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Acoustic environment determines phosphorylation state of the Kv3.1 potassium channel in auditory neurons.

Sound localization by auditory brainstem nuclei relies on the detection of microsecond interaural differences in action potentials that encode sound volume and timing. Neurons in these nuclei express high amounts of the Kv3.1 potassium channel, which allows them to fire at high frequencies with short-duration action potentials. Using computational modeling, we show that high amounts of Kv3.1 current decrease the timing accuracy of action potentials but enable neurons to follow high-frequency stimuli. The Kv3.1b channel is regulated by protein kinase C (PKC), which decreases current amplitude. Here we show that in a quiet environment, Kv3.1b is basally phosphorylated in rat brainstem neurons but is rapidly dephosphorylated in response to high-frequency auditory or synaptic stimulation. Dephosphorylation of the channel produced an increase in Kv3.1 current, facilitating high-frequency spiking. Our results indicate that the intrinsic electrical properties of auditory neurons are rapidly modified to adjust to the ambient acoustic environment.

Acoustic Stimulation↗

Hearing in domestic pigs (Sus scrofa) and goats (Capra hircus).

Behavioral audiograms were determined for three pigs and two goats. The hearing of the pigs ranged from 42 Hz to 40.5 kHz with a region of best sensitivity from 250 Hz to 16 kHz. Hearing in goats ranged from 78 Hz to 37 kHz with a well-defined point of best sensitivity at 2 kHz. Because these animals are unable to localize high-frequency tones, it seems unlikely that selective pressure to use the interaural spectral-difference cue for sound localization is behind their high-frequency hearing. Instead, we suggest that these and other hoofed mammals evolved high-frequency hearing in order to use monaural locus cues which prevent front/back locus reversals.

Acoustic Stimulation↗

Detection of interaural correlation by neurons in the superior olivary complex, inferior colliculus and auditory cortex of the unanesthetized rabbit.

A critical binaural cue important for sound localization and detection of signals in noise is the interaural time difference (ITD), or difference in the time of arrival of sounds at each ear. The ITD can be determined by cross-correlating the sounds at the two ears and finding the ITD where the correlation is maximal. The amount of interaural correlation is affected by properties of spaces and can therefore be used to assess spatial attributes. To examine the neural basis for sensitivity to the overall level of the interaural correlation, we identified subcollicular neurons and neurons in the inferior colliculus (IC) and auditory cortex of unanesthetized rabbits that were sensitive to ITDs and examined their responses as the interaural correlation was varied. Neurons at each brain level could show linear or non-linear responses to changes in interaural correlation. The direction of the non-linearities in most neurons was to increase the slope of the response change for correlations near 1.0. The proportion of neurons with non-linear responses was similar in subcollicular and IC neurons but increased in the auditory cortex. Non-linear response functions to interaural correlation were not related to the type of response as determined by the tuning to ITDs across frequencies. The responses to interaural correlation were also not related to the frequency tuning of the neuron, unlike the responses to ITD, which broadens for neurons tuned to lower frequencies. The neural discriminibility of the ITD using frozen noise in the best neurons was similar to the behavioral acuity in humans at a reference correlation of 1.0. However, for other reference ITDs the neural discriminibility was more linear and generally better than the human discriminibility of the interaural correlation, suggesting that stimulus rather than neural variability is the basis for the decline in human performance at lower levels of interaural correlation.

Animals↗

Mechanically coupled ears for directional hearing in the parasitoid fly Ormia ochracea.

An analysis is presented of the mechanical response to a sound field of the ears of the parasitoid fly Ormia ochracea. This animal shows a remarkable ability to detect the direction of an incident sound stimulus even though its acoustic sensory organs are in very close proximity to each other. This close proximity causes the arrival times of the sound pressures at the two ears to be less than 1 to 2 microseconds depending on the direction of propagation of the sound wave. The small differences in these two pressures must be processed by the animal in order to determine the incident direction of the sound. In this fly, the ears are so close together that they are actually joined by a cuticular structure which couples their motion mechanically and subsequently magnifies interaural differences. The use of a cuticular structure as a means to couple the ears to achieve directional sensitivity is novel and has not been reported in previous studies of directional hearing. An analytical model of the mechanical response of the ear to a sound stimulus is proposed which supports the claim that mechanical interaural coupling is the key to this animal's ability to localize sound sources. Predicted results for sound fields having a range of incident directions are presented and are found to agree very well with measurements.

Animals↗

Psychoacoustical contribution of each lateral lemniscus.

Although each lateral lemniscus is required for sound localization in its contralateral hemifield, no auditory function is yet known for the neural activity evoked in the lemniscus ipsilateral to a sound source. In an attempt to assess the role played by the ipsilateral lemniscus, monaural cats were tested on an array of psychoacoustical tasks before and after surgical section of one or the other lateral lemniscus. The results show that the lemniscus contralateral to the remaining intact ear is either necessary or sufficient for 24 of the 26 tests administered. However, the lemniscus ipsilateral to the intact ear is both necessary and sufficient (or alternatively, the contralateral lemniscus makes no obvious contribution) to normal thresholds in two of the tasks: detection of low-frequency tones (< 4 kHz) and detection of low-frequency AM modulation. Because of their projections to the ipsilateral inferior colliculus via the ipsilateral lemniscus, the anatomical substrate of these two unusual tasks is probably the fibers from the MSO and possibly, the LSO, ipsilateral to the intact ear.

Acoustic Stimulation↗

Classification of response patterns in cochlear nucleus of barn owl: correlation with functional response properties.

Response patterns of neurons in the cochlear nuclei of the barn owl (Tyto alba) were studied by obtaining poststimulus time histograms (PSTHs) and interspike interval histograms for the response to short tone bursts at the neuron's characteristic frequency. The observed response patterns can be classified according to the scheme developed for neurons of the mammalian cochlear nuclear complex (22). Neurons of the magnocellular cochlear nucleus (n. magnocellularis), which respond in a phase-locked manner to sinusoidal signals and do not show large increases in spike discharge rate with changes in stimulus intensity (26), have "primarylike" (PSTH) discharge patterns and broad interspike interval histograms. This indicates that magnocellular neurons have irregular firing patterns, with the timing of individual spikes being dependent on the phase of the stimulus waveform. Neurons of the angular cochlear nucleus (n. angularis), which show little or no phase-locking and large increases in spike rate with increasing intensity (26), had almost exclusively "transient chopper" discharge patterns. The interspike interval histograms of these angular units are sharp, indicating that their discharge is very regular. At the onset of the response where the chopper pattern is observed, both discharge regularity and rate-intensity sensitivity are at their maximum levels. Several "onset" units were isolated in the angular cochlear nucleus, but no "pauser" or "buildup" units were seen. Also, all of the units in the angular nucleus had monotonic rate-intensity functions. Thus no neural response patterns typical of mammalian dorsal cochlear nucleus units were observed. The relationship of response pattern type to neural function is discussed in relation to the acoustic cues used by the owl for two-dimensional sound localization. The primarylike, phase-locked discharge of magnocellular units is undoubtedly involved in the analysis of interaural differences in stimulus phase, which the owl uses for horizontal localization. There is strong evidence suggesting that the angular nucleus is involved in processing stimulus intensity information, which is important for determining sound elevation (due to asymmetries in vertical directionality of the owl's external ears). The predominant chopper patterns seen in the angular nucleus suggest that in the owl, this response type is correlated with stimulus intensity processing. Similarities in both anatomy and physiology suggest that the magnocellular nucleus is analogous to the spherical cell or bushy cell population of the anterior division of the mammalian anteroventral cochlear nucleus.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Aurally aided visual search under virtual and free-field listening conditions.

We examined the minimum latency required to locate and identify a visual target (visual search) in a two-alternative forced-choice paradigm in which the visual target could appear from any azimuth (0 degree to 360 degrees) and from a broad range of elevations (from 90 degrees above to 70 degrees below the horizon) relative to a person's initial line of gaze. Seven people were tested in six conditions: unaided search, three aurally aided search conditions, and two visually aided search conditions. Aurally aided search with both actual and virtual sound localization cues proved to be superior to unaided and visually guided search. Application of synthesized three-dimensional and two-dimensional sound cues in the workstations are discussed.

Acoustic Stimulation↗

Organization of ventrolateral periolivary cells of the cat superior olive as revealed by PEP-19 immunocytochemistry and Nissl stain.

Ventrolateral periolivary cell groups, through their descending projections to the cochlear nucleus (CN) and local projections to principal nuclei of the superior olive, may participate in brainstem mechanisms mediating such tasks as signal detection in noisy environments and sound localization. Understanding the function of these cell groups can be improved by increased knowledge of the organization of their synaptic inputs in relation to their cellular characteristics. Immunocytochemistry for PEP-19 (a putative calcium binding protein) reveals four patterns of immunolabeling within the ventrolateral periolivary region. Three of the patterns, which have distinct fiber and punctate labeling characteristics, help to define three subdivisions of the lateral nucleus of the trapezoid body (LNTB). The fourth pattern defines two other nuclei, the anterolateral periolivary nucleus (rostral) and the posterior periolivary nucleus (caudal), which display many immunoreactive cell bodies but little fiber and punctate labeling. One of the subdivisions of the LNTB contains large PEP-19 immunolabeled puncta arranged in pericellular nests. Analysis of Nissl-stained sections reveals a neuronal population that resembles globular cells of the ventral cochlear nucleus (VCN) and which colocalizes with pericellular nests of large immunolabeled puncta. Cell counts reveal that roughly 10,000 neurons constitute the cat ventrolateral periolivary region, 9,000 of which are found in the LNTB. Three-dimensional reconstructions of auditory brainstem nuclei clarify the complex spatial relationships among these structures.

Animals↗

Frequency dependence of directional amplification at the cat's pinna.

We examined in detail the effects of changing stimulus frequency upon the inclination of the acoustical axis of the pinna and upon the solid angle (area) subtended by isoamplification contours. We measured the relative sound pressure level difference between points on a 1 m radius, coordinate sphere using the cochlear microphonic as an indicator of tympanic sound pressure. The inclination of the acoustical axis for a given frequency was found to vary with the posture of the pinna, and with the pinna in a drooped position (following midline incision) there was a frequency spreading of axial positions such that high frequency axes were inclined progressively more laterally. However, with the pinna in an upright posture the axes for all frequencies tested were relatively tightly clustered. Alternative models for sound localization can be formulated to suit either situation, but it seems likely that the cat can use the frequency spreading effect of its pinna sound transformation as a cue to location. The pinna becomes more directional at higher frequencies, and this is clearly shown when the solid angle of isoamplification contours is plotted against frequency. The inverse relationship formed was shown to be closely matched by a model based upon diffraction by the outer dimension of the pinna.

Acoustics↗

Long-latency auditory evoked potentials in humans and the localization of a sound image.

In the article, we discuss data from an investigation concerning how boundary conditions for the creation of sound-image movement are reflected long-latency auditory evoked potentials and discuss how and important associated with the human localizing function (resistance to interference during the local localization of both a stationary and a moving sound image) appears in long-latency auditory evoked potentials. We establish that a change in the parameters of a signal creating a sensation of sound-image movement results in an exhaltation of t he amplitudes of the N1 and P2 components. The effect of binaural freedom from masking is reflected in these same components of long-latency auditory evoked potentials during movement of spatially shifting signals.

Acoustic Stimulation↗

The influence of underwater data transmission sounds on the displacement behaviour of captive harbour seals (Phoca vitulina).

To prevent grounding of ships and collisions between ships in shallow coastal waters, an underwater data collection and communication network (ACME) using underwater sounds to encode and transmit data is currently under development. Marine mammals might be affected by ACME sounds since they may use sound of a similar frequency (around 12 kHz) for communication, orientation, and prey location. If marine mammals tend to avoid the vicinity of the acoustic transmitters, they may be kept away from ecologically important areas by ACME sounds. One marine mammal species that may be affected in the North Sea is the harbour seal (Phoca vitulina). No information is available on the effects of ACME-like sounds on harbour seals, so this study was carried out as part of an environmental impact assessment program. Nine captive harbour seals were subjected to four sound types, three of which may be used in the underwater acoustic data communication network. The effect of each sound was judged by comparing the animals' location in a pool during test periods to that during baseline periods, during which no sound was produced. Each of the four sounds could be made into a deterrent by increasing its amplitude. The seals reacted by swimming away from the sound source. The sound pressure level (SPL) at the acoustic discomfort threshold was established for each of the four sounds. The acoustic discomfort threshold is defined as the boundary between the areas that the animals generally occupied during the transmission of the sounds and the areas that they generally did not enter during transmission. The SPLs at the acoustic discomfort thresholds were similar for each of the sounds (107 dB re 1 microPa). Based on this discomfort threshold SPL, discomfort zones at sea for several source levels (130-180 dB re 1 microPa) of the sounds were calculated, using a guideline sound propagation model for shallow water. The discomfort zone is defined as the area around a sound source that harbour seals are expected to avoid. The definition of the discomfort zone is based on behavioural discomfort, and does not necessarily coincide with the physical discomfort zone. Based on these results, source levels can be selected that have an acceptable effect on harbour seals in particular areas. The discomfort zone of a communication sound depends on the sound, the source level, and the propagation characteristics of the area in which the sound system is operational. The source level of the communication system should be adapted to each area (taking into account the width of a sea arm, the local sound propagation, and the importance of an area to the affected species). The discomfort zone should not coincide with ecologically important areas (for instance resting, breeding, suckling, and feeding areas), or routes between these areas.

Acoustics↗

Projections of the cochlear nuclei and nucleus laminaris to the inferior colliculus of the barn owl.

The barn owl determines the directions from which sounds emanate by computing the interaural differences in the timing and intensity of sounds. These cues for sound localization are processed in independent channels originating at nucleus magnocellularis (NM) and nucleus angularis (NA), the cochlear nuclei. The cells of NM are specialized for encoding the phase of sounds in the ipsilateral ear. The cells of NA are specialized for encoding the intensity of sounds in the ipsilateral ear. NM projects solely, bilaterally, and tonotopically to nucleus laminaris (NL). NL and NA project to largely nonoverlapping zones in the central nucleus of the inferior colliculus (ICc), thus forming hodological subdivisions in which time and intensity information may be processed. The terminal field of NL occupies a discrete zone in the rostromedial portion of the contralateral ICc, which we have termed the "core" of ICc. The terminal field of NA surrounds the core of ICc and thus forms a "shell" around it. The projection from NL to the core conserves tonotopy. Low-frequency regions of NL project to the dorsal portions of the core whereas higher-frequency regions project to more ventral portions. This innervation pattern is consistent with earlier physiological studies of tonotopy. Physiological studies have also suggested that NL and the core of ICs contain a representation of the location of a sound source along the horizontal axis. Our data suggest that the projection from NL to the core preserves spatiotopy. Thus, the dorsal portion of NL on the left, which contains a representation of eccentric loci in the right hemifield, innervates the area of the right ICc core that represents eccentric right loci. The more ventral portion of the left NL, which represents loci close to the vertical meridian, innervates the more rostral portions of the right core, which also represents loci near the vertical meridian.

Afferent Pathways↗

Disruption of auditory spatial working memory by inactivation of the forebrain archistriatum in barn owls.

Barn owls not only localize auditory stimuli with great accuracy, they also remember the locations of auditory stimuli and can use this remembered spatial information to guide their flight and strike. Although the mechanisms of sound localization have been studied extensively, the neurobiological basis of auditory spatial memory has not. Here we show that the ability of barn owls to orient their gaze towards and fly to the remembered location of auditory targets is lost during pharmacological inactivation of a small region in the forebrain, the anterior archistriatum. In contrast, archistriatal inactivation has no effect on stimulus-guided responses to auditory targets. The memory-dependent deficit is evident only for acoustic events that occur in the hemifield contralateral to the side that is inactivated. The data demonstrate that in the avian archistriatum, as in the mammalian frontal cortex, there exists a region that is essential for the expression of spatial working memory and that, in the barn owl, this region encodes auditory spatial memory.

Animals↗

Comparison of normal and impaired hearing. I. Loudness, localization.

Impaired hearing is characterized by high thresholds and reduced loudness. Loudness, however, may quickly recover as it increases rapidly from an elevated threshold. This rapid growth, known as loudness recruitment, is a sign of cochlear impairment and is generally not seen in conductive or retrocochlear impairment. Loudness recruitment means that the hard-of-hearing person detects small changes in intensity near his elevated threshold but he probably does no better than a normal listener at the same SPLs. Recruitment is often accompanied by reduced loudness summation, which means that the loudness of a band of noise does not increase as much with increasing bandwidth as in normal hearing. This reduced summation of loudness is probably why the cochlearly impaired ear has nearly the same threshold for the acoustic reflex to pure tones as to wide-band noise, whereas the normal ear has a much lower threshold to wide-band noise. Corresponding differences between normal and impaired hearing are not found in auditory localization. Rather, the evidence suggests that persons with residual hearing learn to localize sounds reasonably well. Even the inability of many hearing impaired persons to understand a speaker in a noisy environment may result more from a failure of frequency analysis rather than of localization.

Auditory Threshold↗

Sound delay lines in the nucleus laminaris of the chicken.

Delays of neurophonic potentials (NP) induced by monaural sound stimuli were measured across the three dimensions in the nucleus laminaris (NL) of the anesthetized chicken. Peak latencies and delays in cross-correlograms changed with recording distance. An orderly delay line was observed across the NL thickness, that is, along dendritic trees of individual fusiform cells (FC), where phase lags increased dorso-ventrally during ipsi- and in the opposite direction during contralateral stimuli. Delays along isofrequency FC arrays were variable, with delay ranges being smaller for ipsilateral than for contralateral sound stimuli. Net delays for contralateral sounds were directed medio-laterally and differences between ipsi- and contralateral delays covered, roughly, intercochlear time differences (ITD). The observed delays are thought to contribute to sound localization and frequency analysis.

Animals↗

Proposed mechanisms for coincidence detection in the auditory brainstem.

Sound localization in mammals uses two distinct neural circuits, one for low- and one for high-frequency bands. Recent experiments call for revision of the theory explaining how the direction of incoming sound is calculated. We propose such a revised theory. Our theory is based on probabilistic spiking and probabilistic delay of spikes from both sides. We have applied the mechanism originally proposed as an operation on spike trains resulting in multiplication of firing rates. We have adapted this mechanism for the case of synchronous spike trains. The mechanism has to detect spikes from both sides within a short time window. Therefore, in both circuits neurons act as coincidence detectors. In the excitatory low-frequency circuit we call the mechanism the excitatory coincidence detection, to distinguish it from the mechanism of the inhibitory coincidence detection in the high-frequency circuit. The times to first spike and gains of the two mechanisms are calculated. We show how the output gains of the mechanisms predict the dip within the human frequency sensitivity range. This dip has been described in human psychophysical experiments.

Acoustic Stimulation↗

Localisation of human speech by the newborn baby and the effects of pethidine ('meperidine').

Eight newborn babies were tested to see whether they could make discriminating headturn responses to the sound of a 9-second recording of a human voice coming from 15 degrees, 30 degrees and 80 degrees from the midline, from either the right or left side. The babies showed significant differences in response to sounds coming from the right and the left, and to sounds coming from 15 degrees and 80 degrees from the midline. Pethidine ('Meperidine') given to the mother during labour had a significant effect, both on degree of response and on the babies' apparent ability to localize sound.

Anesthesia, Obstetrical↗

Developmental plasticity and memory.

The cerebral cortex of young kittens is known to be highly malleable during early postnatal development. However, most studies of developmental plasticity have been conducted in primary visual cortex. It has long been unclear to what extent similar plasticity exists in higher cortical areas. We have now studied developmental plasticity in the anterior ectosylvian (AE) region of the cat's parietal association cortex, which receives input from different sensory modalities. One area in this cortical region, which is predominantly visual in normal cats, area AEV, is taken over almost completely by auditory and somatosensory inputs, when cats are binocularly deprived of vision from birth. Furthermore, when single auditory neurons are tested with sound sources in free-field at different locations, they show sharper spatial tuning in visually deprived cats. This compensatory, crossmodal plasticity was explored at the behavioral level by testing visually deprived cats in an auditory localization task, and these cats could indeed localize sound sources more precisely than normal cats. These findings are interpreted as a form of adaptation of the young brain to an altered environment. Similar adaptation is still possible in adult brains by virtue of associative learning and long-term memory. It is argued that the synaptic mechanisms by which associative memories are stored in the cerebral cortex are similar to those in developmental plasticity, only the increment of learning is smaller in adult animals.

Aging↗