Localization of azimuthal sound direction by the great horned owl.
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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.
In this study we have investigated the processing of auditory cues for sound localization in the great horned owl (Bubo virginianus). Previous studies have shown that the barn owl, whose ears are asymmetrically oriented in the vertical plane, has a 2-dimensional, topographic representation of auditory space in the external division of the inferior colliculus (ICx). As in the barn owl, the great horned owl's ICx is anatomically distinct and projects to the optic tectum. Neurons in ICx respond over only a small range of azimuths (mean = 32 degrees), and azimuth is topographically mapped. In contrast to the barn owl, the great horned owl has bilaterally symmetrical ears and its receptive fields are not restricted in elevation. The binaural cues available for sound localization were measured both with cochlear microphonic recordings and with a microphone attached to a probe tube in the auditory canal. Interaural time disparity (ITD) varied monotonically with azimuth. Interaural intensity differences (IID) also changed with azimuth, but the largest IIDs were less than 15 dB, and the variation was not monotonic. Neither ITD nor IID varied systematically with changes in the vertical position of a sound source. We used dichotic stimulation to determine the sensitivity of ICx neurons to these binaural cues. Best ITD of ICx units was topographically mapped and strongly correlated with receptive-field azimuth. The width of ITD tuning curves, measured at 50% of the maximum response, averaged 72 microseconds. All ICx neurons responded only to binaural stimulation and had nonmonotonic IID tuning curves. Best IID was weakly, but significantly, correlated with best ITD (r = 0.39, p less than 0.05). The IID tuning curves, however, were broad (mean 50% width = 24 dB), and 67% of the units had best IIDs within 5 dB of 0 dB IID. ITD tuning was sensitive to variations in IID in the direction opposite to that expected for time-intensity trading, but the magnitude of this effect was only 1.5 microseconds/dB IID. We conclude that, in the great horned owl, the spatial selectivity of ICx neurons arises primarily from their ITD tuning. Except for the absence of elevation selectivity and the narrow range of best IIDs, ICx in the great horned owl appears to be organized much the same as in the barn owl.
1. The behavioral audiograms of four Japanese macaques (Macaca fuscata) were assessed before and after receiving two-stage bilateral lesions of auditory cortex. Thresholds were assessed for each ear with the use of insertion earphones. 2. The bilateral lesions resulted in a large initial hearing loss followed by partial recovery that left the animals with a permanent hearing loss in both ears. 3. The initial hearing loss consisted of a total insensitivity to sound in the ear contralateral to the second lesion with limited hearing in the other ear. However, the animal with the most complete lesion was initially unable to hear sound in either ear. Broadband noise was often more effective in eliciting a behavioral response than tones. 4. Partial recovery occurred in all animals and was observed as early as the first week after surgery. Most of this recovery occurred during the first 3-7 wk after surgery. This rapid phase of recovery was sometimes followed by a more gradual phase although thresholds were still elevated after 94 wk. 5. The permanent hearing loss, which averaged from 30 to 44 dB, was not constant across frequency. Threshold shifts were smallest at 63 Hz and progressively increased with frequency to a maximum loss from 8 to 25 kHz with slightly less loss at 32 kHz. 6. Analysis of the psychophysical functions and threshold stability gave no indication of any nonsensory deficits in attention or vigilance. 7. These results, taken with those of previous experiments, indicate that each hemisphere is primarily involved in the detection of sound in the contralateral ear and secondarily involved in detection in the ipsilateral ear. This arrangement differs from that seen in sound localization where each hemisphere is involved with the contralateral hemifield as opposed to the contralateral ear. Thus it appears that the functional organization of auditory cortex for sound localization is different from that for the detection and identification of sound itself.
The dorsal division of the cochlear nucleus (DCN) is the most complex of its subdivisions in terms of both anatomical organization and physiological response types. Hypotheses about the functional role of the DCN in hearing are as yet primitive, in part because the organizational complexity of the DCN has made development of a comprehensive and predictive model of its input-output processing difficult. The responses of DCN cells to complex stimuli, especially filtered noise, are interesting because they demonstrate properties that cannot be predicted, without further assumptions, from responses to narrow band stimuli, such as tones. In this paper, we discuss the functional organization of the DCN, i.e. the morphological organization of synaptic connections within the nucleus and the nature of synaptic interactions between its cells. We then discuss the responses of DCN principal cells to filtered noise stimuli that model the spectral sound localization cues produced by the pinna. These data imply that the DCN plays a role in interpreting sound localization cues; supporting evidence for such a role is discussed.
1. A currently unresolved question concerning the effects of auditory decortication on sound localization is whether or not operated animals have a normal capacity for discriminating the small interaural differences in phase angle or intensity that result from the spatial separation of sound sources relative to the head. The present experiment was designed to provide data relevant to this question. 2. Four normal and three operated cats (bilateral ablations of AI, AII Ep, SII, I-T), wearing stereo headsets, were tested with an active avoidance procedure to detect reversals in the interaural phase-angle or intensity relations of binaural 1-kHz tones. For both groups of cats, the detection thresholds for interaural intensity and phase angle were found to be close to 1 dB and 5 degrees, respectively. 3. In addition, we found that both unoperated and operated cats exhibited positive transfer from the original lateralization task involving the detection of interaural reversals of phase angle or intensity to a new test, which required the cats to identify, in an absolute sense, which ear received the leading or louder signals. 4. Thus, the present investigation provides additional evidence that the neocortex has no primary sensory role in sound localization.
Rapid control of responding by sound location is obtained in squirrel monkeys when sound stimuli are presented from one of two loudspeakers, each one adjacent to a response key. With this arrangement of loudspeakers and response keys, squirrel monkeys quickly learn to respond on the key near the source of the sound stimulus, and this pattern is the same whether or not responses near the sound source are differentially reinforcedmthis result may depend on a pre-experimental tendency in squirrel monkeys to orient head and eyes toward a sound, which would lead the animal to look at the response key in front of the loudspeaker producing the sound. The present experiment sought to determine whether visual stimuli are necessary for rapid control of responding by sound location. Two monkeys were trained in darkness in a sound-localization task similar to that described above. Results were similar to those obtained from animals trained in light, indicating that visual stimuli are not required for rapid acquisition of sound-localization behavior in monkeys.
The effects of external noise on fetal sheep cerebral glucose utilization were determined with the [14C]deoxyglucose method. Seventeen animals were prepared at 130 days gestation with catheters and electrodes for assessing fetal behavioral state. Five to 7 days later, 7 animals were studied under normal laboratory sound conditions (65-70 dB), 5 animals were exposed to 105-120 dB broadband noise levels produced by two earphones applied to the abdomen of the ewe, and 5 fetuses were stimulated with an electronic artificial larynx (EAL), positioned on the abdomen directly over the fetal head. There were no significant differences between local cerebral glucose utilization in controls and earphone ewes, and no obvious alteration in behavioral states. However, there were marked, significant differences in glucose utilization along the central auditory pathway during EAL stimulation. These autoradiographs revealed isofrequency-like bands in medial geniculate body and irregular darkening of cortex of the temporal lobe. Total time spent in clearly defined high and low voltage electrocortical activity did not change during EAL stimulation.
The influence of sound localization behavior on unit activity in the frontal cortex of awake rhesus monkeys was examined by comparing responses under three behavioral conditions: auditory localization, during which a response was required to the location of a sound (broad-band noise) source; auditory detect, during which a response was required to indicate the occurrence of the sound regardless of location; visual localization, during which no sounds were presented and a response was required to the location of a visual stimulus; and nonperform, presentation of auditory stimuli as in the first two conditions, but with the animal sitting passively. Extracellular microelectrode recordings were made in the periarcuate region and dorsal and ventral prefrontal areas near the principal sulcus. Four monkeys were used with a total of 498 cells studied. Of the total population, only five cells were found to have characteristics similar to those of auditory units in the primary auditory cortex and the surrounding belt area. More typically, units were found that had strong short-latency responses specific to the auditory and/or visual localization tasks. These units had no or weak responses when the same sound stimuli were presented in the auditory detect task or when a monkey received the sound stimuli in a nonperforming condition. Two regions were identified, one medial and/or posterior to the arcuate sulcus, in Brodmann's area 6; the second included parts of areas 8 and 9 within the genu of the arcuate sulcus. Units from these regions are referred to, respectively, as the postarcuate and the prearcuate populations. Both populations responded predominantly during active localization behavior. Sixty-two percent of the postarcuate population responded during auditory localization, 32% responded during auditory detect, and only 18% responded to acoustic stimuli presented in the nonperforming condition. In the prearcuate population percentages in these three conditions were 35, 25, and 12%, respectively. For visual localization, 54% in the postarcuate population responded, whereas 42% in the prearcuate responded. Spatial tuning of units during auditory localization was similar to that seen in units of the primary auditory cortex, with the greatest percentages of units responding to stimuli contralateral to the recording site. Similar tuning was observed for the visual localization task as well. Similarities in spatial tuning between the auditory and visual localization conditions were examined to assess the "bimodal" nature of the units.(ABSTRACT TRUNCATED AT 400 WORDS)
The ability of horses to use binaural time and intensity difference cues to localize sound was assessed in free-field localization tests by using pure tones. The animals were required to discriminate the locus of a single tone pip ranging in frequency from 250 Hz to 25 kHz emitted by loudspeakers located 30 degrees to the left and right of the animals' midline (60 degrees total separation). Three animals were tested with a two-choice procedure; 2 additional animals were tested with a conditioned avoidance procedure. All 5 animals were able to localize 250 Hz, 500 Hz, and 1 kHz but were completely unable to localize 2 kHz and above. Because the frequency of ambiguity for the binaural phase cue delta phi for horses in this test was calculated to be 1.5 kHz, these results indicate that horses can use binaural time differences in the form of delta phi but are unable to use binaural intensity differences. This finding was supported by an unconditioned orientation test involving 4 additional horses, which showed that horses correctly orient to a 500-Hz tone pip but not to an 8-kHz tone pip. Analysis of the superior olivary complex, the brain stem nucleus at which binaural interactions first take place, reveals that the lateral superior olive (LSO) is relatively small in the horse and lacks the laminar arrangement of bipolar cells characteristic of the LSO of most mammals that can use binaural delta I.
A microelectrode survey was made of the midbrain auditory nuclei of the brushtailed possum (Trichosurus vulpecula), a common Australian marsupial. Information was sought on the tuning characteristics of individual neurones, tonotopic organization and mechanisms of sound localization. It was felt that such information would be of use in future studies of the development and evolution of mammalian hearing. Twelve possums were anaesthetized with ketamine and chloralose-urethane, and recordings were made of extracellular unit discharges in the inferior colliculus during monaural and binaural tonal stimulation. The inferior colliculus of the possum consists of a central nucleus - a darkly stained, densely packed group of cells - flanked laterally by an external nucleus with a lower density of paler cells. Tonotopic organization was demonstrated by discretelytuned elements in the central nucleus, but was not observed in the external nucleus. In the latter region broad and irregular tuning was commonly seen. Most units in both divisions were influenced by binaural stimuli, with patterns of binaural interaction similar to those observed in the cat inferior colliculus. Cells influenced by changes in the interaural time and intensity difference were commonly observed, but only a subclass of these were suited in sensitivity for sound localization. In general, the midbrain auditory system of the possum was similar in unit discharge characteristics and organization to those of the eutherian mammals commonly studied.
The role of spectral cues in the sound source to ear transfer function in median plane sound localization is investigated in this paper. At first, transfer functions were measured and analyzed. Then, these transfer functions were used in experiments where sounds from a source on the median plane were simulated and presented to subjects through headphones. In these simulation experiments, the transfer functions were smoothed by ARMA models with different degrees of simplification to investigate the role of microscopic and macroscopic patterns in the transfer functions for median plane localization. The results of the study are summarized as follows: (1) For front-rear judgment, information derived from microscopic peaks and dips in the low-frequency region (below 2 kHz) and the macroscopic patterns in the high-frequency region seems to be utilized; (2) for judgment of elevation angle, major cues exist in the high-frequency region above 5 kHz. The information in macroscopic patterns is utilized instead of that in small peaks and dips.
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