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PubMed2001

Glycinergic/GABAergic synapses in the lateral superior olive are excitatory in neonatal C57Bl/6J mice.

The lateral superior olive (LSO), a nucleus involved in sound localization, receives tonotopically organized, inhibitory input from the medial nucleus of the trapezoid body (MNTB). To better understand the development of this glycinergic/GABAergic pathway, we used Gramicidin-perforated patch clamp recordings to characterize MNTB-evoked postsynaptic potentials in LSO neurons of neonatal C57Bl/6J mice. We found that during the first postnatal week, MNTB-evoked responses change from being depolarizing to being hyperpolarizing. Most interestingly, depolarizing glycinergic/GABAergic synaptic potentials were able to trigger action potentials, demonstrating that the MNTB-LSO pathway can act as a true excitatory pathway. This transient excitatory action of immature MNTB-LSO synapses might play an important role in activity-dependent sharpening of the tonotopic organization of inhibitory connections in the LSO.

Animals↗
PubMed2006

A two-dimensional model of a directional microphone: calculation of the normal force and moment on the diaphragm.

It has been shown that the parasitoid fly Ormia Ochracea exhibits exceptional sound localization ability achieved through the mechanical coupling of its eardrums [R. N. Miles et al., J. Acoust. Soc. Am. 98, 3059-3070 (1995)]. Based on this biological system a new directional microphone has been designed, having as a basic element a special diaphragm undergoing a rocking motion. This paper considers a 2D model of the microphone in which the diaphragm is considered as a 2D plate having slits on the sides. The slits lead to a backing volume limited by an infinite rigid wall parallel to the diaphragm in its neutral position. The reflection and diffraction of an incoming plane wave by this system are studied to determine the resultant force and resultant moment of pressure upon the diaphragm. The results show that such a microphone will be driven better in the case of narrow slits and deep cavities.

Acoustics↗
PubMed1996

Perceptual consequences of cochlear hearing loss and their implications for the design of hearing aids.

This paper provides an overview of changes in the perception of sound that result from cochlear damage. It starts with a brief introduction to the physiology of the cochlea, emphasizing the role of the "active mechanism" and describing how cochlear function is altered by cochlear damage. Then the effects of cochlear damage on various aspects of perception are described, including absolute sensitivity, frequency selectivity, loudness perception and intensity discrimination, temporal resolution, temporal integration, pitch perception and frequency discrimination, and sound localization and other aspects of binaural and spatial hearing. The possible role of each of these aspects of auditory perception in the ability to understand speech in quiet and in noise is discussed and evaluated. It is concluded that, for losses up to about 45 dB, audibility is the single most important factor. However, for greater losses, poor discrimination of suprathreshold (audible) stimuli is also of major importance. The final section of the paper describes applications of the findings to hearing aid design. It is concluded that linear amplification can be of only limited benefit in compensating for the effects of cochlear damage. Hearing aids incorporating compression can help to compensate for the effects of reduced dynamic range. Digital signal processing to enhance spectral contrast may be of some help in compensating for the effects of reduced frequency selectivity.

Cochlea↗
PubMed2004

Anatomical traces of juvenile learning in the auditory system of adult barn owls.

Early experience plays a powerful role in shaping adult neural circuitry and behavior. In barn owls, early experience markedly influences sound localization. Juvenile owls that learn new, abnormal associations between auditory cues and locations in visual space as a result of abnormal visual experience can readapt to the same abnormal experience in adulthood, when plasticity is otherwise limited. Here we show that abnormal anatomical projections acquired during early abnormal sensory experience persist long after normal experience has been restored. These persistent projections are perfectly situated to provide a physical framework for subsequent readaptation in adulthood to the abnormal sensory conditions experienced in early life. Our results show that anatomical changes that support strong learned neural connections early in life can persist even after they are no longer functionally expressed. This maintenance of silenced neural circuitry that was once adaptive may represent an important mechanism by which the brain preserves a record of early experience.

Animals↗
PubMed1998

Psychometric evaluation of the Gothenburg Profile for measurement of experienced hearing disability and handicap: applications with new hearing aid candidates and experienced hearing aid users.

The Gothenburg Profile (GP) for measurement of experienced hearing disability and handicap was developed with content partly taken from the shortened Hearing Measurement Scale (HMS25). The GP consists of 20 items divided into two subscales. The first subscale measures Experienced Disability as to hearing speech (items 1-5) and sound localization (items 6-10). The second subscale targets the Experienced Handicap in social settings (items 11-15) and the personal reactions to the experienced handicap (items 16-20). In this study, data are presented for new hearing aid candidates (NewHA) (n=441) and for experienced hearing aid users (ExpHA) (n=476). Principal components factor analyses were conducted and a three-factor solution was obtained, supporting the two factors of the Experienced Disability subscale, but just confirming one factor in the Experienced Handicap subscale. The internal consistency reliability (coefficient alpha) was good (0.85 to 0.95) for the subscales as was the test-retest reliability. The ExpHA group expressed significantly greater disability (first subscale) as well as experienced handicap (second subscale). However, when controlling for hearing level the differences disappeared. The clinical use of the GP for assessment of rehabilitation needs is recommended.

Adolescent↗
PubMed1993

Demonstration of the precedence effect in an insect.

Field crickets are interesting models for study of auditory phenomena because they solve many of the same acoustic problems as humans, but with simpler nervous systems. Previous work in this lab and others has investigated sound localization, frequency and temporal pattern discrimination, habituation and dishabituation, and categorical perception. This paper demonstrates the precedence effect in crickets, using a standard two-pulse paradigm with a directional escape response to ultrasound. When two pulses of ultrasound are presented form opposite sides with a delay between, crickets respond only to the first pulse for delays of approximately 4 to 75 ms. For delays of 0 to 2 ms, the direction of response is variable (the first wave front does not have precedence); for delays over approximately 75 ms, crickets respond directionally to each of the two pulses. Some neural correlates of the precedence effect were studied by using this paradigm during recordings from a bilateral pair of ascending second-order auditory interneurons known to initiate ultrasound avoidance. There are no ipsilateral-contralateral differences in their responses that could account for the precedence effect; such interactions in the brain must be involved instead. This seems to be the first test of precedence effect in a nonmammal.

Acoustics↗
PubMed1992

On the ability of neurons in the barn owl's inferior colliculus to sense brief appearances of interaural time difference.

This paper investigates the ability of neurons in the barn owl's (Tyto alba) inferior colliculus to sense brief appearances of interaural time difference (ITD), the main cue for azimuthal sound localization in this species. In the experiments, ITD-tuning was measured during presentation of a mask-probe-mask sequence. The probe consisted of a noise having a constant ITD, while the mask consisted of binaurally uncorrelated noise. Collicular neurons discriminated between the probe and masking noise by showing rapid changes from untuned to tuned and back to untuned responses. The curve describing the relation between probe duration and the degree of ITD-tuning resembled a leaky-integration process with a time constant of about 2 ms. Many neurons were ITD-tuned when probe duration was below 1 ms. These extremely short effective probe durations are interpreted as evidence for neuronal convergence within the pathway computing ITD. The minimal probe duration necessary for ITD-tuning was independent of the bandwidth of the neurons' frequency tuning and also of the best frequency of a neuron. Many narrowly tuned neurons having different best frequencies converge to form a broad-band neuron. To yield the short effective probe durations the convergence must occur in strong temporal synchronism.

Acoustic Stimulation↗
PubMed1996

Dynamic temporal processing of nonspeech acoustic information by children with specific language impairment.

The present study investigated whether children with specific language impairment (SLI) differed from children with normal language learning in their ability to process binaural temporal information. The SLI group was matched with peers of the same chronological age, as well as peers with similar language age. All three subject groups were tested with measures of complex sound localization involving the precedence effect phenomenon. Subjects were required to track the apparent motion of a "moving" fused auditory image (FAI). Movement of the FAI was simulated by varying the delay incrementally between pairs of clicks presented, one each, from two matched loudspeakers placed on opposite sides of the child's head. With this task, the SLI subjects' performances were found to be similar to their language age-matched but chronologically younger peers. Both groups exhibited tracking skills that were statistically poorer than that of the chronologically age-matched group. Additional tests indicated this effect was not due to differences in motoric tracking abilities nor to the SLI subjects' abilities to perceive small binaural time cues. Thus, children with SLI appear to be impaired in their ability to use binaural acoustic information in a dynamic ongoing fashion. The requirements for processing such nonlinguistic acoustic information in a "dynamic and ongoing" fashion may be similar to those involved in the ongoing processing of rapid changes in the temporal and spectral components of the speech chain.

Acoustics↗
PubMed2002

Interaural level differences and the level-meter model.

The interaural level difference (ILD) plays a significant role in sound localization. However, the definition of ILD for noise is open to some interpretation because it is not obvious how to deal with the inevitable level fluctuations. In this article, the ILD is interpreted as an energylike (time-integrated) measure of stimulus level, independent of other stimulus details-particularly interaural correlation. This concept is called the "level-meter model." The model was tested by measuring human ILD thresholds for noise stimuli that were interaurally correlated, or anticorrelated, or uncorrelated. An additional test (not involving lateralization) measured the threshold for level discrimination based on loudness. According to the level-meter model, all four thresholds should be the same. The experimental results showed that the predictions of the level-meter model held good to within about half a dB, although thresholds for level discrimination were systematically higher than ILDs. Among the ILDs themselves, thresholds were slightly higher for uncorrelated noise. The latter result could be explained by replacing the level-meter model with a loudness-meter model, incorporating temporal integration. The same model accounted for the bandwidth dependence of the threshold.

Adult↗
PubMed1989

The neural coding of auditory space.

The barn owl's auditory system computes interaural differences in time and amplitude and derives from them the horizontal and vertical coordinates of the sound source, respectively. Within the external nucleus of its inferior colliculus are auditory neurones, called 'space-specific neurones', that have spatial receptive fields. To activate a space-specific neurone, a sound must originate from a circumscribed region of space, or, if the sounds are delivered to each ear separately, using earphones, the stimuli must have the combination of interaural time and amplitude difference that simulates a sound broadcast from their receptive field. The sound-localization cues are processed in parallel, non-overlapping pathways extending from the cochlear nuclei to the subdivision of the inferior colliculus that innervates the space-specific neurones. Processing in the time pathway involves the coding of monaural phase angle, the derivation of sensitivity for interaural phase difference, and the calculation of interaural time difference (ITD) from interaural phase difference. The last process involves groups of neurones in the inferior colliculus whose collective firing signals a unique ITD, even though the activity of each constituent neurone signals multiple ITDs. The projections of these ensembles to the space-specific neurone endow the latter with a selectivity for ITD. Processing in the amplitude channel, about which less is known, initially involves an inhibitory process that sharpens the directional sensitivity of neurones in a lateral lemniscal nucleus. The inhibition is mediated by a commissural projection from the same lemniscal nucleus of the opposite side. At higher levels of the auditory system, neurones that are tuned to a limited range of interaural amplitude differences are found. It is proposed that at these higher stages, interaural amplitude difference, like ITD, is coded amidst an ensemble of neurones.

Animals↗
PubMed2002

Evolutionary convergence in nervous systems: insights from comparative phylogenetic studies.

Over the past 20 years, cladistic analyses have revolutionized our understanding of brain evolution by demonstrating that many structures, some of which had previously been assumed to be homologous, have evolved many times independently. These and other studies demonstrate that evolutionary convergence in brain anatomy and function is widespread. Although there are relatively few neuroethological studies in which brain and behavior have be studied within an evolutionary framework, three relatively well studied cases are reviewed here: electric communication among gymnotiform and mormyriform fishes, prey capture among frogs, and sound localization among owls. These three examples reveal similar patterns of brain evolution. First, it is clear that novel abilities have evolved many times independently in taxa whose common ancestors lack these abilities. Second, it is apparent that small changes in neural pathways can lead to dramatic changes in an organism's abilities. Brain evolution at this small scale is quite common. The behavioral importance of small scale changes on one hand, and the pervasiveness of convergent evolution on the other, have several implications for understanding brain evolution. First, similar abilities may be conferred by convergent rather than homologous circuits, even among closely related species. Furthermore, closely related species may use the same information in different ways, or they may use different means to obtain the same information. One reason that convergence is so common in the biological world may be that the evolutionary appearance of novel functions is associated with constraints, for example in the algorithms used for a given neural computation. Convergence in functional organization may thus reveal basic design features of neural circuits in species that possess unique evolutionary histories but use similar algorithms to solve basic computational problems.

Animals↗
PubMed2006

Spatial orientation in the bushcricket Leptophyes punctatissima (Phaneropterinae; Orthoptera): I. Phonotaxis to elevated and depressed sound sources.

Many species of acoustically interacting insects live in a complex, arboreal or semi-arboreal habitat. Thus mate finding by phonotaxis requires sound localization in the horizontal and vertical plane. Here we investigated the ability of the duetting bushcricket Leptophyes punctatissima to orient to one of three speakers, positioned at different levels in an artificial grid system, where each point in space could be reached by the male with almost equal probability. The system was designed analogous to a spherical calotte model of bismuth, where, once the male arrived at any nodal point had to decide between only three directions: either up or down and/or left and right. This design does not favour any phonotactic path of the males. All 12 males tested reached the three speaker positions (one in the horizontal plane, one elevated by 45 degrees , one depressed by 45 degrees relative to the starting position) with only little deviation from the shortest possible path. There was no significant difference with respect to the whole phonotactic time needed, the number of segments passed, or the number of stimuli received for the different speaker positions. This remarkable spatial orientation is achieved although the insects have no specialized external ear structures such as mammals, or some owls.

Acoustic Stimulation↗
PubMed2006

Deafferentation induces novel axonal projections in the auditory brainstem after hearing onset.

Deafferentation of neural tissue can result in cell death, morphological changes, and/or alterations in sources of innervation. These changes often occur during a limited period of development. In the auditory brainstem, the ventral cochlear nucleus (VCN) projects to the contralateral but not ipsilateral medial nucleus of the trapezoid body (MNTB). This pathway is part of a circuit that computes interaural intensity differences used in sound localization. Previous studies have shown that, after the cochlea is removed early in postnatal development, cells in the VCN on the deafferented side die, and the intact VCN innervates MNTB on both sides of the brain. These changes after cochlea removal are limited to an early postnatal period that preceeds hearing onset. In this study, we lesioned the VCN directly to evaluate plasticity in axonal pathways after hearing onset. We found that novel projections from the intact VCN to ipsilateral MNTB emerge after lesions performed as late as postnatal day 25. The morphological sequence of events is similar to that seen during the initial development of this pathway. These data suggest that plasticity in the auditory brainstem is possible when pathways are challenged with denervation of target nuclei. The results show that the opportunity for plasticity in auditory brainstem circuitry is more prolonged than previously thought and that novel pathways can form after the normal pathways are fully mature and functional. Moreover, sensitive periods for changes in individual pathways are independently regulated.

Aging↗
PubMed1992

A circuit for coding interaural time differences in the chick brainstem.

Third-order auditory neurons in the avian nucleus laminaris (NL) are the first to receive binaural input. In the chick, NL consists of a monolayer of neurons with polarized dendritic arbors oriented dorsally and ventrally. Afferents from second-order neurons in the ipsilateral nucleus magnocellularis (NM) innervate the dorsal dendrites of NL neurons, distributing processes of approximately equal length to NL neurons along an isofrequency band (roughly caudomedial to rostrolateral). Afferents from the contralateral NM innervate the ventral dendrites of NL neurons, distributing collateral branches sequentially as they proceed from caudomedial to rostrolateral along the isofrequency band of neurons. This innervation pattern could be the basis of a "delay line" circuit, as postulated in models of neural networks mediating sound localization. We examined this circuit by analyzing evoked field potentials using a brain slice preparation containing both NL and NM. The results were consistent with the previous anatomical findings. When the ipsilateral auditory nerve or ipsilateral NM was stimulated, there was no consistent variation in the latency of postsynaptic field potentials across the medial-to-lateral extent of NL. In contrast, when the contralateral NM or NM axons in the crossed dorsal cochlear tract were stimulated, a linear increase in the latency of postsynaptic potentials was observed from medial to lateral positions in NL. When stimulation amplitudes for both the ipsilateral and contralateral inputs were adjusted so as to produce little or no postsynaptic field potential, simultaneous bilateral stimulation evoked a pronounced response. Thus, NL neurons can act as "coincidence detectors." The amplitude of the postsynaptic response was dependent on the relative timing of stimulation of the two inputs. The optimal time difference changed systematically across the medial-to-lateral extent of NL. This system of delay lines and coincidence detectors could provide a mechanism for converting interaural time differences into a "place map" within NL.

Animals↗
PubMed1998

Temporal and binaural properties in dorsal cochlear nucleus and its output tract.

The dorsal cochlear nucleus (DCN) is one of three nuclei at the terminal zone of the auditory nerve. Axons of its projection neurons course via the dorsal acoustic stria (DAS) to the inferior colliculus (IC), where their signals are integrated with inputs from various other sources. The DCN presumably conveys sensitivity to spectral features, and it has been hypothesized that it plays a role in sound localization based on pinna cues. To account for its remarkable spectral properties, a DCN circuit scheme was developed in which three inputs converge onto projection neurons: auditory nerve fibers, inhibitory interneurons, and wide-band inhibitors, which possibly consist of Onset-chopper (Oc) cells. We studied temporal and binaural properties in DCN and DAS and examined whether the temporal properties are consistent with the model circuit. Interneurons (type II) and projection (types III and IV) neurons differed from Oc cells by their longer latencies and temporally nonlinear responses to amplitude-modulated tones. They also showed evidence of early inhibition to clicks. All projection neurons examined were inhibited by stimulation of the contralateral ear, particularly by broadband noise, and this inhibition also had short latency. Because Oc cells had short-latency responses and were well driven by broadband stimuli, we propose that they provide short-latency inhibition to DCN for both ipsilateral and contralateral stimuli. These results indicate more complex temporal behavior in DCN than has previously been emphasized, but they are consistent with the recently described nonlinear behavior to spectral manipulations and with the connectivity scheme deduced from such manipulations.

Action Potentials↗
PubMed2004

Cross correlation by neurons of the medial superior olive: a reexamination.

Initial analysis of interaural temporal disparities (ITDs), a cue for sound localization, occurs in the superior olivary complex. The medial superior olive (MSO) receives excitatory input from the left and right cochlear nuclei. Its neurons are believed to be coincidence detectors, discharging when input arrives simultaneously from the two sides. Many current psychophysical models assume a strict version of coincidence, in which neurons of the MSO cross correlate their left and right inputs. However, there have been few tests of this assumption. Here we examine data derived from two earlier studies of the MSO and compare the responses to the output of a computational model. We find that the MSO is not an ideal cross correlator. Ideal cross correlation implies a strict relationship between the precision of phase-locking of the inputs and the range of ITDs to which a neuron responds. This relationship does not appear to be met. Instead, the modeling implies that a neuron responds over a wider range of ITDs than expected from the inferred precision of phase-locking of the inputs. The responses are more consistent with a scheme in which the neuron can also be activated by the input from one side alone. Such activation degrades the tuning of neurons in the MSO to ITDs.

Acoustic Stimulation↗
PubMed1996

Observer weighting of interaural delays in filtered impulses.

Onset dominance in sound localization was examined by estimating observer weighting of interaural delays for each click of a train of high-frequency filtered clicks. The interaural delay of each click was a normal deviate that was sampled independently on each trial of a single-interval design. In Experiment 1, observer weights were derived for trains of n = 2, 4, 8, or 16 clicks as a function of interclick interval (ICI = 1.8, 3.0, or 12.0 msec). For small n and short ICI (1.8 msec), the ratio of onset weight to remaining weights was as large as 10. As ICI increased, the relative onset weight was reduced. For large n and all ICIs, the ongoing train was weighted more heavily than the onset. This diminishing relative onset weight with increasing ICI and n is consistent with optimum distribution of weights among components. Efficiency of weight distribution is near ideal when ICI = 12 msec and n = 2 and very poor for shorter ICIs and larger ns. Further experiments showed that: (1) onset dominance involves both within- and between-frequency-channel mechanisms, and (2) the stimulus configuration (ICI, n, frequency content, and temporal gaps) affects weighting functions in a complex way not explained by cross-correlation analysis or contralateral inhibition (Lindemann, 1986a, 1986b).

Attention↗
PubMed2001

Tuning to interaural time differences across frequency.

Interaural time differences (ITDs) are an important cue for azimuthal sound localization. Sensitivity to this cue depends on temporal synchrony to the waveform (i.e., phase locking) that begins in the hair cells and is relayed to the neural comparators. The synchrony function is low-pass. Therefore, it is expected that neural tuning to ITDs will become narrower with frequency according to a 1/frequency function. To test this, we measured ITD tuning across frequency in neurons from the superior olivary complex, the dorsal nucleus of the lateral lemniscus, the inferior colliculus, the auditory thalamus, and the auditory cortex. For some neurons in each nucleus, the ITD tuning width did become systematically narrower by the expected 1/frequency relationship. However, in other neurons the ITD tuning width was nearly constant across frequency. Constant ITD tuning width was infrequently observed in neurons of the superior olivary complex but was common in neurons in structures above the superior olivary complex. The nearly constant ITD tuning was caused both by sharper ITD tuning at low frequencies and broader tuning at higher frequencies within the low-frequency band. Neurons with nearly constant tuning to ITDs may be the mechanism underlying the perception of ITDs in humans in which just-noticeable differences to changes in ITD decrease by less than the 1/frequency prediction.

Animals↗