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The dichotic frequency following responses (FFR) have been used in studies to infer about dichotic auditory processing. In the present study, we hypothesize that the proximity of the binaural neural generators of the FFR would result in interference of the volume-conducted electrical fields. This might lead to contamination of the scalp-recorded dichotic FFRs due to which it might be difficult to infer about true dichotic processing in the putative neural generators. We investigated this by recording FFRs to binaurally presented 200 Hz pure tone with graded dichotic phase offsets (0°, 90°, 180° and 270°) in normal hearing young adults. Spectral analysis of the FFRs was performed for the estimation of the magnitude and phase at the component frequencies. FFR spectra were compared using non-parametric paired randomizations within the subjects. We found that the brainstem responses to a 200 Hz pure tone consisted of prominent peaks at 200 Hz, and at frequencies corresponding to the harmonics of 200 Hz. The FFR spectral magnitude at 200 Hz diminished with a phase offset of 180°. Phase offsets of 90° and 270° showed reduced spectral magnitudes at 200 Hz than those in the 0° condition. Our findings, in line with the hypothesis, show that the dichotic FFRs do not reflect true dichotic processing and that they are contaminated during volume conduction. Additionally, we found harmonic distortion products (HDP) in the FFRs. We found that the response at 200 Hz and the 3rd HDP systematically varied with a change in phase of the stimulus, while the even HDPs (2nd and 4th) were phase-invariant. Based on our findings, and modeling FFRs using auditory models, we propose a rectification process as the contributors for the generation of HDPs. We also discuss the implications of this HDP generating mechanism in understanding the pitch represented in FFRs.

Many users of bilateral cochlear implants (BiCIs) localize sound sources less accurately than do people with normal hearing. This may be partly due to using two independently functioning CIs with fixed compression, which distorts and/or reduces interaural level differences (ILDs). Here, we investigate the potential benefits of using binaurally coupled, dynamic compression inspired by the medial olivocochlear reflex; an approach termed “the MOC strategy” (Lopez-Poveda et al., 2016, Ear Hear 37:e138-e148). Twelve BiCI users were asked to localize wideband (125–6000 Hz) noise tokens in a virtual horizontal plane. Stimuli were processed through a standard (STD) sound processing strategy (i.e., involving two independently functioning sound processors with fixed compression) and three different implementations of the MOC strategy: one with fast (MOC1) and two with slower contralateral control of compression (MOC2 and MOC3). The MOC1 and MOC2 strategies had effectively greater inhibition in the higher than in the lower frequency channels, while the MOC3 strategy had slightly greater inhibition in the lower than in the higher frequency channels. Localization was most accurate with the MOC1 strategy, presumably because it provided the largest and less ambiguous ILDs. The angle error improved slightly from 25.3° with the STD strategy to 22.7° with the MOC1 strategy. The improvement in localization ability over the STD strategy disappeared when the contralateral control of compression was made slower, presumably because stimuli were too short (200 ms) for the slower contralateral inhibition to enhance ILDs. Results suggest that some MOC implementations hold promise for improving not only speech-in-noise intelligibility, as shown elsewhere, but also sound source lateralization.

Accurate neural representations of acoustic signals under noisy conditions are critical for animals' survival. Detecting signal against background noise can be improved by binaural hearing particularly when an interaural-time-difference (ITD) disparity is introduced between the signal and the noise, a phenomenon known as binaural unmasking. Previous studies have mainly focused on the binaural unmasking effect on response magnitudes, and it is not clear whether binaural unmasking affects the accuracy of central representations of target acoustic signals and the relative contributions of different central auditory structures to this accuracy. Frequency following responses (FFRs), which are sustained phase-locked neural activities, can be used for measuring the accuracy of the representation of signals. Using intracranial recordings of local field potentials, this study aimed to assess whether the binaural unmasking effects include an improvement of the accuracy of neural representations of sound-envelope signals in the rat IC and/or auditory cortex (AC). The results showed that (1) when a narrow-band noise was presented binaurally, the stimulus-response (S-R) coherence of the FFRs to the envelope (FFRenvelope) of the narrow-band noise recorded in the IC was higher than that recorded in the AC. (2) Presenting a broad-band masking noise caused a larger reduction of the S-R coherence for FFRenvelope in the IC than that in the AC. (3) Introducing an ITD disparity between the narrow-band signal noise and the broad-band masking noise did not affect the IC S-R coherence, but enhanced both the AC S-R coherence and the coherence between the IC FFRenvelope and AC FFRenvelope. Thus, although the accuracy of representing envelope signals in the AC is lower than that in the IC, it can be binaurally unmasked, indicating a binaural-unmasking mechanism that is formed during the signal transmission from the IC to the AC.

Binaural hearing allows listeners to take advantage of auditory cues that are necessary for sound localization. Patients with asymmetric or unilateral hearing loss receive degraded or absent binaural cues and often demonstrate decreased sound localization in the horizontal plane. Given the importance of sound localization for everyday function, there is a need for clinically feasible methods to evaluate localization, particularly as validation of intervention for patients with unilateral hearing loss. The present study compared sound localization results obtained for front-facing and rear-facing soundfield systems and for a rear-facing head related transfer function (HRTF) system in two hearing groups, listeners with normal hearing and those with unilateral hearing loss. Results between the three systems were highly correlated and repeatable. Neither age nor a pure-tone average of 0.5, 1, 2 and 4 kHz correlated with localization performance among normal hearing participants. Among the UHL group, age and onset of severe to profound hearing loss were positively correlated with localization results; length of deafness was negatively correlated. Behavioral chance performance was established for localization with rear-facing HRTF and rear-facing soundfield test systems; group mean results for unilateral hearing loss participants were significantly better than chance. Continued work is needed in developing clinically feasible evaluation methods for patients with asymmetric or unilateral hearing loss, particularly as the effects of treatment options are assessed.

The main impairment associated with single-sided deafness (SSD) is the loss of binaural hearing. Currently, the most effective treatment to compensate for this deficit is to supply patients suffering from SSD with a cochlear implant (CI) in the deaf ear. With this approach binaural hearing abilities can be restored to a certain extent, which is expressed in an improvement in such patients with regard to sound source localization and speech comprehension in noise after receipt of a CI. However, binaural performance of these listeners does not reach the level of normal-hearing listeners. One of the reasons for this might be that the electrical stimulation via CI and the physiological stimulation via the intact ear are not synchronized: the CI transmits the information to the auditory nerve with different timing than does the intact inner ear. As a result, there is a timing mismatch of the information transfer between the left and the right side, which may account for the limited binaural performance. The effective mismatch in timing depends on the CI system because of different stimulation strategies implemented in devices from different manufacturers. For the particular CI device used in this study (MED-EL Mi1000/Mi1200) electrical stimulation led to faster activation of the auditory nerve than natural for a wide frequency range. In particular, electrical stimulation was about 1 to up to 2 ms ahead of time for frequencies above 1.5 kHz. Hence, it was hypothesized that information transfer between the left and the right ear can be tuned by delaying the CI signal. The goal of the present study was to investigate whether such a delay in the CI signal affects binaural performance of CI users with SSD. For this purpose, sound source localization and speech perception in noise were tested in a sample of 12 CI users with SSD (mean age 51 ± 12 years). The tests were performed for four different delay times of the CI signal applied spontaneously (0.5, 1, 2 and 4 ms) and for the base line condition “no delay” in the CI signal (i.e. everyday use). It was found that delaying the signal had a significant impact on sound source localization. Speech perception in noise was affected, but less pronounced than was sound localization. Regarding sound source localization, a signal delay of 1 ms applied to this particular CI device produced the best performance in our patients. It is concluded that improving the synchronisation between the CI-transferred signal and the naturally transferred signal could increase binaural hearing performance in CI users with SSD.

The role of the mammalian auditory olivocochlear efferent system in hearing has long been the subject of debate. Its ability to protect against damaging noise exposure is clear, but whether or not this is the primary function of a system that evolved in the absence of industrial noise remains controversial. Here we review the behavioral consequences of olivocochlear activation and diminished olivocochlear function. Attempts to demonstrate a role for hearing in noise have yielded conflicting results in both animal and human studies. A role in selective attention to sounds in the presence of distractors, or attention to visual stimuli in the presence of competing auditory stimuli, has been established in animal models, but again behavioral studies in humans remain equivocal. Auditory processing deficits occur in models of congenital olivocochlear dysfunction, but these deficits likely reflect abnormal central auditory development rather than direct effects of olivocochlear feedback. Additional proposed roles in age-related hearing loss, tinnitus, hyperacusis, and binaural or spatial hearing, are intriguing, but require additional study. These behavioral studies almost exclusively focus on medial olivocochlear effects, and many relied on lesioning techniques that can have unspecific effects. The consequences of lateral olivocochlear and of corticofugal pathway activation for perception remain unknown. As new tools for targeted manipulation of olivocochlear neurons emerge, there is potential for a transformation of our understanding of the role of the olivocochlear system in behavior across species.

Item does not contain fulltext Different amplification options are available for listeners with congenital unilateral conductive hearing loss (UCHL). For example, bone-conduction devices (BCDs) and middle ear implants. The present study investigated whether intervention with an active BCD, the Bonebridge, or a middle ear implant, the Vibrant Soundbridge (VSB), affected sound-localization performance of listeners with congenital UCHL. Listening with a Bonebridge or VSB might provide access to binaural cues. However, when fitted with the Bonebridge, but not with a VSB, binaural processing might be affected through cross stimulation of the contralateral normal hearing ear, and could interfere with processing of binaural cues. In the present study twenty-three listeners with congenital UCHL were included. To assess processing of binaural cues, we investigated localization abilities of broadband (BB, 0.5-20kHz) filtered noise presented at varying sound levels. Sound localization abilities were analyzed separately for stimuli presented at the side of the normal-hearing ear, and for stimuli presented at the side of the hearing-impaired ear. Twenty-six normal hearing children and young adults were tested as control listeners. Sound localization abilities were measured under open-loop conditions by recording head-movement responses. We demonstrate improved sound localization abilities of children with congenital UCHL, when listening with a Bonebridge or VSB, predominantly for stimuli presented at the impaired (aided) side. Our results suggest that the improvement is not related to accurate processing of binaural cues. When listening with the Bonebridge, despite cross stimulation of the contralateral cochlea, localization performance was not deteriorated compared to listening with a VSB. 10 p.

Interaural time and intensity differences (ITD and IID) are important cues in binaural hearing and allow for sound localization, improving speech understanding in noise and reverberation, and integrating sound sources in the auditory scene. Whereas previous research showed that the upper-frequency limit for ITD detection in the fine structure of sound declines in aging, the processing of envelope ITD in low-frequency amplitude modulated (AM) sound and the related brain responses are less understood. This study investigated the cortical processing of envelope ITD and compared the results with previous findings about the fine-structure ITD. In two experiments, participants listened to 40-Hz AM tones containing sudden changes in the envelope ITD. Multiple MEG responses were analyzed, including the auditory evoked N1 responses, elicited both by sound onsets and ITD changes, and 40-Hz responses, elicited by the AM. The first experiment with healthy young adults revealed a substantial decline in the magnitudes of the ITD change N1 response, and the 40-Hz phase resets at higher carrier frequencies, suggesting a similar frequency characteristic as observed for fine structure ITD. The amplitude of the 40-Hz ASSR declined only gradually with increasing carrier frequency, and it was excluded as a confounding factor in the decline in the ITD response. Larger responses to outward ITD changes than inward changes, here first reported for envelope ITD, were another characteristics that were similar to fine-structure ITD. A second experiment with groups of young and older listeners examined the effects of aging and concurrent noise on the cortical envelope ITD responses. One important research question was, whether binaural cues are accessible in noise. Behavioural tests showed an age-related hearing loss in the older group and decreased performance in envelope ITD detection and speech-in-noise (SIN) understanding. Binaural hearing and SIN performance were correlated with one other, but not with hearing loss. The frequency limit for envelope ITD was reduced in older listeners similarly as previously found for fine structure ITD, and older listeners were more susceptible to concurrent multi-talker noise. The similarities between responses to envelope ITD and to fine structure ITD suggest that a common cortical code exists for the envelope and fine structure ITD. The dependency on the carrier frequency suggests that envelope ITD processing at the subcortical level requires stimulus phase locking, which might be reduced in aging.

Over the years, the effect of aging on auditory function has been investigated in animal models and humans in an effort to characterize age-related changes in both perception and physiology. Here, we review how aging may impact neural encoding and processing of binaural and spatial cues in human listeners with a focus on recent work by the authors as well as others. Age-related declines in monaural temporal processing, as estimated from measures of gap detection and temporal fine structure discrimination, have been associated with poorer performance on binaural tasks that require precise temporal processing. In lateralization and localization tasks, as well as in the detection of signals in noise, marked age-related changes have been demonstrated in both behavioral and electrophysiological measures and have been attributed to declines in neural synchrony and reduced central inhibition with advancing age. Evidence for such mechanisms, however, are influenced by the task (passive vs. attending) and the stimulus paradigm (e.g., static vs. continuous with dynamic change). That is, cortical auditory evoked potentials (CAEP) measured in response to static interaural time differences (ITDs) are larger in older versus younger listeners, consistent with reduced inhibition, while continuous stimuli with dynamic ITD changes lead to smaller responses in older compared to younger adults, suggestive of poorer neural synchrony. Additionally, the distribution of cortical activity is broader and less asymmetric in older than younger adults, consistent with the hemispheric asymmetry reduction in older adults model of cognitive aging. When older listeners attend to selected target locations in the free field, their CAEP components (N1, P2, P3) are again consistently smaller relative to younger listeners, and the reduced asymmetry in the distribution of cortical activity is maintained. As this research matures, proper neural biomarkers for changes in spatial hearing can provide objective evidence of impairment and targets for remediation. Future research should focus on the development and evaluation of effective approaches for remediating these spatial processing deficits associated with aging and hearing loss.

While a large body of literature has examined the encoding of binaural spatial cues in the auditory midbrain, studies that ask how quantitative measures of spatial tuning in midbrain neurons compare with an animal's psychoacoustic performance remain rare. Researchers have tried to explain deficits in spatial hearing in certain patient groups, such as binaural cochlear implant users, in terms of declines in apparent reductions in spatial tuning of midbrain neurons of animal models. However, the quality of spatial tuning can be quantified in many different ways, and in the absence of evidence that a given neural tuning measure correlates with psychoacoustic performance, the interpretation of such finding remains very tentative. Here, we characterize ITD tuning in the rat inferior colliculus (IC) to acoustic pulse train stimuli with varying envelopes and at varying rates, and explore whether quality of tuning correlates behavioral performance. We quantified both mutual information (MI) and neural d' as measures of ITD sensitivity. Neural d' values paralleled behavioral ones, declining with increasing click rates or when envelopes changed from rectangular to Hanning windows, and they correlated much better with behavioral performance than MI. Meanwhile, MI values were larger in an older, more experienced cohort of animals than in naive animals, but neural d' did not differ between cohorts. However, the results obtained with neural d' and MI were highly correlated when ITD values were coded simply as left or right ear leading, rather than specific ITD values. Thus, neural measures of lateralization ability (e.g. d' or left/right MI) appear to be highly predictive of psychoacoustic performance in a two-alternative forced choice task.

For deaf patients cochlear implants (CIs) can restore substantial amounts of functional hearing. However, binaural hearing, and in particular, the perception of interaural time differences (ITDs) with current CIs has been found to be notoriously poor, especially in the event of early hearing loss. One popular hypothesis for these deficits posits that a lack of early binaural experience may be a principal cause of poor ITD perception in pre-lingually deaf CI patients. This is supported by previous electrophysiological studies done in neonatally deafened, bilateral CI-stimulated animals showing reduced ITD sensitivity. However, we have recently demonstrated that neonatally deafened CI rats can quickly learn to discriminate microsecond ITDs under optimized stimulation conditions which suggests that the inability of human CI users to make use of ITDs is not due to lack of binaural hearing experience during development. In the study presented here, we characterized ITD sensitivity and tuning of inferior colliculus neurons under bilateral CI stimulation of neonatally deafened and hearing experienced rats. The hearing experienced rats were not deafened prior to implantation. Both cohorts were implanted bilaterally between postnatal days 64-77 and recorded immediately following surgery. Both groups showed comparably large proportions of ITD sensitive multi-units in the inferior colliculus (Deaf: 84.8%, Hearing: 82.5%), and the strength of ITD tuning, quantified as mutual information between response and stimulus ITD, was independent of hearing experience. However, the shapes of tuning curves differed substantially between both groups. We observed four main clusters of tuning curves - trough, contralateral, central, and ipsilateral tuning. Interestingly, over 90% of multi-units for hearing experienced rats showed predominantly contralateral tuning, whereas as many as 50% of multi-units in neonatally deafened rats were centrally tuned. However, when we computed neural d' scores to predict likely limits on performance in sound lateralization tasks, we did not find that these differences in tuning shapes predicted worse psychoacoustic performance for the neonatally deafened animals. We conclude that, at least in rats, substantial amounts of highly precise, "innate" ITD sensitivity can be found even after profound hearing loss throughout infancy. However, ITD tuning curve shapes appear to be strongly influenced by auditory experience although substantial lateralization encoding is present even in its absence.

Previous work has explored novel binaural combinations of reverberation and the resulting perceived reverberation strength (reverberance). The present study examines the perceptual effects of additional binaural combinations of reverberation with the goal of explaining reverberance in terms of basic psychoacoustic principles. Stimuli were generated using virtual space techniques simulating a speech source 3 m to the listener's right in a moderately reverberant environment. Reverberant energy at the ears was varied systematically relative to the natural level for the environment (0-dB gain). The method of magnitude estimation was used to estimate reverberance. Four experiments were conducted. Experiment 1 tested monaural listening conditions for both left and right ears at reverberation gains from -21 dB to 0 dB. Experiment 2 tested a binaural listening condition where only reverberant energy at the ear farther from the source was manipulated (-21 dB to 0 dB). Experiment 3 tested two binaural conditions over a wider range of reverberation gains (-18 dB to +24 dB). In one condition, reverberant energy was manipulated for both ears equally. In the other condition, reverberant energy was manipulated only for the ear nearer the source. In Experiment 4, reverberant tails of the stimuli were removed to test whether listeners were able to use ongoing reverberant information to judge reverberance. The results from all experiments were found to be well predicted by a model of time-varying binaural loudness that focused on “glimpses” in time with relatively high reverberant sound energy and low direct sound energy. These findings suggest that the mechanisms underlying reverberance and loudness may be similar.

For speech in competition with a noise source in the free field, normal-hearing (NH) listeners recognize speech better when listening binaurally than when listening monaurally with the ear that has the better acoustic signal-to-noise ratio (SNR). This benefit from listening binaurally is known as binaural unmasking and indicates that the brain combines information from the two ears to improve intelligibility. Here, we address three questions pertaining to binaural unmasking for NH listeners. First, we investigate if binaural unmasking results from combining the speech and/or the noise from the two ears. In a simulated acoustic free field with speech and noise sources at 0° and 270°azimuth, respectively, we found comparable unmasking regardless of whether the speech was present or absent in the ear with the worse SNR. This indicates that binaural unmasking probably involves combining only the noise at the two ears. Second, we investigate if having binaurally coherent location cues for the noise signal is sufficient for binaural unmasking to occur. We found no unmasking when location cues were coherent but noise signals were generated incoherent or were processed unilaterally through a hearing aid with linear, minimal amplification. This indicates that binaural unmasking requires interaurally coherent noise signals, source location cues, and processing. Third, we investigate if the hypothesized antimasking benefits of the medial olivocochlear reflex (MOCR) contribute to binaural unmasking. We found comparable unmasking regardless of whether speech tokens (words) were sufficiently delayed from the noise onset to fully activate the MOCR or not. Moreover, unmasking was absent when the noise was binaurally incoherent whereas the physiological antimasking effects of the MOCR are similar for coherent and incoherent noises. This indicates that the MOCR is unlikely involved in binaural unmasking.

Pitch is an essential cue that allows the auditory system to distinguish between sound sources. Pitch cues are less useful when listeners are not able to discriminate different pitches between the two ears, a problem encountered by listeners with hearing impairment (HI). Many listeners with HI will fuse the pitch of two dichotically presented tones over a larger range of interaural frequency disparities, i.e., have a broader fusion range, than listeners with normal hearing (NH). One potential explanation for broader fusion in listeners with HI is that hearing aids stimulate at high sound levels. The present study investigated effects of overall sound levels on pitch fusion in listeners with NH. It was hypothesized that if sound level increased, then fusion range would increase. Fusion ranges were measured by presenting a fixed frequency tone to a reference ear simultaneously with a variable frequency tone to the opposite ear and finding the range of frequencies that were fused with the reference frequency. No significant effects of sound level (comfortable level ± 15 decibels) on fusion range were found, even when tested within the range of levels where some listeners with HI show large fusion ranges. Results suggest that increased sound level does not explain increased fusion range in listeners with HI and imply that other factors associated with hearing loss might play a larger role.

Directing gaze towards auditory events is a natural behavior. In addition to the well-known accuracy of auditory elicited gaze responses for normal binaural listening, their latency is a measure of possible clinical interest and methodological importance. The aim was to develop a clinically feasible method to assess sound localization latency (SLL), and to study SLL as a function of simulated unilateral hearing loss (SUHL) and the relationship with accuracy. Eight healthy and normal-hearing adults (18–40 years) participated in this study. Horizontal gaze responses, recorded by non-invasive corneal reflection eye-tracking, were obtained during azimuthal shifts (24 trials) of a 3-min continuous auditory stimulus. In each trial, a sigmoid function was fitted to gaze samples. Latency was estimated by the abscissa corresponding to 50% of the arctangent amplitude. SLL was defined as the mean latency across trials. SLL was measured in normal-hearing and simulated SUHL conditions (SUHL30 and SUHL43: mean threshold of 30 dB HL and 43 dB HL across 0.5, 1, 2, and 4 kHz). In the normal-hearing condition, the mean ± SD SLL was 280 ± 40 ms (n = 8) with a test-retest SD = 20 ms. A linear mixed model showed a statistically significant effect of listening condition on SLL. The SUHL30 and SUHL43 conditions revealed a mean SLL of 370 ± 49 ms and 540 ± 120 ms, respectively. Repeated measures correlation analysis showed a clear relationship between SLL and the average sound localization accuracy (R2 = 0.94). The rapid and reliable method to obtain SLL may be an important clinical tool for evaluation of binaural processing. Future studies in clinical cohorts are needed to assess whether SLL may reveal information about binaural processing abilities beyond that afforded by sound localization accuracy.

Auditory research has a rich history of combining experimental evidence with computational simulations of auditory processing in order to deepen our theoretical understanding of how sound is processed in the ears and in the brain. Despite significant progress in the amount of detail and breadth covered by auditory models, for many components of the auditory pathway there are still different model approaches that are often not equivalent but rather in conflict with each other. Similarly, some experimental studies yield conflicting results which has led to controversies. This can be best resolved by a systematic comparison of multiple experimental data sets and model approaches. Binaural processing is a prominent example of how the development of quantitative theories can advance our understanding of the phenomena, but there remain several unresolved questions for which competing model approaches exist. This article discusses a number of current unresolved or disputed issues in binaural modelling, as well as some of the significant challenges in comparing binaural models with each other and with the experimental data. We introduce an auditory model framework, which we believe can become a useful infrastructure for resolving some of the current controversies. It operates models over the same paradigms that are used experimentally. The core of the proposed framework is an interface that connects three components irrespective of their underlying programming language: The experiment software, an auditory pathway model, and task-dependent decision stages called artificial observers that provide the same output format as the test subject.

The auditory "octave illusion" arises when dichotic tones, presented one octave apart, alternate rapidly between the ears. This study aimed to explore the link between the perception of illusory pitches and brain activity during presentation of dichotic tones. We conducted a behavioral study of how participants perceived binaural dichotic tones of octave illusions and classified them, based on the reported percepts, in an illusion (ILL) group, without an illusion (non-ILL) group, and others. We recorded brain activity using functional magnetic resonance imaging and analyzed the activation due to dichotic illusion tones. The activation in the bilateral planum polare in the auditory cortex was significantly larger in the ILL group than in the non-ILL group. In the right premotor cortex, the non-ILL group showed a significantly larger activation than did the ILL group, suggesting that the sensation of the meter to the stimulus sound was significant in the non-ILL but not in the ILL group. The results indicated that the activity in these areas was related to the occurrence of octave illusions. The nonsignificant sensation of the meter to the stimulus sound in the ILL group may be consistent with the perception of octave illusion.

Complex broadband sounds are decomposed by peripheral auditory filters into a series of relatively narrowband signals, each with a slowly varying envelope (ENV) and a rapidly fluctuating temporal fine structure (TFS). ENV and TFS information at the bilateral ears contribute differentially to auditory perception. However, whether the difference could attribute to mechanisms of binaural integration remains an open question. As a potential neural correlate, subsets of neurons in the central nucleus of the inferior colliculus (ICC) are known to integrate binaural information with binaural inhibition or binaural summation. Therefore, we recorded the frequency-following responses (FFRs) to the ENV and TFS components of narrowband noises in the ICC of anesthetized rats and examined changes in FFR amplitude and stimulus-response coherence under various sound-delivery settings. We showed that binaural FFRENV was predominantly elicited by contralateral inputs and inhibited by ipsilateral inputs, exhibiting a "binaural-inhibition" like property. On the other hand, binaural FFRTFS received a balanced contribution from both sides, echoing the "binaural-summation" mechanism. What is more, binaural FFRENV was significantly correlated with contralateral-evoked but not ipsilateral-evoked FFRENV, while binaural FFRTFS correlated with both contralateral- and ipsilateral-evoked FFRTFS. Overall, these results suggest distinct binaural processing of ENV and TFS information at the midbrain level.

The morphology of the head and pinna shape the spatial and frequency dependence of sound propagation that give rise to the acoustic cues to sound source location. During early development, the physical dimensions of the head and pinna increase rapidly. Thus, the binaural (interaural time and level differences, ITD and ILD) and monaural (spectral shape) cues are also hypothesized to change rapidly. Complex interactions between the size and shape of the head and pinna limit the accuracy of simple acoustical models (e.g. spherical) and necessitate empirical measurements. Here, we measured the cues to location in the developing guinea pig, a precocial species commonly used for studies of the auditory system. We measured directional transfer functions (DTFs) and the dimensions of the head and pinna in guinea pigs from birth (P0) through adulthood. Dimensions of the head and pinna increased by 87% and 48%, respectively, reaching adult values by ~8 weeks (P56). The monaural acoustic gain produced by the head and pinna increased with frequency and age, with maximum gains at higher frequencies (>8 kHz) reaching values of 10–21 dB for all ages. The center frequency of monaural spectral notches also decreased with age, from higher frequencies (~17 kHz) at P0 to lower frequencies (~12 kHz) in adults. In all animals, ILDs and ITDs were dependent on both frequency and spatial location. Over development, the maximum ILD magnitude increased from ~15 dB at P0 to ~30 dB in adults (at frequencies >8 kHz), while the maximum low frequency ITDs increased from ~185 µs at P0 to ~300 µs in adults. These results demonstrate that the changes in the acoustical cues are directly related to changes in head and pinna morphology.

Binaural hearing confers many beneficial functions but our understanding of its underlying neural substrates is limited. This study examines the bilateral synaptic assemblies and binaural computation (or integration) in the central nucleus of the inferior colliculus (ICc) of the auditory midbrain, a key convergent center. Using in-vivo whole-cell patch-clamp, the excitatory and inhibitory postsynaptic potentials (EPSPs/IPSPs) of single ICc neurons to contralateral, ipsilateral and bilateral stimulation were recorded. According to the contralateral and ipsilateral EPSP/IPSP, 7 types of bilateral synaptic assemblies were identified. These include EPSP-EPSP (EE), E-IPSP (EI), E-no response (EO), II, IE, IO and complex-mode (CM) neurons. The CM neurons showed frequency- and/or amplitude-dependent EPSPs/IPSPs to contralateral or ipsilateral stimulation. Bilateral stimulation induced EPSPs/IPSPs that could be larger than (facilitation), similar to (ineffectiveness) or smaller than (suppression) those induced by contralateral stimulation. Our findings have allowed our group to characterize novel neural circuitry for binaural computation in the midbrain.

Subtracting the sum of left and right monaural auditory brainstem responses (ABRs) from the corresponding binaural ABR isolates the binaural interaction component (ABR-BIC). In a previous investigation (Ikeda, 2015), during auditory yet not visual tasks, tone-pips elicited a significant difference in amplitude between summed monaural and binaural ABRs. With click stimulation, this amplitude difference was task-independent. This self-critical reanalysis's purpose was to establish that a difference waveform (i.e., ABR-BIC DN1) reflected an auditory selective attention effect that was isolable from stimulus factors. Regardless of whether stimuli were tone-pips or clicks, effect sizes of the DN1 peak amplitudes relative to zero improved during auditory tasks over visual tasks. Auditory selective attention effects on the monaural and binaural ABR wave-V amplitudes were tone-pip specific. Those wave-V effects thus could not explain the stimulus-universal effect of auditory selective attention on DN1 detectability, which was thus entirely binaural. In a manner isolated from auditory selective attention, multiple mediation analyses indicated that the higher right monaural wave-V amplitudes mediated individual differences in how clicks, relative to tone-pips, augmented DN1 amplitudes. There are implications of these findings for advancing ABR-BIC measurement. acceptedVersion

Cochlear implant (CI) users find it hard and effortful to understand speech in noise with current devices. Binaural CI sound processing inspired by the contralateral medial olivocochlear (MOC) reflex (an approach termed the ‘MOC strategy’) can improve speech-in-noise recognition for CI users. All reported evaluations of this strategy, however, disregarded automatic gain control (AGC) and fine‐structure (FS) processing, two standard features in some current CI devices. To better assess the potential of implementing the MOC strategy in contemporary CIs, here, we compare intelligibility with and without MOC processing in combination with linked AGC and FS processing. Speech reception thresholds (SRTs) were compared for an FS and a MOC-FS strategy for sentences in steady and fluctuating noises, for various speech levels, in bilateral and unilateral listening modes, and for multiple spatial configurations of the speech and noise sources. Word recall scores and verbal response times in a word recognition test (two proxies for listening effort) were also compared for the two strategies in quiet and in steady noise at 5 dB signal-to-noise ratio (SNR) and the individual SRT. In steady noise, mean SRTs were always equal or better with the MOC-FS than with the standard FS strategy, both in bilateral (the mean and largest improvement across spatial configurations and speech levels were 0.8 and 2.2 dB, respectively) and unilateral listening (mean and largest improvement of 1.7 and 2.1 dB, respectively). In fluctuating noise and in bilateral listening, SRTs were equal for the two strategies. Word recall scores and verbal response times were not significantly affected by the test SNR or the processing strategy. Results show that MOC processing can be combined with linked AGC and FS processing. Compared to using FS processing alone, combined MOC-FS processing can improve speech intelligibility in noise without affecting word recall scores or verbal response times.

We have recently proposed a binaural cochlear implant (CI) sound processing strategy inspired by the contralateral medial olivocochlear reflex (the MOC strategy) and shown that it improves intelligibility in steady-state noise (Lopez-Poveda et al., 2016, Ear Hear 37:e138-e148). The aim here was to evaluate possible speech-reception benefits of the MOC strategy for speech maskers, a more natural type of interferer. Speech reception thresholds (SRTs) were measured in six bilateral and two single-sided deaf CI users with the MOC strategy and with a standard (STD) strategy. SRTs were measured in unilateral and bilateral listening conditions, and for target and masker stimuli located at azimuthal angles of (0°, 0°), (−15°, +15°), and (−90°, +90°). Mean SRTs were 2–5 dB better with the MOC than with the STD strategy for spatially separated target and masker sources. For bilateral CI users, the MOC strategy (1) facilitated the intelligibility of speech in competition with spatially separated speech maskers in both unilateral and bilateral listening conditions; and (2) led to an overall improvement in spatial release from masking in the two listening conditions. Insofar as speech is a more natural type of interferer than steady-state noise, the present results suggest that the MOC strategy holds potential for promising outcomes for CI users.

Collin and Lavandier [J. Acoust. Soc. Am. 134, 1146–1159 (2013)] proposed a binaural model predicting speech intelligibility against envelope-modulated noises, evaluated in 24 acoustic conditions, involving similar masker types. The aim of the present study was to test the model robustness modeling 80 additional conditions, and evaluate the influence of its parameters using an approach inspired by a variance-based sensitivity analysis. First, the data from four experiments from the literature and one specifically designed for the present study were used to evaluate the prediction performance of the model, investigate potential interactions between its parameters, and define their values leading to the best predictions. A revision of the model allowed to account for binaural sluggishness. Finally, the optimized model was tested on an additional dataset not used to define its parameters. Overall, one hundred conditions split into six experiments were modeled. Correlation between data and predictions ranged from 0.85 to 0.96 across experiments, and mean absolute prediction errors were between 0.5 and 1.4 dB.

Loudspeaker-based virtual sound environments provide a valuable tool for studying speech perception in realistic, but controllable and reproducible acoustic environments. The evaluation of different loudspeaker reproduction methods with respect to perceptual measures has been rather limited. This study focused on comparing speech intelligibility as measured in a reverberant reference room with virtual versions of that room. Two reproduction methods were based on room acoustic simulations, presented either using mixed-order ambisonics or nearest loudspeaker mapping playback. The third method utilized impulse responses measured with a spherical microphone array and mixed-order ambisonics. Three factors that affect speech intelligibility were varied: reverberation, the spatial configuration and the type of the interferers (speech or noise). Two interferers were placed either colocated with the target, or were symmetrically or asymmetrically separated. The results showed differences between the reference room and the simulation-based reproductions when the target and the interferers were spatially separated but not when they were colocated. The reproduction utilizing the microphone array was most similar to the reference room in terms of measured speech intelligibility. Differences in speech intelligibility could be accounted for using a binaural speech intelligibility model which considers better-ear signal-to-noise ratio differences and binaural unmasking effects. Thus, auditory modeling might be a fast and efficient way to evaluate virtual sound environments.

Asymmetric hearing has been the focus of many studies on brain plasticity in the past. Recently, the topic has gained clinical importance in cases with sequential cochlear implantation or in cases with deafness in one ear and preserved hearing in the other ear. Convergent evidence from animal experiments and from hearing impaired children suggest that asymmetric hearing during early development can reorganize the central auditory representation of the two ears with the consequence of a "stronger" representation of the better hearing ear with a "weaker" representation of the other, more poorly hearing, ear. These changes lead to a persistent aural preference for one ear, demonstrated by asymmetric speech comprehension when each ear is tested separately. Further, binaural integration is compromised as shown by reduced binaural fusion and reduced sensitivity for binaural cues. Finally, the data demonstrate a significant difference in cortical plasticity in response to juvenile monocular deprivation in the visual system and juvenile monaural deafness in the auditory system. The topic represents a unique example of translational research whereby mechanisms explored in animal models are combined with data from children to understand the clinical consequences of asymmetric hearing in early development.

Many neurons in the auditory midbrain are tuned to binaural cues. Two prominent binaural cues are the interaural level difference (ILD) and the interaural time difference (ITD). The ITD cue can further be subdivided into the ongoing envelope ITD cues and transient onset ITD cues. More is known about the sensitivity of single neurons to ongoing envelope ITDs compared to transient onset ITDs in the mammalian auditory system, particularly in bats. The current study examines the response properties of single neurons in the inferior colliculus (IC) of the big brown bat, Eptesicus fuscus, to onset ITDs in response to high frequency pure tones. Measures of neurons’ dynamic ITD response revealed an average change of 36% of its maximum response within the behaviorally relevant range of ITDs (±50 µs). Across all IC neurons, we measured an average time-intensity trading ratio of 30 µs/dB in the sensitivity of the ITD response function to changing ILDs. Minimum and maximum ITD responses were clustered within a narrow range of ITDs. The average peak in the ITD response function was at 268 µs, a finding that is consistent with other non-echolocating mammals. Some ITD-sensitive neurons also showed weak facilitation of maximum response during binaural stimulation, compared to monaural stimulation. These results suggest that echolocating bats possess the potential to use onset ITD cues to assist in the azimuthal sound localization of ultrasonic frequencies.

Spatial hearing skills are essential for children as they grow, learn and play. These skills provide critical cues for determining the locations of sources in the environment, and enable segregation of important sounds, such as speech, from background maskers or interferers. Spatial hearing depends on availability of monaural cues and binaural cues. The latter result from integration of inputs arriving at the two ears from sounds that vary in location. The binaural system has exquisite mechanisms for capturing differences between the ears in both time of arrival and intensity. The major cues that are thus referred to as being vital for binaural hearing are: interaural differences in time (ITDs) and interaural differences in levels (ILDs). In children with normal hearing (NH), spatial hearing abilities are fairly well developed by age 4-5 years. In contrast, most children who are deaf and hear through cochlear implants (CIs) do not have an opportunity to experience normal, binaural acoustic hearing early in life. These children may function by having to utilize auditory cues that are degraded with regard to numerous stimulus features. In recent years there has been a notable increase in the number of children receiving bilateral CIs, and evidence suggests that while having two CIs helps them function better than when listening through a single CI, these children generally perform worse than their NH peers. This paper reviews some of the recent work on bilaterally implanted children. The focus is on measures of spatial hearing, including sound localization, release from masking for speech understanding in noise and binaural sensitivity using research processors. Data from behavioral and electrophysiological studies are included, with a focus on the recent work of the authors and their collaborators. The effects of auditory plasticity and deprivation on the emergence of binaural and spatial hearing are discussed along with evidence for reorganized processing from both behavioral and electrophysiological studies. The consequences of both unilateral and bilateral auditory deprivation during development suggest that the relevant set of issues is highly complex with regard to successes and the limitations experienced by children receiving bilateral cochlear implants. This article is part of a Special Issue entitledAnnual Reviews 2016.

The effect of interaural time difference (ITD) and interaural level difference (ILD) on wave 4 of the binaural and summed monaural auditory brainstem responses (ABRs) as well as on the DN1 component of the binaural interaction component (BIC) of the ABR in young and old Mongolian gerbils (Meriones unguiculatus) was investigated. Measurements were made at a fixed sound pressure level (SPL) and a fixed level above visually detected ABR threshold to compensate for individual hearing threshold differences. In both stimulation modes (fixed SPL and fixed level above visually detected ABR threshold) an effect of ITD on the latency and the amplitude of wave 4 as well as of the BIC was observed. With increasing absolute ITD values BIC latencies were increased and amplitudes were decreased. ILD had a much smaller effect on these measures. Old animals showed a reduced amplitude of the DN1 component. This difference was due to a smaller wave 4 in the summed monaural ABRs of old animals compared to young animals whereas wave 4 in the binaural-evoked ABR showed no age-related difference. In old animals the small amplitude of the DN1 component was correlated with small binaural-evoked wave 1 and wave 3 amplitudes. This suggests that the reduced peripheral input affects central binaural processing which is reflected in the BIC.

Different fields of the auditory cortex can be distinguished by the extent and level tolerance of spatial selectivity. The mechanisms underlying the range of spatial tuning properties observed across cortical fields are unclear. Here, this issue was addressed in the pallid bat because its auditory cortex contains two segregated regions of response selectivity that serve two different behaviors: echolocation for obstacle avoidance and localization of prey-generated noise. This provides the unique opportunity to examine mechanisms of spatial properties in two functionally distinct regions. Previous studies have shown that spatial selectivity of neurons in the region selective for noise (noise-selective region, NSR) is level tolerant and shaped by interaural level difference (ILD) selectivity. In contrast, spatial selectivity of neurons in the echolocation region ('FM sweep-selective region' or FMSR) is strongly level dependent with many neurons responding to multiple distinct spatial locations for louder sounds. To determine the mechanisms underlying such level dependence, frequency, azimuth, rate-level responses and ILD selectivity were measured from the same FMSR neurons. The majority (∼75%) of FMSR neurons were monaural (ILD insensitive). Azimuth tuning curves expanded or split into multiple peaks with increasing sound level in a manner that was predicted by the rate-level response of neurons. These data suggest that azimuth selectivity of FMSR neurons depends more on monaural ear directionality and rate-level responses. The pallid bat cortex utilizes segregated monaural and binaural regions to process echoes and prey-generated noise. Together the pallid bat FMSR/NSR data provide mechanistic explanations for a broad range of spatial tuning properties seen across species.

Sensorineural hearing loss typically results in a steepened loudness function and a reduced dynamic range from elevated thresholds to uncomfortably loud levels for narrowband and broadband signals. Restoring narrowband loudness perception for hearing-impaired (HI) listeners can lead to overly loud perception of broadband signals and it is unclear how binaural presentation affects loudness perception in this case. Here, loudness perception quantified by categorical loudness scaling for nine normal-hearing (NH) and ten HI listeners was compared for signals with different bandwidth and different spectral shape in monaural and in binaural conditions. For the HI listeners, frequency- and level-dependent amplification was used to match the narrowband monaural loudness functions of the NH listeners. The average loudness functions for NH and HI listeners showed good agreement for monaural broadband signals. However, HI listeners showed substantially greater loudness for binaural broadband signals than NH listeners: on average a 14.1 dB lower level was required to reach ""very loud"" (range 30.8 to −3.7 dB). Overall, with narrowband loudness compensation, a given binaural loudness for broadband signals above ""medium loud"" was reached at systematically lower levels for HI than for NH listeners. Such increased binaural loudness summation was not found for loudness categories below ""medium loud"" or for narrowband signals. Large individual variations in the increased loudness summation were observed and could not be explained by the audiogram or the narrowband loudness functions.

Blind people use auditory information to locate sound sources and sound-reflecting objects (echolocation). Sound source localization benefits from the hearing system's ability to suppress distracting sound reflections, whereas echolocation would benefit from “unsuppressing” these reflections. To clarify how these potentially conflicting aspects of spatial hearing interact in blind versus sighted listeners, we measured discrimination thresholds for two binaural location cues: inter-aural level differences (ILDs) and inter-aural time differences (ITDs). The ILDs or ITDs were present in single clicks, in the leading component of click pairs, or in the lagging component of click pairs, exploiting processes related to both sound source localization and echolocation. We tested 23 blind (mean age = 54 y), 23 sighted-age-matched (mean age = 54 y), and 42 sighted-young (mean age = 26 y) listeners. The results suggested greater ILD sensitivity for blind than for sighted listeners. The blind group's superiority was particularly evident for ILD-lag-click discrimination, suggesting not only enhanced ILD sensitivity in general but also increased ability to unsuppress lagging clicks. This may be related to the blind person's experience of localizing reflected sounds, for which ILDs may be more efficient than ITDs. On the ITD-discrimination tasks, the blind listeners performed better than the sighted age-matched listeners, but not better than the sighted young listeners. ITD sensitivity declines with age, and the equal performance of the blind listeners compared to a group of substantially younger listeners is consistent with the notion that blind people's experience may offset age-related decline in ITD sensitivity.

Bilateral acoustic hearing in cochlear implant (CI) recipients with hearing preservation may allow access to binaural cues. Sensitivity to acoustic binaural cues has been shown in some listeners combining electric and acoustic stimulation (EAS), yet remains poorly understood and may be subject to limitations imposed by the electrical stimulation and/or amplification asymmetries. The purpose of this study was to investigate the effect of stimulus level, frequency-dependent gain, and the addition of unilateral electrical stimulation on sensitivity to low-frequency binaural cues. Thresholds were measured for interaural time and level differences (ITD and ILD) carried by a low-frequency, bandpass noise (100–800 Hz). 16 adult CI EAS listeners (mean age = 50.2 years) each participated in three listening conditions: acoustic hearing only at 90 dB SPL, acoustic hearing only at 60 dB SPL with frequency-dependent gain, and acoustic hearing plus unilateral CI at 60 dB SPL with frequency-dependent gain applied to the acoustic channels only. Results revealed thresholds within the ecologically relevant ITD and/or ILD range for most EAS listeners. No significant effects of presentation level, frequency-dependent gain, or the addition of unilateral electrical stimulation on the resultant thresholds for ITDs or ILDs were observed at the group level. Correlational analyses related ITD and ILD thresholds to the degree of EAS benefit (i.e., advantage of acoustic hearing in the implanted ear) for speech recognition in diffuse noise. There was a significant relationship between EAS benefit and ITD thresholds, but no statistically significant relationship between EAS benefit and ILD thresholds. In summary, the results of this study are not consistent with our previous data obtained with simulated EAS in normal-hearing listeners, which showed significant binaural interference by a unilateral electrical “distractor” (Van Ginkel et al., 2019). The difference between studies suggests that chronic exposure to unilateral electrical stimulation combined with bilateral acoustic stimulation may reduce interference effects, perhaps because listeners adapt to the presence of the constant but binaurally incongruous CI stimulus. These results are consistent with past studies that demonstrated no interference in spatial hearing tasks due to the addition of a unilateral CI in adult EAS listeners.

This study aimed to characterize binaural hearing abilities with bone conduction stimulation in simulated conductive hearing loss. Bone conduction hearing devices (BCDs) are a common method of rehabilitating conductive hearing loss. However, little is known about the access these devices provide to binaural cues. To study the ability of BCDs to restore access to binaural cues in conductive loss, normal hearing listeners were plugged unilaterally and bilaterally and localization ability was assessed using a non-surgical BCD attached to the mastoid/s via an adhesive (MED-EL, Corp). The results demonstrate that 1) application of the BCD in simulated unilateral conductive hearing loss does not restore access to binaural cues, evidenced by poor localization abilities. 2) bilateral application of BCDs in simulated bilateral conductive hearing loss provides access to binaural cues, 2) unilateral application of BCDs in simulated bilateral conductive hearing loss disrupts these cues and impairs localization performance, The transcutaneous stimulation of the adhesive BCD resulted in decreased access to sound compared to the normal open ear, resulting in asymmetries in aided versus non-aided hearing thresholds. Symmetrical hearing results in improved localization abilities, while asymmetric hearing disrupts sound localization abilities.

Binaural integration of interaural temporal information is essential for sound source localization and segregation. Current models of binaural interaction have shown that accurate sound localization in the horizontal plane depends on the resolution of phase ambiguous information by across-frequency integration. However, as such models are mostly static, it is not clear how proximate in time binaural events in different frequency channels should occur to form an auditory object with a unique lateral position. The present study examined the spectrotemporal window required for effective integration of binaural cues across frequency to form the perception of a stationary position. In Experiment 1, listeners judged whether dichotic frequency-modulated (FM) sweeps with a constant large nominal interaural delay (1500 μs), whose perceived laterality was ambiguous depending on the sweep rate (1500, 3000, 6000, and 12,000 Hz/s), produced a percept of continuous motion or a stationary image. Motion detection performance, indexed by d-prime (d') values, showed a clear effect of sweep rate, with auditory motion effects most pronounced for low sweep rates, and a punctate stationary image at high rates. Experiment 2 examined the effect of modulation rate (0.5, 3, 20, and 50 Hz) on lateralizing sinusoidally frequency-modulated (SFM) tones to confirm the effect of sweep rate on motion detection, independent of signal duration. Lateralization accuracy increased with increasing modulation rate up to 20 Hz and saturated at 50 Hz, with poorest performance occurring below 3 Hz depending on modulator phase. Using the transition point where percepts changed from motion to stationary images, we estimated a spectrotemporal integration window of approximately 150 ms per octave required for effective integration of interaural temporal cues across frequency channels. A Monte Carlo simulation based on a cross-correlation model of binaural interaction predicted 90% of the variance on perceptual motion detection performance as a function of FM sweep rate. Findings suggest that the rate of frequency channel convergence of binaural cues is essential to binaural lateralization.

Despite the common use of guinea pigs in investigations of the neural mechanisms of binaural and spatial hearing, their behavioral capabilities in spatial hearing tasks have surprisingly not been thoroughly investigated. To begin to fill this void, we tested the spatial hearing of adult male guinea pigs in several experiments using a paradigm based on the prepulse inhibition (PPI) of the acoustic startle response. In the first experiment, we presented continuous broadband noise from one speaker location and switched to a second speaker location (the “prepulse”) along the azimuth prior to presenting a brief, ∼110 dB SPL startle-eliciting stimulus. We found that the startle response amplitude was systematically reduced for larger changes in speaker swap angle (i.e., greater PPI), indicating that using the speaker “swap” paradigm is sufficient to assess stimulus detection of spatially separated sounds. In a second set of experiments, we swapped low- and high-pass noise across the midline to estimate their ability to utilize interaural time- and level-difference cues, respectively. The results reveal that guinea pigs can utilize both binaural cues to discriminate azimuthal sound sources. A third set of experiments examined spatial release from masking using a continuous broadband noise masker and a broadband chirp signal, both presented concurrently at various speaker locations. In general, animals displayed an increase in startle amplitude (i.e., lower PPI) when the masker was presented at speaker locations near that of the chirp signal, and reduced startle amplitudes (increased PPI) indicating lower detection thresholds when the noise was presented from more distant speaker locations. In summary, these results indicate that guinea pigs can: 1) discriminate changes in source location within a hemifield as well as across the midline, 2) discriminate sources of low- and high-pass sounds, demonstrating that they can effectively utilize both low-frequency interaural time and high-frequency level difference sound localization cues, and 3) utilize spatial release from masking to discriminate sound sources. This report confirms the guinea pig as a suitable spatial hearing model and reinforces prior estimates of guinea pig hearing ability from acoustical and physiological measurements.

This longitudinal study aimed to identify auditory plasticity promoted by a cochlear implant in children with asymmetric hearing loss. Participants included 10 children who experienced (mean ± SD) 3.1 ± 3.6 years of asymmetric hearing (difference of 47.2 ± 47.6 dB) before receiving an implant at age 8.7 ± 5.1 years. Multi-channel electroencephalography was measured at initial implant use (5.8 ± 3.2 days) and after 10.2 ± 4.1 months in each child. Monaurally presented stimuli consisted of 36 ms trains of 9 acoustic clicks/biphasic electric pulses at a rate of 250 Hz, repeated at 1 Hz. The time-restricted artifact and coherent source suppression (TRACS) beamformer was used to locate sources underlying peak amplitudes of cortical responses. Results indicated consistent activity from the non-implanted ear but significant implant-driven changes to the auditory cortices. Initially, the newly implanted ear evoked activity which strongly lateralized to the ipsilateral auditory cortex and contributed to a significant aural preference for implant stimulation in children with limited acoustic experience pre-implantation. Cochlear implant use reversed these abnormalities, but the resolution was limited in children with longer periods of asymmetric hearing. These findings suggest that early implantation of children with asymmetric hearing rapidly restores hemispheric representations of bilateral auditory input in the auditory cortex. Most recorded changes were isolated to pathways stimulated by the cochlear implant, potentially reflecting an abnormal independence of the bilateral pathways with possible consequences for binaural integration in these bimodal listeners.

Binaural interaction in the auditory brainstem response (ABR) represents the discrepancy between the binaural waveform and the sum of monaural ones. A typical ABR binaural interaction in humans is a reduction of the binaural amplitude compared to the monaural sum at the wave-V latency, i.e., the DN1 component. It has been considered that the DN1 is mainly elicited by high frequency components of stimuli whereas some studies have shown the contribution of low-to-middle frequency components to the DN1. To examine this issue, the present study compared the ABR binaural interaction elicited by tone pips (1 kHz, 10-ms duration) with the one by clicks (a rectangular wave, 0.1-ms duration) presented at 80 dB peak equivalent SPL and a fixed stimulus onset interval (180 ms). The DN1 due to tone pips was vulnerable compared to the click-evoked DN1. The pip-evoked DN1 was significantly detected under auditory attention whereas it failed to reach significance under visual attention. The click-evoked DN1 was robustly present for the two attention conditions. The current results might confirm the high frequency sound contribution to the DN1 elicitation.

Under certain conditions, sighted and blind humans can use echoes to discern characteristics of otherwise silent objects. Previous research concluded that robust horizontal-plane object localisation ability, without using head movement, depends on information above 2 kHz. While a strong interaural level difference (ILD) cue is available, it was not clear if listeners were using that or the monaural level cue that necessarily accompanies ILD. In this experiment, 13 sighted and normal-hearing listeners were asked to identify the right-vs.-left position of an object in virtual auditory space. Sounds were manipulated to remove binaural cues (binaural vs. diotic presentation) and prevent the use of monaural level cues (using level roving). With low- (2 kHz) frequency bands of noise, performance with binaural presentation and level rove exceeded that expected from use of monaural level cues and that with diotic presentation. It is argued that a high-frequency binaural cue (most likely ILD), and not a monaural level cue, is crucial for robust object localisation without head movement.

Long-term experience enhances neural representation of temporal attributes of pitch in the brainstem and auditory cortex in favorable listening conditions. Herein we examine whether cortical pitch mechanisms shaped by language experience are more resilient to degradation in background noise, and exhibit greater binaural release from masking (BRM). Cortical pitch responses (CPR) were recorded from Mandarin- and English-speaking natives using a Mandarin word exhibiting a high rising pitch (/yi2/). Stimuli were presented diotically in Quiet, and in noise at +5, and 0 dB SNR. CPRs were also recorded in binaural conditions, SONO (where signal and noise were in phase at both ears); or S0Nπ (where signal was in phase and noise 180° out of phase at each ear), using 0 dB SNR. At Fz, both groups showed increase in CPR peak latency and decrease in amplitude with increasing noise level. A language-dependent enhancement of Na–Pb amplitude (Chinese > English) was restricted to Quiet and +5 dB SNR conditions. At T7/T8 electrode sites, Chinese natives exhibited a rightward asymmetry for both CPR components. A language-dependent effect (Chinese > English) was restricted to T8. Regarding BRM, both CPR components showed greater response amplitude for the S0Nπ condition compared to S0N0 across groups. Rightward asymmetry for BRM in the Chinese group indicates experience-dependent recruitment of right auditory cortex. Restriction of the advantage in pitch representation to the quiet and +5 SNR conditions, and the absence of group differences in the binaural release from masking, suggest that language experience affords limited advantage in the neural representation of pitch-relevant information in the auditory cortex under adverse listening conditions.

Unilateral hearing loss (UHL) occurs in 25% of cases of congenital sensorineural hearing loss. Due to the unilaterally reduced audibility associated with UHL, everyday demanding listening situations may be disrupted despite normal hearing in one ear. The aim of this study was to quantify acute changes in recognition of speech in spatially separate competing speech and sound localization accuracy, and relate those changes to two levels of temporary induced UHL (UHL30 and UHL43; suffixes denote the average hearing threshold across 0.5, 1, 2, and 4 kHz) for 8 normal-hearing adults. A within-subject repeated-measures design was used (normal binaural conditions, UHL30 and UHL43). The main outcome measures were the threshold for 40% correct speech recognition and the overall variance in sound localization accuracy quantified by an Error Index (0 = perfect performance, 1.0 = random performance). Distinct and statistically significant deterioration in speech recognition (2.0 dB increase in threshold, p 0.05), while sound localization was additionally impaired (Error Index increase of 0.33, p < 0.01) with an associated large increase in individual variability. Qualitative analyses on a subject-by-subject basis showed that high-frequency audibility was important for speech recognition, while low-frequency audibility was important for horizontal sound localization accuracy. While the data might not be entirely applicable to individuals with long-standing UHL, the results suggest a need for intervention for mild-to-moderate UHL.

Hearing aid (HA) users can differ markedly in their benefit from directional processing (or beamforming) algorithms. The current study therefore investigated candidacy for different bilateral directional processing schemes. Groups of elderly listeners with symmetric (N = 20) or asymmetric (N = 19) hearing thresholds for frequencies below 2 kHz, a large spread in the binaural intelligibility level difference (BILD), and no difference in age, overall degree of hearing loss, or performance on a measure of selective attention took part. Aided speech reception was measured using virtual acoustics together with a simulation of a linked pair of completely occluding behind-the-ear HAs. Five processing schemes and three acoustic scenarios were used. The processing schemes differed in the tradeoff between signal-to-noise ratio (SNR) improvement and binaural cue preservation. The acoustic scenarios consisted of a frontal target talker presented against two speech maskers from ±60° azimuth or spatially diffuse cafeteria noise. For both groups, a significant interaction between BILD, processing scheme, and acoustic scenario was found. This interaction implied that, in situations with lateral speech maskers, HA users with BILDs larger than about 2 dB profited more from preserved low-frequency binaural cues than from greater SNR improvement, whereas for smaller BILDs the opposite was true. Audiometric asymmetry reduced the influence of binaural hearing. In spatially diffuse noise, the maximal SNR improvement was generally beneficial. N0Sπ detection performance at 500 Hz predicted the benefit from low-frequency binaural cues. Together, these findings provide a basis for adapting bilateral directional processing to individual and situational influences. Further research is needed to investigate their generalizability to more realistic HA conditions (e.g., with low-frequency vent-transmitted sound).

Although the tonotopic organisation of the human primary auditory cortex (PAC) has already been studied, the question how its responses are affected in sensorineural hearing loss remains open. Twenty six patients (aged 38.1 ± 9.1 years; 12 men) with symmetrical sloping sensorineural hearing loss (SNHL) and 32 age- and gender-matched controls (NH) participated in an fMRI study using a sparse protocol. The stimuli were binaural 8s complex tones with central frequencies of 400 HzCF, 800 HzCF, 1600 HzCF, 3200 HzCF, or 6400 HzCF, presented at 80 dB(C). In NH responses to all frequency ranges were found in bilateral auditory cortices. The outcomes of a winnermap approach, showing a relative arrangement of active frequency-specific areas, was in line with the existing literature and revealed a V-shape high-frequency gradient surrounding areas that responded to low frequencies in the auditory cortex. In SNHL frequency-specific auditory cortex responses were observed only for sounds from 400 HzCF to 1600 HzCF, due to the severe or profound hearing loss in higher frequency ranges. Using a stringent statistical threshold (p

There are three main acoustical cues to sound location, each attributable to space-and frequency-dependent filtering of the propagating sound waves by the outer ears, head, and torso: Interaural differences in time (ITD) and level (ILD) as well as monaural spectral shape cues. While the guinea pig has been a common model for studying the anatomy, physiology, and behavior of binaural and spatial hearing, extensive measurements of their available acoustical cues are lacking. Here, these cues were determined from directional transfer functions (DTFs), the directional components of the head-related transfer functions, for eleven adult guinea pigs. In the frontal hemisphere, monaural spectral notches were present for frequencies from ~10 to 20 kHz; in general, the notch frequency increased with increasing sound source elevation and in azimuth toward the contralateral ear. The maximum ITDs calculated from low-pass filtered (2 kHz cutoff frequency) DTFs were ~250 µs, whereas the maximum ITD measured with low frequency tone pips was over 320 µs. A spherical head model underestimates ITD magnitude under normal conditions, but closely approximates values when the pinnae were removed. Interaural level differences (ILDs) strongly depended on location and frequency; maximum ILDs were < 10 dB for frequencies < 4 kHz and were as large as 40 dB for frequencies > 10 kHz. Removal of the pinna reduced the depth and sharpness of spectral notches, altered the acoustical axis, and reduced the acoustical gain, ITDs, and ILDs; however, spectral shape features and acoustical gain were not completely eliminated, suggesting a substantial contribution of the head and torso in altering the sounds present at the tympanic membrane.

The count-comparison principle in binaural auditory modeling is based on the assumption that there are nuclei in the mammalian auditory pathway that encode the directional cues in the rate of the output. When this principle is applied, the outputs of the modeled nuclei do not directly result in a topographically organized map of the auditory space that could be monitored as such. Therefore, this article presents a method for visualizing the information from the outputs as well as the nucleus models. The functionality of the auditory model presented here is tested in various binaural listening scenarios, including localization tasks and the discrimination of a target in the presence of distracting sound as well as sound scenarios consisting of multiple simultaneous sound sources. The performance of the model is illustrated with binaural activity maps. The activations seen in the maps are compared to human performance in similar scenarios, and it is shown that the performance of the model is in accordance with the psychoacoustical data.

Neuroimaging studies have provided important information regarding how and where pitch is coded and processed in the human brain. Recordings of the frequency-following response (FFR), an electrophysiological measure of neural temporal coding in the brainstem, have shown that the precision of temporal pitch information is dependent on linguistic and musical experience, and can even be modified by short-term training. However, the FFR does not seem to represent the output of a pitch extraction process, and this raises questions regarding how the peripheral neural signal is processed to produce a unified sensation. Since stimuli with a wide variety of spectral and binaural characteristics can produce the same pitch, it has been suggested that there is a place in the ascending auditory pathway at which the representations converge. There is evidence from many different human neuroimaging studies that certain areas of auditory cortex are specifically sensitive to pitch, although the location is still a matter of debate. Taken together, the results suggest that the initial temporal pitch code in the auditory periphery is converted to a code based on neural firing rate in the brainstem. In the upper brainstem or auditory cortex, the information from the individual harmonics of complex tones is combined to form a general representation of pitch. This article is part of a Special Issue entitled Human Auditory Neuroimaging.

Binaural notched-noise experiments indicate a reduced frequency selectivity of the binaural system compared to monaural processing. The present study investigates how auditory efferent activation (via the medial olivocochlear system) affects binaural frequency selectivity in normal-hearing listeners. Thresholds were measured for a 1-kHz signal embedded in a diotic notched-noise masker for various notch widths. The signal was either presented in phase (diotic) or in antiphase (dichotic), gated with the noise. Stimulus duration was 25 ms, in order to avoid efferent activation due to the masker or the signal. A bandpass-filtered noise precursor was presented prior to the masker and signal stimuli to activate the efferent system. The silent interval between the precursor and the masker-signal complex was 50 ms. For comparison, thresholds for detectability of the masked signal were also measured in a baseline condition without the precursor and, in addition, without the masker. On average, the results of the baseline condition indicate an effectively wider binaural filter, as expected. For both signal phases, the addition of the precursor results in effectively wider filters, which is in agreement with the hypothesis that cochlear gain is reduced due to the presence of the precursor.

In cochlear implants, severe limitations arise from electrical crosstalk between channels. Therefore, the current trend in cochlear implants is to increase stimulation rates to encode signals with higher temporal precision. However, the fundamental question: "What is the limit of temporal precision due to inherent neuronal dynamics of the stimulated neurons?" has not yet been resolved. In this study we have developed a double-pulse method and, for the first time, reversed stimulus polarity systematically between consecutive pulses to elucidate subthreshold-induced temporal interaction effects. This method allowed us to determine the time-course of subthreshold temporal interaction in human subjects which identifies the limits of encoded temporal precision. Our results show significant temporal interaction up to 600 μs inter-pulse interval. In all the cases tested we saw a facilitation effect on threshold. Interaction effects at a 20% below threshold pre-conditioning stimulation showed up to 38% ± 6% threshold reduction. These results imply that there is significant temporal interaction between two subsequent pulses. This interaction diminishes the precision of amplitude coding. We predict interaction effects on temporal precision and channel interaction. For (interleaved) stimulation with short inter-pulse intervals it is interesting to consider our interaction results; and it may become important to consider them for future coding strategies where high temporal precision is required. In an increasing group of binaural implanted patients this will be the case when interaural time differences are encoded with μs precision.

The detection of low-rate frequency modulation (FM) carried by a low-frequency tone has been employed as a means of assessing the fidelity of temporal fine structure coding. Detection of low-rate FM can be made more acute, relative to the monaural case, by the addition of a pure tone to the contralateral ear. This study examined whether FM detection in the 500-Hz region could be further improved by using a binaural stimulation mode where the modulator was antiphasic across the two ears. The study also sought to determine whether these dichotic FM conditions were beneficial in identifying the emergence of a temporal fine structure processing deficiency relatively early in the aging process. Young, mid-aged, and older listeners (n = 12 per group) were tested. The results demonstrated better FM acuity in the dichotic task irrespective of listener age. Dichotic FM detection also differentiated between age groups more definitively than diotic detection, especially in terms of distinguishing mid-aged from older listeners. In the group of older listeners, dichotic FM detection was weakly associated with absolute sensitivity to the carrier. In addition, this group failed to show a dichotic benefit in the presence of a marked asymmetry in sensation level across ears. The overall pattern of results suggests that dichotic FM measurements have advantages over monaural measurements for the purposes of assessing age-related temporal processing effects, although a marked asymmetry in absolute thresholds across ears could undermine these advantages.

In a forward-masked intensity discrimination task, we manipulated the perceived lateralization of the masker via variation of the interaural time difference (ITD). The maskers and targets were 500 Hz pure tones with a duration of 30 ms. Standards of 30 and 60 dB SPL were combined with 60 or 90 dB SPL maskers. As expected, the presentation of a forward masker perceived as lateralized to the other side of the head as the target resulted in a significantly smaller elevation of the intensity difference limen than a masker lateralized ipsilaterally. This binaural release from masking in forward-masked intensity discrimination cannot be explained by peripheral mechanisms because varying the ITD leaves the neural representation in the monaural channels (i.e., in the auditory nerve) unaltered. Instead, our results are compatible with the assumption that lateralization differences between masker and target promote object segregation and therefore facilitate object-based selective attention to the target.