The Stria Vascularis: Renewed Attention on a Key Player in Age-Related Hearing Loss.

Age-related hearing loss (HL), or presbycusis, is a complex and heterogeneous condition, affecting a significant portion of older adults and involving various interacting mechanisms. Metabolic presbycusis, a type of age-related HL, is characterized by the dysfunction of the stria vascularis, which is crucial for maintaining the endocochlear potential necessary for hearing. Although attention on metabolic presbycusis has waned in recent years, research continues to identify strial pathology as a key factor in age-related HL. This narrative review integrates past and recent research, bridging findings from animal models and human studies, to examine the contributions of the stria vascularis to age-related HL. It provides a brief overview of the structure and function of the stria vascularis and then examines mechanisms contributing to age-related strial dysfunction, including altered ion transport, changes in pigmentation, inflammatory responses, and vascular atrophy. Importantly, this review outlines the contribution of metabolic mechanisms to age-related HL, highlighting areas for future research. It emphasizes the complex interdependence of metabolic and sensorineural mechanisms in the pathology of age-related HL and highlights the importance of animal models in understanding the underlying mechanisms. The comprehensive and mechanistic investigation of all factors contributing to age-related HL, including cochlear metabolic dysfunction, remains crucial to identifying the underlying mechanisms and developing personalized, protective, and restorative treatments.

Cochlear Ribbon Synapses in Aged Gerbils.

In mammalian hearing, type-I afferent auditory nerve fibers comprise the basis of the afferent auditory pathway. They are connected to inner hair cells of the cochlea via specialized ribbon synapses. Auditory nerve fibers of different physiological types differ subtly in their synaptic location and morphology. Low-spontaneous-rate auditory nerve fibers typically connect on the modiolar side of the inner hair cell, while high-spontaneous-rate fibers are typically found on the pillar side. In aging and noise-damaged ears, this fine-tuned balance between auditory nerve fiber populations can be disrupted and the functional consequences are currently unclear. Here, using immunofluorescent labeling of presynaptic ribbons and postsynaptic glutamate receptor patches, we investigated changes in synaptic morphology at three different tonotopic locations along the cochlea of aging gerbils compared to those of young adults. Quiet-aged gerbils showed about 20% loss of afferent ribbon synapses. While the loss was random at apical, low-frequency cochlear locations, at the basal, high-frequency location it almost exclusively affected the modiolar-located synapses. The subtle differences in volumes of pre- and postsynaptic elements located on the inner hair cell's modiolar versus pillar side were unaffected by age. This is consistent with known physiology and suggests a predominant, age-related loss in the low-spontaneous-rate auditory nerve population in the cochlear base, but not the apex.

Age-Related Deficits in Binaural Hearing: Contribution of Peripheral and Central Effects.

Pure-tone audiograms often poorly predict elderly humans' ability to communicate in everyday complex acoustic scenes. Binaural processing is crucial for discriminating sound sources in such complex acoustic scenes. The compromised perception of communication signals presented above hearing threshold has been linked to both peripheral and central age-related changes in the auditory system. Investigating young and old Mongolian gerbils of both sexes, an established model for human hearing, we demonstrate age-related supra-threshold deficits in binaural hearing using behavioral, electrophysiological, anatomical, and imaging methods. Binaural processing ability was measured as the binaural masking level difference (BMLD), an established measure in human psychophysics. We tested gerbils behaviorally with "virtual headphones," recorded single-unit responses in the auditory midbrain and evaluated gross midbrain and cortical responses using positron emission tomography (PET) imaging. Furthermore, we obtained additional measures of auditory function based on auditory brainstem responses, auditory-nerve synapse counts, and evidence for central inhibitory processing revealed by PET. BMLD deteriorates already in middle-aged animals having normal audiometric thresholds and is even worse in old animals with hearing loss. The magnitude of auditory brainstem response measures related to auditory-nerve function and binaural processing in the auditory brainstem also deteriorate. Furthermore, central GABAergic inhibition is affected by age. Because the number of synapses in the apical turn of the inner ear was not reduced in middle-aged animals, we conclude that peripheral synaptopathy contributes little to binaural processing deficits. Exploratory analyses suggest increased hearing thresholds, altered binaural processing in the brainstem and changed central GABAergic inhibition as potential contributors.

The Relative Contribution of Cochlear Synaptopathy and Reduced Inhibition to Age-Related Hearing Impairment for People With Normal Audiograms.

Older people often show auditory temporal processing deficits and speech-in-noise intelligibility difficulties even when their audiogram is clinically normal. The causes of such problems remain unclear. Some studies have suggested that for people with normal audiograms, age-related hearing impairments may be due to a cognitive decline, while others have suggested that they may be caused by cochlear synaptopathy. Here, we explore an alternative hypothesis, namely that age-related hearing deficits are associated with decreased inhibition. For human adults (N = 30) selected to cover a reasonably wide age range (25-59 years), with normal audiograms and normal cognitive function, we measured speech reception thresholds in noise (SRTNs) for disyllabic words, gap detection thresholds (GDTs), and frequency modulation detection thresholds (FMDTs). We also measured the rate of growth (slope) of auditory brainstem response wave-I amplitude with increasing level as an indirect indicator of cochlear synaptopathy, and the interference inhibition score in the Stroop color and word test (SCWT) as a proxy for inhibition. As expected, performance in the auditory tasks worsened (SRTNs, GDTs, and FMDTs increased), and wave-I slope and SCWT inhibition scores decreased with ageing. Importantly, SRTNs, GDTs, and FMDTs were not related to wave-I slope but worsened with decreasing SCWT inhibition. Furthermore, after partialling out the effect of SCWT inhibition, age was no longer related to SRTNs or GDTs and became less strongly related to FMDTs. Altogether, results suggest that for people with normal audiograms, age-related deficits in auditory temporal processing and speech-in-noise intelligibility are mediated by decreased inhibition rather than cochlear synaptopathy.

Altered neural encoding of vowels in noise does not affect behavioral vowel discrimination in gerbils with age-related hearing loss.

Introduction: Understanding speech in a noisy environment, as opposed to speech in quiet, becomes increasingly more difficult with increasing age. Using the quiet-aged gerbil, we studied the effects of aging on speech-in-noise processing. Specifically, behavioral vowel discrimination and the encoding of these vowels by single auditory-nerve fibers were compared, to elucidate some of the underlying mechanisms of age-related speech-in-noise perception deficits. Methods: Young-adult and quiet-aged Mongolian gerbils, of either sex, were trained to discriminate a deviant naturally-spoken vowel in a sequence of vowel standards against a speech-like background noise. In addition, we recorded responses from single auditory-nerve fibers of young-adult and quiet-aged gerbils while presenting the same speech stimuli. Results: Behavioral vowel discrimination was not significantly affected by aging. For both young-adult and quiet-aged gerbils, the behavioral discrimination between /eː/ and /iː/ was more difficult to make than /eː/ vs. /aː/ or /iː/ vs. /aː/, as evidenced by longer response times and lower d' values. In young-adults, spike timing-based vowel discrimination agreed with the behavioral vowel discrimination, while in quiet-aged gerbils it did not. Paradoxically, discrimination between vowels based on temporal responses was enhanced in aged gerbils for all vowel comparisons. Representation schemes, based on the spectrum of the inter-spike interval histogram, revealed stronger encoding of both the fundamental and the lower formant frequencies in fibers of quiet-aged gerbils, but no qualitative changes in vowel encoding. Elevated thresholds in combination with a fixed stimulus level, i.e., lower sensation levels of the stimuli for old individuals, can explain the enhanced temporal coding of the vowels in noise. Discussion: These results suggest that the altered auditory-nerve discrimination metrics in old gerbils may mask age-related deterioration in the central (auditory) system to the extent that behavioral vowel discrimination matches that of the young adults.

Masking release at 4 and 32 kHz in harbor seals associated with sinusoidal amplitude-modulated masking noise.

Masking can reduce the efficiency of communication and prey and predator detection. Most underwater sounds fluctuate in amplitude, which may influence the amount of masking experienced by marine mammals. The hearing thresholds of two harbor seals for tonal sweeps (centered at 4 and 32 kHz) masked by sinusoidal amplitude modulated (SAM) Gaussian one-third octave noise bands centered around the narrow-band test sweep frequencies, were studied with a psychoacoustic technique. Masking was assessed in relation to signal duration, (500, 1000, and 2000 ms) and masker level, at eight amplitude modulation rates (1-90 Hz). Masking release (MR) due to SAM compared thresholds in modulated and unmodulated maskers. Unmodulated maskers resulted in critical ratios of 21 dB at 4 kHz and 31 dB at 32 kHz. Masked thresholds were similarly affected by SAM rate with the lowest thresholds and the largest MR being observed for SAM rates of 1 and 2 Hz at higher masker levels. MR was higher for 32-kHz maskers than for 4-kHz maskers. Increasing signal duration from 500 ms to 2000 ms had minimal effect on MR. The results are discussed with respect to MR resulting from envelope variation and the impact of noise in the environment on target signal detection.

Audio-Visual Interference During Motion Discrimination in Starlings.

Motion discrimination is essential for animals to avoid collisions, to escape from predators, to catch prey or to communicate. Although most terrestrial vertebrates can benefit by combining concurrent stimuli from sound and vision to obtain a most salient percept of the moving object, there is little research on the mechanisms involved in such cross-modal motion discrimination. We used European starlings as a model with a well-studied visual and auditory system. In a behavioural motion discrimination task with visual and acoustic stimuli, we investigated the effects of cross-modal interference and attentional processes. Our results showed an impairment of motion discrimination when the visual and acoustic stimuli moved in opposite directions as compared to congruent motion direction. By presenting an acoustic stimulus of very short duration, thus lacking directional motion information, an additional alerting effect of the acoustic stimulus became evident. Finally, we show that a temporally leading acoustic stimulus did not improve the response behaviour compared to the synchronous presentation of the stimuli as would have been expected in case of major alerting effects. This further supports the importance of congruency and synchronicity in the current test paradigm with a minor role of attentional processes elicited by the acoustic stimulus. Together, our data clearly show cross-modal interference effects in an audio-visual motion discrimination paradigm when carefully selecting real-life stimuli under parameter conditions that meet the known criteria for cross-modal binding.

Neural processing and perception of Schroeder-phase harmonic tone complexes in the gerbil: Relating single-unit neurophysiology to behavior.

Schroeder-phase harmonic tone complexes have been used in physiological and psychophysical studies in several species to gain insight into cochlear function. Each pitch period of the Schroeder stimulus contains a linear frequency sweep; the duty cycle, sweep velocity, and direction are controlled by parameters of the phase spectrum. Here, responses to a range of Schroeder-phase harmonic tone complexes were studied both behaviorally and in neural recordings from the auditory nerve and inferior colliculus of Mongolian gerbils. Gerbils were able to discriminate Schroeder-phase harmonic tone complexes based on sweep direction, duty cycle, and/or velocity for fundamental frequencies up to 200 Hz. Temporal representation in neural responses based on the van Rossum spike-distance metric, with time constants of either 1 ms or related to the stimulus' period, was compared with average discharge rates. Neural responses and behavioral performance were both expressed in terms of sensitivity, d', to allow direct comparisons. Our results suggest that in the auditory nerve, stimulus fine structure is represented by spike timing, whereas envelope is represented by rate. In the inferior colliculus, both temporal fine structure and envelope appear to be represented best by rate. However, correlations between neural d' values and behavioral sensitivity for sweep direction were strongest for both temporal metrics, for both auditory nerve and inferior colliculus. Furthermore, the high sensitivity observed in the inferior colliculus neural rate-based discrimination suggests that these neurons integrate across multiple inputs arising from the auditory periphery.

Chickens have excellent sound localization ability.

The mechanisms of sound localization are actively debated, especially which cues are predominately used and why. Our study provides behavioural data in chickens (Gallus gallus) and relates these to estimates of the perceived physical cues. Sound localization acuity was quantified as the minimum audible angle (MAA) in azimuth. Pure-tone MAA was 12.3, 9.3, 8.9 and 14.5 deg for frequencies of 500, 1000, 2000 and 4000 Hz, respectively. Broadband-noise MAA was 12.2 deg, which indicates excellent behavioural acuity. We determined 'external cues' from head-related transfer functions of chickens. These were used to derive 'internal cues', taking into account published data on the effect of the coupled middle ears. Our estimates of the internal cues indicate that chickens likely relied on interaural time difference cues alone at low frequencies of 500 and 1000 Hz, whereas at 2000 and 4000 Hz, interaural level differences may be the dominant cue.

Age-related decline in cochlear ribbon synapses and its relation to different metrics of auditory-nerve activity.

Loss of inner hair cell-auditory nerve fiber synapses is considered to be an important early stage of neural presbyacusis. Mass potentials, recorded at the cochlear round window, can be used to derive the neural index (NI), a sensitive measure for pharmacologically-induced synapse loss. Here, we investigate the applicability of the NI for measuring age-related auditory synapse loss in young-adult, middle-aged, and old Mongolian gerbils. Synapse loss, which was progressively evident in the 2 aged groups, correlated weakly with NI when measured at a fixed sound level of 60 dB SPL. However, the NI was confounded by decreases in single-unit firing rates at 60 dB SPL. NI at 30 dB above threshold, when firing rates were similar between age groups, did not correlate with synapse loss. Our results show that synapse loss is poorly reflected in the NI of aged gerbils, particularly if further peripheral pathologies are present. The NI may therefore not be a reliable clinical tool to assess synapse loss in aged humans with peripheral hearing loss.

Uncertainty-based informational masking in a vowel discrimination task for young and old Mongolian gerbils.

Informational masking emerges with processing of complex sounds in the central auditory system and can be affected by uncertainty emerging from trial-to-trial variation of stimulus features. Uncertainty can be non-informative but confusing and thus mask otherwise salient stimulus changes resulting in increased discrimination thresholds. With increasing age, the ability for processing of such complex sound scenes degrades. Here, 6 young and 4 old gerbils were tested behaviorally in a vowel discrimination task. Animals were trained to discriminate between sequentially presented target and reference vowels of the vowel pair/I/-/i/. Reference and target vowels were generated shifting the three formants of the reference vowel in steps towards the formants of the target vowels. Non-informative but distracting uncertainty was introduced by random changes in location, level, fundamental frequency or all three features combined. Young gerbils tested with uncertainty for the target or target and reference vowels showed similar informational masking effects for both conditions. Young and old gerbils were tested with uncertainty for the target vowels only. Old gerbils showed no threshold increase discriminating vowels without uncertainty in comparison with young gerbils. Introducing uncertainty, vowel discrimination thresholds increased for young and old gerbils and vowel discrimination thresholds increased most when presenting all three uncertainty features combined. Old gerbils were more susceptible to non-informative uncertainty and their thresholds increased more than thresholds of young gerbils. Gerbils' vowel discrimination thresholds are compared to human performance in the same task (Eipert et al., 2019).

Interaction of spatial source separation, fundamental frequency, and vowel pairing in a sequential informational masking paradigm in Mongolian gerbils.

Informational masking (IM) defines the compromised ability to perceive and analyze signals from a single source in a clutter of other sounds even if there is no interference between these signals' excitation patterns in the inner ear. IM is affected by the similarity between target and masker and the variation of stimulus features from trial to trial, that is, stimulus uncertainty, both modulating discrimination thresholds. We applied a sequential IM paradigm measuring Mongolian gerbils' sensitivity to detect level increments between constant-level standard (reference) and deviant (target) vowels with a level increase in a background of level-varying distracting (masker) vowels. Different combinations of vowels (/I/, /i/, /æ/, /ε/) and fundamental frequencies (101 Hz, 127 Hz) as well as sound source position (colocated, 90° separated) were presented, and the effect of target and masker similarity on IM in a condition of high stimulus uncertainty was determined. We observed a release from IM, that is, lower level increment thresholds, by differences in vowel type, fundamental frequency, or spatial separation between standard/deviant and distractor vowels only. The effects of vowel type and fundamental frequency interacted, such as the release from IM by fundamental frequency was stronger for similar than for different vowel types. The spatial separation of vowels did not interact with vowel type and fundamental frequency but offered an additional release from IM. If two of the cues supported stream segregation, the release from IM was nearly complete. (PsycINFO Database Record (c) 2020 APA, all rights reserved).

Evidence for the origin of the binaural interaction component of the auditory brainstem response.

The binaural interaction component (BIC) represents the mismatch between auditory brainstem responses (ABR) obtained with binaural stimulation and the sum of ABRs obtained with monaural left and right stimulation. It is generally assumed that the BIC reflects binaural integration. Its potential use as a diagnostic tool, however, is hampered by the lack of direct evidence about its origin. While an origin at the initial site of binaural integration seems likely, there is no general agreement on the contribution of the two primary candidate nuclei, the lateral and medial superior olives (LSO and MSO, respectively). Here, we recorded local field potentials (LFP) and responses of units in the LSO and MSO of Mongolian gerbils (Meriones unguiculatus), presenting clicks with an interaural time or level difference (ITD and ILD, respectively), while simultaneously recording ABR. We determined the BIC from the ABR and, importantly, from LFP and responses of units in the LSO and MSO. If stimulus-induced changes in the ABR-derived BIC have their source in the LSO and/or MSO, we expect coherent changes in the unit-derived and the ABR-derived BIC. We find that BIC obtained from LSO units exhibits the same ITD and ILD dependence as the ABR-derived BIC. Neither BIC obtained from MSO units nor LFP-derived BIC recorded in either LSO or MSO did. The data thus strongly suggest that it is the activity of LSO units in the gerbil that is decisive for the generation of the ABR-derived BIC, determining its properties.

Interaction of spatial and non-spatial cues in auditory stream segregation in the European starling.

Integrating sounds from the same source and segregating sounds from different sources in an acoustic scene are an essential function of the auditory system. Naturally, the auditory system simultaneously makes use of multiple cues. Here, we investigate the interaction between spatial cues and frequency cues in stream segregation of European starlings (Sturnus vulgaris) using an objective measure of perception. Neural responses to streaming sounds were recorded, while the bird was performing a behavioural task that results in a higher sensitivity during a one-stream than a two-stream percept. Birds were trained to detect an onset time shift of a B tone in an ABA- triplet sequence in which A and B could differ in frequency and/or spatial location. If the frequency difference or spatial separation between the signal sources or both were increased, the behavioural time shift detection performance deteriorated. Spatial separation had a smaller effect on the performance compared to the frequency difference and both cues additively affected the performance. Neural responses in the primary auditory forebrain were affected by the frequency and spatial cues. However, frequency and spatial cue differences being sufficiently large to elicit behavioural effects did not reveal correlated neural response differences. The difference between the neuronal response pattern and behavioural response is discussed with relation to the task given to the bird. Perceptual effects of combining different cues in auditory scene analysis indicate that these cues are analysed independently and given different weights suggesting that the streaming percept arises consecutively to initial cue analysis.

Release from informational masking by auditory stream segregation: perception and its neural correlate.

In the analysis of acoustic scenes, we easily miss sounds or are insensitive to sound features that are salient if presented in isolation. This insensitivity that is not due to interference in the inner ear is termed informational masking (IM). So far, the cellular mechanisms underlying IM remained elusive. Here, we apply a sequential IM paradigm to humans and gerbils using a sound level increment detection task determining the sensitivity to target tones in a background of standard (same frequency) and distracting tones (varying in level and frequency). The amount of IM that was indicated by the level increment thresholds depended on the frequency separation between the distracting and the standard and target tones. In humans and gerbils, we observed similar perceptual thresholds. A release from IM of more than 20 dB was observed in both species if the distracting tones were well segregated in frequency from the other tones. Neuronal rate responses elicited by similar sequences in gerbil inferior colliculus and auditory cortex were recorded. At both levels of the auditory pathway, the neuronal thresholds obtained with a signal-detection-theoretic approach deducing the sensitivity from the analysis of the neurons' receiver operating characteristics matched the psychophysical thresholds revealing that IM already emerges at midbrain level. By applying objective response measures in physiology and psychophysics, we demonstrated that the population of neurons has a sufficient sensitivity for explaining the perceptual level increment thresholds indicating IM. There was a good correspondence between the neuronal and perceptual release from IM being related to auditory stream segregation.

The barn owls' Minimum Audible Angle.

Interaural time differences (ITD) and interaural level differences (ILD) are physical cues that enable the auditory system to pinpoint the position of a sound source in space. This ability is crucial for animal communication and predator-prey interactions. The barn owl has evolved an exceptional sense of hearing and shows abilities of sound localisation that outperform most other species. So far, behavioural studies in the barn owl often used reflexive responses to investigate aspects of sound localisation. Furthermore, they predominately probed the higher frequencies of the owl's hearing range (> 3 kHz). In the present study we used a Go/NoGo paradigm to measure the barn owl's behavioural sound localisation acuity (expressed as the Minimum Audible Angle, MAA) as a function of stimulus type (narrow-band noise centred at 500, 1000, 2000, 4000 and 8000 Hz, and broad-band noise) and sound source position. We found significant effects of both stimulus type and sound source position on the barn owls' MAA. The MAA improved with increasing stimulus frequency, from 14° at 500 Hz to 6° at 8000 Hz. The smallest MAA of 4° was found for broadband noise stimuli. Comparing different sound source positions revealed smaller MAAs for frontal compared to lateral stimulus presentation, irrespective of stimulus type. These results are consistent with both the known variations in physical ITDs and variation in the width of neural ITD tuning curves with azimuth and frequency. Physical and neural characteristics combine to result in better spatial acuity for frontal compared to lateral sounds and reduced localisation acuity at lower frequencies.

Uncertainty in location, level and fundamental frequency results in informational masking in a vowel discrimination task for young and elderly subjects.

Informational masking (IM) is defined as the compromised ability to perceive and analyze signals from a single sound source in a cacophony of sounds from other sources even if the excitation patterns produced by these signals in the auditory periphery are well separated from those produced by the sounds from the other sources. IM that causes an elevation of discrimination thresholds is affected by the similarity between target and masker and by stimulus uncertainty. Here, six young and six elderly subjects were asked to discriminate between sequentially presented reference and target vowels of the vowel pairs /I/-/i/, /æ/-/ε/, and /α/-/Λ/. Psychometric functions were collected characterizing the discrimination of target vowels from reference vowels. Target vowels differed from the reference by one of seven steps shifting the three formants of a reference vowel towards the formants of the corresponding target vowel. Stimulus statistics were varied, generating uncertainty by non-informative but potentially distracting location, level, and fundamental frequency changes or all three combined. Young subjects tested with distracting changes applied to the target vowels only, the reference vowels only, or the target and reference vowels showed similar amounts of IM for all three conditions. Elderly subjects were tested with distracting changes applied to target vowels only. Applying uncertainty only to the target vowels led to worse vowel discrimination thresholds for young and elderly subjects and thresholds increased most for the three distracting changes combined. Elderly subjects showed higher vowel discrimination thresholds than young subjects, but the increase in vowel discrimination thresholds due to IM did not differ between young and elderly subjects. The temporal fine structure processing of elderly subjects was degraded in comparison to young subjects, but it was only correlated with the discrimination threshold for vowel pair /I/-/i/.

Violation of the Unity Assumption Disrupts Temporal Ventriloquism Effect in Starlings.

When stimuli from different sensory modalities are received, they may be combined by the brain to form a multisensory percept. One key mechanism for multisensory binding is the unity assumption under which multisensory stimuli that share certain physical properties like temporal and/or spatial correspondence are grouped together as deriving from one object. In humans, evidence for a role of the unity assumption has been found in spatial tasks and also in temporal tasks using stimuli that share physical properties (speech-related stimuli, musical and synesthetically congruent stimuli). In our study, we investigate the role of the unity assumption in an animal model in a temporal order judgment task. When subjects are asked to indicate which of two spatially separated visual stimuli appeared first in time, performance improves when the visual stimuli are paired (in time) with spatially non-informative acoustic cues, a phenomenon known as the temporal ventriloquism effect. Here, we show that European starlings perform better when one singleton acoustic cue is paired with the first visual stimulus as compared to pairing with the second visual stimulus. This shows, in combination with our previous study, that a non-informative singleton acoustic cue, when temporally paired with the first visual stimulus, triggers alerting while, when temporally pairing with the second visual stimulus, it prevents a temporal ventriloquism effect because the unity assumption is violated. Thus, the unity assumption influences sensory perception not only in humans but also in an animal model. The importance of the unity assumption in this task supports the idea that the temporal ventriloquism effect, similar to the spatial ventriloquism effect, is based on multisensory binding and integration but not on alerting effects.

Processing of interaural phase differences in components of harmonic and mistuned complexes in the inferior colliculus of the Mongolian gerbil.

Harmonicity and spatial location provide eminent cues for the perceptual grouping of sounds. In general, harmonicity is a strong grouping cue. In contrast, spatial cues such as interaural phase or time difference provide for strong grouping of stimulus sequences but weak grouping for simultaneously presented sounds. By studying the neuronal basis underlying the interaction of these cues in processing simultaneous sounds using van Rossum spike train distance measures, we aim at explaining the interaction observed in psychophysical experiments. Responses to interaural phase differences imposed on single components of harmonic and mistuned complex tones as well as noise delay functions were recorded as multiunit responses from the inferior colliculus of Mongolian gerbils. Results revealed a better representation of interaural phase differences if imposed on a harmonic rather than a mistuned frequency component of a complex tone. The representation of interaural phase differences was better for long integration-time windows approximately reflecting firing rates rather than short integration-time windows reflecting the temporal pattern of the stimulus-driven response. We found only a weak impact of interaural phase differences if combined with mistuning of a component in a harmonic tone complex.

Interaction of interaural cues and their contribution to the lateralisation of Mongolian gerbils (Meriones unguiculatus).

The main sound localisation cues in the horizontal plane are interaural time and level differences (ITDs and ILDs, respectively). ITDs are thought to be the dominant cue in the low-frequency range, ILDs the dominant cue in the high-frequency range. ITDs and ILDs co-occur. Their interaction and contribution to the lateralisation of pure tones by Mongolian gerbils was investigated behaviourally using cross-talk cancellation techniques for presenting ITDs and ILDs independently. First, ITDs were applied to pure tones with frequencies ≤ 2 kHz to the ongoing waveform, at the onsets and offsets, or in both the ongoing waveform and at the onsets and offsets. Gerbils could lateralise tones only if ongoing ITDs were present indicating that ongoing ITDs are decisive for the lateralisation of low-frequency tones. Second, an ITD was added to 2-to-6-kHz tones with varying ILD. Gerbils' lateralisation was unaffected by the ITD indicating that a large ILD provides a strong lateralisation cue at those frequencies. Finally, small ILDs were applied to 2-kHz tones with an ongoing ITD, pointing either to the same or opposing sides as the ITD. Gerbils' lateralisation was driven by the ITD but strongly affected by the ILD indicating that both interaural cues contribute to the lateralisation.