhance perceptual fusion between the two sources. Stimuli were presented over headphones. Experiments 1 and 2 presented high-fidelity spatial cues with unprocessed and vocoded speech. Experiment 3 maintained faithful long-term average interaural level differences but presented scrambled interaural time differences with vocoded speech. Results show a robust spatial release from masking in Experiments 1 and 2, and a greatly reduced spatial release in Experiment 3. Faithful long-term average interaural level differences were insufficient for producing spatial release from masking. This suggests that appropriate interaural time differences are necessary for restoring spatial release from masking, at least for a situation where there are few viable alternative segregation cues.     

Critical ratio: effects of contralateral masking on normal and pathological ears]

In order to evaluate the central interferences on the auditory efficiency, the effect of contralateral masking noise on Critical Ratio (CR) has been studied in 14 normal ears (control group), in 12 ears with cochlear disorders and in 6 with retrocochlear lesion (acoustic neuromas); in all cases the impairment was unilateral, while the other ear was normal. CR values were calculated for 1 KHz pulsed tones (duration 500 ms, rise/fall 25 ms, duty cycle of 50%); the masking noise was a wide band (90-20,000 Hz) delivered at 40 dB SL. The results have demonstrated that in the presence of contralateral masking noise, CRs don't modify both in the normal ears and in those with retrocochlear disorder, whereas they increase, almost always, in ears with cochlear deafness. These results demonstrate that the involvement of central auditory pathways, because of contralateral noise, makes the auditory efficiency worse, only in ears with cochlear dysfunctions; this behaviour seems to confirm the peripheral origin of CR.     

Sound localisation in crickets

Intracellular recordings were made in the brain of the cricket Gryllus bimaculatus from an ascending auditory interneuron (AN1). Acoustic stimuli with calling song temporal pattern were delivered via earphones in a preparation with the acoustic trachea cut (attenuation of crossing sound > 30 dB). The input-output function of this cell was then determined by recording its responses to stimulation of the ipsilateral ear alone, of the contralateral ear alone and to stimulation of both ears simultaneously with the same or different carrier frequencies and intensities.This interneuron was excited by the ear ipsilateral to its axon and dendritic field and unresponsive to stimuli presented to the axon-contralateral ear alone. However, in binaural stimulation experiments, the response to a constant ipsilateral stimulus was progressively reduced as the intensity of a simultaneous contralateral stimulus was increased, above a threshold intensity.Tuning curves for threshold of this inhibition, determined in binaural stimulation experiments, indicated significant inhibition in the range 3–20 kHz with lowest threshold at 4–5 kHz. The inhibition was unaffected by sectioning of the contralateral circumoesophageal or neck connective, indicating that the inhibitory influence crosses the midline at the level of the prothoracic ganglion. Intracellular recordings from AN1 in the prothoracic ganglion confirmed that it was indeed neurally inhibited by inputs from the contralateral ear.Tuning curves for excitation of an omega neuron (ON1) by the ear ipsilateral to its soma and also the tuning of inhibition of ON1 by its contralateral ON1 partner, closely match the tuning of inhibition of AN1 and to a lesser extent, of AN2. This was taken as evidence that each AN1 is inhibited by the contralateral ON1. The significance of this interaction for directional hearing and phonotaxis is discussed.Abbreviations AP/CHP action potentials per chirp - AN1, AN2 ascending auditory interneurons 1, 2 - ON1 omega neuron 1 - ipsi ipsilateral contra contralateral - PTG prothoracic ganglion loc lateral ocellar nerve - On optic nerve an antennal nerve - coc circum-oesophageal connective so sound off     

Contralateral Noise Stimulation Delays P300 Latency in School-Aged Children

Background and Objective

The auditory cortex modulates auditory afferents through the olivocochlear system, which innervates the outer hair cells and the afferent neurons under the inner hair cells in the cochlea. Most of the studies that investigated the efferent activity in humans focused on evaluating the suppression of the otoacoustic emissions by stimulating the contralateral ear with noise, which assesses the activation of the medial olivocochlear bundle. The neurophysiology and the mechanisms involving efferent activity on higher regions of the auditory pathway, however, are still unknown. Also, the lack of studies investigating the effects of noise on human auditory cortex, especially in peadiatric population, points to the need for recording the late auditory potentials in noise conditions. Assessing the auditory efferents in schoolaged children is highly important due to some of its attributed functions such as selective attention and signal detection in noise, which are important abilities related to the development of language and academic skills. For this reason, the aim of the present study was to evaluate the effects of noise on P300 responses of children with normal hearing.

Methods

P300 was recorded in 27 children aged from 8 to 14 years with normal hearing in two conditions: with and whitout contralateral white noise stimulation.

Results

P300 latencies were significantly longer at the presence of contralateral noise. No significant changes were observed for the amplitude values.

Conclusion

Contralateral white noise stimulation delayed P300 latency in a group of school-aged children with normal hearing. These results suggest a possible influence of the medial olivocochlear activation on P300 responses under noise condition.     

Tinnitus and Other Auditory Problems – Occupational Noise Exposure below Risk Limits May Cause Inner Ear Dysfunction

The aim of the investigation was to study if dysfunctions associated to the cochlea or its regulatory system can be found, and possibly explain hearing problems in subjects with normal or near-normal audiograms. The design was a prospective study of subjects recruited from the general population. The included subjects were persons with auditory problems who had normal, or near-normal, pure tone hearing thresholds, who could be included in one of three subgroups: teachers, Education; people working with music, Music; and people with moderate or negligible noise exposure, Other. A fourth group included people with poorer pure tone hearing thresholds and a history of severe occupational noise, Industry. N_total = 193. The following hearing tests were used:− pure tone audiometry with Békésy technique,− transient evoked otoacoustic emissions and distortion product otoacoustic emissions, without and with contralateral noise;− psychoacoustical modulation transfer function,− forward masking,− speech recognition in noise,− tinnitus matching.A questionnaire about occupations, noise exposure, stress/anxiety, muscular problems, medication, and heredity, was addressed to the participants. Forward masking results were significantly worse for Education and Industry than for the other groups, possibly associated to the inner hair cell area. Forward masking results were significantly correlated to louder matched tinnitus. For many subjects speech recognition in noise, left ear, did not increase in a normal way when the listening level was increased. Subjects hypersensitive to loud sound had significantly better speech recognition in noise at the lower test level than subjects not hypersensitive. Self-reported stress/anxiety was similar for all groups. In conclusion, hearing dysfunctions were found in subjects with tinnitus and other auditory problems, combined with normal or near-normal pure tone thresholds. The teachers, mostly regarded as a group exposed to noise below risk levels, had dysfunctions almost identical to those of the more exposed Industry group.     

Aided and Unaided Speech Perception by Older Hearing Impaired Listeners

The most common complaint of older hearing impaired (OHI) listeners is difficulty understanding speech in the presence of noise. However, tests of consonant-identification and sentence reception threshold (SeRT) provide different perspectives on the magnitude of impairment. Here we quantified speech perception difficulties in 24 OHI listeners in unaided and aided conditions by analyzing (1) consonant-identification thresholds and consonant confusions for 20 onset and 20 coda consonants in consonant-vowel-consonant (CVC) syllables presented at consonant-specific signal-to-noise (SNR) levels, and (2) SeRTs obtained with the Quick Speech in Noise Test (QSIN) and the Hearing in Noise Test (HINT). Compared to older normal hearing (ONH) listeners, nearly all unaided OHI listeners showed abnormal consonant-identification thresholds, abnormal consonant confusions, and reduced psychometric function slopes. Average elevations in consonant-identification thresholds exceeded 35 dB, correlated strongly with impairments in mid-frequency hearing, and were greater for hard-to-identify consonants. Advanced digital hearing aids (HAs) improved average consonant-identification thresholds by more than 17 dB, with significant HA benefit seen in 83% of OHI listeners. HAs partially normalized consonant-identification thresholds, reduced abnormal consonant confusions, and increased the slope of psychometric functions. Unaided OHI listeners showed much smaller elevations in SeRTs (mean 6.9 dB) than in consonant-identification thresholds and SeRTs in unaided listening conditions correlated strongly (r = 0.91) with identification thresholds of easily identified consonants. HAs produced minimal SeRT benefit (2.0 dB), with only 38% of OHI listeners showing significant improvement. HA benefit on SeRTs was accurately predicted (r = 0.86) by HA benefit on easily identified consonants. Consonant-identification tests can accurately predict sentence processing deficits and HA benefit in OHI listeners.     

先天性小耳畸形手术治疗的研究进展

先天性小耳畸形是发病率较高的头面部畸形之一,常为耳廓及中耳腔同时存在,内耳发育不良相对少见,这一疾病所致的缺陷不仅影响到患者的容貌,更重要的是导致患者听觉功能的障碍,严重影响到患者的日常工作学习和生活。部分患者还伴有或多或少的心理影响,这在双侧小耳畸形的患者中更多见。手术是其主要的治疗方法,要求不仅重建外形正常的耳廓,同时还拥有正常或接近正常的听力。外耳廓再造和听力重建手术不仅使先天性小耳畸形患者的耳部外观明显改善,还能使其听觉功能进一步提高。本文主要综述了先天性小耳畸形的病因和流行病学、分类、手术时机和方式的选择、听力重建、耳廓再造的方法及组织工程学耳再造技术,重点介绍了耳廓再造材料的选择及手术注意点,以期为先天性小耳畸形的临床治疗提供更多理论依据。     

Spatial tuning of neurons in the inferior colliculus of the big brown bat: effects of sound level, stimulus type and multiple sound sources

We examined factors that affect spatial receptive fields of single units in the central nucleus of the inferior colliculus of Eptesicus fuscus. Pure tones, frequency- or amplitude-modulated sounds, or noise bursts were presented in the free-field, and responses were recorded extracellularly. For 58 neurons that were tested over a 30 dB range of sound levels, 7 (12%) exhibited a change of less than 10° in the center point and medial border of their receptive field. For 28 neurons that were tested with more than one stimulus type, 5 (18%) exhibited a change of less than 10° in the center point and medial border of their receptive field.The azimuthal response ranges of 19 neurons were measured in the presence of a continuous broadband noise presented from a second loudspeaker set at different fixed azimuthal positions. For 3 neurons driven by a contralateral stimulus only, the effect of the noise was simple masking. For 11 neurons driven by sound at either side, 8 were unaffected by the noise and 1 showed a simple masking effect. For the remaining 2, as well as for 5 neurons that were excited by contralateral sound and inhibited by ipsilateral sound, the peak of the azimuthal response range shifted toward the direction of the noise.Abbreviations E/E excitation at either ear - I/E inhibition at the ipsilateral ear, excitation at the contralateral ear - O/E no effect from the ipsilateral ear, excitation at the contralateral ear - FM downward frequency modulation - FM upward frequency modulation - IC inferior colliculus - ICC central nucleus of the inferior colliculus - ILD interaural level difference - ITD interaural time difference - PT pure tone - SAM sinusoidally amplitude modulated sounds - SFM sinusoidally frequency modulated sounds     

Impact of a Moving Noise Masker on Speech Perception in Cochlear Implant Users

ObjectivesPrevious studies investigating speech perception in noise have typically been conducted with static masker positions. The aim of this study was to investigate the effect of spatial separation of source and masker (spatial release from masking, SRM) in a moving masker setup and to evaluate the impact of adaptive beamforming in comparison with fixed directional microphones in cochlear implant (CI) users.DesignSpeech reception thresholds (SRT) were measured in S₀N₀ and in a moving masker setup (S₀N_move) in 12 normal hearing participants and 14 CI users (7 subjects bilateral, 7 bimodal with a hearing aid in the contralateral ear). Speech processor settings were a moderately directional microphone, a fixed beamformer, or an adaptive beamformer. The moving noise source was generated by means of wave field synthesis and was smoothly moved in a shape of a half-circle from one ear to the contralateral ear. Noise was presented in either of two conditions: continuous or modulated.ResultsSRTs in the S₀N_move setup were significantly improved compared to the S₀N₀ setup for both the normal hearing control group and the bilateral group in continuous noise, and for the control group in modulated noise. There was no effect of subject group. A significant effect of directional sensitivity was found in the S₀N_move setup. In the bilateral group, the adaptive beamformer achieved lower SRTs than the fixed beamformer setting. Adaptive beamforming improved SRT in both CI user groups substantially by about 3 dB (bimodal group) and 8 dB (bilateral group) depending on masker type.ConclusionsCI users showed SRM that was comparable to normal hearing subjects. In listening situations of everyday life with spatial separation of source and masker, directional microphones significantly improved speech perception with individual improvements of up to 15 dB SNR. Users of bilateral speech processors with both directional microphones obtained the highest benefit.     

Attentional Modulation of Auditory Steady-State Responses

Auditory selective attention enables task-relevant auditory events to be enhanced and irrelevant ones suppressed. In the present study we used a frequency tagging paradigm to investigate the effects of attention on auditory steady state responses (ASSR). The ASSR was elicited by simultaneously presenting two different streams of white noise, amplitude modulated at either 16 and 23.5 Hz or 32.5 and 40 Hz. The two different frequencies were presented to each ear and participants were instructed to selectively attend to one ear or the other (confirmed by behavioral evidence). The results revealed that modulation of ASSR by selective attention depended on the modulation frequencies used and whether the activation was contralateral or ipsilateral. Attention enhanced the ASSR for contralateral activation from either ear for 16 Hz and suppressed the ASSR for ipsilateral activation for 16 Hz and 23.5 Hz. For modulation frequencies of 32.5 or 40 Hz attention did not affect the ASSR. We propose that the pattern of enhancement and inhibition may be due to binaural suppressive effects on ipsilateral stimulation and the dominance of contralateral hemisphere during dichotic listening. In addition to the influence of cortical processing asymmetries, these results may also reflect a bias towards inhibitory ipsilateral and excitatory contralateral activation present at the level of inferior colliculus. That the effect of attention was clearest for the lower modulation frequencies suggests that such effects are likely mediated by cortical brain structures or by those in close proximity to cortex.     

Bilateral brainstem connections of the rat supratrigeminal region

Efferent and afferent connections of the supratrigeminal region were studied in the rat using iontophoretically delivered horseradish peroxidase and Phaseolus vulgaris leuco-agglutinin. Projections of supratrigeminal efferents were found to the contralateral supratrigeminal region, to the ipsi- and contralateral trigeminal motor nuclei and the medullary reticular formation, and to the ipsilateral facial and hypoglossal motor nuclei. Neurons projecting to the supratrigeminal region were located in the contralateral supratrigeminal nucleus, in the ipsilateral mesencephalic trigeminal nucleus and bilaterally in the medullary reticular formation. This organization is discussed with respect to bilateral oral motor control mechanisms.     

Neuromagnetic responses elicited by auditory stimuli in dichotic listening

We measured N1m and P2m components of the magnetic field responses that were elicited by random series of a tone burst given to the left ear and a monosyllabic speech sound given to the right ear. The magnetic responses had smaller amplitudes and/or longer peak latencies of the N1m and the P2m when the stimulus was preceded by a stimulus at the same ear than when preceded by a stimulus at the different ear. This reduction of the response by preceding stimulation of the same ear was significant over the hemisphere contralateral, but not ipsilateral, to the ear stimulated. The peak latencies of N1m and P2m were significantly longer in the response over the hemisphere contralateral than ipsilateral to the stimulated ear.     

Retinal projections in the catfish,Mystus vittatus (Bloch) as revealed by tracer studies with horseradish peroxidase

Summary The retinal efferents of the catfish, Mystus vittatus, were investigated with the use of the horseradish peroxidase (HRP) technique. Most retinal fibres extended contralateral to the eye that had received HRP label, while a few fascicles projected to the ipsilateral side without decussation in the optic chiasma. The contralateral fibres projected to the suprachiasmatic nucleus, the nucleus opticus dorsolateralis, the nucleus of the posterior commissure, the nucleus geniculatus lateralis, pretectal nuclear complex, and to two layers of the optic tectum, i.e., stratum fibrosum et griseum superficiale and stratum griseum centrale. The accessory optic tract arose from the inner area of the optic tract and extended ventromedially to the accessory optic nucleus. The ipsilateral fascicles projected to almost all the above mentioned nuclei, but these projections were comparatively sparse. The ipsilateral retinal projection was restricted to the rostral tectum.     

Corticofugal modulation of initial neural processing of sound information from the ipsilateral ear in the mouse

Background

Cortical neurons implement a high frequency-specific modulation of subcortical nuclei that includes the cochlear nucleus. Anatomical studies show that corticofugal fibers terminating in the auditory thalamus and midbrain are mostly ipsilateral. Differently, corticofugal fibers terminating in the cochlear nucleus are bilateral, which fits to the needs of binaural hearing that improves hearing quality. This leads to our hypothesis that corticofugal modulation of initial neural processing of sound information from the contralateral and ipsilateral ears could be equivalent or coordinated at the first sound processing level.

Methodology/Principal Findings

With the focal electrical stimulation of the auditory cortex and single unit recording, this study examined corticofugal modulation of the ipsilateral cochlear nucleus. The same methods and procedures as described in our previous study of corticofugal modulation of contralateral cochlear nucleus were employed simply for comparison. We found that focal electrical stimulation of cortical neurons induced substantial changes in the response magnitude, response latency and receptive field of ipsilateral cochlear nucleus neurons. Cortical stimulation facilitated auditory response and shortened the response latency of physiologically matched neurons whereas it inhibited auditory response and lengthened the response latency of unmatched neurons. Finally, cortical stimulation shifted the best frequencies of cochlear neurons towards those of stimulated cortical neurons.

Conclusion

Our data suggest that cortical neurons enable a high frequency-specific remodelling of sound information processing in the ipsilateral cochlear nucleus in the same manner as that in the contralateral cochlear nucleus.     

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.     

Reversible induction of phantom auditory sensations through simulated unilateral hearing loss

Tinnitus, a phantom auditory sensation, is associated with hearing loss in most cases, but it is unclear if hearing loss causes tinnitus. Phantom auditory sensations can be induced in normal hearing listeners when they experience severe auditory deprivation such as confinement in an anechoic chamber, which can be regarded as somewhat analogous to a profound bilateral hearing loss. As this condition is relatively uncommon among tinnitus patients, induction of phantom sounds by a lesser degree of auditory deprivation could advance our understanding of the mechanisms of tinnitus. In this study, we therefore investigated the reporting of phantom sounds after continuous use of an earplug. 18 healthy volunteers with normal hearing wore a silicone earplug continuously in one ear for 7 days. The attenuation provided by the earplugs simulated a mild high-frequency hearing loss, mean attenuation increased from <10 dB at 0.25 kHz to >30 dB at 3 and 4 kHz. 14 out of 18 participants reported phantom sounds during earplug use. 11 participants presented with stable phantom sounds on day 7 and underwent tinnitus spectrum characterization with the earplug still in place. The spectra showed that the phantom sounds were perceived predominantly as high-pitched, corresponding to the frequency range most affected by the earplug. In all cases, the auditory phantom disappeared when the earplug was removed, indicating a causal relation between auditory deprivation and phantom sounds. This relation matches the predictions of our computational model of tinnitus development, which proposes a possible mechanism by which a stabilization of neuronal activity through homeostatic plasticity in the central auditory system could lead to the development of a neuronal correlate of tinnitus when auditory nerve activity is reduced due to the earplug.