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PubMed1991

Sound localization by human listeners.

In keeping with our promise earlier in this review, we summarize here the process by which we believe spatial cues are used for localizing a sound source in a free-field listening situation. We believe it entails two parallel processes: 1. The azimuth of the source is determined using differences in interaural time or interaural intensity, whichever is present. Wightman and colleagues (1989) believe the low-frequency temporal information is dominant if both are present. 2. The elevation of the source is determined from spectral shape cues. The received sound spectrum, as modified by the pinna, is in effect compared with a stored set of directional transfer functions. These are actually the spectra of a nearly flat source heard at various elevations. The elevation that corresponds to the best-matching transfer function is selected as the locus of the sound. Pinnae are similar enough between people that certain general rules (e.g. Blauert's boosted bands or Butler's covert peaks) can describe this process. Head motion is probably not a critical part of the localization process, except in cases where time permits a very detailed assessment of location, in which case one tries to localize the source by turning the head toward the putative location. Sound localization is only moderately more precise when the listener points directly toward the source. The process is not analogous to localizing a visual source on the fovea of the retina. Thus, head motion provides only a moderate increase in localization accuracy. Finally, current evidence does not support the view that auditory motion perception is anything more than detection of changes in static location over time.

Attention↗
PubMed1988

Sound localization acuity in the cat: effect of azimuth, signal duration, and test procedure.

The sound localization acuity of cats was determined at 0 degrees, 30 degrees, 60 degrees, and 90 degrees from the median sagittal plane for four durations of noise bursts using two behavioral procedures. Similar thresholds were also obtained for humans. The cats' average thresholds for a 40 ms noise burst ranged from 4.8 degrees at 0 degrees azimuth to 9.0 degrees at 90 degrees azimuth. Comparable thresholds for humans ranged from 1.3 degrees at 0 degrees to 9.7 degrees at 90 degrees. The fact that humans and cats had similar thresholds at 90 degrees azimuth suggests that the mobile pinnae of the cat are not an advantage in localizing sound in the lateral fields. Varying the duration of the stimulus from 10 ms to 'continuous' had little effect on threshold. No difference was found between the results of the two behavioral procedures.

Adolescent↗
PubMed2001

Spatial attention modulates sound localization in barn owls.

Attentional influence on sound-localization behavior of barn owls was investigated in a cross-modal spatial cuing paradigm. After being cued to the most probable target side with a visual cuing stimulus, owls localized upcoming auditory target stimuli with a head turn toward the position of the sound source. In 80% of the trials, cuing stimuli pointed toward the side of the upcoming target stimulus (valid configuration), and in 20% they pointed toward the opposite side (invalid configuration). We found that owls initiated the head turns by a mean of 37.4 ms earlier in valid trials, i.e., mean response latencies of head turns were reduced by 16% after a valid cuing stimulus. Thus auditory stimuli appearing at the cued side were processed faster than stimuli appearing at the uncued side, indicating the influence of a spatial-selective attention mechanism. Turning angles were not different when owls turned their head toward a cued or an uncued location. Other types of attention influencing sound localization, e.g., a reduction of response latency as a function of the duration of cue-target delay, could not be observed. This study is the first attempt to investigate attentional influences on sound localization in an animal model.

Acoustic Stimulation↗
PubMed2006

Sound localization during homotopic and heterotopic bilateral cooling deactivation of primary and nonprimary auditory cortical areas in the cat.

Although the contributions of primary auditory cortex (AI) to sound localization have been extensively studied in a large number of mammals, little is known of the contributions of nonprimary auditory cortex to sound localization. Therefore the purpose of this study was to examine the contributions of both primary and all the recognized regions of acoustically responsive nonprimary auditory cortex to sound localization during both bilateral and unilateral reversible deactivation. The cats learned to make an orienting response (head movement and approach) to a 100-ms broad-band noise stimulus emitted from a central speaker or one of 12 peripheral sites (located in front of the animal, from left 90 degrees to right 90 degrees , at 15 degrees intervals) along the horizontal plane after attending to a central visual stimulus. Twenty-one cats had one or two bilateral pairs of cryoloops chronically implanted over one of ten regions of auditory cortex. We examined AI [which included the dorsal zone (DZ)], the three other tonotopic fields [anterior auditory field (AAF), posterior auditory field (PAF), ventral posterior auditory field (VPAF)], as well as six nontonotopic regions that included second auditory cortex (AII), the anterior ectosylvian sulcus (AES), the insular (IN) region, the temporal (T) region [which included the ventral auditory field (VAF)], the dorsal posterior ectosylvian (dPE) gyrus [which included the intermediate posterior ectosylvian (iPE) gyrus], and the ventral posterior ectosylvian (vPE) gyrus. In accord with earlier studies, unilateral deactivation of AI/DZ caused sound localization deficits in the contralateral field. Bilateral deactivation of AI/DZ resulted in bilateral sound localization deficits throughout the 180 degrees field examined. Of the three other tonotopically organized fields, only deactivation of PAF resulted in sound localization deficits. These deficits were virtually identical to the unilateral and bilateral deactivation results obtained during AI/DZ deactivation. Of the six nontonotopic regions examined, only deactivation of AES resulted in sound localization deficits in the contralateral hemifield during unilateral deactivation. Although bilateral deactivation of AI/DZ, PAF, or AES resulted in profound sound localization deficits throughout the entire field, the cats were generally able to orient toward the hemifield that contained the acoustic stimulus, but not accurately identify the location of the stimulus. Neither unilateral nor bilateral deactivation of areas AAF, VPAF, AII, IN, T, dPE, nor vPE had any effect on the sound localization task. Finally, bilateral heterotopic deactivations of AI/DZ, PAF, or AES yielded deficits that were as profound as bilateral homotopic cooling of any of these sites. The fact that deactivation of any one region (AI/DZ, PAF, or AES) was sufficient to produce a deficit indicated that normal function of all three regions was necessary for normal sound localization. Neither unilateral nor bilateral deactivation of AI/DZ, PAF, or AES affected the accurate localization of a visual target. The results suggest that hemispheric deactivations contribute independently to sound localization deficits.

Acoustic Stimulation↗
PubMed2004

Sound localization in bilateral users of MED-EL COMBI 40/40+ cochlear implants.

OBJECTIVE: The purpose of the study was to investigate sound localization with bilateral and unilateral cochlear implants. DESIGN: Sound localization tests were performed on 20 bilaterally implanted MED-EL COMBI 40/40+ users. All subjects were bilaterally implanted during adolescence or later. Sound localization was tested in the frontal horizontal plane by using 9 equally spaced loudspeakers and speech-shaped noise bursts at randomized levels. RESULTS: The group of subjects who were bilaterally deafened after 5 to 6 yr of age (18 subjects) showed a statistically significant improvement in sound localization when using both implants, compared with when using only one. The mean deviation between the presentation azimuth and the response azimuth was 16.6 degrees when using both implants, which was on average 37.1 degrees smaller than when using one implant only. When adjusted for the localization error that was constant across loudspeakers, the mean deviation was 15.9 degrees for bilateral implant use, representing an improvement of 30.1 degrees over unilateral implant use. Statistical analysis showed that in this group, performance measures were not correlated with subject details such as age at onset of deafness or duration of unilateral implant use. In contrast, subjects who were bilaterally deafened before 6 yr of age (2 subjects) did not show a benefit in sound localization from bilateral implants. CONCLUSIONS: Bilateral cochlear implants offer a substantial benefit in sound localization to late-deafened, late-implanted subjects. The very limited data from early-deafened subjects implanted at a later age could suggest that these subjects may not benefit in sound localization from bilateral cochlear implants. It is possible that early implantation for early deafened subjects might allow better acquisition of spatial hearing, thus leading to improved localization performance.

Adolescent↗
PubMed1990

Effects of aging on the precedence effect in sound localization.

The precedence effect in sound localization can be evoked by presenting identical sounds (e.g., clicks) from pairs of loudspeakers placed on opposite sides of a subject's head. With appropriate inter-loudspeaker delays, normal subjects perceive a fused image originating from the side of the leading loudspeaker. Separate tests at loudspeaker delays ranging from 0 to 8 ms were presented to groups of young and elderly subjects. At 0 ms delay, young subjects perceived the fused image to be located halfway between the loudspeakers; at progressively longer delays, the image was perceived closer to the leading loudspeaker. Significant numbers of elderly subjects exhibited discrimination difficulties with delays below 0.7 ms.

Adult↗
PubMed2006

Monkey sound localization: head-restrained versus head-unrestrained orienting.

The sound localization abilities of three rhesus monkeys were tested under head-restrained and head-unrestrained conditions. Operant conditioning and the magnetic search coil technique were used to measure eye and head movements to sound sources. Whereas the results support previous findings that monkeys localize sounds very poorly with their heads restrained, the data also reveal for the first time that monkeys localize sounds much more accurately and with less variability when their heads are allowed to move. Control experiments using acoustic stimuli known to produce spatial auditory illusions such as summing localization confirmed that the monkeys based their orienting on localizing the sound sources and not on remembering spatial locations that resulted in rewards. Overall, the importance of using ecologically valid behaviors for studies of sensory processes is confirmed, and the potential of the rhesus monkey, the model closest to human, for studies of spatial auditory function, is established.

Acoustic Stimulation↗
PubMed1993

Degenerate hearing and sound localization in naked mole rats (Heterocephalus glaber), with an overview of central auditory structures.

Behavioral tests of absolute sensitivity and sound localization in African naked mole rats show that, despite their communal social structure and large vocal repertoire, their hearing has degenerated much like that of other subterranean species. First, their ability to detect sound is limited, with their maximum sensitivity being only 35 dB (occurring at 4 kHz). Second, their high-frequency hearing is severely limited, with their hearing range (at 60 dB sound pressure level [SPL]) extending from 65 Hz to only 12.8 kHz. Third, determination of the effect of duration on noise thresholds indicates that, compared with other animals, mole rats require a sound to be present for a much longer duration before reaching asymptotic threshold. Finally, they are unable consistently to localize sounds shorter than 400 ms and cannot accurately localize sounds of longer duration, raising the possibility that they are unable to use binaural locus cues. Thus, it seems that the essentially one-dimensional burrow system of a subterranean habitat produces severe changes in hearing comparable to the changes in vision that result from the absence of light. To explore the relation between vision and sound-localization acuity, retinal ganglion cell densities were determined. The results indicate that naked mole rats have a broad area of best (albeit poor) vision, with maximum acuity estimated at 44 cycles/degree. That mammals with wide fields of best vision have poorer sound-localization acuity than those with narrower fields is consistent with the thesis that a major function of sound localization is to direct the gaze to the source of a sound. However, the fact that subterranean mammals have little use for vision in a lightless environment suggests that they represent an extreme case in this relationship and may explain the fact that, unlike surface-dwelling mammals, they have virtually lost the ability to localize brief sounds. Finally, despite their very limited auditory abilities, the major brainstem auditory nuclei, although relatively small, appear to be present.

Animals↗
PubMed1991

Sound localization and sensitivity to interaural time differences in human infants.

The role of interaural time differences in infants' sound localization was investigated. One experiment on free-field sound localization corroborated previous findings that the minimum audible angle changes substantially toward the end of the first half year after birth, and 3 experiments explored interaural time discrimination in that age range. The first of these 3 experiments used an adaptive psychophysical procedure, showing that infants responded appropriately in a sound lateralization task that provides a direct measure of interaural time discrimination. The other 2 experiments improved on the psychophysical procedure by taking into account the ceiling level on performance in the task. Infants aged 16, 20, and 28 weeks had thresholds in the range of 50 to 75 microsec, with no apparent age difference. These thresholds were much lower than would be predicted from studies of free-field sound localization, indicating that sensitivity to interaural time differences is not a limiting factor for the precision of sound localization in this age range. Instead, age-related changes in free-field sound localization may reflect the need to integrate across different localization cues and to calibrate the changing values of cues due to head growth.

Acoustic Stimulation↗
PubMed1993

Passive sound localization of prey by the pallid bat (Antrozous p. pallidus).

The pallid bat (Antrozous p. pallidus) uses passive sound localization to capture terrestrial prey. This study of captive pallid bats examined the roles of echolocation and passive sound localization in prey capture, and focused on their spectral requirements for accurate passive sound localization. Crickets were used as prey throughout these studies. All tests were conducted in dim, red light in an effort to preclude the use of vision. Hunting performance did not differ significantly in red light and total darkness, nor did it differ when visual contrast between the terrestrial prey and the substrate was varied, demonstrating that the bats did not use vision to locate prey. Our bats apparently used echolocation for general orientation, but not to locate prey. They did not increase their pulse emission rate prior to prey capture, suggesting that they were not actively scanning prey. Instead, they required prey-generated sounds for localization. The bats attended to the sound of walking crickets for localization, and also attacked small, inanimate objects dragged across the floor. Stationary and/or anesthetized crickets were ignored, as were crickets walking on substrates that greatly attenuated walking sounds. Cricket communication sounds were not used in prey localization; the bats never captured stationary, calling crickets. The accuracy of their passive sound localization was tested with an open-loop passive sound localization task that required them to land upon an anesthetized cricket tossed on the floor. The impact of a cricket produced a single 10-20 ms duration sound, yet with this information, the bats were able to land within 7.6 cm of the cricket from a maximum distance of 4.9 m. This performance suggests a sound localization accuracy of approximately +/- 1 degree in the horizontal and vertical dimensions of auditory space. The lower frequency limit for accurate sound localization was between 3-8 kHz. A physiological survey of frequency representation in the pallid bat inferior colliculus suggests that this lower frequency limit is around 5 kHz.

Animals↗
PubMed1997

Comparative study of sound localization and its anatomical correlates in mammals.

One of the fundamental features of hearing is the ability to localize the sources of sounds, particularly brief sounds, which may warn of nearby animals. Yet not all mammals localize sound equally well with threshold acuity ranging from about 1 degree for elephants and humans to more than 25 degrees for gerbils and horses and a near absence of localization in some subterranean species. During the past decade evidence has accumulated that this variation cannot be accounted for simply by the availability of the physical cues for locus. Nor does it appear to be a function of an animal's lifestyle. Rather sound-localization acuity in mammals appears to be a function of the precision required of the visual orienting response to sound. Thus the neural integration of hearing and vision in cortex, as well as in multimodal subcortical structures, is a reflection of their behavioral integration and evolutionary coupling.

Animals↗
PubMed2004

Contribution of head shadow and pinna cues to chronic monaural sound localization.

Monaurally deaf people lack the binaural acoustic difference cues in sound level and timing that are needed to encode sound location in the horizontal plane (azimuth). It has been proposed that these people therefore rely on spectral pinna cues of their normal ear to localize sounds. However, the acoustic head-shadow effect (HSE) might also serve as an azimuth cue, despite its ambiguity when absolute sound levels are unknown. Here, we assess the contribution of either cue in the monaural deaf to two-dimensional (2D) sound localization. In a localization test with randomly interleaved sound levels, we show that all monaurally deaf listeners relied heavily on the HSE, whereas binaural control listeners ignore this cue. However, some monaural listeners responded partly to actual sound-source azimuth, regardless of sound level. We show that these listeners extracted azimuth information from their pinna cues. The better monaural listeners were able to localize azimuth on the basis of spectral cues, the better their ability to also localize sound-source elevation. In a subsequent localization experiment with one fixed sound level, monaural listeners rapidly adopted a strategy on the basis of the HSE. We conclude that monaural spectral cues are not sufficient for adequate 2D sound localization under unfamiliar acoustic conditions. Thus, monaural listeners strongly rely on the ambiguous HSE, which may help them to cope with familiar acoustic environments.

Acoustic Stimulation↗
PubMed2000

Study of sound localization by owls and its relevance to humans.

Human psychoacoustical studies have been the main sources of information from which the brain mechanisms of sound localization are inferred. The value of animal models would be limited, if humans and the animals did not share the same perceptual experience and the neural mechanisms for it. Barn owls and humans use the same method of computing interaural time differences for localization in the horizontal plane. The behavioral performance of owls and its neural bases are consistent with some of the theories developed for human sound localization. Neural theories of sound localization largely owe their origin to the study of sound localization by humans, even though little is known about the physiological properties of the human auditory system. One of these ideas is binaural cross-correlation which assumes that the human brain performs a process similar to mathematical cross-correlation to measure the interaural time difference for localization in the horizontal plane. The most complete set of neural evidence for this theory comes from the study of sound localization and its brain mechanisms in barn owls, although partial support is also available from studies on laboratory mammals. Animal models of human sensory perception make two implicit assumptions; animals and humans experience the same percept and the same neural mechanism underlies the creation of the percept. These assumptions are hard to prove for obvious reason. This article reviews several lines of evidence that similar neural mechanisms must underlie the perception of sound locations in humans and owls.

Animals↗
PubMed2005

Relearning sound localization with a new ear.

Human sound localization results primarily from the processing of binaural differences in sound level and arrival time for locations in the horizontal plane (azimuth) and of spectral shape cues generated by the head and pinnae for positions in the vertical plane (elevation). The latter mechanism incorporates two processing stages: a spectral-to-spatial mapping stage and a binaural weighting stage that determines the contribution of each ear to perceived elevation as function of sound azimuth. We demonstrated recently that binaural pinna molds virtually abolish the ability to localize sound-source elevation, but, after several weeks, subjects regained normal localization performance. It is not clear which processing stage underlies this remarkable plasticity, because the auditory system could have learned the new spectral cues separately for each ear (spatial-mapping adaptation) or for one ear only, while extending its contribution into the contralateral hemifield (binaural-weighting adaptation). To dissociate these possibilities, we applied a long-term monaural spectral perturbation in 13 subjects. Our results show that, in eight experiments, listeners learned to localize accurately with new spectral cues that differed substantially from those provided by their own ears. Interestingly, five subjects, whose spectral cues were not sufficiently perturbed, never yielded stable localization performance. Our findings indicate that the analysis of spectral cues may involve a correlation process between the sensory input and a stored spectral representation of the subject's ears and that learning acts predominantly at a spectral-to-spatial mapping level rather than at the level of binaural weighting.

Acoustic Stimulation↗
PubMed2006

A review of learning with normal and altered sound-localization cues in human adults.

Humans use several cues to determine the position of sound sources in space. Here we provide a review of the literature on how practice influences sound-localization performance in human adults. We divided this literature into two categories: adaptation to alterations in sound-localization cues and learning with normal, unaltered cues. For altered cues, partial adaptation has been reported with a variety of cue transformations, except in the extreme case in which the cues were reversed between the two ears. For normal cues, improvement has been observed in some but not all experiments, though the most recent data generally show learning. Thus, the reviewed data indicated that human adults can recalibrate, as well as refine the use of, sound-localization cues, and that training regimens can be developed to enhance sound-localization performance in individuals with normal and impaired localization abilities.

Adult↗
PubMed1975

Contribution of auditory cortex to sound localization in the monkey (Macaca mulatta).

Monkeys with lesions of auditory cortex were tested for their ability to localize the source of brief sounds. Although those deprived of primary auditory cortex bilaterally were able to indicate the direction of a sound with near-normal acuity, they were unable to locate its source. This dissociation of abilities suggest that the role of auditory cortex in sound localization is not so much sensory or perceptual as it is auditomotor or associative. Thus, sound localization joins loudness, pitch, and most other traditional attributes of sound as dimensions whose discrimination does not depend on auditory cortex. The question would now seem to turn to whether or not auditory cortex is necessary for any sensory discrimination whatever.

Acoustic Stimulation↗
PubMed1983

Sound localization in subjects with impaired hearing. Spatial-discrimination and interaural-discrimination tests.

In order to get a systematic picture of how various hearing impairments and neurologic disorders may affect sound localization, psychophysical spatial- and lateralization-discrimination measurements were performed on 140 subjects, including 69 with different types of hearing impairments, 32 with neurological diseases, and 39 with normal hearing. The quantities measured were: in the freefield, the horizontal minimum audible angle (MAA) at eight reference azimuths around the head and the vertical MAA straight ahead; with headphones, the just-noticeable difference (JND) in interaural time delay and the JND in interaural intensity difference. The standard stimulus was broadband (0.25-10 kHz), pulsed (1-sec), noise presented at a suprathreshold level for both ears (65-100 dB SPL). The results show that there exist characteristic impairments of sound localization in the different types of hearing impairments tested. On a general level, the results are consistent with the concept that the localization of sound relies on a decision made by the central auditory system based on a number of cues present in the acoustic signal at the two ears. The cues tested in our study are: 1) the interaural time difference, 2) the interaural intensity difference, and 3) the spectrum of the received signal at each ear. At a more specific level, the sound localization impairments found in conductive hearing losses are interpreted as bone-conduction effects, the results found in sensorineural hearing losses are interpreted as consequences of impaired or preserved spectral processing, the results in neurinomas are interpreted as impaired signal transmission in the auditory nerves, and the results of subjects with central involvements suggest that separate processors exist at some level in the central auditory system for the different localization cues. Finally, comments are made about the practical clinical significance of sound localization tests in the audiological and neurological evaluation of patients.

Adolescent↗
PubMed1989

Vision calibrates sound localization in developing barn owls.

This study demonstrates that continuous exposure of baby barn owls to a displaced visual field causes a shift in sound localization in the direction of the visual displacement. This implies an innate dominance of vision over audition in the development and maintenance of sound localization. Twelve owls were raised from the first day of eye opening wearing binocular prisms that displaced the visual field to the right by 11 degrees, 23 degrees, or 34 degrees. The prisms were worn for periods of up to 7 months. Consistent with previous results (Knudsen and Knudsen, 1989a), owls reared with displacing prisms did not adjust head orientation to visual stimuli. While wearing prisms, owls consistently oriented the head to the right of visual targets, and, as soon as the prisms were removed, they oriented the head directly at visual targets, as do normal owls. In contrast, prism-reared owls did change head orientation to sound sources even though auditory cues were not altered significantly. Birds reared wearing 11 degrees or 23 degrees prisms oriented the head to the right of acoustic targets by an amount approximately equal to the optical displacement induced by the prisms. Birds raised wearing 34 degrees prisms adjusted sound localization by only about 50% of the optical displacement. Thus, visually guided adjustment of sound localization appears to be limited to about 20 degrees in azimuth. The data indicate that when confronted with consistently discordant localization information from the auditory and visual systems, developing owls use vision to calibrate associations of auditory localization cues with locations in space in an attempt to bring into alignment the perceived locations of auditory and visual stimuli emanating from a common source. Vision exerts this instructive influence on sound localization whether or not visual information is accurate.

Animals↗