The neural algorithm for sound localization in the owl.
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The physical measurements reported here test whether the European starling (Sturnus vulgaris) evaluates the azimuth direction of a sound source with a peripheral auditory system composed of two acoustically coupled pressure-difference receivers (1) or of two decoupled pressure receivers (2). A directional pattern of sound intensity in the free-field was measured at the entrance of the auditory meatus using a probe microphone, and at the tympanum using laser vibrometry. The maximum differences in the sound-pressure level measured with the microphone between various speaker positions and the frontal speaker position were 2.4 dB at 1 and 2 kHz, 7.3 dB at 4 kHz, 9.2 dB at 6 kHz, and 10.9 dB at 8 kHz. The directional amplitude pattern measured by laser vibrometry did not differ from that measured with the microphone. Neither did the directional pattern of travel times to the ear. Measurements of the amplitude and phase transfer function of the starling's interaural pathway using a closed sound system were in accord with the results of the free-field measurements. In conclusion, although some sound transmission via the interaural canal occurred, the present experiments support the hypothesis 2 above that the starling's peripheral auditory system is best described as consisting of two functionally decoupled pressure receivers.
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1. Binaurally time-shifted and intensity-unbalanced noise, delivered through earphones, induced owls to respond with a head-orienting behavior similar to that which occurs to free field auditory stimuli. 2. Owls derived the azimuthal and elevational coordinates of a sound from a combination of interaural time difference (ITD) and interaural intensity difference (IID). 3. IID and ITD each contained information about the azimuth and elevation of the signal. Thus, IID and ITD formed a coordinate system in which the axes were non-orthogonal. 4. ITD was a strong determinant of azimuth, and IID was a strong determinant of elevation, of elicited head turn.
Listeners were tested under free head/torso movement conditions, using a brief pulsed broadband noise signal presented from any 1 of 20 loudspeakers in two intersecting hemicircumferential arrays: one in the horizontal plane, one in the vertical. The listeners' task was to judge the whereabouts of each signal while listening in normal, open-ears conditions or while wearing various types of standard or electronic earmuffs or compressible foam earplugs. Results showed that listeners could accomplish the task successfully in normal (open-ear) conditions. Results using earplugs, nonelectronic earmuffs, and two types of dichotic (dual microphone) electronic earmuffs showed decrements in performance similar to each other. These decrements took the general form of loss of appreciation of the vertical plane and of front-rear differences in the horizontal plane. Lateral discrimination was blurred but otherwise intact. Of two diotic (single microphone) electronic earmuffs, one completely disrupted localization function and the other came close to it.
Bimodal units in the barn owl's optic tectum are tuned to the location of auditory and visual stimuli, and are systematically organized according to their spatial tuning to form mutually aligned maps of auditory and visual space. Map alignment results from the fact that, normally, units are tuned to the values of interaural level difference (ILD) and interaural time difference (ITD) produced by a sound source at the location of their visual receptive fields (VRFs). Monaural occlusion alters the correspondence of ILD and ITD values with locations in space. We investigated the effect that raising owls with a chronic monaural occlusion has on the tuning of tectal units to ILD and ITD. Owls were monaurally occluded beginning at 1 month of age. The effects of monaural occlusion were assessed 2-4 months later by comparing the ILD and ITD tuning of units in monaurally occluded owls with the ILD and ITD tuning of units with equivalent VRFs in normal owls. ILD and ITD tuning was shifted substantially and in the direction of the unoccluded ear (the adaptive direction) in owls raised with a monaural occlusion. In most tecta, the mapped representations of ILD and ITD were shifted systematically. In addition, in some tecta, monaural occlusion induced a change in the topography of the ILD map such that ILD tuning remained essentially constant at values near 0 dB over abnormally large portions of the tectum. Across all recording sites, the average shift in ILD tuning was 9 dB (n = 396) and the average shift in ITD tuning was 40 microseconds (n = 414). In four of five animals, the magnitude of the effect was not equivalent on the two sides of the brain, the adjustments being significantly larger and more systematic on the side ipsilateral to the occlusion. Such differences in the altered ILD and ITD maps on the two sides of the brain in individual animals indicate that, although a component of the adaptive adjustment might be due to regulation of the gain and phase response of the monaural signals early in the auditory pathway, a major component of the adjustment must occur at or beyond the level where the encoding of ILDs and ITDs for left and right space separates.
1. Bilateral recording of cochlear potentials was used to measure the variations in interaural time differences (ITDs) and interaural intensity differences (IIDs) as a free-field auditory stimulus was moved to different positions around a barn owl's head. 2. ITD varied smoothly with stimulus azimuth across a broad frequency range. 3. ITD varied minimally with stimulus elevation, except at extreme angles from the horizontal. 4. IID varied with both stimulus elevation and stimulus azimuth. Lower frequencies were more sensitive to variations in azimuth, whereas higher frequencies were more sensitive to variations in elevation. 5. The loci of spatial coordinates that form iso-IID contours and iso-ITD contours form a non-orthogonal grid that relates binaural disparity cues to sound location.
Good directional hearing ability demands good and symmetrical hearing in both ears. We report the effect of impaired hearing on the directional hearing ability of 98 patients, especially of patients with bilateral asymmetrical hearing loss. The directional testing device included 12 loudspeakers placed at 30 degree intervals in a circle with a diameter of 3.25 m, whose centre lay between the ears of the patient. In included an audiometer for producing the signals, an amplifier and a PDP11/23 computer interfaced to a loudspeaker switch bank. The subject's answers to 60 directionally randomized stimuli were recorded. During the presentation of the signal the patients were not allowed to turn their head. The patients had to name the number of the loudspeaker on the circle that they thought was producing the sound. In addition to the directional hearing test a pure-tone audiogram was done, and the middle- and high-frequency hearing loss estimated. The records of the directional hearing test were analysed in two new ways: firstly, vector analysis of the errors; secondly, averaging of the difference between the true interaural time delay and the virtual time difference, which was implicated in the possibly incorrect answer of the patient (effective delta-t-parameter). This average gives a score for the uncertainty in defining the correct "cone of confusion". In addition to the statistical analysis, two cases are reported showing the directional hearing ability of two patients with neuromas treated by transtemporal surgery, with some residual hearing.(ABSTRACT TRUNCATED AT 250 WORDS)
Noise localization thresholds and the ability to localize pure tones at 60 degrees separation were determined for three domestic pigs using a conditioned avoidance procedure. The average threshold for localizing a brief noise burst was 4.5 degrees which is much more accurate than the thresholds of other hoofed mammals, such as horses, cattle and goats. The ability of pigs to localize low-frequency tones indicates that they can use the binaural phase-difference cue. However, they were unable to localize tones of 4 kHz and higher, indicating that, like other hoofed mammals, their ability to use binaural intensity cues is greatly restricted if not completely absent. An examination of the superior olivary complex of pigs revealed that in relative size, shape and cell density it is more like that of cats than that of other hoofed mammals.
In the previous paper the directional response characteristics of the ferret auditory periphery were examined. In this study further measurements of the spectral transfer functions (STFs) of the auditory periphery were obtained at locations close to the tympanic membrane. There was considerable variation in the STFs recorded from different animals and between recordings made at each end of the auditory canal in the same animal. However, calculation of the so called "location dependency function" demonstrated that changes in the location of the stimulus produced the same pattern of changes in the STFs in all recordings. Changes in the spectral transformation for azimuth locations in the ipsilateral auditory field were examined by calculating the horizon STF. The gain transformations of frequencies below 20 kHz were found to be asymmetrical about the interaural axis so that maximum gain was obtained for anterior stimulus locations. In contrast, the maximum gain for frequencies above 20 kHz was obtained for stimulus locations about the interaural axis, and movement of the stimulus location into either the anterior or posterior fields produced symmetrical reductions in gain. These changes were related to the directional properties of the periphery examined in the previous paper [S. Carlile, J. Acoust. Soc. Am. 88, 2180-2195 (1990)]. The spatial resolution of the monaural information provided by the peripheral STFs is dependent on the rate of change of the transformations as a function of azimuthal displacement of the stimulus location. This was examined by calculating the unsigned first spatial derivative for each frequency in the horizon STF.(ABSTRACT TRUNCATED AT 250 WORDS)
Neurons in the developing optic tectum adjust their tuning to auditory localization cues in response to chronic monaural occlusion so that auditory spatial fields align with visual receptive fields (VRFs). We tested whether this adaptive adjustment of auditory tuning requires visual instruction. Both eyelids were sutured closed at the same time that one ear was occluded in two barn owls that were 1 month old. After 70 and 100 d, respectively, the tuning of units to interaural level difference (ILD) and to interaural time difference (ITD) was measured. These data were compared with equivalent data from 15 normal owls. Unit tuning to ITD was shifted from normal in both of the monaurally occluded owls. In one owl, ILD tuning was also clearly shifted. In the other owl, the map of ILD was flipped upside down and adaptive adjustments in ILD tuning could not be assessed. Instead, adjustments in ILD tuning were observed following removal of the earplug with the eyelids kept closed. Unit tuning was monitored at several sites in the tectum for 1 month after earplug removal using chronically implanted electrodes. Then, ILD tuning was resampled across the entire tectum. Both measures indicated shifts in ILD tuning in response to removal of the earplug in the second blind owl. In both animals, the magnitude of the shifts in ILD tuning and ITD tuning was smaller than has been observed previously in monaurally occluded but sighted owls. The results demonstrate that the brain can make adaptive adjustments in ILD and ITD tuning in response to early monaural occlusion even without the guiding influence of vision.
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Three groups of hearing-impaired listeners who had been fitted binaurally with behind-the-ear (BTE), in-the-ear (ITE), or in-the-ear-canal (ITC) hearing aids were tested on spatial localization function for sources in the frontal horizontal and vertical planes. No significant differences in unaided performance were observed between the groups, nor between that and aided performance in the BTE and ITE wearers. ITC wearers, by contrast, showed a deterioration in aided over unaided performance. From observations of performance when each group wore temporary fittings of the other two types of system, and from the performance of a non-impaired control group, it appears that the performance decrement for the ITC wearers was due to their own particular systems although specific reasons for this decrement could not be identified. In all conditions, aided and unaided, vertical plane localization was markedly disrupted in all the hearing impaired groups. It was also disrupted, to a lesser but still substantial extent, in aided conditions for the non-impaired listeners.
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