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At least 19 recordsLinked to original sources

Precognitive and cognitive elements in sound localization.

Sound localization behavior is of great importance for an animal's survival. To localize a sound, animals have to detect a sound source and assign a location to it. In this review we discuss recent results on the underlying mechanisms and on modulatory influences in the barn owl, an auditory specialist with very well developed capabilities to localize sound. Information processing in the barn owl auditory pathway underlying the computations of detection and localization is well understood. This analysis of the sensory information primarily determines the following orienting behavior towards the sound source. However, orienting behavior may be modulated by cognitive (top-down) influences such as attention. We show how advanced stimulation techniques can be used to determine the importance of different cues for sound localization in quasi-realistic stimulation situations, how attentional influences can improve the response to behaviorally relevant stimuli, and how attention can modulate related neural responses. Taken together, these data indicate how sound localization might function in the usually complex natural environment.

Journal Article↗

Midline and lateral field sound localization in the ferret (Mustela putorius): contribution of the superior olivary complex.

1. The ability of ferrets to localize sound in space was determined before and after unilateral or bilateral lesions of the superior olivary complex (SOC). Lesions were made by pressure injection of kainic acid into the SOC through a stereotaxically positioned glass micropipette. The lesions destroyed the cell bodies in the superior olive without disrupting fibers of passage in the trapezoid body or other pathways in the auditory brain stem. The integrity of fibers was demonstrated by protargol staining of axonal processes and by the retrograde transport of horseradish peroxidase (HRP) from the inferior colliculus to other auditory brain stem nuclei. Behavioral tests were carried out separately for sound localization at midline and lateral field positions. Minimum audible angles were determined for single 45-ms noise bursts presented through paired loudspeakers positioned symmetrically around 0, -60, and +60 degrees azimuth. 2. Four ferrets received complete lesions of the left SOC, and two received complete lesions of the right SOC. In general, unilateral destruction of the superior olive resulted in impairments in sound localization in both left and right lateral fields. In some cases, deficits were also apparent on midline. Four additional animals received unilateral lesions that spared cells within the SOC. In most cases, deficits were apparent despite incomplete lesions of the SOC. The pattern of deficits was generally consistent with that found in animals with complete lesions. Most animals had difficulty localizing sounds in the lateral fields. 3. Four animals received bilateral lesions of the SOC. Three had complete or near-complete destruction of the superior olive on one side of the brain with relatively minor damage on the other side. Each of these animals exhibited behavioral deficits that were particularly severe ipsilateral to the more extensively damaged superior olive. One animal with complete bilateral destruction of the SOC was incapable of sound localization, even with 2-s noise bursts. This animal, however, suffered severe motor impairments after surgery that might have contributed to the apparent inability to localize sound. 4. Two animals with kainic acid lesions that caused little or no damage to the SOC were still capable of high levels of performance in tests of sound localization and had no elevation in minimum audible angles. These cases served as controls for the possible effects of nonspecific brain damage and demonstrated that kainic acid injections per se resulted in no obvious deficits in our test situation.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Distortion and adaptation in underwater sound localization.

Sounds should be localized at more medial positions under water than in air, because the values of the interaural temporal and intensity differences are reduced in water. Thirteen blindfolded divers were required to move a pointer towards the apparent location of a sound source under water, and they showed significant errors towards the median plane. In a second experiment, five divers were tested in air before and after underwater training at swimming towards a sound source. The search patterns of these divers showed systematic errors towards the median plane. A comparison of the first and second air tests showed some evidence of an aftereffect away from the median plane, discounting the most lateral angles (80 degrees left and right) where little effect would be expected. It is concluded that both a distortion of localization and some adaptation to the distortion occur under water.

Auditory Perception↗

Sound localization after unilateral lesions of inferior colliculus in the ferret (Mustela putorius).

1. The ability of ferrets to localize sounds in space was determined before and after large unilateral lesions of the inferior colliculus. Three animals sustained restricted lesions that destroyed the inferior colliculus caudally but spared some tissue rostrally. These restricted lesions did not extend substantially beyond the cytoarchitectonic boundaries of the inferior colliculus. Four animals received more extensive lesions that destroyed all of the inferior colliculus, including the rostral pole. These lesions also involved to various degrees other midbrain structures, including the dorsal nucleus of the lateral lemniscus and the lateral tegmentum. Psychophysical curves and minimum audible angles were obtained for midline and left and right lateral fields. Comparisons were made of deficits ipsilateral and contralateral to the lesion. 2. The animals were tested in a semicircular apparatus with loudspeakers located at various positions around the periphery. Separate sound localization tests were conducted for midline, left, and right field positions. In each test the animals were trained first to discriminate between speakers separated by 60 degrees. They were then tested with progressively smaller angles of speaker separation to obtain their psychophysical threshold (minimum audible angle). All critical tests of sound localization were based on the ability of animals to identify the location of a 45-ms noise burst presented at the beginning of each trial. Correct responses were rewarded by delivery of a small quantity of water from spouts located around the perimeter of the apparatus. 3. Unilateral lesions restricted to the inferior colliculus had relatively little effect on sound localization. All three animals with restricted lesions were still capable of localizing a single noise burst and minimum audible angles were similar before and after surgery for midline, left, and right field tests. There was no difference in postoperative performance ipsilateral and contralateral to the lesion site. 4. More extensive unilateral lesions that included the dorsal nucleus of the lateral lemniscus and the lateral tegmentum as well as the inferior colliculus resulted in severe deficits in sound localization. Each of the four animals with large unilateral lesions had a substantial impairment in sound localization in the field contralateral to the lesion site. Postoperative performance in the contralateral field was too low to obtain a minimum audible angle for these animals. Midline sound localization was also affected in some cases but ipsilateral sound localization was only slightly affected.

Animals↗

Sound localization after transection of the commissure of Probst in the albino rat.

1. The ability of rats to localize sounds in space was determined before and after cutting of the commissure of Probst. The commissure of Probst was transected at its midline decussation with a microknife inserted into the brain according to stereotaxic coordinates. Six animals were tested after extensive lesions that destroyed all of the commissure of Probst fibers. An additional animal was tested after a smaller lesion that destroyed most of the commissure of Probst but left some fibers intact. Three control animals were tested before and after surgical intervention that did not involve the commissure of Probst. 2. The animals were tested in a semicircular apparatus with loudspeakers located on the right or left of midline. They were trained to make a response toward the left or right in the direction of the active loudspeaker. Correct reponses were rewarded by delivery of a small quantity of water from spouts located at +30 and -30 degrees azimuth. Tests of sound localization were conducted with a single broadband noise burst, 45 ms in duration, presented at the beginning of each trial. The position of the active loudspeakers was varied from trial to trial and performance at different speaker angles was calculated to determine psychometric curves. Minimum audible angles were estimated by interpolation from a performance level of 75% correct. 3. After postoperative testing was completed, the effectiveness of the lesions was confirmed by cell counts to determine the extent of retrograde degeneration in the dorsal nucleus of the lateral lemniscus (DNLL). These data showed that most of the contralaterally projecting neurons in DNLL underwent retrograde degeneration and the number of neurons was reduced by 60-65%. Ninety to 95% of the contralaterally projecting neurons in the DNLL disappeared within 7 wk after transection of the commissure of Probst. 4. The condition of damaged commissural fibers was further confirmed by tract tracing methods. A unilateral Fluoro-Gold injection was made into the inferior colliculus and the auditory brain stem was examined for retrograde label. No labeled neurons were seen in the contralateral DNLL in cases with complete transection of the commissure of Probst. In addition, the distribution of Fluoro-Gold labeling in other brain stem auditory structures was similar to that seen in normal animals. This result confirmed that the transection of the commissure of Probst was successful and that projections to other auditory structures remained intact. 5. Transection of the commissure of Probst produced marked deficits in midline sound localization. Although sound localization was still possible, there was a degradation in the ability of rats to localize sounds in the horizontal plane. The mean minimum audible angle was elevated 22.1 degrees after destruction of the commissure of Probst compared with a shift of only 2.0 degrees in control animals.

Animals↗

A functional neuroimaging study of sound localization: visual cortex activity predicts performance in early-blind individuals.

Blind individuals often demonstrate enhanced nonvisual perceptual abilities. However, the neural substrate that underlies this improved performance remains to be fully understood. An earlier behavioral study demonstrated that some early-blind people localize sounds more accurately than sighted controls using monaural cues. In order to investigate the neural basis of these behavioral differences in humans, we carried out functional imaging studies using positron emission tomography and a speaker array that permitted pseudo-free-field presentations within the scanner. During binaural sound localization, a sighted control group showed decreased cerebral blood flow in the occipital lobe, which was not seen in early-blind individuals. During monaural sound localization (one ear plugged), the subgroup of early-blind subjects who were behaviorally superior at sound localization displayed two activation foci in the occipital cortex. This effect was not seen in blind persons who did not have superior monaural sound localization abilities, nor in sighted individuals. The degree of activation of one of these foci was strongly correlated with sound localization accuracy across the entire group of blind subjects. The results show that those blind persons who perform better than sighted persons recruit occipital areas to carry out auditory localization under monaural conditions. We therefore conclude that computations carried out in the occipital cortex specifically underlie the enhanced capacity to use monaural cues. Our findings shed light not only on intermodal compensatory mechanisms, but also on individual differences in these mechanisms and on inhibitory patterns that differ between sighted individuals and those deprived of vision early in life.

Auditory Perception↗

Sound localization in a predatory rodent, the northern grasshopper mouse (Onychomys leucogaster).

A comparison of the ability of mammals to localize sound revealed that among the animals examined to date, none of the rodents have been able to localize as accurately as the carnivores. Because all of these rodents are prey animals, the question arises as to whether their poor localization acuity is a phyletic trait of Rodentia or whether it is a trait common to prey species that may be under less selective pressure than predators to localize sound accurately. To answer this question, sound localization acuity was determined in a species that is both predatory and a rodent, the northern grasshopper mouse. Localization thresholds for a single 100-ms noise burst were determined for three grasshopper mice using a conditioned avoidance procedure. Their 50% discrimination threshold of 19 degrees is larger than that of any of the previously tested carnivores and well within the range of other rodents. However, calculations of the binaural sound localization cues available to rodents (based on their head size) suggest that the grasshopper mouse may make more efficient use of the available locus cues than other rodents. Thus, although the grasshopper mouse cannot localize as accurately as carnivores, it appears to be more accurate than predicted for a nonpredatory rodent of its size.

Animals↗

EEG coherence in early-blind humans during sound localization.

Human blind individuals have demonstrated cross-modal plasticity in research over the past decade. In one such study, we showed that early-blind subjects were able to localize sound sources accurately despite the lack of visual input for the calibration of their auditory space. A further ERP study with these subjects also revealed N1 and P3 components during a sound localization task to be more posteriorly distributed than for sighted controls, indicating an involvement of posterior regions in sound localization for blind subjects not present for sighted subjects. In the current study, we analyzed these data for EEG power and coherence in theta, alpha, beta, and gamma frequency bands to see whether blind individuals would show increased coherence reflecting increased connectivity between the central and posterior cortical regions. Blind and sighted subjects did not differ with respect to overall EEG power in any frequency range. However, EEG coherence was significantly increased in blind subjects compared to sighted in the theta, alpha, and beta frequency bands. These results have implications for cortical plasticity affected by sensory deprivation in humans.

Acoustic Stimulation↗

Sound localization in chinchillas. II. Front/back and vertical localization.

The ability of chinchillas to make front/back and vertical locus discriminations was examined behaviorally using a conditioned avoidance procedure. Their minimum audible angle for localizing single broadband noise bursts was 36 degrees for front/back localization and 23 degrees for vertical localization. Sound localization tests using filtered noise demonstrated that the signal must contain high frequencies in order for chinchillas to make front/back and vertical locus judgements and that frequencies in their highest audible octave (i.e., above 16 kHz) contribute to localization. These results support the view that a major selective advantage of high-frequency hearing in mammalian evolution was its utility for monaural as well as binaural sound localization.

Acoustic Stimulation↗

Sound localization. Part II: a clinical procedure.

Difficulty in localizing sound has been shown to be associated with severe unilateral hearing impairment, middle-ear conditions and lesions of the cerebello-pontine angle. This was in contrast to the performance of subjects with cochlear lesions and certain brain conditions who had mainly normal ability to localize sound. The investigation appeared to indicate that a sound localization test in the horizontal plane using low frequency stimuli may be a useful addition to the test battery employed in the differential diagnosis of cochlear and retrocochlear lesions involving the cerebello-pontine angle.

Adolescent↗

Effects of auditory cortical lesions on sound localization by the rat.

1. Studies with cats, dogs, and monkeys have shown that bilateral ablation of auditory cortex can result in severe deficits in the ability to localize sounds in space. In the present series of studies we sought to extend this observation to include the laboratory rat. 2. Rats were tested in a two-choice sound-localization task, which required a spatial response to a distant goal box. Although the test conditions were very similar to those employed with cat, dog, and monkey, deficits following cortical ablation were minimal. Indeed, following bilateral ablation of both primary and secondary auditory projection areas, rats were still capable of localization at small angles. 3. Several possibilities were considered to explain the apparent species difference in the effect of cortical ablations. It seemed unlikely that the difference was due simply to testing procedures or details of stimulus presentation. Central nervous system factors were discussed, including the possibility that the degree of impairment was related to the extent of cortical development in different species. It was concluded that the effects of auditory cortical ablation are not the same for all mammals and that for some species auditory cortex is not essential for sound localization.

Animals↗

Sound localization in wild Norway rats (Rattus norvegicus).

The ability of three wild Norway rats to localize sound was determined for single clicks and 100-ms white noise bursts. Chance level localization thresholds were 12 degrees for clicks and 9.7 degrees for white noise. A comparison of these results with published localization thresholds for the domestic albino rat yielded no significant differences. It appears that the combined effects of domestication and albinism have not affected the ability of the laboratory rat to localize sound. Instead, the relatively poor localization acuity of these rats appears to be part of the normal variation in sound localization acuity found among different species of mammals.

Animals↗

Visual factors in sound localization in mammals.

The ability of mammals to localize sound varies widely among species. During the past decade, evidence has accumulated that this variation cannot be accounted for simply on the basis of the availability of the physical cues for locus. Evidence is presented that a major factor in sound localization is the need to direct the field of best vision to a sound source for further scrutiny. Thus, species with broad fields of best vision (such as visual streaks) require less accurate information regarding the location of a sound source than do species with very narrow fields of best vision (such as foveae). To support this suggestion, data are reported for the width of the field of best vision in the form of retinal ganglion cell isodensity contours for thirteen species of mammals. The possible contribution of other factors including binocular fields, visual acuity, and the degree to which a species is predatory in lifestyle, is also examined.

Animals↗

[Neural mechanisms of sound localization in owls].

Barn owls localize sound by using the interaural time difference of the horizontal plane and the interaural intensity difference for the vertical plane. The owl's auditory system possesses the two binaural cues in separate pathways in the brainstem. Owls use a process similar to cross-correlation to derive interaural time differences. Convergence of different frequency bands in the inferior colliculus solves the problems of phase-ambiguity which is inherent in cross-correlating periodic signals. The two pathways converge in the external nucleus of the inferior colliculus to give rise to neurons that are selective for combinations of the two cues. These neurons form a map of auditory space. The map projects to the optic tectum to form a bimodal map which, in turn, projects to a motor map for head turning. The visual system calibrates the auditory space map during ontogeny in which acoustic variables change. In addition to this tectal pathway, the forebrain can also control the sound-localizing behaviour.

Animals↗

Sound localization in large mammals: localization of complex sounds by horses.

The idea that large mammals localize sounds more accurately than small mammals has been noted frequently and is usually explained by reference to their large interaural distance and the correspondingly broad binaural time (delta t) and spectral (delta fi) differences between their two ears. Sound-localization thresholds for single clicks and 100-ms noise bursts were determined for horses, and the magnitude of the binaural time (delta t) and spectral (delta fi) cues for sound direction were measured on a horse. Although horses have relatively large interaural distances and physically broad binaural-localization cues available to them, their sound direction thresholds were markedly poorer than those of other large mammals--averaging 22 degrees for noise and 30 degrees for clicks. It appears that sound-localization acuity is not determined simply by the physical availability of binaural cues.

Animals↗

Developmental refinement of inhibitory sound-localization circuits.

The ability to localize sound rapidly and accurately depends on the precise organization of inhibitory neuronal circuits in the auditory brainstem. However, the rules and mechanisms by which this precision is established during development are still poorly understood. Although activity-dependent reorganization has been known for over a decade to have a central role in this process, more recent studies have revealed an unanticipated degree of reorganization that occurs on levels ranging from cellular phenotype to network connectivity. These results suggest novel mechanisms by which immature inhibitory sound-localization circuits become optimized. Lessons from auditory brainstem circuits thus could provide insight into inhibitory development in other brain areas, where inhibitory networks are less experimentally accessible.

Age Factors↗

Sound localization with hearing protectors.

A preliminary investigation was conducted of the ability of normal-hearing subjects to localize sound while wearing hearing protective devices. Performance with Bilsom 2392 level-dependent stereophonic muffs with limited amplification was compared with that of conventional level-independent E-A-R 3000 muffs and E-A-R plugs. Sound localization was assessed using an array of six speakers, positioned in the horizontal plane, 1 m from the subject at azimuth angles of 30 degrees, 90 degrees, 150 degrees, 210 degrees, 270 degrees and 330 degrees. The stimulus was an 80 dB SPL 300-ms one-third octave noise band centered at 500 Hz or 4000 Hz. The experiment was conducted in quiet and in a 65 dB SPL white noise background. The results indicated that at 500 Hz, the wearing of protectors resulted in a 20% decrease in the accuracy of speaker identification but the three types of device were indistinguishable. At 4000 Hz, performance was best in the unoccluded condition and relatively worst with the level-dependent muff. Error patterns depended on the interaction of stimulus frequency, speaker azimuth and protector. Front/back errors in localizing the high-frequency stimulus were greatest with the conventional muff, whereas right/left errors were greatest with the level-dependent muff. Protected listening was generally unaffected by the presence of a noise background for both of the frequencies tested.

Adult↗

Hearing and sound localization in blind mole rats (Spalax ehrenbergi).

Two blind mole rats were tested for their ability to detect and localize sound. The results indicate that blind mole rats have severely limited, and probably degenerate, auditory abilities. Although their 60-dB low-frequency hearing limit of 54 Hz is within the range for other rodents, the highest frequency they can hear at a level of 60 dB SPL is only 5.9 kHz, giving them the poorest high-frequency sensitivity yet observed in any mammal. In addition they have poor sensitivity as indicated by the fact that their lowest threshold is only 32 dB SPL (at 1 kHz). Finally, they are unable to localize brief sounds but retain a rudimentary ability to localize sounds of 0.5 s or longer. These results, combined with those of previous studies of subterranean species (i.e., blind mole rats, naked mole rats, and pocket gophers), suggest that poor auditory sensitivity, the loss of high-frequency hearing, and an inability to localize brief sounds is a degenerate state which may be characteristic of subterranean mammals. Thus it appears that an exclusive adaptation to a subterranean lifestyle (where airborne sound propagates poorly and where directional responses are limited by the tunnels) can result in vestigial auditory abilities just as the absence of light results in vestigial vision.

Acoustic Stimulation↗