Sensitivity to interaural level and envelope time differences of two bilateral cochlear implant listeners using clinical sound processors

OBJECTIVES: To assess the sensitivity of two bilateral cochlear implant users to interaural level and time differences (ILDs and ITDs) for various signals presented through the auxiliary inputs of clinical sound processors that discard fine timing information and only preserve envelope cues. DESIGN: In a lateralization discrimination experiment, the just noticeable difference (JND) for ILDs and envelope ITDs was measured by means of an adaptive 2-AFC method. Different stimuli were used, including click trains at varying repetition rates, a speech fragment, and noise bursts. For one cochlear implant listener and one stimulus, the sensitivity to envelope ITDs was also determined with the method of constant stimuli. The dependency of ILD-JNDs on the interaural place difference was studied with stimulation at single electrode pairs by using sinusoidal input signals in combination with appropriate single-channel processor fittings. In a lateralization position experiment, subjects were required to use a visual pointer on a computer screen to indicate in-the-head positions for blocks of stimuli containing either ILD or ITD cues. All stimuli were loudness balanced (before applying ILD) and fed directly into the auxiliary inputs of the BTE processors (TEMPO+, Med-El Corp.). The automatic gain control and the processors' microphones were deactivated. RESULTS: Both cochlear implant listeners were highly sensitive to ILDs in all broadband stimuli used; JNDs approached those of normal-hearing listeners. Pitch-matched single electrode pairs showed significantly lower ILD-JNDs than pitch-mismatched electrode pairs. Envelope ITD-JNDs of cochlear implant listeners obtained with the adaptive method were substantially higher and showed a higher test-retest variability than waveform ITD-JNDs of normal-hearing control listeners and envelope ITD-JNDs of normal-hearing listeners reported in the literature for comparable signals. The envelope ITD-JNDs for the click trains were significantly lower than for the speech token or the noise bursts. The best envelope ITD-JND measured was ca. 250 mus for the click train at 100 cycles per sec. Direct measurement of the psychometric function for envelope ITD by the method of constant stimuli showed discrimination above chance level down to 150 micros. The lateralization position experiment showed that both ILDs and envelope ITDs can lead to monotonic changes in lateral percept. CONCLUSIONS: The two cochlear implant users tested showed strong effects of ILDs in various broadband stimuli with respect to JNDs as well as lateralization position. The high dependency of ILD-JNDs on the interaural pitch difference suggests the potential importance of pitch-matched assignment of electrodes in the two ears by the speech processors. Envelope ITDs appear to be more ambiguous cues than ILDs, as reflected by the higher and more variable JNDs compared with normal-hearing listeners. The envelope ITD-JNDs of cochlear implant listeners depended on the stimulus.     

Are Interaural Time and Level Differences Represented by Independent or Integrated Codes in the Human Auditory Cortex?

Sound localization is important for orienting and focusing attention and for segregating sounds from different sources in the environment. In humans, horizontal sound localization mainly relies on interaural differences in sound arrival time and sound level. Despite their perceptual importance, the neural processing of interaural time and level differences (ITDs and ILDs) remains poorly understood. Animal studies suggest that, in the brainstem, ITDs and ILDs are processed independently by different specialized circuits. The aim of the current study was to investigate whether, at higher processing levels, they remain independent or are integrated into a common code of sound laterality. For that, we measured late auditory cortical potentials in response to changes in sound lateralization elicited by perceptually matched changes in ITD and/or ILD. The responses to the ITD and ILD changes exhibited significant morphological differences. At the same time, however, they originated from overlapping areas of the cortex and showed clear evidence for functional coupling. These results suggest that the auditory cortex contains an integrated code of sound laterality, but also retains independent information about ITD and ILD cues. This cue-related information might be used to assess how consistent the cues are, and thus, how likely they would have arisen from the same source.     

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.     

Modulation Enhancement in the Electrical Signal Improves Perception of Interaural Time Differences with Bimodal Stimulation

Interaural timing cues are important for sound source localization and for binaural unmasking of speech that is spatially separated from interfering sounds. Users of a cochlear implant (CI) with residual hearing in the non-implanted ear (bimodal listeners) can only make very limited use of interaural timing cues with their clinical devices. Previous studies showed that bimodal listeners can be sensitive to interaural time differences (ITDs) for simple single- and three-channel stimuli. The modulation enhancement strategy (MEnS) was developed to improve the ITD perception of bimodal listeners. It enhances temporal modulations on all stimulated electrodes, synchronously with modulations in the acoustic signal presented to the non-implanted ear, based on measurement of the amplitude peaks occurring at the rate of the fundamental frequency in voiced phonemes. In the first experiment, ITD detection thresholds were measured using the method of constant stimuli for five bimodal listeners for an artificial vowel, processed with either the advanced combination encoder (ACE) strategy or with MEnS. With MEnS, detection thresholds were significantly lower, and for four subjects well within the physically relevant range. In the second experiment, the extent of lateralization was measured in three subjects with both strategies, and ITD sensitivity was determined using an adaptive procedure. All subjects could lateralize sounds based on ITD and sensitivity was significantly better with MEnS than with ACE. The current results indicate that ITD cues can be provided to bimodal listeners with modified sound processing.     

Interaural time and level difference thresholds for acoustically presented signals in post-lingually deafened adults fitted with bilateral cochlear implants using CIS+ processing

OBJECTIVES: The main purpose of the study was to measure thresholds for interaural time differences (ITDs) and interaural level differences (ILDs) for acoustically presented noise signals in adults with bilateral cochlear implants (CIs). A secondary purpose was to assess the correlation between the ILD and ITD thresholds and error scores in a horizontal-plane localization task, to test the hypothesis that localization by individuals with bilateral implants is mediated by the processing of ILD cues. DESIGN: Eleven adults, all postlingually deafened and all bilaterally fitted with MED-EL COMBI 40+ CIs, were tested in ITD and ILD discrimination tasks in which signals were presented acoustically through headphones that fit over their two devices. The stimulus was a 200-msec burst of Gaussian noise bandpass filtered from 100 to 4000 Hz. A two-interval forced-choice adaptive procedure was used in which the subject had to respond on each trial whether the lateral positions of the two sound images (with the interaural difference favoring the left and right sides in the two intervals) moved from left-to-right or right-to-left. RESULTS: In agreement with previously reported data, ITD thresholds for the subjects with bilateral implants were poor. The best threshold was approximately 400 microsec, and only five of 11 subjects tested achieved thresholds <1000 microsec. In contrast, ILD thresholds were relatively good; mean threshold was 3.8 dB with the initial compression circuit on the implant devices activated and 1.9 dB with the compression deactivated. The ILD and ITD thresholds were higher than previously reported thresholds obtained with direct electrical stimulation (generally, <1.0 dB and 100 to 200 microsec, respectively). When the data from two outlying subjects were omitted, ILD thresholds were highly correlated with total error score in a horizontal-plane localization task, computed for sources near midline (r = 0.87, p < 0.01). CONCLUSIONS: The higher ILD and ITD thresholds obtained in this study with acoustically presented signals (when compared with prior data with direct electrical stimulation) can be attributed-at least partially-to the signal processing carried out by the CI in the former case. The processing strategy effectively leaves only envelope information as a basis for ITD discrimination, which, for the acoustically presented noise stimuli, is mainly coded in the onset information. The operation of the compression circuit reduces the ILDs in the signal, leading to elevated ILD thresholds for the acoustically presented signals in this condition. The large magnitude of the ITD thresholds indicates that ITDs could not have contributed to the performance in the horizontal-plane localization task. Overall, the results suggest that for subjects using bilateral implants, localization of noise signals is mediated entirely by ILD cues, with little or no contribution from ITD information.     

Influence of dynamic compression on directional hearing in the horizontal plane

OBJECTIVE: The purpose of this study was to assess the influence of dynamic compression on directional hearing in the frontal horizontal plane. DESIGN: Compression schemes with various compression ratios and attack times were created by using a digital signal processor. The influence of compression on isolated interaural level differences (ILDs) and interaural time differences (ITDs) was examined in discrimination experiments. The combination of ILDs and ITDs was investigated with measurements of localization based on head-related transfer functions. Five normal listeners and seven hearing-impaired subjects participated in the study. RESULTS: Dynamic compression revealed substantial effects on discrimination of ILDs but not on ITDs. With higher compression ratios and shorter attack times, just-noticeable differences for ILDs increased. Lateralization via head-related transfer functions was predominantly affected by compression in the higher frequency range because for lower frequencies, interaural time cues dominate over interaural level cues. The effect of compression was similar for normal and hearing-impaired listeners, but the latter performed worse on almost all tasks, especially with ITD discrimination at 4000 Hz. CONCLUSIONS: The influence of compression ratio and attack time could be shown for ILDs, but the effect was rather small, at least for the low compression ratios commonly used in hearing aids. The influence further decreased when attack time was prolonged. The dramatic impairment of the discrimination of ITDs with the hearing-impaired subjects in the high-frequency range suggests that they primarily rely on ILD cues.     

The effects of experimentally induced conductive hearing loss on spectral and temporal aspects of sound transmission through the ear

Conductive hearing loss (CHL) is known to produce hearing deficits, including deficits in sound localization ability. The differences in sound intensities and timing experienced between the two tympanic membranes are important cues to sound localization (ILD and ITD, respectively). Although much is known about the effect of CHL on hearing levels, little investigation has been conducted into the actual impact of CHL on sound location cues. This study investigated effects of CHL induced by earplugs on cochlear microphonic (CM) amplitude and timing and their corresponding effect on the ILD and ITD location cues. Acoustic and CM measurements were made in 5 chinchillas before and after earplug insertion, and again after earplug removal using pure tones (500?Hz to 24?kHz). ILDs in the unoccluded condition demonstrated position and frequency dependence where peak far-lateral ILDs approached 30?dB for high frequencies. Unoccluded ear ITD cues demonstrated positional and frequency dependence with increased ITD cue for both decreasing frequency (±420?μs at 500?Hz, ±310?μs for 1-4?kHz) and increasingly lateral sound source locations. Occlusion of the ear canal with foam plugs resulted in a mild, frequency-dependent conductive hearing loss of 10-38?dB (mean 31?±?3.9?dB) leading to a concomitant frequency dependent increase in ILDs at all source locations. The effective ITDs increased in a frequency dependent manner with ear occlusion as a direct result of the acoustic properties of the plugging material, the latter confirmed via acoustical measurements using a model ear canal with varying volumes of acoustic foam. Upon ear plugging with acoustic foam, a mild CHL is induced. Furthermore, the CHL induced by acoustic foam results in substantial changes in the magnitudes of both the ITD and ILD cues to sound location.     

Sensitivity of the auditory middle latency response of the guinea pig to interaural level and time differences

This study examines the extent to which the auditory middle latency response (MLR) of the guinea pig is sensitive to sound localization cues such as interaural level and time differences (ILD and ITD, respectively). The MLR was recorded with an epidural electrode placed over the auditory cortex of an anesthetized guinea pig. Click stimuli were presented monaurally or binaurally with various ILDs and ITDs. The MLR was much larger for contralateral stimulation than for ipsilateral stimulation, and its amplitude was intermediate for diotic stimulation. The MLR amplitude was sensitive to both ILD and ITD: it decreased as the ipsilateral stimulus increased in level or advanced in time relative to the contralateral stimulus. The steep slope of the amplitude-versus-ITD function fell within an ITD range of +/-330 micros, namely the guinea pig's physiological ITD range. The response reduction that resulted from increasing the relative level of the ipsilateral level could be cancelled out by advancing the contralateral onset time relative to the ipsilateral onset time. This parallels the "time-intensity trading" in sound lateralization exhibited in human psychophysics. The results imply that the binaural interaction in the guinea pig MLR reflects aspects of neural processes that are involved in sound localization.     

Channel Interaction and Current Level Affect Across-Electrode Integration of Interaural Time Differences in Bilateral Cochlear-Implant Listeners

Sensitivity to interaural time differences (ITDs) is important for sound localization. Normal-hearing listeners benefit from across-frequency processing, as seen with improved ITD thresholds when consistent ITD cues are presented over a range of frequency channels compared with when ITD information is only presented in a single frequency channel. This study aimed to clarify whether cochlear-implant (CI) listeners can make use of similar processing when being stimulated with multiple interaural electrode pairs transmitting consistent ITD information. ITD thresholds for unmodulated, 100-pulse-per-second pulse trains were measured in seven bilateral CI listeners using research interfaces. Consistent ITDs were presented at either one or two electrode pairs at different current levels, allowing for comparisons at either constant level per component electrode or equal overall loudness. Different tonotopic distances between the pairs were tested in order to clarify the potential influence of channel interaction. Comparison of ITD thresholds between double pairs and the respective single pairs revealed systematic effects of tonotopic separation and current level. At constant levels, performance with double-pair stimulation improved compared with single-pair stimulation but only for large tonotopic separation. Comparisons at equal overall loudness revealed no benefit from presenting ITD information at two electrode pairs for any tonotopic spacing. Irrespective of electrode-pair configuration, ITD sensitivity improved with increasing current level. Hence, the improved ITD sensitivity for double pairs found for a large tonotopic separation and constant current levels seems to be due to increased loudness. The overall data suggest that CI listeners can benefit from combining consistent ITD information across multiple electrodes, provided sufficient stimulus levels and that stimulating electrode pairs are widely spaced.     

The influence of externalization and spatial cues on the generation of auditory brainstem responses and middle latency responses

The effect of externalization and spatial cues on the generation of auditory brainstem responses (ABRs) and middle latency responses (MLRs) was investigated in this study. Most previous evoked potential studies used click stimuli with variations of interaural time (ITDs) and interaural level differences (ILDs) which merely led to a lateralization of sound inside the subject's head. In contrast, in the present study potentials were elicited by a virtual acoustics stimulus paradigm with 'natural' spatial cues and compared to responses to a diotic, non-externalized reference stimulus. Spatial sound directions were situated on the horizontal plane (corresponding to variations in ITD, ILD, and spectral cues) or the midsagittal plane (variation of spectral cues only). An optimized chirp was used which had proven to be advantageous over the click since it compensates for basilar membrane dispersion. ABRs and MLRs were recorded from 32 scalp electrodes and both binaural potentials (B) and binaural difference potentials (BD, i.e., the difference between binaural and summed monaural responses) were investigated. The amplitudes of B and BD to spatial stimuli were not higher than those to the diotic reference. ABR amplitudes decreased and latencies increased with increasing laterality of the sound source. A rotating dipole source exhibited characteristic patterns in dependence on the stimulus laterality. For the MLR data, stimulus laterality was reflected in the latency of component N(a). In addition, dipole source analysis revealed a systematic magnitude increase for the dipole contralateral to the azimuthal position of the sound source. For the variation of elevation, the right dipole source showed a stronger activation for stimuli away from the horizontal plane. The results indicate that at the level of the brainstem and primary auditory cortex binaural interaction is mostly affected by interaural cues (ITD, ILD). Potentials evoked by stimuli with natural combinations of ITD, ILD, and spectral cues were not larger than those elicited by diotic chirps.     

Mapping lateralization of click trains in younger and older populations

The main purpose of this study was to describe and compare lateralization of earphone-presented stimuli in younger and older individuals. Lateralization functions, relating perceived location to either interaural time differences (ITDs) or interaural level differences (ILDs) were determined for 78 subjects, aged 21-88 years, who responded by pressing one of nine keys to indicate the perceived location of the stimulus. All subjects were healthy, without any history of hearing loss or ear surgery and within the normal pure tone audiometric range for their age group. Interaural pure tone and click thresholds did not differ by more than 5 dB across ears. The ILD lateralization functions, ranging from 10 dB favoring the left ear to 10 dB favoring the right ear were linear. In contrast, the ITD lateralization functions were S-shaped with a clear linear component ranging from 750 micros favoring one ear to 750 micros favoring the other ear and with an asymptote from 750 micros to 1 ms. The same general shape of the ITD and ILD lateralization functions was found at all ages, but the linear slope of the ITD lateralization function became shallower with age. The ability to discriminate midline-located click trains (ITD and ILD=0) from ITD-lateralized click trains deteriorated with age, while the comparable ability to discriminate ILD-lateralized click trains did not change significantly with age. The data support two general conclusions. First there seems to be an overall reduction in the range of ITD-based lateralization due to aging. Second, there is a greater reduction in sensitivity due to aging in changes from the perceived midline position (ITD and ILD=0) when ITD is manipulated than when ILD is manipulated.     

Interaural Time Difference Perception with a Cochlear Implant and a Normal Ear

Currently there is a growing population of cochlear-implant (CI) users with (near) normal hearing in the non-implanted ear. This configuration is often called SSD (single-sided deafness) CI. The goal of the CI is often to improve spatial perception, so the question raises to what extent SSD CI listeners are sensitive to interaural time differences (ITDs). In a controlled lab setup, sensitivity to ITDs was investigated in 11 SSD CI listeners. The stimuli were 100-pps pulse trains on the CI side and band-limited click trains on the acoustic side. After determining level balance and the delay needed to achieve synchronous stimulation of the two ears, the just noticeable difference in ITD was measured using an adaptive procedure. Seven out of 11 listeners were sensitive to ITDs, with a median just noticeable difference of 438 μs. Out of the four listeners who were not sensitive to ITD, one listener reported binaural fusion, and three listeners reported no binaural fusion. To enable ITD sensitivity, a frequency-dependent delay of the electrical stimulus was required to synchronize the electric and acoustic signals at the level of the auditory nerve. Using subjective fusion measures and refined by ITD sensitivity, it was possible to match a CI electrode to an acoustic frequency range. This shows the feasibility of these measures for the allocation of acoustic frequency ranges to electrodes when fitting a CI to a subject with (near) normal hearing in the contralateral ear.     

Sound-Localization Experiments with Barn Owls in Virtual Space: Influence of Interaural Time Difference on Head-Turning Behavior

Specific cues in a sound signal are naturally linked to certain parameters in acoustic space. In the barn owl, interaural time difference (ITD) varies mainly with azimuth, while interaural level difference (ILD) varies mainly with elevation. Previous data suggested that ITD is indeed the main cue for azimuthal sound localization in this species, while ILD is an important cue for elevational sound localization. The exact contributions of these parameters could be tested only indirectly because it was not possible to generate a stimulus that contained all relevant spatial information on the one hand, and allowed for a clean separation of these parameters on the other hand. Virtual auditory worlds offer this opportunity. Here we show that barn owls responded to azimuthal variations in virtual space in the same way as to variations in free-field stimuli. We interpret the increase of turning angle with sound-source azimuths (up to +/- 140 degrees) such that the owls did not experience front/back confusions with virtual stimuli. We then separated the influence of ITD from the influence of all other stimulus parameters by fixing the overall ITD in virtual stimuli to a constant value (+100 micros or +100 micros) while leaving all other sound characteristics unchanged. This manipulation influenced both azimuthal and elevational components of head arms. Since the owls' azimuthal head-turn amplitude always resembled the value signified by the ITD, these data demonstrated that azimuthal sound localization is influenced only by ITD both in the frontal hemisphere and in large parts of the rear hemisphere. ILDs did not have an influence on azimuthal components of head turns. While response latency to normal virtual stimuli was found to be largely independent of stimulus position, response delays of the head turns became longer if the ITD information pointed into a different hemisphere as the other cues of the sounds.     

Context Effects in the Discriminability of Spatial Cues

In order to investigate whether performance in an auditory spatial discrimination task depends on the prevailing listening conditions, we tested the ability of human listeners to discriminate target sounds with and without presentation of a preceding sound. Target sounds were either lateralized by means of interaural time differences (ITDs) of +400, 0, or −400 μs or interaural level differences (ILDs) with the same subjective intracranial locations. The preceding sound was always lateralized by means of ITD. This allowed for testing whether the effects of a preceding sound were location- or cue-specific. Preceding sounds and target sounds were randomly paired across trials. Listeners had to discriminate whether they perceived the target sounds as coming from the same or different intracranial locations. Finally, stimuli were selected so that, without any preceding sound, ITD and ILD cues were equally discriminable at all target lateralizations. Stimuli were 800 Hz-wide, 400-ms duration bands of noise centered at 500 Hz, presented over headphones. The duration of the preceding sound was randomly selected from a uniform distribution spanning from 1s to 2s. Results show that discriminability of both binaural cues was improved for midline target positions when preceding sound and targets were co-located, whereas it was impaired when preceding sound and targets came from different positions. No effect of the preceding sound was found for left or right target positions. These results are compatible with a purely bottom–up mechanism based on adaptive coding of ITD around the midline that may be combined with top–down mechanisms to increase localization accuracy in realistic listening conditions.     

Interaction in the perceptual processing of interaural time and level differences

Phillips and Hall [Psychophysical evidence for adaptation of central auditory processors for interaural differences in time and level, Hear. Res., 202 (2005) 188-199.] recently described the frequency-specific, selective adaptation of perceptual channels for interaural differences in level (ILD) and time (ITD). Psychometric functions for laterality based on ITD or ILD were obtained before and after exposure to adaptor tones of two frequencies presented alternately and highly lateralized to opposite sides. Following adaptation, points of perceived centrality (PPCs) were displaced towards the sides of the adaptor tones, and in opposite directions for the two frequencies. That is, laterality judgements showed a shift away from the adapted side, particularly for test cue values near the middle of the range. These data were congruent with a two-channel, opponent-process model of sound laterality coding. The present study used the same general paradigm to explore the independence of perceptual ITD and ILD processing. Psychometric functions for laterality based on ITD or ILD were obtained for each of two frequencies concurrently, before and after exposure to adaptor tones lateralized using the complementary cue. Once again, PPCs derived from the psychometric functions were displaced towards the sides of the adaptor tones, consistent with an opponent-process account of sound laterality coding. The size of the adaptation effect was at least as great as that described in the earlier study. Thus, a quarter cycle ITD adapting stimulus effected a 3 dB shift in the mean ILD-based PPC, and a 12 dB ILD adapting stimulus effected a 100 micros shift in the mean ITD-based PPC. These data offer new evidence concerning interaction in the processing of ITDs and ILDs.     

Interaural Correlation Fails to Account for Detection in a Classic Binaural Task: Dynamic ITDs Dominate N0Sπ Detection

Binaural signal detection in an NoSπ task relies on interaural disparities introduced by adding an antiphasic signal to diotic noise. What metric of interaural disparity best predicts performance? Some models use interaural correlation; others differentiate between dynamic interaural time differences (ITDs) and interaural level differences (ILDs) of the effective stimulus. To examine the relative contributions of ITDs and ILDs in binaural detection, we developed a novel signal processing technique that selectively degrades different aspects (potential cues) of binaural stimuli (e.g., only ITDs are scrambled). Degrading a particular cue will affect performance only if that cue is relevant to the binaural processing underlying detection. This selective scrambling technique was applied to the stimuli of a classic N0Sπ task in which the listener had to detect an antiphasic 500-Hz signal in the presence of a diotic wideband noise masker. Data obtained from five listeners showed that (1) selective scrambling of ILDs had little effect on binaural detection, (2) selective scrambling of ITDs significantly degraded detection, and (3) combined scrambling of ILDs and ITDs had the same effect as exclusive scrambling of ITDs. Regarding the question which stimulus properties determine detection, we conclude that for this binaural task (1) dynamic ITDs dominate detection performance, (2) ILDs are largely irrelevant, and (3) interaural correlation of the stimulus is a poor predictor of detection. Two simple stimulus-based models that each reproduce all binaural aspects of the data quite well are described: (1) a single-parameter detection model using ITD variance as detection criterion and (2) a compressive transformation followed by a crosscorrelation analysis. The success of both of these contrasting models shows that our data alone cannot reveal the mechanisms underlying the dominance of ITD cues. The physiological implications of our findings are discussed.     

Behavioral sensitivity to interaural time differences in the rabbit

An important cue for sound localization and separation of signals from noise is the interaural time difference (ITD). Humans are able to localize sounds within 1-2 degrees and can detect very small changes in the ITD (10-20micros). In contrast, many animals localize sounds with less precision than humans. Rabbits, for example, have sound localization thresholds of approximately 22 degrees . There is only limited information about behavioral ITD discrimination in animals with poor sound localization acuity that are typically used for the neural recordings. For this study, we measured behavioral discrimination of ITDs in the rabbit for a range of reference ITDs from 0 to +/-300micros. The behavioral task was conditioned avoidance and the stimulus was band-limited noise (500-1500Hz). Across animals, the average discrimination threshold was 50-60micros for reference ITDs of 0 to +/-200micros. There was no trend in the thresholds across this range of reference ITDs. For a reference ITD of +/-300micros, which is near the limit of the physiological window defined by the head width in this species, the discrimination threshold increased to approximately 100micros. The ITD discrimination in rabbits less acute than in cats, which have a similar head size. This result supports the suggestion that ITD discrimination, like sound localization [see Heffner, 1997. Acta Otolaryngol. 532 (Suppl.), 46-53] is determined by factors other than head size.     

Concurrent Development of the Head and Pinnae and the Acoustical Cues to Sound Location in a Precocious Species,the Chinchilla (<Emphasis Type="Italic">Chinchilla lanigera)</Emphasis>

Sounds are filtered in a spatial- and frequency-dependent manner by the head and pinna giving rise to the acoustical cues to sound source location. These spectral and temporal transformations are dependent on the physical dimensions of the head and pinna. Therefore, the magnitudes of binaural sound location cues—the interaural time (ITD) and level (ILD) differences—are hypothesized to systematically increase while the lower frequency limit of substantial ILD production is expected to decrease due to the increase in head and pinna size during development. The frequency ranges of the monaural spectral notch cues to source elevation are also expected to decrease. This hypothesis was tested here by measuring directional transfer functions (DTFs), the directional components of head-related transfer functions, and the linear dimensions of the head and pinnae for chinchillas from birth through adulthood. Dimensions of the head and pinna increased by factors of 1.8 and 2.42, respectively, reaching adult values by ~6 weeks. From the DTFs, the ITDs, ILDs, and spectral shape cues were computed. Maximum ITDs increased by a factor of 1.75, from ~160 μs at birth (P0-1, first postnatal day) to 280 μs in adults. ILDs depended on source location and frequency exhibiting a shift in the frequency range of substantial ILD (>10 dB) from higher to lower frequencies with increasing head and pinnae size. Similar trends were observed for the spectral notch frequencies which ranged from 14.7–33.4 kHz at P0-1 to 5.3–19.1 kHz in adults. The development of the spectral notch cues, the spatial- and frequency-dependent distributions of DTF amplitude gain, acoustic directionality, maximum gain, and the acoustic axis were systematically related to the dimensions of the head and pinnae. The dimension of the head and pinnae in the chinchilla as well as the acoustical properties associated with them are mature by ~6 weeks.     

Sound pressure transformations by the head and pinnae of the adult Chinchilla (Chinchilla lanigera)

There are three main cues to sound location: the interaural differences in time (ITD) and level (ILD) as well as the monaural spectral shape cues. These cues are generated by the spatial- and frequency-dependent filtering of propagating sound waves by the head and external ears. Although the chinchilla has been used for decades to study the anatomy, physiology, and psychophysics of audition, including binaural and spatial hearing, little is actually known about the sound pressure transformations by the head and pinnae and the resulting sound localization cues available to them. Here, we measured the directional transfer functions (DTFs), the directional components of the head-related transfer functions, for 9 adult chinchillas. The resulting localization cues were computed from the DTFs. In the frontal hemisphere, spectral notch cues were present for frequencies from ～6-18?kHz. In general, the frequency corresponding to the notch increased with increases in source elevation as well as in azimuth towards the ipsilateral ear. The ILDs demonstrated a strong correlation with source azimuth and frequency. The maximum ILDs were <10?dB for frequencies <5?kHz, and ranged from 10-30?dB for the frequencies >5?kHz. The maximum ITDs were dependent on frequency, yielding 236?μs at 4?kHz and 336?μs at 250?Hz. Removal of the pinnae eliminated the spectral notch cues, reduced the acoustic gain and the ILDs, altered the acoustic axis, and reduced the ITDs.     

Sound localization and sensitivity to interaural cues in bilateral users of the Med-El Combi 40/40+cochlear implant system.

OBJECTIVE: The purpose of this study was to investigate sound localization in subjects bilaterally implanted with MED-EL COMBI 40/40+ cochlear implants. In addition, the sensitivity to interaural cues was assessed. METHODS: In the localization test (11 subjects), CCITT noise (500 ms, original and HRTF-filtered, 70/75/80 dB sound pressure level) was presented from one of seven loudspeakers between -90 degrees and 90 degrees azimuth. The subject had to indicate which loudspeaker the noise was presented from. Sensitivity to interaural level differences (ILD) was assessed by performing localization tests (4 subjects) with the loudness of the two speech processors unbalanced to various degrees. To investigate the subjects' sensitivity to interaural time differences (ITD), lateralization was measured (7 subjects) as a function of the time difference between two Gaussian-like pulses, each directed to one of the subject's speech processor microphones by way of headphones. RESULTS: The judgments of all subjects significantly correlated with the positions of the loudspeakers. The scatter in the judged azimuth measured by the standard deviation of the responses was on average 27.5 degrees . Unbalanced loudness of the speech processors produced a bias in azimuth toward the speech processor with the louder volume setting. The mean rate of shift was 1.4 degrees per unit on the Wuerzburg loudness scale. Six of seven subjects showed a significant sensitivity to ITDs with the approximate time difference required for complete lateralization being 1,200 micros on an average. The one subject not showing a sensitivity to ITDs performed worst in the localization test. CONCLUSIONS: Bilateral cochlear implantation can restore spatial hearing in cochlear implant users. Both ILDs and ITDs are used by bilateral cochlear implant users in sound localization with ILDs appearing to be the dominant cue.