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Spatial coordinates of human auditory working memory.

The accuracy of localizing remembered sound sources was investigated by employing a delayed-response task, where a small light spot, projected onto a screen by a laser diode attached to the head, had to be spatially aligned with either actual or remembered stimulus positions. Systematic errors indicated overestimation of the eccentricity of remembered targets compared to direct stimulus localization. This overestimation increased with prolonged response delay, suggesting that the coordinates of memorized space are distorted with respect to perceived actual sound location and that this distortion increases as a function of time.

Acoustic Stimulation↗

Directional selectivity and frequency tuning of midbrain cells in the oyster toadfish, Opsanus tau.

Single-unit recordings were made from areas in the midbrain (torus semicircularis) of the oyster toadfish. We evaluated frequency tuning and directional responses using whole-body oscillation to simulate auditory stimulation by particle motion along axes in the horizontal and mid-sagittal planes. We also tested for bimodality in responses to auditory and hydrodynamic stimuli. One recording location in each animal was marked by a neurobiotin injection to confirm the recording site. Recordings were made in nucleus centralis, nucleus ventrolateralis, and the deep cell layer. Most units were frequency-selective with best frequencies between 50 and 141 Hz. Suppression of activity was apparent in 10% of the cells. Bimodality was common, including inhibition and suppression of background activity by auditory or hydrodynamic stimulation. The majority of the cells were directionally selective with directional response patterns that were sharpened compared with those of primary saccular afferents. The best directional axes were arrayed widely in spherical space, covering most azimuths and elevations. This representation is adequate for the computation of the motional axis of an auditory stimulus for sound source localization.

Acoustic Stimulation↗

Localization of marine mammals near Hawaii using an acoustic propagation model.

Humpback whale songs were recorded on six widely spaced receivers of the Pacific Missile Range Facility (PMRF) hydrophone network near Hawaii during March of 2001. These recordings were used to test a new approach to localizing the whales that exploits the time-difference of arrival (time lag) of their calls as measured between receiver pairs in the PMRF network. The usual technique for estimating source position uses the intersection of hyperbolic curves of constant time lag, but a drawback of this approach is its assumption of a constant wave speed and straight-line propagation to associate acoustic travel time with range. In contrast to hyperbolic fixing, the algorithm described here uses an acoustic propagation model to account for waveguide and multipath effects when estimating travel time from hypothesized source positions. A comparison between predicted and measured time lags forms an ambiguity surface, or visual representation of the most probable whale position in a horizontal plane around the array. This is an important benefit because it allows for automated peak extraction to provide a location estimate. Examples of whale localizations using real and simulated data in algorithms of increasing complexity are provided.

Algorithms↗

The effects of perceived motion on sound-source lateralization.

Lateralization of dichotic click pairs differing in time of onset was studied under conditions of angular acceleration, optokinetic stimulation, and gaze fixation. Data obtained from 20 subjects with normal hearing indicate poorer left-right judgment performance for small time differences, as well as shifts in subjective simultaneity, for all experimental conditions relative to control conditions. In addition, response times increased for the experimental conditions. The results suggest that real or apparent motion may affect an individual's ability to process one of the major cues for sound-source localization--binaural time difference--under conditions similar to those encountered in vehicular motion.

Acceleration↗

Neuromagnetic responses associated with perceptual segregation of pitch.

In recent EEG investigations [Johnson, 2003] [Hautus, 2005], we described a novel late negative ERP component associated with binaural processing of auditory pitch based solely on interaural timing differences ("dichotic pitch"), an acoustic phenomenon that is closely analogous to visual perception of stereoscopic depth based on retinal disparities. The present study extends this research with neuromagnetic recordings of auditory evoked fields (AEFs) elicited by dichotically-embedded pitches. Eight healthy adult subjects listened to control stimuli consisting of 500 ms segments of broadband acoustic noise presented identically to both ears via earphones, and dichotic pitch stimuli created by introducing a dichotic delay to a narrow frequency region of the same noise segments and resulting in a perception of a pitch lateralized to the left or right of auditory space. Auditory-evoked fields (AEFs) were recorded using a 151 channel whole-head MEG system. Comparison of control and dichotic-pitch AEFs showed reliable amplitude differences during a time window of 150-350 ms. AEFs over the left hemisphere showed larger effects for contralateral than ipsilateral pitches, while the right hemisphere showed no differences for differently lateralized sources. The results indicate a relatively late stage of neural processing of binaurally-derived cues for the perceptual segregation of concurrent sound sources and support a right-hemisphere dominance for the processing of sound-source localization.

Acoustic Stimulation↗

Enhanced sensitivity to echo cues in blind subjects.

Many studies have reported that blind people compensate for their visual deficit by sharpening auditory processes. Here we compare the sensitivity to echo cues between blind and sighted subjects. In the first experiment, the blind subjects were more accurate than the sighted subjects in localizing an object on the basis of echo cues. To ensure that enhanced echolocalization abilities were not only due to the fact that blind individuals are more used to consciously paying attention to echo cues and are more familiar with this kind of tasks than sighted subjects, we tested both groups of subjects in a simple azimuthal localization task of auditory stimuli. In this second experiment, we evaluated the influence of irrelevant echo signals on auditory localization by placing the subjects and the sound sources at different positions in a sound reverberant room. Results revealed that blind subjects exhibit a higher sensitivity to echo signals than sighted subjects.

Acoustic Stimulation↗

Right hemisphere specialization for intensity discrimination of musical and speech sounds.

Sound intensity is the primary and most elementary feature of auditory signals. Its discrimination plays a fundamental role in different behaviours related to auditory perception such as sound source localization, motion detection, and recognition of speech sounds. This study was aimed at investigating hemispheric asymmetries for processing intensity of complex tones and consonant-vowel syllables. Forty-four right-handed non-musicians were presented with two dichotic matching-to-sample tests with focused attention: one with complex tones of different intensities (musical test) and the other with consonant-vowel syllables of different intensities (speech test). Intensity differences (60, 70, and 80 dBA) were obtained by altering the gain of a synthesized harmonic tone (260 Hz fundamental frequency) and of a consonant-vowel syllable (/ba/) recorded from a natural voice. Dependent variables were accuracy and reaction time. Results showed a significant clear-cut left ear advantage in both tests for both dependent variables. A monaural control experiment ruled out possible attentional biases. This study provides behavioural evidence of a right hemisphere specialization for the perception of the intensity of musical and speech sounds in healthy subjects.

Adult↗

Horizontal localization with bilateral hearing aids: without is better than with.

This paper studies the effect of bilateral hearing aids on directional hearing in the frontal horizontal plane. Localization tests evaluated bilateral hearing aid users using different stimuli and different noise scenarios. Normal hearing subjects were used as a reference. The main research questions raised in this paper are: (i) How do bilateral hearing aid users perform on a localization task, relative to normal hearing subjects? (ii) Do bilateral hearing aids preserve localization cues, and (iii) Is there an influence of state of the art noise reduction algorithms, more in particular an adaptive directional microphone configuration, on localization performance? The hearing aid users were tested without and with their hearing aids, using both a standard omnidirectional microphone configuration and an adaptive directional microphone configuration. The following main conclusions are drawn. (i) Bilateral hearing aid users perform worse than normal hearing subjects in a localization task, although more than one-half of the subjects reach normal hearing performance when tested unaided. For both groups, localization performance drops significantly when acoustical scenarios become more complex. (ii) Bilateral, i.e., independently operating hearing aids do not preserve localization cues. (iii) Overall, adaptive directional noise reduction can have an additional and significant negative impact on localization performance.

Acoustic Stimulation↗

Directional response properties of saccular afferents of the toadfish, Opsanus tau.

The displacement sensitivity, frequency response, and directional response properties of primary saccular afferents of toadfish (Opsanus tau) were studied in response to a simulation of acoustic particle motion for which displacement magnitudes and directions were manipulated in azimuth and elevation. Stimuli were 50, 100, and 200 Hz sinusoidal, translatory oscillations of the animal at various axes in the horizontal and midsagittal planes. Thresholds in these planes defined a cell's characteristic axis (the axis having the lowest threshold) in spherical coordinates. Recordings were made from afferents in rostral, middle, and caudal bundles of the saccular nerve. The most sensitive saccular afferents responded with a phase-locked response to displacements as small as 0.1 nm. This sensitivity rivals that of the mammalian cochlea and is probably common to the sacculi and other otolith organs of most fishes. Most afferents showed lower thresholds at 100 Hz than at 50 or 200 Hz. Eighty percent of afferents have three-dimensional directional properties that would be expected if they innervated a group of hair cells having the same directional orientation on the saccular epithelium. Of the afferents that are not perfectly directional, most appear to innervate just two groups of hair cells having different orientations. The directional characteristics of afferents are qualitatively correlated with anatomically defined patterns of hair cell orientation on the saccule. In general, azimuths of best sensitivity tend to lie parallel to the plane of the otolith and sensory epithelium. Elevations of best sensitivity correspond well with hair cell orientation patterns in different regions of the saccular epithelium. Directional hearing in the horizontal plane probably depends upon the processing of interaural differences in overall response magnitude. These response differences arise from the gross orientations of the sacculi and are represented, in part, as time differences among nonspontaneous afferents that show level-dependent phase angles of synchronization. Directional hearing in the vertical plane may be derived from the processing of across-afferent profiles of activity within each saccule. Fishes were probably the first vertebrates to solve problems in sound source localization, and we suggest that their solutions formed a model for those of their terrestrial inheritors.

Acoustic Stimulation↗

Investigations of the precedence effect in budgerigars: effects of stimulus type, intensity, duration, and location.

Auditory experiments on the localization of sounds in the presence of reflections, or echoes, that arrive later and from different directions are important to understanding hearing in natural environments. The perceived location of the auditory image can change with the time delay between the presentations of a leading and lagging sound. These changes in perceived location, encompassing the precedence effect, have been examined behaviorally or physiologically in humans and a number of animals. Here, these results are extended to include budgerigars. Behavioral methods were used to measure the discrimination performance between a stimulus presented at + and - 90 degrees azimuth with a delay (left-right), from the same two stimuli presented with the opposite delay (right-left). At short delays, where humans experience summing localization, budgerigars have difficulty discriminating between the two presentation types. With increasing delays, where humans experience localization dominance, budgerigars show improved discrimination performance. At even longer delays, where echo thresholds are found in humans, discrimination performance worsens again. The shapes of the discrimination functions are affected by the intensity, locations, and durations of the stimuli, and are subject to a buildup effect. These results show that budgerigars exhibit the phases of the precedence effect similar to humans and other animals.

Animal Communication↗

Processing of acoustic signals in the auditory system of bony fish.

In order to determine unambiguously the bearing of a sound source, a fish must be able to resolve acoustic pressure and the components of the acoustic displacement vector from the signals detected by the otolithic organs. A new hypothesis for the processing of acoustical information by bony fish is presented. It is demonstrated that much of the processing required to do this may be implicit in the structure of the ear and its associated neural innervation. Possible algorithms are presented that the central nervous system might use to further process the derived information to localize a sound source and discriminate frequency and range. The hypothesis is shown to be consistent with much of what is known of the morphology and physiology of the auditory system of bony fishes.

Algorithms↗

Linking the sounds of dolphins to their locations and behavior using video and multichannel acoustic recordings.

It is difficult to attribute underwater animal sounds to the individuals producing them. This paper presents a system developed to solve this problem for dolphins by linking acoustic locations of the sounds of captive bottlenose dolphins with an overhead video image. A time-delay beamforming algorithm localized dolphin sounds obtained from an array of hydrophones dispersed around a lagoon. The localized positions of vocalizing dolphins were projected onto video images. The performance of the system was measured for artificial calibration signals as well as for dolphin sounds. The performance of the system for calibration signals was analyzed in terms of acoustic localization error, video projection error, and combined acoustic localization and video error. The 95% confidence bounds for these were 1.5, 2.1, and 2.1 m, respectively. Performance of the system was analyzed for three types of dolphin sounds: echolocation clicks, whistles, and burst-pulsed sounds. The mean errors for these were 0.8, 1.3, and 1.3 m, respectively. The 95% confidence bound for all vocalizations was 2.8 m, roughly the length of an adult bottlenose dolphin. This system represents a significant advance for studying the function of vocalizations of marine animals in relation to their context, as the sounds can be identified to the vocalizing dolphin and linked to its concurrent behavior.

Acoustics↗

Statistics and vertical directionality of low-frequency ambient noise at the North Pacific Acoustic Laboratory site.

We examine statistical and directional properties of the ambient noise in the 10-100 Hz frequency band from the NPAL array. Marginal probability densities are estimated as well as mean square levels, skewness and kurtoses in third octave bands. The kurotoses are markedly different from Gaussian except when only distant shipping is present. Extremal levels reached approximately 150 dB re 1 micro Pa, suggesting levels 60dB greater than the mean ambient were common in the NPAL data sets. Generally, these were passing ships. We select four examples: i) quiescent noise, ii) nearby shipping, iii) whale vocalizations and iv) a micro earthquake for the vertical directional properties of the NPAL noise since they are representative of the phenomena encountered. We find there is modest broadband coherence for most of these cases in their occupancy band across the NPAL aperture. Narrowband coherence analysis from VLA to VLA was not successful due to ambiguities. Examples of localizing sources based upon this coherence are included. kw diagrams allow us to use data above the vertical aliasing frequency. Ducted propagation for both the quiescent and micro earthquake (T phase) are identified and the arrival angles of nearby shipping and whale vocalizations. MFP localizations were modestly successful for nearby sources, but long range ones could not be identified, most likely because of signal mismatch in the MFP replica.

Acoustic Stimulation↗

Evolution of hearing in vertebrates: the inner ears and processing.

This paper considers aspects of the evolution of the vertebrate auditory system from an 'ichthyocentric' perspective. It is argued that all vertebrate auditory systems are required to do certain basic tasks including acoustic feature discrimination, sound source localization, frequency analysis, and auditory scene analysis, among others. These sorts of capabilities arose very early in the evolution of the vertebrates and have been modified by selection in different species. In some cases the same structures have been involved in detection and analysis throughout the vertebrates, while in other cases the mechanism by which the same type of analysis takes place may have changed.

Animals↗

Modeling individual differences in ferret external ear transfer functions.

Individual variations in head and outer ear size, as well as growth of these structures during development, can markedly alter the values of the binaural and monaural cues which form the basis for auditory localization. This study investigated individual differences in the directional component of the head-related transfer function of both adult and juvenile ferrets. In line with previous studies in humans and cats, intersubject spectral differences were found to be reduced by scaling one of the directional transfer functions on a log-frequency axis. The optimal scale factor correlated most highly with pinna cavity height. Optimal frequency scaling reduced interear spectral difference equally well for adult-juvenile comparisons as for comparisons between pairs of adult ears. This illustrates that the developmental changes in localization cue values should be at least partly predictable on the basis of the expected growth rate of the outer ear structures. Predictions of interaural time differences (ITDs) were also derived from the physical dimensions of the head. ITDs were found to be poorly fitted by the spherical head model, while much better predictions could be derived from a model based on von Mises spherical basis functions. Together, these findings show how more accurate estimates of spatial cue values can be made from knowledge of the dimensions of the head and outer ears, and may facilitate the generation of virtual acoustic space stimuli in the absence of acoustical measurements from individual subjects.

Age Factors↗

Synaptic depression improves coincidence detection in the nucleus laminaris in brainstem slices of the chick embryo.

Neurons in the nucleus laminaris detect the coincidence of binaural signals, and are the first neurons to calculate the interaural time difference for the sound source localization in birds. In this paper, we have studied contributions of synaptic depression to the coincidence detection in the nucleus laminaris in a slice preparation of the chick embryo (E16-18), using the whole-cell patch recording technique. Under voltage clamp, EPSCs decreased progressively in their amplitude during the course of tetanic stimuli. This synaptic depression was primarily ascribed to the reduction of transmitter release from the presynaptic terminal, because the depression was decreased by reducing transmitter release with 2.5 microm Cd2+ but was not affected by reducing desensitization of postsynaptic AMPA receptors with 20 microm cyclothiazide. Under current clamp, trains of 10 stimuli of 100 Hz were applied bilaterally with changing the time intervals systematically between both sides. Response window, defined as the time interval corresponding to the half-maximum firing probability, was narrowed during the course of the stimulus train, and this occurred in parallel with a decrease in the EPSP amplitude. In addition, the reduction of the EPSP amplitude due to 2.5 microm Cd2+ or 2 microm CNQX improved the accuracy of coincidence detection. These results indicate that the synaptic depression may improve the coincidence detection in the chick laminaris neurons.

Action Potentials↗