Interaural Level Differences Sensitivity Research Articles

Impact of processing-latency induced interaural delay and level discrepancy on sensitivity to interaural level differences in cochlear implant users.

This study investigated whether an interaural delay, e.g. caused by the processing latency of a hearing device, can affect sensitivity to interaural level differences (ILDs) in normal hearing subjects or cochlear implant (CI) users with contralateral normal hearing (SSD-CI). Sensitivity to ILD was measured in 10SSD-CI subjects and in 24normal hearing subjects. The stimulus was a noise burst presented via headphones and via a direct cable connection (CI). ILD sensitivity was measured for different interaural delays in the range induced by hearing devices. ILD sensitivity was correlated with results obtained in a sound localization task using seven loudspeakers in the frontal horizontal plane. In the normal hearing subjects the sensitivity to interaural level differences deteriorated significantly with increasing interaural delays. In the CI group, no significant effect of interaural delays on ILD sensitivity was found. The NH subjects were significantly more sensitive to ILDs. The mean localization error in the CI group was 10.8° higher than in the normal hearing group. No correlation between sound localization ability and ILD sensitivity was found. Interaural delays influence the perception of ILDs. For normal hearing subjects a significant decrement in sensitivity to ILD was measured. The effect could not be confirmed in the tested SSD-CI group, probably due to a small subject group with large variations. The temporal matching of the two sides may be beneficial for ILD processing and thus sound localization for CI patients. However, further studies are needed for verification.

Effects of interaural decoherence on sensitivity to interaural level differences across frequency.

The interaural level difference (ILD) is a robust indicator of sound source azimuth, and human ILD sensitivity persists under conditions that degrade normally-dominant interaural time difference (ITD) cues. Nonetheless, ILD sensitivity varies somewhat with both stimulus frequency and interaural correlation (coherence). To further investigate the combined binaural perceptual influence of these variables, the present study assessed ILD sensitivity at frequencies 250-4000 Hz using stimuli of varied interaural correlation. In the first of two experiments, ILD discrimination thresholds were modestly elevated, and subjective lateralization slightly reduced, for both half-correlated and uncorrelated narrowband noise tokens relative to correlated tokens. Different from thresholds in the correlated condition, which were worst at 1000 Hz [Grantham, D.W. (1984). J. Acoust. Soc. Am. 75, 1191-1194], thresholds in the decorrelated conditions were independent of frequency. However, intrinsic envelope fluctuations in narrowband stimuli caused moment-to-moment variation of the nominal ILD, complicating interpretation of measured thresholds. Thus, a second experiment employed low-fluctuation noise tokens, revealing a clear effect of interaural decoherence per se that was strongly frequency-dependent, decreasing in magnitude from low to high frequencies. Measurements are consistent with known integration times in ILD-sensitive neurons and also suggest persistent influences of covert ITD cues in putative "ILD" tasks.

Comparing Methods for Pairing Electrodes Across Ears With Cochlear Implants.

Currently, bilateral cochlear implants (CIs) are independently programmed in clinics using frequency allocations based on the relative location of a given electrode from the end of each electrode array. By pairing electrodes based on this method, bilateral CI recipients may have decreased sensitivity to interaural time differences (ITD) and/or interaural level differences (ILD), two cues critical for binaural tasks. There are multiple different binaural measures that can potentially be used to determine the optimal way to pair electrodes across the ears. Previous studies suggest that the optimal electrode pairing between the left and right ears may vary depending on the binaural task used. These studies, however, have only used one reference location or a single bilateral CI user. In both instances, it is difficult to determine if the results that were obtained reflect a measurement error or a systematic difference across binaural tasks. It is also difficult to determine from these studies if the differences between the three cues vary across electrode regions, which could result from differences in the availability of binaural cues across frequency regions. The purpose of this study was to determine if, after experience-dependent adaptation, there are systematic differences in the optimal pairing of electrodes at different points along the array for the optimal perception of ITD, ILD, and pitch. Data from seven bilateral Nucleus users was collected and analyzed. Participants were tested with ITD, ILD, and pitch-matching tasks using five different reference electrodes in one ear, spaced across the array. Comparisons were conducted to determine if the optimal bilateral electrode pairs systematically differed in different regions depending on whether they were measured based on ITD sensitivity, ILD sensitivity, or pitch matching, and how those pairs differed from the pairing in the participants' clinical programs. Results indicate that there was a significant difference in the optimal pairing depending on the cue measured, but only at the basal end of the array. The results suggest that optimal electrode pairings differ depending on the cue measured to determine optimal pairing, at least for the basal end of the array. This also suggests that the improvements seen when using optimally paired electrodes may be tied to the particular percept being measured both to determine electrode pairing and to assess performance, at least for the basal end of the array.

Spatial Hearing by Bilateral Cochlear Implant Users With Temporal Fine-Structure Processing.

Several studies have demonstrated the advantages of the bilateral vs. unilateral cochlear implantation in listeners with bilateral severe to profound hearing loss. However, it remains unclear to what extent bilaterally implanted listeners have access to binaural cues, e.g., accurate processing of interaural timing differences (ITDs) for low-frequency sounds (<1.5 kHz) and interaural level differences (ILDs) for high frequencies (>3 kHz). We tested 25 adult listeners, bilaterally implanted with MED-EL cochlear implant (CI) devices, with and without fine-structure (FS) temporal processing as encoding strategy in the low-frequency channels. In order to assess whether the ability to process binaural cues was affected by fine-structure processing, we performed psychophysical ILD and ITD sensitivity measurements and free-field sound localization experiments. We compared the results of the bilaterally implanted listeners with different numbers of FS channels. All CI listeners demonstrated good sensitivity to ILDs, but relatively poor to ITD cues. Although there was a large variability in performance, some bilateral CI users showed remarkably good localization skills. The FS coding strategy for bilateral CI hearing did not improve fine-structure ITD processing for spatial hearing on a group level. However, some CI listeners were able to exploit weakly informative temporal cues to improve their low-frequency spatial perception.

Cortical Processing of Level Cues for Spatial Hearing is Impaired in Children with Prelingual Deafness Despite Early Bilateral Access to Sound.

Bilateral cochlear implantation aims to restore binaural hearing, important for spatial hearing, to children who are deaf. Improvements over unilateral implant use are attributed largely to the detection of interaural level differences (ILDs) but emerging evidence of impaired sound localization and binaural fusion suggest that these binaural cues are abnormally coded by the auditory system. We used multichannel electroencephalography (EEG) to assess cortical responses to ILDs in two groups: 13 children who received early bilateral cochlear implants (CIs) simultaneously, known to protect the developing auditory cortices from unilaterally driven reorganization, and 15 age matched peers with normal hearing. EEG source analyses indicated a dominance of right auditory cortex in both groups. Expected reductions in activity to ipsilaterally weighted ILDs were evident in the right hemisphere of children with normal hearing. By contrast, cortical activity in children with CIs showed: (1) limited ILD sensitivity in either cortical hemisphere, (2) limited correlation with reliable behavioral right-left lateralization of ILDs (in 10/12 CI users), and (3) deficits in parieto-occipital areas and the cerebellum. Thus, expected cortical ILD coding develops with normal hearing but is affected by developmental deafness despite early and simultaneous bilateral implantation. Findings suggest that impoverished fidelity of ILDs in independently functioning CIs may be impeding development of cortical ILD sensitivity in children who are deaf but do not altogether limit benefits of listening with bilateral CIs. Future efforts to provide consistent/accurate ILDs through auditory prostheses including CIs could improve binaural hearing for children with hearing loss.

Temporal effects in interaural and sequential level difference perception.

Temporal effects in interaural level difference (ILD) perception are not well understood. While it is often assumed that ILD sensitivity is independent of the temporal stimulus properties, a reduction of ILD sensitivity for stimuli with a high modulation rate has been reported (known under the term binaural adaptation). Experiment 1 compared ILD thresholds and sequential-level-difference (SLD) thresholds using 300-ms bandpass-filtered pulse trains (centered at 4 kHz) with rates of 100, 400, and 800 pulses per second (pps). In contrast to the SLD thresholds, ILD thresholds were elevated at 800 pps, consistent with literature data that had previously been attributed to binaural adaptation. Experiment 2 showed better ILD sensitivity for pulse trains than for pure tones, suggesting that amplitude modulation enhances ILD sensitivity. The present ILD data and binaural adaptation data from the literature were predicted by a model combining well-established auditory periphery front-ends with an interaural comparison stage. The model also accounted for other published ILD data, including target ILD thresholds in diotic forward and backward fringes and ILD thresholds with different amounts of interaural correlation. Overall, a variety of temporal effects in ILD perception, including binaural adaptation, appear to be largely attributable to monaural peripheral auditory processing.

Interaural level difference processing as a function of frequency

To localize in the azimuthal plane, humans utilize interaural time differences and interaural level differences (ILDs). For sounds presented in the free field at different azimuthal locations, ILDs generated by the head are complex functions that are frequency dependent and non-monotonic, and are thought to contribute towards sound localization primarily at frequencies above 1500 Hz. However, previous studies have shown that ILD sensitivity is relatively frequency independent for tones presented over headphones, even at frequencies below 1500 Hz. The purpose of this study was to carefully re-investigate the frequency dependency of ILD sensitivity in normal-hearing listeners. Ten normal-hearing listeners were presented ILDs using insert earphones at 31 frequencies between 100 and 11,000 Hz. A 20-dB range of level roving was used to limit listeners’ ability to use monaural level cues to perform the task. On average, ILD sensitivity had several high and low sensitivity frequency regions, was best at 7000 Hz (1.5 dB) and worst at 1000 Hz (2.5 dB). Individuals also showed some unique frequency dependent patterns. These results suggest that ILD sensitivity is frequency dependent and neural models of ILD processing need to accommodate this dependency.

Temporary unilateral hearing loss during development impairs behavioral and neural sensitivity to interaural level difference cues for sound localization

Children who experience persistent conductive hearing loss (CHL) early in life often display binaural hearing impairments that persist long after CHL is resolved, suggesting abnormal central auditory development. Abnormal sensitivity to interaural level differences (ILDs) is particularly likely as a CHL (such as an ear infection) can attenuate sound in the affected ear by >30 dB, dramatically distorting ILD cues. Here, we quantified the effects of unilateral CHL on (1) behavioral spatial acuity and (2) neural information processing of ILD cues in the guinea pig auditory midbrain (inferior colliculus, IC) using the mathematical framework of Fisher information (FI). Animals raised with unilateral CHL displayed larger minimum audible angles for high-pass noise compared to age-matched controls, suggesting impaired ILD sensitivity. Based on acoustic directional transfer function measurements, ILD discrimination thresholds were elevated by ~3–6 dB. Following behavior, extracellular recordings were made in the I...

Blind people are more sensitive than sighted people to binaural sound-location cues, particularly inter-aural level differences

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.

Effect of background noise on neuronal coding of interaural level difference cues in rat inferior colliculus.

Humans can accurately localize sounds even in unfavourable signal-to-noise conditions. To investigate the neural mechanisms underlying this, we studied the effect of background wide-band noise on neural sensitivity to variations in interaural level difference (ILD), the predominant cue for sound localization in azimuth for high-frequency sounds, at the characteristic frequency of cells in rat inferior colliculus (IC). Binaural noise at high levels generally resulted in suppression of responses (55.8%), but at lower levels resulted in enhancement (34.8%) as well as suppression (30.3%). When recording conditions permitted, we then examined if any binaural noise effects were related to selective noise effects at each of the two ears, which we interpreted in light of well-known differences in input type (excitation and inhibition) from each ear shaping particular forms of ILD sensitivity in the IC. At high signal-to-noise ratios (SNR), in most ILD functions (41%), the effect of background noise appeared to be due to effects on inputs from both ears, while for a large percentage (35.8%) appeared to be accounted for by effects on excitatory input. However, as SNR decreased, change in excitation became the dominant contributor to the change due to binaural background noise (63.6%). These novel findings shed light on the IC neural mechanisms for sound localization in the presence of continuous background noise. They also suggest that some effects of background noise on encoding of sound location reported to be emergent in upstream auditory areas can also be observed at the level of the midbrain.

Responses of neurons in the marmoset primary auditory cortex to interaural level differences: comparison of pure tones and vocalizations.

Interaural level differences (ILDs) are the dominant cue for localizing the sources of high frequency sounds that differ in azimuth. Neurons in the primary auditory cortex (A1) respond differentially to ILDs of simple stimuli such as tones and noise bands, but the extent to which this applies to complex natural sounds, such as vocalizations, is not known. In sufentanil/N2O anesthetized marmosets, we compared the responses of 76 A1 neurons to three vocalizations (Ock, Tsik, and Twitter) and pure tones at cells' characteristic frequency. Each stimulus was presented with ILDs ranging from 20 dB favoring the contralateral ear to 20 dB favoring the ipsilateral ear to cover most of the frontal azimuthal space. The response to each stimulus was tested at three average binaural levels (ABLs). Most neurons were sensitive to ILDs of vocalizations and pure tones. For all stimuli, the majority of cells had monotonic ILD sensitivity functions favoring the contralateral ear, but we also observed ILD sensitivity functions that peaked near the midline and functions favoring the ipsilateral ear. Representation of ILD in A1 was better for pure tones and the Ock vocalization in comparison to the Tsik and Twitter calls; this was reflected by higher discrimination indices and greater modulation ranges. ILD sensitivity was heavily dependent on ABL: changes in ABL by ±20 dB SPL from the optimal level for ILD sensitivity led to significant decreases in ILD sensitivity for all stimuli, although ILD sensitivity to pure tones and Ock calls was most robust to such ABL changes. Our results demonstrate differences in ILD coding for pure tones and vocalizations, showing that ILD sensitivity in A1 to complex sounds cannot be simply extrapolated from that to pure tones. They also show A1 neurons do not show level-invariant representation of ILD, suggesting that such a representation of auditory space is likely to require population coding, and further processing at subsequent hierarchical stages.

Rate effects in interaural and sequential level difference perception

The relative weighting of post-onset interaural level difference (ILD) cues decreases with increasing modulation rate of a signal. It is unclear, however, whether overall ILD sensitivity decreases with increasing rate, particularly if stimulus duration and loudness are kept constant. Moreover, it is unclear if the rate effect arises also in monaural sequential level discrimination. In experiment 1, ILD-based lateralization discrimination thresholds and sequential level difference (SLD) discrimination thresholds were measured using bandpass-filtered pulse trains (4 kHz) with rates of 100, 400, and 800 pulses/s. From 100 to 400 pulses/s, both ILD and SLD thresholds decreased, contrary to a rate limitation. From 400 to 800 pulses/s ILD thresholds increased while SLD thresholds remained constant. The latter result is consistent with a high-rate limitation being specific to binaural hearing. Experiment 2 evaluated whether this rate limitation is due to the loss of transmitted modulation at high rates. The ILD thresholds for an unmodulated 4-kHz pure tone were indeed higher than those for pulse-trains. An auditory nerve model (Zilany et al., 2014, JASA 135, 283–286) combined with an interaural discharge rate comparison stage qualitatively predicted the nonmonotonic pattern of ILD thresholds. Overall, the results suggest that modulation can be beneficial for ILD perception.

Linear coding of complex sound spectra by discharge rate in neurons of the medial nucleus of the trapezoid body (MNTB) and its inputs

The interaural level difference (ILD) cue to sound location is first encoded in the lateral superior olive (LSO). ILD sensitivity results because the LSO receives excitatory input from the ipsilateral cochlear nucleus and inhibitory input indirectly from the contralateral cochlear nucleus via glycinergic neurons of the ipsilateral medial nucleus of the trapezoid body (MNTB). It is hypothesized that in order for LSO neurons to encode ILDs, the sound spectra at both ears must be accurately encoded via spike rate by their afferents. This spectral-coding hypothesis has not been directly tested in MNTB, likely because MNTB neurons have been mostly described and studied recently in regards to their abilities to encode temporal aspects of sounds, not spectral. Here, we test the hypothesis that MNTB neurons and their inputs from the cochlear nucleus and auditory nerve code sound spectra via discharge rate. The Random Spectral Shape (RSS) method was used to estimate how the levels of 100-ms duration spectrally stationary stimuli were weighted, both linearly and non-linearly, across a wide band of frequencies. In general, MNTB neurons, and their globular bushy cell inputs, were found to be well-modeled by a linear weighting of spectra demonstrating that the pathways through the MNTB can accurately encode sound spectra including those resulting from the acoustical cues to sound location provided by head-related directional transfer functions (DTFs). Together with the anatomical and biophysical specializations for timing in the MNTB-LSO complex, these mechanisms may allow ILDs to be computed for complex stimuli with rapid spectrotemporally-modulated envelopes such as speech and animal vocalizations and moving sound sources.

Synaptic mechanisms underlying interaural level difference selectivity in rat auditory cortex.

The interaural level difference (ILD) is a sound localization cue that is extensively processed in the auditory brain stem and midbrain and is also represented in the auditory cortex. Here, we asked whether neurons in the auditory cortex passively inherit their ILD tuning from subcortical sources or whether their spiking preferences were actively shaped by local inhibition. If inherited, the ILD selectivity of spiking output should match that of excitatory synaptic input. If shaped by local inhibition, by contrast, excitation should be more broadly tuned than spiking output with inhibition suppressing spiking for nonpreferred stimuli. To distinguish between these two processing strategies, we compared spiking responses with excitation and inhibition in the same neurons across a range of ILDs and average binaural sound levels. We found that cells preferring contralateral ILDs (often called EI cells) followed the inheritance strategy. In contrast, cells that were unresponsive to monaural sounds but responded predominantly to near-zero ILDs (PB cells) instead showed evidence of the local processing strategy. These PB cells received excitatory inputs that were similar to those received by the EI cells. However, contralateral monaural sounds and ILDs >0 dB elicited strong inhibition, quenching the spiking output. These results suggest that in the rat auditory cortex, EI cells do not utilize inhibition to shape ILD sensitivity, whereas PB cells do. We conclude that an auditory cortical circuit computes sensitivity for near-zero ILDs.

Spiking models of interaural level difference encoding - beyond the rate subtraction code

Interaural Level Difference (ILD) provides an important cue for the location of a sound source in the azimuthal plane. Typically, ILD decoding in the brainstem is modeled as a subtraction of spike rates, with inhibitory inputs from one ear subtracted from the excitatory inputs from the other [1-3]. The inferior colliculus (IC) is known to receive input from this circuit, and to encode the spatial location of sounds. Recent experimental evidence suggests that inhibitory input for ILD doesn’t provide subtraction, but instead provides a gain adjustment [4]. In addition, the exact mechanism of the creation of spatial receptive fields in the inferior colliculus remains unclear, and may also be a gain mechanism [5]. The excitatory input to the IC from neurons that decode ILD may contain spike timing cues for location. These spike-timing cues may be initiated in the ILD encoding cells even if the cues are absent from the inputs from the cochlear nucleus. In this study we used a spiking neuron model to recreate and model the full circuit of ILD sensitivity, and explore both the issue of ILD decoding, and the representation of sound source location in the IC. The auditory periphery was modeled as a gammatone filterbank which provided inputs directly to a leaky integrate-and-fire model representing the cells of the cochlear nucleus. These cells are known to lock to the envelope of the sound stimulus, and this behavior was recreated by low-pass filtering of the gammatone filterbank inputs to the cells, and use of a dynamic spike threshold mechanism [6]. The ILD sensitive cells and IC cells were both modeled as simple leaky integrate-and-fire neurons. The model was able to recreate important experimental results regarding ILD encoding cells, particularly the variation of sensitivity with source intensity, and successfully created spatial receptive fields like those found in the IC. The results will be helpful in the future understanding of the binaural mechanisms of the auditory brainstem.

Neuronal interaural level difference response shifts are level-dependent in the rat auditory cortex

How does the brain accomplish sound localization with invariance to total sound level? Sensitivity to interaural level differences (ILDs) is first computed at the lateral superior olive (LSO) and is observed at multiple levels of the auditory pathway, including the central nucleus of inferior colliculus (ICC) and auditory cortex. In LSO, this ILD sensitivity is level-dependent, such that ILD response functions shift toward the ipsilateral (excitatory) ear with increasing sound level. Thus early in the processing pathway changes in firing rate could indicate changes in sound location, sound level, or both. In ICC, while ILD responses can shift toward either ear in individual neurons, there is no net ILD response shift at the population level. In behavioral studies of human sound localization acuity, ILD sensitivity is invariant to increasing sound levels. Level-invariant sound localization would suggest transformation in level sensitivity between LSO and perception of sound sources. Whether this transformation is completed at the level of the ICC or continued at higher levels remains unclear. It also remains unknown whether perceptual sound localization is level-invariant in rats, as it is in humans. We asked whether ILD sensitivity is level-invariant in rat auditory cortex. We performed single-unit and whole cell recordings in rat auditory cortex under ketamine anesthesia and measured responses to white noise bursts presented through sealed earphones at a range of ILDs. Surprisingly, we found that with increasing sound levels ILD responses shifted toward the ipsilateral ear (which is typically inhibitory), regardless of whether cells preferred ipsilateral, contralateral, or binaural stimuli. Voltage-clamp recordings suggest that synaptic inhibition does not contribute substantially to this transformation in level sensitivity. We conclude that the level invariance of ILD sensitivity seen in behavioral studies is not present in rat auditory cortex.

Behavioural sensitivity to binaural spatial cues in ferrets: evidence for plasticity in the duplex theory of sound localization.

For over a century, the duplex theory has guided our understanding of human sound localization in the horizontal plane. According to this theory, the auditory system uses interaural time differences (ITDs) and interaural level differences (ILDs) to localize low-frequency and high-frequency sounds, respectively. Whilst this theory successfully accounts for the localization of tones by humans, some species show very different behaviour. Ferrets are widely used for studying both clinical and fundamental aspects of spatial hearing, but it is not known whether the duplex theory applies to this species or, if so, to what extent the frequency range over which each binaural cue is used depends on acoustical or neurophysiological factors. To address these issues, we trained ferrets to lateralize tones presented over earphones and found that the frequency dependence of ITD and ILD sensitivity broadly paralleled that observed in humans. Compared with humans, however, the transition between ITD and ILD sensitivity was shifted toward higher frequencies. We found that the frequency dependence of ITD sensitivity in ferrets can partially be accounted for by acoustical factors, although neurophysiological mechanisms are also likely to be involved. Moreover, we show that binaural cue sensitivity can be shaped by experience, as training ferrets on a 1-kHz ILD task resulted in significant improvements in thresholds that were specific to the trained cue and frequency. Our results provide new insights into the factors limiting the use of different sound localization cues and highlight the importance of sensory experience in shaping the underlying neural mechanisms.

The discrimination of interaural level difference sensitivity functions: development of a taxonomic data template for modelling

BackgroundA major cue for the position of a high-frequency sound source in azimuth is the difference in sound pressure levels in the two ears, Interaural Level Differences (ILDs), as a sound is presented from different positions around the head. This study aims to use data classification techniques to build a descriptive model of electro-physiologically determined neuronal sensitivity functions for ILDs. The ILDs were recorded from neurons in the central nucleus of the Inferior Colliculus (ICc), an obligatory midbrain auditory relay nucleus. The majority of ICc neurons (~ 85%) show sensitivity to ILDs but with a variety of different forms that are often difficult to unambiguously separate into different information-bearing types. Thus, this division is often based on laboratory-specific and relatively subjective criteria. Given the subjectivity and non-uniformity of ILD classification methods in use, we examined if objective data classification techniques for this purpose. Our key objectives were to determine if we could find an analytical method (A) to validate the presence of four typical ILD sensitivity functions as is commonly assumed in the field, and (B) whether this method produced classifications that mapped on to the physiologically observed results.MethodsThe three-step data classification procedure forms the basic methodology of this manuscript. In this three-step procedure, several data normalization techniques were first tested to select a suitable normalization technique to our data. This was then followed by PCA to reduce data dimensionality without losing the core characteristics of the data. Finally Cluster Analysis technique was applied to determine the number of clustered data with the aid of the CCC and Inconsistency Coefficient values.ResultsThe outcome of a three-step analytical data classification process was the identification of seven distinctive forms of ILD functions. These seven ILD function classes were found to map to the four “known” ideal ILD sensitivity function types, namely: Sigmoidal-EI, Sigmoidal-IE, Peaked, and Insensitive, ILD functions, and variations within these classes. This indicates that these seven templates can be utilized in future modelling studies.ConclusionsWe developed a taxonomy of ILD sensitivity functions using a methodological data classification approach. The number and types of generic ILD function patterns found with this method mapped well on to our electrophysiologically determined ILD sensitivity functions. While a larger data set of the latter functions may bring a more robust outcome, this good mapping is encouraging in providing a principled method for classifying such data sets, and could be well extended to other such neuronal sensitivity functions, such as contrast tuning in vision.

Behavioral Sensitivity to Broadband Binaural Localization Cues in the Ferret

Although the ferret has become an important model species for studying both fundamental and clinical aspects of spatial hearing, previous behavioral work has focused on studies of sound localization and spatial release from masking in the free field. This makes it difficult to tease apart the role played by different spatial cues. In humans and other species, interaural time differences (ITDs) and interaural level differences (ILDs) play a critical role in sound localization in the azimuthal plane and also facilitate sound source separation in noisy environments. In this study, we used a range of broadband noise stimuli presented via customized earphones to measure ITD and ILD sensitivity in the ferret. Our behavioral data show that ferrets are extremely sensitive to changes in either binaural cue, with levels of performance approximating that found in humans. The measured thresholds were relatively stable despite extensive and prolonged (>16 weeks) testing on ITD and ILD tasks with broadband stimuli. For both cues, sensitivity was reduced at shorter durations. In addition, subtle effects of changing the stimulus envelope were observed on ITD, but not ILD, thresholds. Sensitivity to these cues also differed in other ways. Whereas ILD sensitivity was unaffected by changes in average binaural level or interaural correlation, the same manipulations produced much larger effects on ITD sensitivity, with thresholds declining when either of these parameters was reduced. The binaural sensitivity measured in this study can largely account for the ability of ferrets to localize broadband stimuli in the azimuthal plane. Our results are also broadly consistent with data from humans and confirm the ferret as an excellent experimental model for studying spatial hearing.

Onset- and offset-specific effects in interaural level difference discrimination

The relative sensitivity of human listeners to interaural level differences (ILDs) carried by the onsets, offsets, and interior portions of brief sounds was examined. Stimuli consisted of single 4000-Hz Gabor clicks (Gaussian-windowed tone bursts) or trains of 16 such clicks repeating at an interclick interval (ICI) of 2 or 5 ms. In separate conditions, ILDs favored the right ear by a constant amount for all clicks (condition RRRR) or a changing amount that was maximal at sound onset (condition R000), offset (condition 000R), both onset and offset (condition R00R), or at the temporal midpoint of the stimulus (condition 0RR0). ILD increases and decreases were implemented as linear decibel sweeps across four clicks to minimize transient distortion. Threshold ILDs were determined adaptively for each of these conditions and for single clicks. Thresholds were similar for ILDs presented near sound onset or offset (condition R000 vs 000R) but lower when ILDs were carried by both onset and offset clicks (condition R00R) than for ILDs carried by interior clicks alone (condition 0RR0). The results suggest that similar sensitivity to onset and offset ILD does not reflect uniform temporal weighting; instead, ILD sensitivity favors onsets and offsets over the interior portions of sounds.