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PubMed2003

[Role of phasic changes in sound signal in localization of auditory image].

The work presents experimental data on certain changes in electrical responses of the auditory system's midbrain centre in a contraphasic binaural presentation of sound impulse series. Neuronal cortical activity is selective in respect to dynamic interaural changes of signals' phasic spectre which may serve as a basis for the mechanisms of localising a moving source of sound. Human auditory evoked potentials reveal a manifestation of memorizing the auditory image movement direction as shown by appearance of stimuli deviant from standard mismatch negativity.

Acoustic Stimulation↗

PubMed1993

Responses of inferior colliculus neurons to time-varying interaural phase disparity: effects of shifting the locus of virtual motion.

1. Motion of sound sources results in temporal variation of the binaural cues for sound localization. We evaluated the influence of virtual motion on neural tuning to one of these cues, interaural phase disparity (IPD). Responses to dichotic stimuli were recorded from single units in the inferior colliculus of the anesthetized cat and gerbil (Meriones unguiculatus). Static IPDs were generated by presenting dichotic tone pairs with a constant phase offset maintained for the duration of the stimulus. Time-varying IPDs were generated by simultaneously presenting a pure tone to one ear and a phase-modulated tone to the other ear. Sets of time-varying stimuli consisted of modulations through partially overlapping ranges of IPD, corresponding to movement of a sound source through partially overlapping arcs in the horizontal plane. 2. In agreement with previous results, neuronal discharge was typically a peaked function of static IPD resulting from both binaural facilitation at favorable IPDs and binaural suppression at unfavorable IPDs. Responses to time-varying IPD stimuli appeared to be shaped by the same facilitative and inhibitory mechanisms that underlie static IPD tuning. Modulation toward the peak of binaural facilitation increased the probability of discharge, and modulation toward the peak of binaural suppression decreased the probability of discharge. However, it was also clear that IPD tuning could be significantly altered by the temporal context of the stimulus. For the vast majority of units in response to modulation through partially overlapping ranges of IPD the discharge rate profiles were generally nonoverlapping. This shift in IPD tuning induced by the virtual motion reflects the fact that the binaural interaction associated with a given IPD depends on the recent history of stimulation. In addition, modulation in opposite directions through the same range of IPDs often elicited asymmetric responses. These nonlinearities imply that most inferior colliculus neurons do not unambiguously encode a specific IPD, but instead may encode small changes of IPD occurring virtually anywhere within their receptive fields. In a few cases modulation through overlapping ranges of IPD elicited contiguous response profiles, indicating that for these units responses were determined entirely by instantaneous IPD. 3. The nonlinearity of responses to time-varying IPD stimuli could not be attributed to monaural entrainment to the phase-modulated signals, did not depend on the phase modulation waveform, and occurred irrespective of which ear received the phase-modulated signal. Responses were similar in cats and gerbils, suggesting that the underlying mechanisms are common to binaural processing in diverse mammalian species.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗

PubMed2002

Necessary conditions for a maximum likelihood estimate to become asymptotically unbiased and attain the Cramer-Rao lower bound. II. Range and depth localization of a sound source in an ocean waveguide.

Analytic expressions for the first order bias and second order covariance of a maximum-likelihood estimate (MLE) are applied to the problem of localizing an acoustic source in range and depth in a shallow water waveguide with a vertical hydrophone array. These expressions are then used to determine necessary conditions on sample size, or equivalently signal-to-noise ratio (SNR), for the localization MLE to become asymptotically unbiased and attain minimum variance as expressed by the Cramer-Rao lower bound (CRLB). These analytic expressions can be applied in a similar fashion to any ocean-acoustic inverse problem involving random data. Both deterministic and completely randomized signals embedded in independent and additive waveguide noise are investigated. As the energy ratio of received signal to additive noise (SANR) descends to the lower operational range of a typical passive localization system, source range and depth estimates exhibit significant biases and have variances that can exceed the CRLB by orders of magnitude. The spatial structure of the bias suggests that acoustic range and depth estimates tend to converge around particular range and depth cells for moderate SANR values.

Environment↗

PubMed1998

Ultrastructure of neurons and large synaptic terminals in the lateral nucleus of the trapezoid body of the cat.

Neurons of the lateral nucleus of the trapezoid body (LNTB), the most prominent periolivary nucleus of the cat superior olivary complex, form an important component of the descending auditory pathways and also innervate the medial superior olive. Cells forming the posteroventral subnucleus (pvLNTB), when investigated by light microscopy, exhibit morphological similarities with globular bushy cells of the cochlear nucleus and principal cells of the medial nucleus of the trapezoid body. These latter two cell types are integral components of brainstem circuitry mediating the early stages of sound localization. In this report, ultrastructural features of LNTB neurons are described. pvLNTB cell bodies are characterized by a round to oval shape, smooth nuclear membrane, and the relative paucity of stacks of rough endoplasmic reticulum. In addition, pvLNTB cell bodies and proximal dendrites are contacted by large synaptic terminals which contain round synaptic vesicles and form multiple asymmetric synaptic junctions. These ultrastructural characteristics are similar to those previously described for globular and principal cells and distinguish pvLNTB cells from cells of the main subnucleus. Large terminals contacting pvLNTB cells contain a specialized organelle assembly, including an adherens plaque associated by filamentous strands with a mitochondrion. We name this organelle assembly the mitochondria-associated adherens complex (MAC) and note its proximity to synaptic junctions. Because high activity rates are characteristic of large terminals in the lower auditory system, the MAC may play a specialized role in membrane stabilization at synapses which generate high rates of vesicle membrane turnover.

Animals↗

PubMed1999

Topographic organization of the dorsal nucleus of the lateral lemniscus in the cat.

The dorsal nucleus of the lateral lemniscus (DNLL) is an auditory structure of the brainstem. It plays an important role in binaural processing and sound localization and it provides the inferior colliculus with an inhibitory projection. The DNLL is a highly conserved auditory structure across mammals, but differences among species in its detailed organization have been reported. The main goal of this study was to analyze the topographic organization of the cat DNLL. Single, small iontophoretic injections of biotinylated dextran amine were made at different loci in the central nucleus of the inferior colliculus (CNIC). The distribution of the labeled structures in the ipsi- and contralateral DNLL was computer reconstructed in three dimensions. In individual sections, a band of labeling is seen in the DNLL on both sides. These two labeled bands occupy symmetric locations and are made of retrogradely labeled neurons with flattened dendritic arbors oriented parallel to each other. Moreover, the ipsilateral labeled band contains labeled terminal fibers parallel to the labeled dendrites. With three-dimensional reconstructions, it becomes evident that the labeled band seen in each individual DNLL section represents a slice through a rostrocaudally oriented lamina. The shape, size, orientation, and location of this lamina change as the injection site is shifted along the tonotopic axis of the CNIC. An injection in the low-frequency region of the CNIC, produces a lamina that resembles a flattened tube located in the dorsolateral corner of the DNLL. An injection in the high-frequency region of the CNIC, by contrast, results in a lamina that is an elongated sheet located at the ventromedial surface of the DNLL. The laminae of the DNLL might constitute the structural basis for its tonotopical organization. Previous studies (Merchan MA, et al. 1994. J Comp Neurol 342:259-278) in conjunction with our current results suggest that the laminar organization in the DNLL might be common among mammals.

Animals↗

PubMed1999

Structure, development, and evolution of insect auditory systems.

This paper provides an overview of insect peripheral auditory systems focusing on tympanate ears (pressure detectors) and emphasizing research during the last 15 years. The theme throughout is the evolution of hearing in insects. Ears have appeared independently no fewer than 19 times in the class Insecta and are located on various thoracic and abdominal body segments, on legs, on wings, and on mouth parts. All have fundamentally similar structures-a tympanum backed by a tracheal sac and a tympanal chordotonal organ-though they vary widely in size, ancillary structures, and number of chordotonal sensilla. Novel ears have recently been discovered in praying mantids, two families of beetles, and two families of flies. The tachinid flies are especially notable because they use a previously unknown mechanism for sound localization. Developmental and comparative studies have identified the evolutionary precursors of the tympanal chordotonal organs in several insects; they are uniformly chordotonal proprioceptors. Tympanate species fall into clusters determined by which of the embryologically defined chordotonal organ groups in each body segment served as precursor for the tympanal organ. This suggests that the many appearances of hearing could arise from changes in a small number of developmental modules. The nature of those developmental changes that lead to a functional insect ear is not yet known.

Animals↗

PubMed2001

Expression of AMPA receptor subunit flip/flop splice variants in the rat auditory brainstem and inferior colliculus.

The expression of alpha-amino-3-hydroxy-5-methyl-4-isoxazolepropionate (AMPA) receptor subunit mRNAs and their flip/flop splice variants was evaluated in the rat auditory brainstem and inferior colliculus employing in situ hybridization with radiolabeled oligonucleotide probes. A differential expression of AMPA receptor subunits in auditory nuclei was observed. In general, neurons in all nuclei of the auditory brainstem express high levels of GluR-C flop and GluR-D flop mRNA, but low to very low levels of GluR-A and GluR-B mRNA. The strongest GluR-C and -D flop expression is found in the ventral and medial part of the anteroventral cochlear nucleus, the posteroventral cochlear nucleus, and the medial and the lateral superior olive. These nuclei are part of the binaural auditory pathway which is important for sound localization in space. In contrast, neurons in the central nucleus of the inferior colliculus express high levels of GluR-B flip but only low levels of the other AMPA receptor subunits. From our data, we conclude that neurons of nuclei involved in binaural processing exhibit a specific "auditory AMPA receptor" which consists primarily of GluR-C flop and -D flop and often lacks GluR-B subunits; this indicates fast kinetics and high Ca(2+) permeability of AMPA receptor currents. In contrast, neurons in the central nucleus of the inferior colliculus contain large amounts of GluR-B flip subunits resulting in Ca(2+) impermeable AMPA receptors with slow kinetics.

Animals↗

PubMed2000

Plasticity of the superior olivary complex.

The superior olivary complex (SOC) is part of the auditory brainstem of the vertebrate brain. Residing ventrally in the rhombencephalon, it receives sensory signals from both cochleae through multisynaptic pathways. Neurons of the SOC are also a target of bilateral descending projections. Ascending and descending efferents of the SOC affect the processing of auditory signals on both sides of the brainstem and in both organs of Corti. The pattern of connectivity indicates that the SOC fulfills functions of binaural signal integration serving sound localization. But whereas many of these connectional features are shared with the inferior colliculus (with the important exception of a projection to the inner ear), cellular and molecular investigations have shown that cells residing in SOC are unique in several respects. Unlike those of other auditory brainstem nuclei, they specifically express molecules known to be involved in development, plasticity, and learning (e.g., GAP-43 mRNA, specific subunits of integrin). Moreover, neurons of the SOC in adult mammals respond to various kinds of hearing impairment with the expression of plasticity-related substances (e.g., GAP-43, c-Jun, c-Fos, cytoskeletal elements), indicative of a restructuring of auditory connectivity. These observations suggest that the SOC is pivotal in the developmental and adaptive tuning of binaural processing in young and adult vertebrates.

Animals↗

PubMed2005

Comparative anatomy of the facial motor nucleus in mammals, with an analysis of neuron numbers in primates.

The facial motor nucleus (VII) contains motoneurons that innervate the facial muscles of expression. In this review, the comparative anatomy of this brainstem nucleus is examined. Several aspects of the anatomical organization of the VII appear to be common across mammals, such as the distribution of neuron types, general topography of muscle representation, and afferent connections from the midbrain and brainstem. Phylogenetic specializations are apparent in the proportion of neurons allocated to the representation of subsets of muscles and the degree of differentiation among subnuclei. These interspecific differences may be related to the elaboration of certain facial muscles in the context of socioecological adaptations such as whisking behavior, sound localization, vocalization, and facial expression. Furthermore, current evidence indicates that direct descending corticomotoneuron projections in the VII are present only in catarrhine primates, suggesting that this connectivity is an important substrate for the evolution of enhanced mobility and flexibility in facial expression. Data are also presented from a stereologic analysis of VII neuron numbers in 18 primate species and a scandentian. Using phylogenetic comparative statistics, it is shown that there is not a correlation between group size and VII neuron number (adjusted for medulla volume) among primates. Great apes and humans, however, display moderately more VII neurons that expected for their medulla size.

Anatomy, Comparative↗

PubMed2006

Comparative gene expression analysis reveals a characteristic molecular profile of the superior olivary complex.

The superior olivary complex (SOC) is a very conspicuous structure in the mammalian auditory brainstem. It represents the first binaural processing center and is important for sound localization in the azimuth and in feedback regulation of cochlear function. In order to define molecular determinants of the SOC, which are of potential functional relevance, we have performed a comprehensive analysis of its transcriptome by serial analysis of gene expression in adult rats. Here, we performed a detailed analysis of the SOC's gene expression profile compared to that of two other neural tissues, the striatum and the hippocampus, and with extraocular muscle tissue. This tested the hypothesis that SOC-specific or significantly upregulated transcripts provide candidates for the specific function of auditory neurons. Thirty-three genes were significantly upregulated in the SOC when compared to the two other neural tissues. Thirteen encoded proteins involved in neurotransmission, including action potential propagation, exocytosis, and myelination; five genes are important for the energy metabolism, and five transcripts are unknown or poorly characterized and have yet to be described in the nervous system. The comparison of functional gene classes indicates that the SOC has the highest energy demand of the three neural tissues, yet protein turnover is apparently not increased. This suggests a high energy demand for fueling auditory neurotransmission. Such a demand may have implications on auditory-specific tasks and relate to central auditory processing disorders. Ultimately, these data provide new avenues to foster investigations of auditory function and to advance molecular physiology in the central auditory system.

Animals↗

PubMed2003

Axonal pathways to the lateral superior olive labeled with biotinylated dextran amine injections in the dorsal cochlear nucleus of rats.

The lateral superior olive (LSO) contains cells that are sensitive to intensity differences between the two ears, a feature used by the brain to localize sounds in space. This report describes a source of input to the LSO that complements bushy cell projections from the ventral cochlear nucleus (VCN). Injections of biotinylated dextran amine (BDA) into the dorsal cochlear nucleus (DCN) of the rat label axons and swellings in several brainstem structures, including the ipsilateral LSO. Labeling in the ipsilateral LSO was confined to a thin band that extended throughout the length of the structure such that it resembled an LSO isofrequency lamina. The source of this labeled pathway was not obvious, because DCN neurons do not project to the LSO, and VCN bushy cells were not filled by these injections. Filled neurons in several brainstem structures emerged as possible sources. Three observations suggest that most of the axonal labeling in the LSO derives from a single source. First, the number of labeled VCN planar multipolar cells and the amount of labeling in the LSO were consistent and robust across animals. In contrast, the number of labeled cells in most other structures was small and highly variable. Second, the locations of planar cells and filled axons in the LSO were related topographically to the position of the DCN injection site. Third, labeled terminal arborizations in the LSO arose from collaterals of axons in the trapezoid body (output tract of planar cells). We infer that planar multipolar cells, in addition to bushy cells, are a source of ascending input from the cochlear nucleus to the LSO.

Animals↗

PubMed2006

Formation of the avian nucleus magnocellularis from the auditory anlage.

In the avian auditory system, the neural network for computing the localization of sound in space begins with bilateral innervation of nucleus laminaris (NL) by nucleus magnocellularis (NM) neurons. We used antibodies against the neural specific markers Hu C/D, neurofilament, and SV2 together with retrograde fluorescent dextran labeling from the contralateral hindbrain to identify NM neurons within the anlage and follow their development. NM neurons could be identified by retrograde labeling as early as embryonic day (E) 6. While the auditory anlage organized itself into NM and NL in a rostral-to-caudal fashion between E6 and E8, labeled NM neurons were visible throughout the extent of the anlage at E6. By observing the pattern of neuronal rearrangements together with the pattern of contralaterally projecting NM fibers, we could identify NL in the ventral anlage. Ipsilateral NM fibers contacted the developing NL at E8, well after NM collaterals had projected contralaterally. Furthermore, the formation of ipsilateral connections between NM and NL neurons appeared to coincide with the arrival of VIIIth nerve fibers in NM. By E10, immunoreactivity for SV2 was heavily concentrated in the dorsal and ventral neuropils of NL. Thus, extensive pathfinding and morphological rearrangement of central auditory nuclei occurs well before the arrival of cochlear afferents. Our results suggest that NM neurons may play a central role in formation of tonotopic connections in the auditory system.

Animals↗

PubMed1975

Organization and development of brain stem auditory nuclei of the chicken: organization of projections from n. magnocellularis to n. laminaris.

The tonotopic and topographic organization of the bilateral projection from second-order auditory neurons of nucleus magnocellularis (NM) to nucleus laminaris (NL) was examined in young chickens. In one group of birds, the NM axons which innvervate the contralateral NL were severed by cutting the crossed dorsal cochlear tract at the midline. Heavy terminal degeneration in NL was confined to the neuropil area immediately ventral to the perikaryl lamina. Very little degeneration was seen in the dorsal neuropil region. In a second series of animals, the charactertistic frequency (CF) of cells in an area of NM was first determined by microelectrode recording techniques and then a small electrolytic lesion was made through the recording electrode. Following survival periods of 24-48 hours, the distribution of projections from the lesioned area to the ipsilateral and contralateral NL was examined using the Fink-Heimer method. As previously described in the pigeon, projections from NM terminate densely in the neuropil region immediately dorsal to the ipsilateral NL cell bodies and ventral to the perikaryl layer on the contralateral side, providing each NL neuron with segregated binaural innervation. Lesions in any area of the NM produced degeneration confined to a limited caudo-rostral and medio-lateral portion of both laminar nuclei. To investigate this topographic relationship, the cuado-rostral extents of the lesion in NM and of the resulting degeneration in both NL were determined. Linear regression and correlation analyses then related these positional values to each other and to the CF found at the center of each lesion. All correlations were highly significant and ranged from 0.78 between the position of the lesion in NM and CF to 0.91 between the caudo-rostral position of degeneration in the NL ipsilateral and contralateral to the lesion. It is concluded that neurons in NM project in a very discrete topographic, tonotopic and symmetrical fashion to NL on both sides of the brain, contributing to the binaural response properties and tonotopic organization of neurons in NL. The results also suggest that the organization of projections from NM to NL could provide a mechanism for the differential transmission delay required by a "place" model of low-frequency sound localization.

Animals↗

PubMed1986

Ultrastructural features of the ventromedial region of the laminar nucleus in the red-eared turtle (Chrysemys scripta elegans).

The nuclei of the torus semicircularis, in particular the laminar nucleus, have been functionally implicated in sound localization, vocalization, and mating behavior. In the red-eared turtle the ventromedial region of the laminar nucleus (containing two discrete dense cellular areas and the surrounding neuropil) was examined electron microscopically in the present study. Neuronal cell bodies in the two cellular areas were different in size, shape, cytoplasmic constituents, and their relationship to each other. Cell bodies in the layer beneath the ependymal surface were almost always surrounded by lamellae while cell bodies in the layer above the central nucleus were in close apposition. We are speculating that cell bodies in the superficial layer may be neuroendocrine in nature. This speculation is based on the presence of lipidlike droplets within cell bodies, and previous findings indicating steroid binding and synthesis in this region in reptilian brains. Cell bodies located above the central nucleus were characteristic in that they contained lamellar bodies and extensive well-developed Golgi regions. Closely apposed groups of these cell bodies were frequently surrounded by or in close apposition to large axonal profiles. These profiles were filled with clear core vesicles, numerous mitochondria, and clusters of small dense core particles. Synaptic contacts between large axonal profiles and cell bodies were not observed. The neuropil surrounding cell bodies of the inner layer of this region of the laminar nucleus contained glomeruli composed of a central axon surrounded, and in synaptic contact, with dendrites. Throughout the area of the laminar nucleus studied, synapses appeared to be primarily of the asymmetrical type.

Animals↗

PubMed1991

Classification of the principal cells of the medial nucleus of the trapezoid body.

Cells in the medial nucleus of the trapezoid body were intracellularly labeled in brainstem tissue slices of two bat and two rodent species. The main cell type found in this nucleus, the principal cell, is an important link in the relay of ascending projections from the contralateral cochlear nucleus to the lateral superior olive, completing an essential pathway for sound localization. Principal cells are often viewed as a highly homogeneous group with a consistent morphology as well as a common function. Intracellular labeling has revealed a number of new axonal and dendritic features of principal cells. Some of these features vary widely from cell to cell, suggesting that the population of principal cells contains several morphologically distinct subgroups. Similar subsets of principal cells were recognized in all species examined. Five subgroups were distinguished on the basis of the position of dendritic fields. Although the dendrites of most labeled cells were confined to the medial nucleus of the trapezoid body, some principal cells had dendrites that spread outside the nucleus to one of several adjacent periolivary cell groups. Cells were also found that had dendrites that spread medially across the midline and into the contralateral medial nucleus of the trapezoid body. Axonal projections were used to distinguish two additional subgroups of principal cells. All principal cells project to the lateral superior olive and virtually all have one or more secondary projections. There are two subgroups with unusual collateral projections: one with collaterals that extended to the lateral lemniscus and one with recurrent collateral axons.

Animals↗

PubMed1992

Acoustic chiasm V: inhibition and excitation in the ipsilateral and contralateral projections of LSO.

When this series of experiments was begun in 1984, the activity of each lateral superior olive (LSO) in the mammalian hindbrain was known to encode the hemifield of acoustic space containing a sound source. However, the almost random bilaterality of its ascending projections seemed to jumble that identification before reaching the midbrain. At the same time, electrophysiological studies of LSO and its efferent target in the inferior colliculus, along with the strictly contralateral deficits in sound localization resulting from unilateral lesions above the level of the superior olives, indicated that hemifield allegiance was largely maintained (though reversed) at the midbrain. Here we present seven lines of biochemical evidence, some combined with prior ablations, supporting the notion that the anatomical segregation of the ipsilateral and contralateral fibers ascending from the LSO is accompanied by a corresponding segregation of their neurotransmitters: most of the ascending ipsilateral projection is probably glycinergic and, hence, inhibitory in effect, while most of the contralateral projection is probably glutamatergic/aspartergic and, hence, excitatory in effect. Taken together, the inhibitory ipsilateral projections and the excitatory contralateral projections serve to amplify functional contralaterality at the higher levels of the auditory system.

Acoustic Stimulation↗

PubMed1995

Contributions of experience to the development of musical processing in infancy.

Full-term infants' performance in detection of melodic alterations appeared to be influenced by perceptual experience from 6 months to 1 year of age, and an experiment with infants born prematurely supported the hypothesis that experience affects music processing in infancy. These findings suggest parallel developmental tendencies in the perception of music and speech that may reflect general acquisition of perceptual abilities for processing of complex auditory patterns. This acquisition may contribute to the cultural enfranchisement of infants through perceptual experience.

Attention↗

PubMed1977

Morphology of the macula neglecta in sharks of the genus Carcharhinus.

Ears from several species of carcharhinid sharks were studied by gross dissection, light microscopy, transmission electron microscopy, and scanning electron microscopy. Structures along a possible sound transmission path to the ear are described, but main consideration is given to the structure of the macula neglecta. The macula neglecta is composed of two patches of sensory epithelium which line part of the posterior canal duct. In an adult shark the larger of these contains 224,000 sensory hair cells oriented so as to detect forces directed posteroventrolaterally in the duct. The smaller patch contains 43,000 hair cells oriented so as to detect oppositely directed forces. These receptor cells project through numerous small terminals to a total for both patches of 4,700 myelinated nerve fibers. Cytostructural variations throughout the hair cell population are also reported. Estimated acoustic properties of the tissues in this complex and the processing potential of the neural elements are interpreted as suggestive of auditory function. A mechanism based on the geometry of the receptor arrays is proposed to explain behaviorally observed instantaneous sound localization from the farfield. Evolution of the macula neglecta is reviewed, and evidence for homology of the macula neglecta and amphibian papilla is presented.

Animals↗