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Hyperacute directional hearing in a microscale auditory system.

The physics of sound propagation imposes fundamental constraints on sound localization: for a given frequency, the smaller the receiver, the smaller the available cues. Thus, the creation of nanoscale acoustic microphones with directional sensitivity is very difficult. The fly Ormia ochracea possesses an unusual 'ear' that largely overcomes these physical constraints; attempts to exploit principles derived from O. ochracea for improved hearing aids are now in progress. Here we report that O. ochracea can behaviourally localize a salient sound source with a precision equal to that of humans. Despite its small size and minuscule interaural cues, the fly localizes sound sources to within 2 degrees azimuth. As the fly's eardrums are less than 0.5 mm apart, localization cues are around 50 ns. Directional information is represented in the auditory system by the relative timing of receptor responses in the two ears. Low-jitter, phasic receptor responses are pooled to achieve hyperacute timecoding. These results demonstrate that nanoscale/microscale directional microphones patterned after O. ochracea have the potential for highly accurate directional sensitivity, independent of their size. Notably, in the fly itself this performance is dependent on a newly discovered set of specific coding strategies employed by the nervous system.

Animal Communication↗

Rapid adaptation to auditory-visual spatial disparity.

The so-called ventriloquism aftereffect is a remarkable example of rapid adaptative changes in spatial localization caused by visual stimuli. After exposure to a consistent spatial disparity of auditory and visual stimuli, localization of sound sources is systematically shifted to correct for the deviation of the sound from visual positions during the previous adaptation period. In the present study, this aftereffect was induced by presenting, within 17 min, 1800 repetitive noise or pure-tone bursts in combination with synchronized, and 20 degrees disparate flashing light spots, in total darkness. Post-adaptive sound localization, measured by a method of manual pointing, was significantly shifted 2.4 degrees (noise), 3.1 degrees (1 kHz tones), or 5.8 degrees (4 kHz tones) compared with the pre-adaptation condition. There was no transfer across frequencies; that is, shifts in localization were insignificant when the frequencies used for adaptation and the post-adaptation localization test were different. It is hypothesized that these aftereffects may rely on shifts in neural representations of auditory space with respect to those of visual space, induced by intersensory spatial disparity, and may thus reflect a phenomenon of neural short-term plasticity.

Acoustic Stimulation↗

Localization and identification of concurrent sounds in the owl's auditory space map.

In nature, sounds from multiple sources sum at the eardrums, generating complex cues for sound localization and identification. In this clutter, the auditory system must determine "what is where." We examined this process in the auditory space map of the barn owl's (Tyto alba) inferior colliculus using two spatially separated sources simultaneously emitting uncorrelated noise bursts, which were uniquely identified by different frequencies of sinusoidal amplitude modulation. Spatial response profiles of isolated neurons were constructed by testing the source-pair centered at various locations in virtual auditory space. The neurons responded whenever a source was placed within the receptive field, generating two clearly segregated foci of activity at appropriate loci. The spike trains were locked strongly to the amplitude modulation of the source within the receptive field, whereas the other source had minimal influence. Two sources amplitude modulated at the same rate were resolved successfully, suggesting that source separation is based on differences of fine structure. The spike rate and synchrony were stronger for whichever source had the stronger average binaural level. A computational model showed that neuronal activity was primarily proportional to the degree of matching between the momentary binaural cues and the preferred values of the neuron. The model showed that individual neurons respond to and synchronize with sources in their receptive field if there are frequencies having an average binaural-level advantage over a second source. Frequencies with interaural phase differences that are shared by both sources may also evoke activity, which may be synchronized with the amplitude modulations from either source.

Acoustic Stimulation↗

Effects of interaural decorrelation on neural and behavioral detection of spatial cues.

The detection of interaural time differences (ITDs) for sound localization critically depends on the similarity between the left and right ear signals (interaural correlation). We show that, like humans, owls can localize phantom sound sources well until the correlation declines to a very low value, below which their performance rapidly deteriorates. Decreasing interaural correlation also causes the response of the owl's tectal auditory neurons to decline nonlinearly, with a rapid drop followed by a more gradual reduction. A detection-theoretic analysis of the statistical properties of neuronal responses could account for the variance of behavioral responses as interaural correlation is decreased. Finally, cross-correlation analysis suggests that low interaural correlations cause misalignment of cross-correlation peaks across different frequencies, contributing heavily to the nonlinear decline in neural and ultimately behavioral performance.

Animals↗

Speech perception, localization, and lateralization with bilateral cochlear implants.

Five bilateral cochlear implant users were tested for their localization abilities and speech understanding in noise, for both monaural and binaural listening conditions. They also participated in lateralization tasks to assess the impact of variations in interaural time delays (ITDs) and interaural level differences (ILDs) for electrical pulse trains under direct computer control. The localization task used pink noise bursts presented from an eight-loudspeaker array spanning an arc of approximately 108 degrees in front of the listeners at ear level (0-degree elevation). Subjects showed large benefits from bilateral device use compared to either side alone. Typical root-mean-square (rms) averaged errors across all eight loudspeakers in the array were about 10 degrees for bilateral device use and ranged from 20 degrees to 60 degrees using either ear alone. Speech reception thresholds (SRTs) were measured for sentences presented from directly in front of the listeners (0 degrees) in spectrally matching speech-weighted noise at either 0 degrees, +90 degrees or -90 degrees for four subjects out of five tested who could perform the task. For noise to either side, bilateral device use showed a substantial benefit over unilateral device use when noise was ipsilateral to the unilateral device. This was primarily because of monaural head-shadow effects, which resulted in robust SRT improvements (P<0.001) of about 4 to 5 dB when ipsilateral and contralateral noise positions were compared. The additional benefit of using both ears compared to the shadowed ear (i.e., binaural unmasking) was only 1 or 2 dB and less robust (P = 0.04). Results from the lateralization studies showed consistently good sensitivity to ILDs; better than the smallest level adjustment available in the implants (0.17 dB) for some subjects. Sensitivity to ITDs was moderate on the other hand, typically of the order of 100 micros. ITD sensitivity deteriorated rapidly when stimulation rates for unmodulated pulse-trains increased above a few hundred Hz but at 800 pps showed sensitivity comparable to 50-pps pulse-trains when a 50-Hz modulation was applied. In our opinion, these results clearly demonstrate important benefits are available from bilateral implantation, both for localizing sounds (in quiet) and for listening in noise when signal and noise sources are spatially separated. The data do indicate, however, that effects of interaural timing cues are weaker than those from interaural level cues and according to our psychophysical findings rely on the availability of low-rate information below a few hundred Hz.

Adult↗

Interaural time sensitivity dominated by cochlea-induced envelope patterns.

To localize sounds in space, humans heavily depend on minute interaural time differences (ITDs) generated by path-length differences to the two ears. Physiological studies of ITD sensitivity have mostly used deterministic, periodic sounds, in which either the waveform fine structure or a sinusoidal envelope is delayed interaurally. For natural broadband stimuli, however, auditory frequency selectivity causes individual channels to have their own envelopes; the temporal code in these channels is thus a mixture of fine structure and envelope. This study introduces a method to disentangle the contributions of fine structure and envelope in both binaural and monaural responses to broadband noise. In the inferior colliculus (IC) of the cat, a population of neurons was found in which envelope fluctuations dominate ITD sensitivity. This population extends over a surprisingly wide range of frequencies, including low frequencies for which fine-structure information is also available. A comparison with the auditory nerve suggests that an elaboration of envelope coding occurs between the nerve and the IC. These results suggest that internally generated envelopes play a more important role in binaural hearing than is commonly thought.

Acoustic Stimulation↗

Development and preliminary evaluation of an earmold sound-to-tactile aid for the hearing-impaired.

A binaural earmold sound-to-tactile aid was constructed by inserting a vibrating element into a Lucite earmold. The earmold could be vibrated at either 80 Hz (when incoming acoustic signals were below 2000 Hz), at 300 Hz (when incoming acoustic signals were above 2000 Hz), or both (when incoming acoustic signals were broadband). Subjects were fitted with one of these bimodal vibrating earmolds in each ear. Normal-hearing and hearing-impaired subjects were tested in three tasks: sound localization, environmental sound identification, and syllable rhythm and stress. The device provided some benefit to performance, although the amounts of improvement varied across tasks and subjects. Possible modifications in device design, and potential combinations of auditory and tactile input via earmold systems, are discussed.

Adult↗

The role of timing in the brain stem auditory nuclei of vertebrates.

Vertebrate animals gain biologically important information from environmental sounds. Localization of sound sources enables animals to detect and respond appropriately to danger, and it allows predators to detect and localize prey. In many species, rapidly fluctuating sounds are also the basis of communication between conspecifics. This information is not provided directly by the output of the ear but requires processing of the temporal pattern of firing in the tonotopic array of auditory nerve fibers. The auditory nerve feeds information through several parallel ascending pathways. Anatomical and electrophysiological specializations for conveying precise timing, including calyceal synaptic terminals and matching axonal conduction times, are evident in several of the major ascending auditory pathways through the ventral cochlear nucleus and its nonmammalian homologues. One pathway that is shared by all higher vertebrates makes an ongoing comparison of interaural phase for the localization of sound in the azimuth. Another pathway is specifically associated with higher frequency hearing in mammals and is thought to make use of interaural intensity differences for localizing high-frequency sounds. Balancing excitation from one ear with inhibition from the other in rapidly fluctuating signals requires that the timing of these synaptic inputs be matched and constant for widely varying sound stimuli in this pathway. The monaural nuclei of the lateral lemniscus, whose roles are not understood (although they are ubiquitous in higher vertebrates), receive input from multiple pathways that encode timing with precision, some through calyceal endings.

Animals↗

The function of the medial superior olive in small mammals: temporal receptive fields in auditory analysis.

Traditionally, the medial superior olive, a mammalian auditory brainstem structure, is considered to encode interaural time differences, the main cue for localizing low-frequency sounds. Detection of binaural excitatory and inhibitory inputs are considered as an underlying mechanism. Most small mammals, however, hear high frequencies well beyond 50 kHz and have small interaural distances. Therefore, they can not use interaural time differences for sound localization and yet possess a medial superior olive. Physiological studies in bats revealed that medial superior olive cells show similar interaural time difference coding as in larger mammals tuned to low-frequency hearing. Their interaural time difference sensitivity, however, is far too coarse to serve in sound localization. Thus, interaural time difference sensitivity in medial superior olive of small mammals is an epiphenomenon. We propose that the original function of the medial superior olive is a binaural cooperation causing facilitation due to binaural excitation. Lagging inhibitory inputs, however, suppress reverberations and echoes from the acoustic background. Thereby, generation of antagonistically organized temporal fields is the basic and original function of the mammalian medial superior olive. Only later in evolution with the advent of larger mammals did interaural distances, and hence interaural time differences, became large enough to be used as cues for sound localization of low-frequency stimuli.

Animals↗

Auditory localization: role of auditory pathways in brain stem of the cat.

Cats were trained to localize sound in space. The animals' localization accuracy was determined before and after one of the following operations: 1) transection of the trapezoid body, 2) unilateral and 3) bilateral transection of the lateral lemniscus, 4) unilateral and 5) bilateral transection of the brachium of the inferior colliculus. The results after bilateral transections of the lateral lemniscus and the one deep bilateral transection of the brachium of the inferior colliculus indicate that some portion of the ascending auditory system must be intact above the medulla for an animal to be able to localize sound. A small loss in accuracy of localization was found after unilateral transection of the lateral lemniscus or brachium of the inferior colliculus. This loss, when compared with the much larger loss that monaural animals show, is an indication that binaural analysis, important for sound localization, occurs at the level of the medulla. Some transections of the trapezoid body resulted in a deficit in localization ability that appeared to be complete and permanent. The position of the lesions in the trapezoid body indicated that important encoding of the binaural cues to localization most likely occurs at the superior olivary complex, probably at the medial superior olive. But the trapezoid body or other commissures of the brain stem auditory system are probably also involved in transmission of information necessary for localization to higher centers.

Animals↗

Binaural and monaural localization of sound in two-dimensional space.

Two experiments were conducted. In experiment 1, part 1, binaural and monaural localization of sounds originating in the left hemifield was investigated. 104 loudspeakers were arranged in a 13 x 8 matrix with 15 degrees separating adjacent loudspeakers in each column and in each row. In the horizontal plane (HP), the loudspeakers extended from 0 degrees to 180 degrees; in the vertical plane (VP), they extended from -45 degrees to 60 degrees with respect to the interaural axis. Findings of special interest were: (i) binaural listeners identified the VP coordinate of the sound source more accurately than did monaural listeners, and (ii) monaural listeners identified the VP coordinate of the sound source more accurately than its HP coordinate. In part 2, it was found that foreknowledge of the HP coordinate of the sound source aided monaural listeners in identifying its VP coordinate, but the converse did not hold. In experiment 2, part 1, localization performances were evaluated when the sound originated from consecutive 45 degrees segments of the HP, with the VP segments extending from -22.5 degrees to 22.5 degrees. Part 2 consisted of measuring, on the same subjects, head-related transfer functions by means of a miniature microphone placed at the entrance of their external ear canal. From these data, the 'covert' peaks (defined and illustrated in text) of the sound spectrum were extracted. This spectral cue was advanced to explain why monaural listeners in this study as well as in other studies performed better when locating VP-positioned sounds than when locating HP-positioned sounds. It is not claimed that there is inherent advantage for localizing sound in the VP; rather, monaural localization proficiency, whether in the VP or HP, depends on the availability of covert peaks which, in turn, rests on the spatial arrangement of the sound sources.

Attention↗

Localization of aerial broadband noise by pinnipeds.

Although many pinnipeds (seals, sea lions, and walruses) emit broadband calls on land as part of their communication system, few studies have addressed these animals' ability to localize aerial broadband sounds. In this study, the aerial sound localization acuities of a female northern elephant seal (Mirounga angustirostris), a male harbor seal (Phoca vitulina), and a female California sea lion (Zalophus californianus) were measured in the horizontal plane. The stimulus was broadband white noise that was band pass filtered between 1.2 and 15 kHz. Testing was conducted in a hemi-anechoic chamber using a left/right forced choice procedure to measure the minimum audible angle (MAA) for each subject. MAAs were defined as half the angular separation of two sound sources bisected by a subject's midline that corresponded to 75% correct discrimination. MAAs were 4.7 degrees, 3.6 degrees, and 4.2 degrees for the northern elephant seal, harbor seal, and California sea lion, respectively. These results demonstrate that individuals of these pinniped species have sound localization abilities comparable to the domestic cat and rhesus macaque. The acuity differences between our subjects were small and not predicted by head size. These results likely reflect the relatively acute general abilities of pinnipeds to localize aerial broadband signals.

Animals↗

Bone-anchored hearing aids in unilateral inner ear deafness: an evaluation of audiometric and patient outcome measurements.

OBJECTIVE: To evaluate the benefit of a bone-anchored hearing aid contralateral routing of sound hearing aid (BAHA CROS hearing aid) in 29 patients with unilateral inner ear deafness. STUDY DESIGN: Prospective clinical follow-up study. SETTING: Tertiary referral center. PATIENTS: Thirty patients were recruited. There were 19 patients with a history of acoustic neuroma surgery and 11 patients with unilateral inner ear deafness due to other causes; 1 patient was excluded. The first 21 patients had also participated in a previous evaluation. INTERVENTION: Audiometric measurements were taken before intervention, when fitted with a conventional CROS, and after BAHA implementation. Patients' subjective benefit was quantified with four different hearing aid-specific instruments: the Abbreviated Profile of Hearing Aid Benefit, the Glasgow Hearing Aid Benefit Profile, the International Outcome Inventory for Hearing Aids, and the Single-Sided Deafness questionnaire. MAIN OUTCOME MEASURES: The same instruments were used at a mean long-term follow-up of 1 year after BAHA implantation. RESULTS: Sound localization in an audiologic test setting was no different from chance level. The main effect of the BAHA CROS that was found was the "lift the head shadow" effect in the speech-in-noise measurements. All instruments also showed positive results in favor of the BAHA CROS at long-term follow-up. CONCLUSIONS: The poor sound-localization results in an audiologic test setting illustrated the inability of patients with unilateral inner ear deafness to localize sounds. The speech-in-noise measurements demonstrated the efficacy of the BAHA CROS to lift the head shadow. Patients were still satisfied at 1-year follow-up, according to the four instruments.

Adult↗

Envelope representations of pinna impulse responses relating to three-dimensional localization of sound sources.

Acoustical impulse responses of the external ear system are measured in order to investigate human three-dimensional auditory localization. A sound source of electric spark discharge is presented in nine directions for each of four vertical planes, 1.5 m from the center of the subject's head. The results show that a human pinna works as a compound sound reflector which produces major reflected components within 350 microseconds. The relationship between the direction of the sound source and the impulse response is clearly shown by the use of an envelope formation technique and radial ray displays.

Auditory Perception↗

Primate hearing from a mammalian perspective.

This review discusses hearing performance in primates and selective pressures that may influence it. The hearing sensitivity and sound-localization abilities of primates, as indicated by behavioral tests, are reviewed and compared to hearing and sound localization among mammals in general. Primates fit the mammalian pattern with small species hearing higher frequencies than larger species in order to use spectral/intensity cues for sound localization. In this broader comparative context, the restricted high-frequency hearing of humans is not unusual. All of the primates tested so far are able to hear frequencies below 125 Hz, placing them among the majority of mammals. Sound-localization acuity has been determined for only three primates, and here also they have relatively good localization acuity (with a minimum audible angle roughly similar to other mammals such as cats, pigs, and opossums). This is in keeping with the pattern among mammals in general, in which species with narrow fields of best vision, such as a fovea, are better localizers than those with broad fields of best vision. Multiple lines of evidence support the view that sound localization is the selective pressure on smaller primates and on other mammals with short interaural distances for hearing high frequencies.

Adaptation, Physiological↗

[Capacity for monaural localization of sound sources in the horizontal plane].

Using special procedures, we investigated monaural spatial sound as perceived in the horizontal plane in 30 otologically healthy subjects, aged 17 to 23 years. It was found that on the masked side (over 50 dB) the localization of sound sources in this plane was disturbed and the numbers of loudspeakers as indicated by the test subjects were shifted towards the open ear. On the open side they showed the ability to locate a loudspeaker positioned at an angle of 18 degrees and failure to locate loudspeakers positioned at angles of 9 degrees and 27 degrees. Also, 60 audiologically healthy people in the age group from 21 to 70 years were examined. Most subjects displayed a similar pattern of response which seems to be independent of their age, sex or the state of concha auriculae.

Adolescent↗

The sensitive period for auditory localization in barn owls is limited by age, not by experience.

Early in life, the barn owl passes through a sensitive period during which it can interpret and make use of abnormal auditory cues for accurate sound localization. This capacity is lost at about 8 weeks of age, just after the head and ears reach adult size (knudsen et al. 1984a). The end of the sensitive period could be triggered either by an age-dependent process or by the exposure of the auditory system to stable or adult-like cues. To distinguish between these alternatives, we subjected baby owls to constant abnormal cues (chronic monaural occlusion) or to frequently changing abnormal cues (alternating monaural occlusion) throughout the sensitive period. In the first group of animals (n = 2), one ear was plugged continuously until 73 or 79 d of age, respectively, and then the earplug was switched to the opposite ear. Although these animals adjusted sound localization accuracy during the initial chronic monaural occlusion, they could not localize sounds at all after the earplug was switched to the opposite ear, and they remained unable to localize sounds as long as the opposite ear remained occluded (7 and 27 weeks, respectively). When the second monaural occlusion was finally removed, both birds localized sounds with errors that were similar to the errors they exhibited immediately after removal of the first monaural occlusion. One bird that was 127-d-old at the time the second earplug was removed corrected its localization error; the other bird, 250-d-old when the second earplug was removed, did not.(ABSTRACT TRUNCATED AT 250 WORDS)

Aging↗

Spectral cues explain illusory elevation effects with stereo sounds in cats.

Mammals localize sound sources in azimuth based on two binaural cues, interaural differences in the time of arrival and level of the sounds at the ears. In contrast, the cue for elevation is based on patterns of the broadband power spectra at each ear that result from the direction-dependent acoustic filtering properties of the head and pinnae. Although the exact form of this "spectral shape" cue is unknown, most attention has been directed toward a prominent direction-dependent energy minimum, or "notch," because its location in frequency, for both humans and cats, moves predictably from low to high as a source is moved from low to high elevations. However, there is little direct evidence that these spectral notches are important elevational cues in animals other than humans. Here we demonstrate a striking illusion in the localization of sounds in elevation by cats using stimulus configurations that elicit summing localization and the precedence effect that can be explained by spectral shape cues.

Animals↗