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Directional hearing in the gray tree frog Hyla versicolor: eardrum vibrations and phonotaxis.

1. We used laser vibrometry to study the vibrational frequency response of the eardrum of female gray tree frogs for different positions of the sound source in three-dimensional space. Furthermore, we studied the accuracy of 3-D phonotaxis in the same species for sounds with different frequency contents. 2. The directionality of the eardrum was most pronounced in a narrow frequency range between 1.3 and 1.8 kHz. 3. The average 3-D, horizontal and vertical jump error angles for phonotactic approaches with a sound similar to the natural advertisement call (1.1 and 2.2 kHz frequency components) were 23 degrees, 19 degrees and 12 degrees, respectively. 4. 3-D jump error angle distributions for the 1.4 + 2.2 kHz, 1.0 kHz and 2.0 kHz sounds were not significantly different from that for the 1.1 + 2.2 kHz sound. 5. The average 3-D jump error angle for the 1.4 kHz sound was 36 degrees, and the distribution was significantly different from that for the 1.1 + 2.2 kHz sound. Hence, phonotactic accuracy was poorer in the frequency range of maximum eardrum directionality. 6. Head scanning was not observed and is apparently unnecessary for accurate sound localization in three-dimensional space. 7. Changes in overall sound pressure level experienced by the frog during phonotactic approaches are not an important cue for sound localization.

Acoustic Stimulation↗

Ventriloquism in patients with unilateral visual neglect.

Can visual stimuli that go undetected, because they are presented in the extinguished region of neglect patients' visual field, nevertheless shift in their direction the apparent location of simultaneous sounds (the well-known 'ventriloquist effect')? This issue was examined using a situation in which each trial involved the simultaneous presentation of a tone over loudspeakers, together with a bright square area on either the left, the right or both sides of fixation. Participants were required to report the presence of squares, and indicate by hand pointing the apparent location of the tone. Five patients with left hemineglect consistently failed to detect the left square, either presented alone or together with another square on the right. Nevertheless, on bimodal trials with a single undetected square to the left, their sound localization was significantly shifted in the direction of that undetected square. By contrast, in bimodal trials with either a single square on the right or a square on each side, their sound localization showed only small and non-significant shifts. This particular result might be due to a combination of low discrimination of lateral sound deviations with variable individual strategies triggered by conscious detection of the right square. The important finding is the crossmodal bias produced by the undetected left visual distractors. It provides a new example of implicit processing of inputs affected by unilateral visual neglect, and on the other hand is consistent with earlier demonstrations of the automaticity of crossmodal bias.

Aged↗

Utility of monaural spectral cues is enhanced in the presence of cues to sound-source lateral angle.

The contention that normally binaural listeners can localize sound under monaural conditions has been challenged by Wightman and Kistler (J. Acoust. Soc. Am. 101:1050-1063, 1997), who found that listeners are almost completely unable to localize virtual sources of sound when sound is presented to only one ear. Wightman and Kistler's results raise the question of whether monaural spectral cues are used by listeners to localize sound under binaural conditions. We have examined the possibility that monaural spectral cues provide useful information regarding sound-source elevation and front-back hemifield when interaural time differences are available to specify sound-source lateral angle. The accuracy with which elevation and front-back hemifield could be determined was compared between a monaural condition and a binaural condition in which a wide-band signal was presented to the near ear and a version of the signal that had been lowpass-filtered at 2.5 kHz was presented to the far ear. It was found that accuracy was substantially greater in the latter condition, suggesting that information regarding sound-source lateral angle is required for monaural spectral cues to elevation and front-back hemifield to be correctly interpreted.

Acoustic Stimulation↗

Update on bilateral cochlear implantation.

PURPOSE OF REVIEW: Providing a unilateral cochlear implant in a patient with a profound bilateral hearing loss has now been a standard clinical practice for more than a decade. Although results are often very good, normal hearing has not been restored. One exciting opportunity to improve hearing in this population is to provide a second implant. However, it is not obvious that bilateral electrical stimulation can be integrated by the central nervous system. This article describes binaural hearing and reviews currently published articles on binaural cochlear implants. RECENT FINDINGS: Controlled laboratory trials have focused on distinguishing different categories of potential binaural advantages. A potential summation effect occurs when the same stimulus is available at two ears. Listening in noise with two ears should be better than listening with one ear when the additional ear is away from the noise. This head shadow benefit results from acoustic effects, not physiologic ones. When the second ear is added near the noise source, a binaural squelch benefit can occur, requiring neural integration from both sides. Finally, two ears may improve sound localization. Binaural implantees generally benefit from head shadow effects. Only some benefit from summation and squelch effects. Most, but not all, show improved horizontal plane localization. SUMMARY: It is now appropriate to begin experimental studies of binaural cochlear implants. Preliminary results show promise to improve head shadow, a physical advantage, and sound localization. Some benefits have been observed for improved summation and squelch. These findings have demonstrated that the brain can integrate electrical stimulation from the two ears. Future studies will be required to maximize this binaural hearing.

Cochlear Implantation↗

Bilateral hearing aids--effects and consequences from a user perspective.

The present study assesses the experiences with bilateral amplification in a group of 144 people provided with two hearing aids between May 1994 and July 1997. Two-thirds of the participants preferred to use both prescribed hearing aids. The participants reported that they used their hearing aids frequently. Bilateral amplification was appreciated especially for recognizing speech, for sound localization and for superior sound quality. Situations in which the subjects showed the highest appreciation of two hearing aids were when attending lectures and theatre performances and for communication in noisy background situations. There were statistically significant differences between users of bilateral and unilateral hearing aids regarding judgement of sound quality of hearing aid processed signals and localization ability in favour of bilateral hearing aid amplification. The self-assessed advantages of bilateral amplification demonstrated in this study should motivate a generous prescription strategy of two hearing aids.

Acoustic Stimulation↗

Localization of brief sounds: effects of level and background noise.

Listeners show systematic errors in vertical-plane localization of wide-band sounds when tested with brief-duration stimuli at high intensities, but long-duration sounds at any comfortable level do not produce such errors. Improvements in high-level sound localization associated with increased stimulus duration might result from temporal integration or from adaptation that might allow reliable processing of later portions of the stimulus. Free-field localization judgments were obtained for clicks and for 3- and 100-ms noise bursts presented at sensation levels from 30 to 55 dB. For the brief (clicks and 3-ms) stimuli, listeners showed compression of elevation judgments and increased rates and unusual patterns of front/back confusion at sensation levels higher than 40-45 dB. At lower sensation levels, brief sounds were localized accurately. The localization task was repeated using 3-ms noise burst targets in a background of spatially diffuse, wide-band noise intended to pre-adapt the system prior to the target onset. For high-level targets, the addition of background noise afforded mild release from the elevation compression effect. Finally, a train of identical, high-level, 3-ms bursts was found to be localized more accurately than a single burst. These results support the adaptation hypothesis.

Adolescent↗

Receptive fields and binaural interactions for virtual-space stimuli in the cat inferior colliculus.

Sound localization depends on multiple acoustic cues such as interaural differences in time (ITD) and level (ILD) and spectral features introduced by the pinnae. Although many neurons in the inferior colliculus (IC) are sensitive to the direction of sound sources in free field, the acoustic cues underlying this sensitivity are unknown. To approach this question, we recorded the responses of IC cells in anesthetized cats to virtual space (VS) stimuli synthesized by filtering noise through head-related transfer functions measured in one cat. These stimuli not only possess natural combinations of ITD, ILD, and spectral cues as in free field but also allow precise control over each cue. VS receptive fields were measured in the horizontal and median vertical planes. The vast majority of cells were sensitive to the azimuth of VS stimuli in the horizontal plane for low to moderate stimulus levels. Two-thirds showed a "contra-preference" receptive field, with a vigorous response on the contralateral side of an edge azimuth. The other third of receptive fields were tuned around a best azimuth. Although edge azimuths of contra-preference cells had a broad distribution, best azimuths of tuned cells were near the midline. About half the cells tested were sensitive to the elevation of VS stimuli along the median sagittal plane by showing either a peak or a trough at a particular elevation. In general receptive fields for VS stimuli were similar to those found in free-field studies of IC neurons, suggesting that VS stimulation provided the essential cues for sound localization. Binaural interactions for VS stimuli were studied by comparing responses to binaural stimulation with responses to monaural stimulation of the contralateral ear. A majority of cells showed either purely inhibitory (BI) or mixed facilitatory/inhibitory (BF&I) interactions. Others showed purely facilitatory (BF) or no interactions (monaural). Binaural interactions were correlated with azimuth sensitivity: most contra-preference cells had either BI or BF&I interactions, whereas tuned cells were usually BF. These correlations demonstrate the importance of binaural interactions for azimuth sensitivity. Nevertheless most monaural cells were azimuth-sensitive, suggesting that monaural cues also play a role. These results suggest that the azimuth of a high-frequency sound source is coded primarily by edges in azimuth receptive fields of a population of ILD-sensitive cells.

Acoustic Stimulation↗

Localization of virtual sound at 4 Gz.

INTRODUCTION: Acceleration directed along the body's z-axis (Gz) leads to misperception of the elevation of visual objects (the "elevator illusion"), most probably as a result of errors in the transformation from eye-centered to head-centered coordinates. We have investigated whether the location of sound sources is misperceived under increased Gz. METHOD: Visually guided localization responses were made, using a remotely controlled laser pointer, to virtual auditory targets under conditions of 1 and 4 Gz induced in a human centrifuge. As these responses would be expected to be affected by the elevator illusion, we also measured the effect of Gz on the accuracy with which subjects could point to the horizon. RESULTS: Horizon judgments were lower at 4 Gz than at 1 Gz, so sound localization responses at 4 Gz were corrected for this error in the transformation from eye-centered to head-centered coordinates. We found that the accuracy and bias of sound localization are not significantly affected by increased Gz. CONCLUSION: The auditory modality is likely to provide a reliable means of conveying spatial information to operators in dynamic environments in which Gz can vary.

Acceleration↗

A precedence effect in the perception of inter-aural cross correlation.

Does the precedence effect, well known in the field of sound localization or lateralization, also apply to other percepts based on binaural processing? We have compared, with one and the same experimental paradigm, a manifestation of the traditional precedence effect in lateralization with a possible similar effect in the perception of diffuseness or compactness of a sound image. With dichotic headphone stimulation, lateralization was controlled by the inter-aural time delay (IATD), and diffuseness/compactness by the inter-aural cross correlation (IACC). The experimental paradigm rests on the principle of estimating the over-all sensation of a 20-ms noise burst, which was subdivided in two parts, with the relevant dichotic information (IATD or IACC) in the leading part being opposite to that in the trailing part. When each part is 10 ms, it is found that the overall sensation is slightly dominated by the information in the leading part, both for lateralization and for compactness/diffuseness. This dominance of the leading part can be compensated by a certain decrease of its duration and/or amplitude relative to that of the trailing part. It is found that this quantitative measure for the 'strength' of the precedence effect for the present stimulus is essentially the same for IATD and IACC, suggesting that the precedence effect does not apply exclusively to sound localization or lateralization, but to at least one other percept based on binaural processing as well, namely the processing of inter-aural cross correlation.

Acoustic Stimulation↗

Cochlear microphonic measurements of interaural time differences in the chick.

The major cues for the sound localization are the interaural differences in the timing and intensity of acoustic information. This poses a difficult coding problem for animals with relatively small heads, such as birds, because interaural time differences (ITDs) would have a small range and magnitude and interaural intensity differences (IIDs) would be significant for only high frequency sounds. It has been suggested that this coding problem is mitigated in birds by an enhancement of ITDs and IIDs resulting from the acoustic coupling of the two middle ear cavities through an interaural canal. In this report, the functional ITDs for sounds at different azimuthal locations were recorded in young chicks, and the contribution of middle ear acoustic coupling was evaluated. ITDs were calculated from simultaneous cochlear microphonic (CM) recordings evoked by pure tone stimuli. These effective ITDs were larger than predicted by the physical separation of the two ears, and this enhancement was more pronounced at low (0.8 and 1 kHz) than at high (2 and 4 kHz) frequencies, reaching maximum values of approximately 180 and 100 microseconds, respectively. The amplitude of the CM also varied as a function of sound source location. This variation was as much as +/- 30%, even for the low frequency tones. This suggests that IID cues are also available to the chick. To determine the contribution of middle ear acoustic coupling to the timing and amplitude of the CM response, the CM in one ear was measured prior to, and following occlusion of the contralateral external auditory canal. The cochlear microphonic from the ear distal to the sound source advanced in time and increased in amplitude when the ear proximal to the sound source was sealed. These effects were more pronounced for low frequency sounds. These findings confirm that acoustic coupling of the middle ear cavities plays a role in enhancing sound localization cues in the chick.

Acoustic Stimulation↗

Spatial processing in the primate auditory cortex.

Spatial localization of auditory stimuli is dependent on the cerebral cortex, yet it remains unclear how cortical activity gives rise to spatial percepts. It has recently been proposed that spatial information is processed serially within the primate auditory cortex, initially in the primary auditory cortex (AI) through the auditory areas caudal to AI, particularly the caudomedial (CM) and caudolateral fields, and onward to the parietal lobe. The activity of single neurons in AI and CM supports this hypothesis, where a greater percentage of CM neurons are sensitive to the spatial location of acoustic stimuli than AI neurons, and the spatial sensitivity of CM neurons extends across a broader range of the stimulus spectrum compared to AI neurons. Further, populations of CM neurons are better able to predict sound localization ability than are populations of AI neurons. We have recently explored the effects of stimulus intensity on both sound localization performance and the spatial sensitivity of auditory cortical neurons. The preliminary results of these experiments again indicate that spatial information is serially processed between AI and the caudal fields. The effects of visual stimulation on auditory localization have also been investigated. Under the appropriate circumstances, visual stimuli can "capture" the spatial location of auditory stimuli in both humans and monkeys. This perceptual illusion suggests that there is a plastic shift in auditory spatial perception. Where the representation of this shift resides is unknown, although two likely candidates are the multimodal regions of the parietal lobe and the superior temporal sulcus.

Acoustic Stimulation↗

Azimuthal tuning of human perceptual channels for sound location.

Human sound localization is acute for frontal locations, but relatively poor in the lateral hemifields. Previous studies in man have not, however, provided evidence on the tuning of the perceptual channels for auditory space that subserve this pattern of acuity. The spatial tuning of perceptual channels used in human azimuthal sound localization was determined using a between-channel auditory temporal gap detection paradigm. In this paradigm, gap thresholds are low when the markers bounding the silent period (gap) activate the same perceptual channel but are elevated when the two markers activate different channels. To determine the tuning of spatial channels, gap thresholds were obtained in an anechoic room with white noise markers coming from each combination of 12 leading marker locations and 18 trailing marker locations throughout the full 360 degrees of azimuth in the horizontal plane through the interaural axis. Gap thresholds remained low (2-4 ms) for all combinations of leading and trailing markers between 30 degrees and 150 degrees in both lateral hemifields. When the leading marker was located deep in one hemifield, and the trailing marker was in the opposite hemifield, gap thresholds rose to 8-16 ms. For leading marker locations at 30 degrees from the midline, gap thresholds were low for all trailing marker locations in the ipsilateral hemifield and locations near the midline in the contralateral hemifield, and were elevated (6-8 ms) only near the contralateral pole. Finally, for leading marker locations at 0 degree or 180 degrees, gap thresholds were low for any trailing location within 30 degrees of the midline at the front or back, and thresholds were elevated for trailing locations at the lateral poles. These data are accountable in terms of two broadly tuned perceptual channels, occupying the left and right auditory hemifields, respectively, each extending 30 degrees across the midline. These channels have widths and locations similar to the spatial receptive fields previously described for central auditory neurons in animals. The data suggest a model of spatial acuity based on the rates of activation of two spatially overlapping channels, rather than the selective activation of members of a large population of finely tuned channels.

Adult↗

Bone-anchored hearing aids in unilateral inner ear deafness.

In nine patients with unilateral deafness and normal hearing in the contralateral ear, measurements of sound localization and speech perception were obtained before intervention, with a conventional contralateral routing of sound (CROS) hearing aid and later with a bone-anchored hearing aid (BAHA) implanted in the deaf ear. Sound localization did not show any differences between the three conditions. Speech perception using short, everyday sentences showed a reduction in the head-shadow effect of 2 dB for both the conventional CROS hearing aid and the BAHA in comparison to the unaided condition. Patients' real-life experiences of the three conditions were evaluated using the Abbreviated Profile of Hearing Aid Benefit (APHAB) questionnaire. The results showed a significant benefit with the BAHA in situations involving background noise and reverberation and a reduced aversion to loud sounds in comparison to the unaided and conventional CROS conditions.

Audiometry, Speech↗

Head-related transfer functions of the Rhesus monkey.

Head-related transfer functions (HRTFs) are direction-specific acoustic filters formed by the head, the pinnae and the ear canals. They can be used to assess acoustical cues available for sound localization and to construct virtual auditory environments. We measured the HRTFs of three anesthetized Rhesus monkeys (Macaca mulatta) from 591 locations in the frontal hemisphere ranging from -90 degrees (left) to 90 degrees (right) in azimuth and -60 degrees (down) to 90 degrees (up) in elevation for frequencies between 0.5 and 15 kHz. Acoustic validation of the HRTFs shows good agreement between free field and virtual sound sources. Monaural spectra exhibit deep notches at frequencies above 9 kHz, providing putative cues for elevation discrimination. Interaural level differences (ILDs) and interaural time differences (ITDs) generally vary monotonically with azimuth between 0.5 and 8 kHz, suggesting that these two cues can be used to discriminate azimuthal position. Comparison with published subsets of HRTFs from squirrel monkeys (Saimiri sciureus) shows good agreement. Comparison with published human HRTFs from the frontal hemisphere demonstrates overall similarity in the patterns of ILD and ITD, suggesting that the Rhesus monkey is a good acoustic model for these two sound localization cues in humans. Finally, the measured ITDs in the horizontal plane agree well between -40 degrees and 40 degrees in azimuth with those calculated from a spherical head model with a radius of 52 mm, one-half the interaural distance of the monkey.

Animals↗