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Location coding by opponent neural populations in the auditory cortex.

Although the auditory cortex plays a necessary role in sound localization, physiological investigations in the cortex reveal inhomogeneous sampling of auditory space that is difficult to reconcile with localization behavior under the assumption of local spatial coding. Most neurons respond maximally to sounds located far to the left or right side, with few neurons tuned to the frontal midline. Paradoxically, psychophysical studies show optimal spatial acuity across the frontal midline. In this paper, we revisit the problem of inhomogeneous spatial sampling in three fields of cat auditory cortex. In each field, we confirm that neural responses tend to be greatest for lateral positions, but show the greatest modulation for near-midline source locations. Moreover, identification of source locations based on cortical responses shows sharp discrimination of left from right but relatively inaccurate discrimination of locations within each half of space. Motivated by these findings, we explore an opponent-process theory in which sound-source locations are represented by differences in the activity of two broadly tuned channels formed by contra- and ipsilaterally preferring neurons. Finally, we demonstrate a simple model, based on spike-count differences across cortical populations, that provides bias-free, level-invariant localization-and thus also a solution to the "binding problem" of associating spatial information with other nonspatial attributes of sounds.

Acoustic Stimulation↗

Evidence for a sound movement area in the human cerebral cortex.

Human listeners can localize sounds by the difference in both arrival time (phase) and loudness between the two ears. Movement of the sound source modulates these cues, and responses to moving sounds have been detected in animals in primary auditory cortex and in humans in other cortical areas. Here we show that detection of changes in the interaural phase or amplitude difference occurs through a mechanism distinct from that used to detect changes in one ear alone. Moreover, a patient with a right hemisphere stroke is unable to detect sound movement, regardless of whether it is defined by phase or by loudness cues. We propose that this deficit reflects damage to a distinct cortical area, outside the classical auditory areas, that is specialized for the detection of sound motion. The deficit is analagous to cerebral akinotopsia (motion blindness) in the visual system, and so the auditory system may, like the visual system, show localization of specialized functions to different cortical regions.

Aged↗

Mismatch negativity to change in spatial location of an auditory stimulus.

Auditory stimulus blocks were presented to 12 reading subjects. Each block consisted of 2 types, standard (P = 90%) and deviant stimuli (P = 10%), delivered in a random order. The only difference between these stimuli was their spatial location of origin. The subject always heard the standards as coming straight in front and the deviants from an angle of either 10, 45, or 90 degrees to the right of the standards. The spatial locations were produced via earphones by introducing for low-frequency (600 Hz) tones an interaural phase difference and for high-frequency (3000 Hz) tones an interaural intensity difference. Standard and deviant stimuli were also delivered in more natural, free-field, conditions via differently positioned loudspeakers. The deviant tones elicited an event-related brain potential component called the mismatch negativity (MMN), followed by a P3a component. Thus changes in spatial location of an auditory stimulus produced by following either one of the two main principles of human sound localization elicited the MMN. Consequently, it was concluded that the spatial location of a sound source is coded in the hypothesized neuronal stimulus traces reflected by the MMN and, further, that a change in this location is automatically detected by the brain by means of the MMN generator process.

Adult↗

Auditory localization: effects of reflecting surfaces.

Based on anatomical and evolutionary conceptions of the human ear, an experiment was conducted in which forty-eight human subjects were asked to localize sounds (a human voice) emitted by one of twenty-seven stationary loudspeakers in an anechoic chamber. The position of the active loudspeaker varied with respect to azimuth, distance, and elevation in three steps each. The position of a single sound-reflecting surface (about 6 m2) was varied: on the floor, on the ceiling, to the left, and to the right. The accuracy of identifying the active loudspeaker for each position of the sound-reflecting surface was compared intraindividually with the absence of reflection. The results show an overall increase in correct localizations with a sound-reflecting surface on the floor. Especially the elevation of the sound source can be detected with greater precision. Additionally, the percentage of correct localizations decreased systematically with the presence of a sound-reflecting ceiling, while the presence of sound-reflecting walls did not systematically affect the localization performance. Judgments in the horizontal plane and those of distance were not systematically influenced by the presence of a sound-reflecting surface.

Adult↗

Directional hearing by mechanical coupling in the parasitoid fly Ormia ochracea.

Sound localization is a basic processing task of the auditory system. The directional detection of an incident sound impinging on the ears relies on two acoustic cues: interaural amplitude and interaural time differences. In small animals, with short interaural distances both amplitude and time cues can become very small, challenging the directional sensitivity of the auditory system. The ears of a parasitoid fly Ormia ochracea, are unusual in that both acoustic sensors are separated by only 520 microns and are contained within an undivided air-filled chamber. This anatomy results in minuscule differences in interaural time cues (ca. 2 microseconds) and no measurable difference in interaural intensity cues generated from an incident sound wave. The tympana of both ears are anatomically coupled by a cuticular bridge. This bridge also mechanically couples the tympanana, providing a basis for directional sensitivity. Using laser vibrometry, it is shown that the mechanical response of the tympanal membranes has a pronounced directional sensitivity. Interaural time and intensity differences in the mechanical response of the ears are significantly larger than those available in the acoustic field. The tympanal membranes vibrate with amplitude differences of about 12 dB and time differences on the order of 50 microseconds to sounds at 90 degrees off the longitudinal body axis. The analysis of the deflection shapes of the tympanal vibrations shows that the interaural differences in the mechanical response are due to the dynamic properties of the tympanal system and reflect its intrinsic sensitivity to the direction of a sound source. Using probe microphones and extracellular recording techniques, we show that the primary auditory afferents encode sound direction with a time delay of about 300 microseconds. Our data point to a novel mechanism for directional hearing in O. ochracea based on intertympanal mechanical coupling, a process that amplifies small acoustic cues into interaural time and amplitude differences that can be reliably processed at the neural level. An intuitive description of the mechanism is proposed using a simple mechanical model in which the ears are coupled through a flexible lever.

Acoustics↗

Vibratory-coded directional analysis: evaluation of a three-microphone/four-vibrator DSP system.

A sound localization aid based on eyeglasses with three microphones and four vibrators was tested in a sound-treated acoustic test room and in an ordinary office. A digital signal-processing algorithm provided a determination of the source angle, which was transformed into eight vibrator codes each corresponding to a 45 degrees sector. The instrument was tested on nine deaf and three deaf-blind individuals. The results show an average hit rate of about 80% in a sound-treated room with 100% for the front 135 degrees sector. The results in a realistic communication situation in an ordinary office room were 70% correct based on single presentations and 95% correct when more realistic criteria for an adequate reaction were used. Ten of the twelve subjects were interested in participating in field tests using a planned miniaturized version.

Adult↗

Azimuthal processing in the posterior auditory thalamus of cats.

The responses to free-field acoustic stimuli of 157 units in the auditory thalamus of anesthetized cats were studied in relation to the localization of pure tone stimuli in the azimuthal plane. Units were classified as 'directional' if their firing rates at sound levels in excess of 20 dB above threshold varied by more than 50% as a function of azimuth. Sixty-five % of the units in the nucleus of the brachium of the inferior colliculus and 30% in the ventral division of the medial geniculate body were found to be directional, suggesting different processing channels for sound localization between colliculus and cortex.

Acoustic Stimulation↗

Lateralized auditory spatial perception and the contralaterality of cortical processing as studied with functional magnetic resonance imaging and magnetoencephalography.

Functional magnetic resonance imaging (fMRI) and magnetoencephalography (MEG) were used to study the relationships between lateralized auditory perception in humans and the contralaterality of processing in auditory cortex. Subjects listened to rapidly presented streams of short FM-sweep tone bursts to detect infrequent, slightly deviant tone bursts. The stimulus streams consisted of either monaural stimuli to one ear or the other or binaural stimuli with brief interaural onset delays. The onset delay gives the binaural sounds a lateralized auditory perception and is thought to be a key component of how our brains localize sounds in space. For the monaural stimuli, fMRI revealed a clear contralaterality in auditory cortex, with a contralaterality index (contralateral activity divided by the sum of contralateral and ipsilateral activity) of 67%. In contrast, the fMRI activations from the laterally perceived binaural stimuli indicated little or no contralaterality (index of 51%). The MEG recordings from the same subjects performing the same task converged qualitatively with the fMRI data, confirming a clear monaural contralaterality, with no contralaterality for the laterally perceived binaurals. However, the MEG monaural contralaterality (55%) was less than the fMRI and decreased across the several hundred millisecond poststimulus time period, going from 57% in the M50 latency range (20-70 ms) to 53% in the M200 range (170-250 ms). These data sets provide both quantification of the degree of contralaterality in the auditory pathways and insight into the locus and mechanism of the lateralized perception of spatially lateralized sounds.

Acoustic Stimulation↗

Altitude-dependent changes of directional hearing in mountaineers.

This study demonstrates apparent deterioration in the ability to localize sound associated with acute exposure to high altitude in ten subjects on three mountaineering expeditions. Furthermore, the auditory localization errors improved to sea level values after a period of acclimatization. Occurring at altitudes where overt neurological symptoms are not usually seen, impairment of sensory perception may explain the increase in accidental deaths associated with altitude exposure due to disorientation and misjudgment but before hypoxia is evident.

Acclimatization↗

Codes for sound-source location in nontonotopic auditory cortex.

We evaluated two hypothetical codes for sound-source location in the auditory cortex. The topographical code assumed that single neurons are selective for particular locations and that sound-source locations are coded by the cortical location of small populations of maximally activated neurons. The distributed code assumed that the responses of individual neurons can carry information about locations throughout 360 degrees of azimuth and that accurate sound localization derives from information that is distributed across large populations of such panoramic neurons. We recorded from single units in the anterior ectosylvian sulcus area (area AES) and in area A2 of alpha-chloralose-anesthetized cats. Results obtained in the two areas were essentially equivalent. Noise bursts were presented from loudspeakers spaced in 20 degrees intervals of azimuth throughout 360 degrees of the horizontal plane. Spike counts of the majority of units were modulated >50% by changes in sound-source azimuth. Nevertheless, sound-source locations that produced greater than half-maximal spike counts often spanned >180 degrees of azimuth. The spatial selectivity of units tended to broaden and, often, to shift in azimuth as sound pressure levels (SPLs) were increased to a moderate level. We sometimes saw systematic changes in spatial tuning along segments of electrode tracks as long as 1.5 mm but such progressions were not evident at higher sound levels. Moderate-level sounds presented anywhere in the contralateral hemifield produced greater than half-maximal activation of nearly all units. These results are not consistent with the hypothesis of a topographic code. We used an artificial-neural-network algorithm to recognize spike patterns and, thereby, infer the locations of sound sources. Network input consisted of spike density functions formed by averages of responses to eight stimulus repetitions. Information carried in the responses of single units permitted reasonable estimates of sound-source locations throughout 360 degrees of azimuth. The most accurate units exhibited median errors in localization of <25 degrees, meaning that the network output fell within 25 degrees of the correct location on half of the trials. Spike patterns tended to vary with stimulus SPL, but level-invariant features of patterns permitted estimates of locations of sound sources that varied through 20-dB ranges. Sound localization based on spike patterns that preserved details of spike timing consistently was more accurate than localization based on spike counts alone. These results support the hypothesis that sound-source locations are represented by a distributed code and that individual neurons are, in effect, panoramic localizers.

Action Potentials↗

Hearing in the ferret (Mustela putorius): effects of primary auditory cortical lesions on thresholds for pure tone detection.

1. Pure tone thresholds were determined for five adult male ferrets before and after bilateral ablation of primary auditory cortex. Complete audiograms ranging from 0.016 to 48 kHz were obtained for two animals. The remaining three animals were tested at five frequencies selected to assess hearing throughout the audible range (0.125, 0.5, 2.0, 8.0, and 32.0 kHz). 2. Shortly after surgery one animal had elevated thresholds across the entire frequency range with the most pronounced hearing loss above 12.0 kHz. Four other animals had no elevation of thresholds at low and midrange frequencies but suffered a hearing loss at very high frequencies (32 kHz). 3. Repeated testing over a period of several months revealed substantial recovery of sensitivity. There was complete recovery of sensitivity in the low- and middle-frequency range of the audiogram. Some hearing loss persisted at the extreme upper end of the audiogram (32 kHz), but in two cases there was evidence of recovery at this frequency as well. 4. Following determination of absolute thresholds all animals were assessed for their ability to localize sound in space. Minimum audible angles were obtained on midline as well as within both left and right hemifields, i.e., around 0, -60 and +60 degrees azimuth. All animals had severe and persistent deficits in their ability to localize brief sounds within the lateral fields, but were still capable of midline localization.

Animals↗

Localization of noise, use of binaural cues, and a description of the superior olivary complex in the smallest carnivore, the least weasel (Mustela nivalis).

Cats and dogs have relatively good sound-localization acuity, and the question arises as to whether this trait is a characteristic of all carnivores or whether it is due to the fact that they have large heads and correspondingly large binaural localization cues available to them. The localization acuity of the least weasel, the smallest extant carnivore, was found to be less accurate than larger carnivores but more accurate than other small mammals. This suggests that carnivores may be under strong selective pressure to localize accurately but that interaural distance may be a limiting factor. The least weasel is capable of using both binaural phase differences and intensity differences to localize, but has a relatively broad mid-frequency range for which neither cue is optimal. Finally, the superior olivary complex of the least weasel is well developed and resembles that of larger carnivores more than that of small rodents.

Animals↗

The auditory periphery of the ferret: postnatal development of acoustic properties.

The development of the acoustics of the auditory periphery of the ferret was examined by measuring the spectral transfer functions (STFs) and the directional characteristics of the outer ears of animals ranging in age from postnatal day 32 (P32) to P54. Using an impulse response technique the STFs were obtained from up to 250 locations throughout free space. The directional responses were calculated for frequencies between 1 kHz and 30 kHz. The low frequency roll-off of the STF decreased with increasing age from around 15 kHz at P32 to an adult value of around 8 kHz by P51. The directional responses of the outer ear of the immature ferrets differed significantly from adult animals in a fashion that was consistent with the smaller size of the auditory periphery. However, by P51 the responses were generally within the normal adult range. The implications of the relatively rapid development of the acoustics of the auditory periphery are discussed in terms of the development of mechanisms subserving sound localization.

Acoustic Stimulation↗

The effects of hearing protectors on auditory localization: evidence from audio-visual target acquisition.

Response times (RT) in an audio-visual target acquisition task were collected from 3 participants while wearing either circumaural earmuffs, foam earplugs, or no hearing protection. Analyses revealed that participants took significantly longer to locate and identify an audio-visual target in both hearing protector conditions than they did in the unoccluded condition, suggesting a disturbance of the cues used by listeners to localize sounds in space. RTs were significantly faster in both hearing protector conditions than in a non-audio control condition, indicating that auditory localization was not completely disrupted. Results are discussed in terms of safety issues involved with wearing hearing protectors in an occupational environment.

Adult↗

Auditory processing of interaural timing information: new insights.

Differences in the time-of-arrival of sounds at the two ears, or interaural temporal disparities (ITDs), constitute one of the major binaural cues that underlie our ability to localize sounds in space. In addition, ITDs contribute to our ability to detect and to discriminate sounds, such as speech, in noisy environments. For low-frequency signals, ITDs are conveyed primarily by "cycle-by-cycle" disparities present in the fine-structure of the waveform. For high-frequency signals, ITDs are conveyed by disparities within the time-varying amplitude, or envelope, of the waveform. The results of laboratory studies conducted over the past few decades indicate that ITDs within the envelopes of high-frequency are less potent than those within the fine-structure of low-frequency stimuli. This is true for both measures of sensitivity to changes in ITD and for measures of the extent of the perceived lateral displacement of sounds containing ITDs. Colburn and Esquissaud (1976) hypothesized that it is differences in the specific aspects of the waveform that are coded neurally within each monaural (single ear) channel that account for the greater potency of ITDs at low frequencies rather than any differences in the more central binaural mechanisms that serve these different frequency regions. In this review, the results of new studies are reported that employed special high-frequency "transposed" stimuli that were designed to provide the high-frequency channels of the binaural processor with envelope-based information that mimics waveform-based information normally available only in low-frequency channels. The results demonstrate that these high-frequency transposed stimuli (1) yield sensitivity to ITDs that approaches, or is equivalent to, that obtained with "conventional" low-frequency stimuli and (2) yield large extents of laterality that are similar to those measured with conventional low-frequency stimuli. These findings suggest that by providing the high-frequency channels of the binaural processor with information that mimics that normally available only at low frequencies, the potency of ITDs in the two frequency regions can be made to be similar, if not identical. These outcomes provide strong support for Colburn and Esquissaud's (1976) hypothesis. The use of high-frequency transposed stimuli, in both behavioral and physiological investigations offers the promise of new and important insights into the nature of binaural processing.

Acoustic Stimulation↗

[Objective determinations of individual sound protectors (author's transl)].

A method for objective measurements of the sound attenuation provided by ear protectors is described, utilizing a head model which contains all normal human sound properties. The difficulties experienced in localizing sound sources when wearing ear protectors is explained by changing the characteristics of the individual frequency bands.

Ear Protective Devices↗

Sound lateralization test in adolescent blind individuals.

Blind individuals require to compensate for the lack of visual information by other sensory inputs. In particular, auditory inputs are crucial to such individuals. To investigate whether blind individuals localize sound in space better than sighted individuals, we tested the auditory ability of adolescent blind individuals using a sound lateralization method. The interaural time difference discrimination thresholds of blind individuals were statistically significantly shorter than those of blind individuals with residual vision and controls. These findings suggest that blind individuals have better auditory spatial ability than individuals with visual cues; therefore, some perceptual compensation occurred in the former.

Acoustic Stimulation↗

[Role of higher divisions of the auditory system in localizing a moving sound source].

Impairment of the ability to localize a moving acoustic image was studied in animal (dogs) following experimental ablation of the auditory cortical areas, and in patients following unilateral electro-shock seizures and focal injuries of the temporal cortex. Unilateral ablation of AI, AII and Ep areas in animals produces a disturbance in differentiation of parameters of a moving acoustic image, while a bilateral ablation results in disappearance of the ability to localize the image. After left-side electro-shock seizures in the patients, localization of movement of the acoustic image did not differ from the normal, while after right-side seizures, the trajectory of the image movement was sharply shortened and shifted to the right. In the case of a focal injury of the patient's left hemisphere, localization changes were of a diffuse nature (shortened trajectory of movement both on the right and left side). In the case of a right-side focus, the trajectory of the movement of the acoustic image on the side of the lesion did not differ from the normal, while on the opposite side it was sharply shortened and shifted towards the mid-line of the head. The conclusion has been made that there is a specialization in the human right hemisphere in achieving spatial hearing, while it is absent in animals.

Animals↗