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Pathology: whales, sonar and decompression sickness.

We do not yet know why whales occasionally strand after sonar has been deployed nearby, but such information is important for both naval undersea activities and the protection of marine mammals. Jepson et al. suggest that a peculiar gas-forming disease afflicting some stranded cetaceans could be a type of decompression sickness (DCS) resulting from exposure to mid-range sonar. However, neither decompression theory nor observation support the existence of a naturally occurring DCS in whales that is characterized by encapsulated, gas-filled cavities in the liver. Although gas-bubble formation may be aggravated by acoustic energy, more rigorous investigation is needed before sonar can be firmly linked to bubble formation in whales.

Acoustic Stimulation↗

Synaptic computation.

Neurons are often considered to be the computational engines of the brain, with synapses acting solely as conveyers of information. But the diverse types of synaptic plasticity and the range of timescales over which they operate suggest that synapses have a more active role in information processing. Long-term changes in the transmission properties of synapses provide a physiological substrate for learning and memory, whereas short-term changes support a variety of computations. By expressing several forms of synaptic plasticity, a single neuron can convey an array of different signals to the neural circuit in which it operates.

Animals↗

Top-down gain control of the auditory space map by gaze control circuitry in the barn owl.

High-level circuits in the brain that control the direction of gaze are intimately linked with the control of visual spatial attention. Immediately before an animal directs its gaze towards a stimulus, both psychophysical sensitivity to that visual stimulus and the responsiveness of high-order neurons in the cerebral cortex that represent the stimulus increase dramatically. Equivalent effects on behavioural sensitivity and neuronal responsiveness to visual stimuli result from focal electrical microstimulation of gaze control centres in monkeys. Whether the gaze control system modulates neuronal responsiveness in sensory modalities other than vision is unknown. Here we show that electrical microstimulation applied to gaze control circuitry in the forebrain of barn owls regulates the gain of midbrain auditory responses in an attention-like manner. When the forebrain circuit was activated, midbrain responses to auditory stimuli at the location encoded by the forebrain site were enhanced and spatial selectivity was sharpened. The same stimulation suppressed responses to auditory stimuli represented at other locations in the midbrain map. Such space-specific, top-down regulation of auditory responses by gaze control circuitry in the barn owl suggests that the central nervous system uses a common strategy for dynamically regulating sensory gain that applies across modalities, brain areas and classes of vertebrate species. This approach provides a path for discovering mechanisms that underlie top-down gain control in the central nervous system.

Acoustic Stimulation↗

Axonal site of spike initiation enhances auditory coincidence detection.

Neurons initiate spikes in the axon initial segment or at the first node in the axon. However, it is not yet understood how the site of spike initiation affects neuronal activity and function. In nucleus laminaris of birds, neurons behave as coincidence detectors for sound source localization and encode interaural time differences (ITDs) separately at each characteristic frequency (CF). Here we show, in nucleus laminaris of the chick, that the site of spike initiation in the axon is arranged at a distance from the soma, so as to achieve the highest ITD sensitivity at each CF. Na+ channels were not found in the soma of high-CF (2.5-3.3 kHz) and middle-CF (1.0-2.5 kHz) neurons but were clustered within a short segment of the axon separated by 20-50 microm from the soma; in low-CF (0.4-1.0 kHz) neurons they were clustered in a longer stretch of the axon closer to the soma. Thus, neurons initiate spikes at a more remote site as the CF of neurons increases. Consequently, the somatic amplitudes of both orthodromic and antidromic spikes were small in high-CF and middle-CF neurons and were large in low-CF neurons. Computer simulation showed that the geometry of the initiation site was optimized to reduce the threshold of spike generation and to increase the ITD sensitivity at each CF. Especially in high-CF neurons, a distant localization of the spike initiation site improved the ITD sensitivity because of electrical isolation of the initiation site from the soma and dendrites, and because of reduction of Na+-channel inactivation by attenuating the temporal summation of synaptic potentials through the low-pass filtering along the axon.

Action Potentials↗

Dynamic shifts in the owl's auditory space map predict moving sound location.

The optic tectum of the barn owl contains a map of auditory space. We found that, in response to moving sounds, the locations of receptive fields that make up the map shifted toward the approaching sound. The magnitude of the receptive field shifts increased systematically with increasing stimulus velocity and, therefore, was appropriate to compensate for sensory and motor delays inherent to auditory orienting behavior. Thus, the auditory space map is not static, but shifts adaptively and dynamically in response to stimulus motion. We provide a computational model to account for these results. Because the model derives predictive responses from processes that are known to occur commonly in neural networks, we hypothesize that analogous predictive responses will be found to exist widely in the central nervous system. This hypothesis is consistent with perceptions of stimulus motion in humans for many sensory parameters.

Adaptation, Physiological↗

Passive eye displacement alters auditory spatial receptive fields of cat superior colliculus neurons.

The superior colliculus (SC) is thought to use a set of superimposed, topographically organized neural maps of visual, auditory, somatosensory and motor space to direct the eyes toward novel stimuli. Auditory spatial response fields (SRFs) of SC neurons may change when an animal moves its eyes, presumably to compensate for the resulting misalignment of visual and auditory sensory spatial reference frames, but the mechanisms responsible for these SRF changes remain unknown. Here we report that passive deviation of the eye in anesthetized, paralyzed animals can profoundly affect the auditory responsiveness of SC neurons, but seems insufficient by itself to provide adaptive shifts of auditory SRFs.

Acoustic Stimulation↗

Behavioral auditory assessment of young infants: methodological limitations or natural lack of auditory responsiveness?

The determination of auditory thresholds by means of behavioral techniques in young infants can be difficult. This could be the result of limitations in methodology, a lack of observable auditory responsiveness, or both. In the current study, 2- and 4-month old infants were tested under enhanced conditions for obtaining behavioral responses (i.e., salient auditory stimuli, reduced visual distractions, reinforced correct responses). A two-interval, forced-choice task with four intensity levels was used. Although a behavioral threshold was obtained for the 4-month-olds, threshold determination for the 2-month-olds remained elusive. In light of the current findings and previous studies of visual acuity of infants, these results suggest a lack of behavioral responsiveness to auditory stimuli for the younger infants rather than methodological limitations. With infants in the 2-month-old age range, clinical audiologists should expect few behavioral responses to auditory stimuli at intensity levels below those that elicit startle responses.

Attention↗

Audiological assessment, rehabilitation, and spatial hearing considerations associated with visual impairment in adults: an overview.

The purpose of this paper is to increase awareness and interest among audiologists regarding the needs of adults who have both hearing loss and visual impairment, particularly people who are blind and travel independently. Case history, audiometric testing, and rehabilitation issues are considered, and extensive discussion of spatial hearing research as it relates to orientation and mobility is provided.

Adult↗

Motion processing in the auditory cortex of the rufous horseshoe bat: role of GABAergic inhibition.

This study examined the influence of inhibition on motion-direction-sensitive responses of neurons in the dorsal fields of auditory cortex of the rufous horseshoe bat. Responses to auditory apparent motion stimuli were recorded extracellularly from neurons while microiontophoretically applying gamma-aminobutyric acid (GABA) and the GABAA receptor antagonist bicuculline methiodide (BMI). Neurons could respond with a directional preference exhibiting stronger responses to one direction of motion or a shift of receptive field (RF) borders depending on direction of motion. BMI influenced the motion direction sensitivity of 53% of neurons. In 21% of neurons the motion-direction sensitivity was decreased by BMI by decreasing either directional preference or RF shift. In neurons with a directional preference, BMI increased the spike number for the preferred direction by a similar amount as for the nonpreferred direction. Thus, inhibition was not direction specific. BMI increased motion-direction sensitivity by either increasing directional preference or magnitude of RF shifts in 22% of neurons. Ten percent of neurons changed their response from a RF shift to a directional preference under BMI. In these neurons, the observed effects could often be better explained by adaptation of excitation rather than inhibition. The results suggest, that adaptation of excitation, as well as cortex specific GABAergic inhibition, contribute to motion-direction sensitivity in the auditory cortex of the rufous horseshoe bat.

Acoustic Stimulation↗

Spectral envelope coding in cat primary auditory cortex: linear and non-linear effects of stimulus characteristics.

Electrophysiological studies in mammal primary auditory cortex have demonstrated neuronal tuning and cortical spatial organization based upon spectral and temporal qualities of the stimulus including: its frequency, intensity, amplitude modulation and frequency modulation. Although communication and other behaviourally relevant sounds are usually complex, most response characterizations have used tonal stimuli. To better understand the mechanisms necessary to process complex sounds, we investigated neuronal responses to a specific class of broadband stimuli, auditory gratings or ripple stimuli, and compared the responses with single tone responses. Ripple stimuli consisted of 150-200 frequency components with the intensity of each component adjusted such that the envelope of the frequency spectrum is sinusoidal. It has been demonstrated that neurons are tuned to specific characteristics of those ripple stimulus including the intensity, the spacing of the peaks, and the location of the peaks and valleys (C. E. Schreiner and B. M. Calhoun, Auditory Neurosci., 1994; 1: 39-61). Although previous results showed that neuronal response strength varied with the intensity and the fundamental frequency of the stimulus, it is shown here that the relative response to different ripple spacings remains essentially constant with changes in the intensity and the fundamental frequency. These findings support a close relationship between pure-tone receptive fields and ripple transfer functions. However, variations of other stimulus characteristics, such as spectral modulation depth, result in non-linear alterations in the ripple transformation. The processing between the basilar membrane and the primary auditory cortex of broadband stimuli appears generally to be non-linear, although specific stimulus qualities, including the phase of the spectral envelope, are processed in a nearly linear manner.

Acoustic Stimulation↗

Effect of GABA on the processing of interaural time differences in nucleus laminaris neurons in the chick.

Neurons in the avian nucleus laminaris (NL) are the first to receive binaural information and are presumed to play a role in encoding interaural time differences (ITD). NL not only receives excitatory projections from the ipsi- and contralateral nucleus magnocellularis, but also receives inhibitory (GABAergic) input. This study investigates how GABA (gamma-aminobutyric acid) influences ITD coding in NL. Intracellular responses of chick NL neurons were studied in a brain slice preparation. Both excitatory inputs to NL were electrically activated and the delay between trains of bilateral stimuli (simulated-interaural time difference [s-ITD]) was varied. The resulting s-ITD functions were recorded in the presence of 0-75 microM GABA. The discharge rate of NL neurons varied with s-ITD. Cells responded maximally using s-ITDs at which the peak of the ipsi- and contralateral excitatory postsynaptic potentials occurred simultaneously (favourable s-ITD). At unfavourable s-ITDs, the discharge rates usually fell below unilateral levels. GABA had contrary effects on the s-ITD functions depending on the drug concentration. A low GABA dose enhanced excitability at favourable s-ITD, but not at unfavourable s-ITDs. In contrast, higher GABA concentrations diminished excitability. Moderate GABA concentrations had no consistent effect. These results suggest that the GABAergic input to NL will either increase or decrease the excitability of the NL neuron depending on the degree to which this GABAergic input is activated. A gain control hypothesis is presented in which the GABAergic input makes ITD processing in NL independent of the stimulus intensity by adjusting the excitability of NL neurons.

Animals↗

Temporal integration vs. parallel processing: coping with the variability of neuronal messages in directional hearing of insects.

The behaviour of many animals indicates a high precision in the processing of sensory signals, which often seems at odds with the large variability of individual neuronal responses. Using the directional hearing of the grasshopper Chorthippus biguttulus (Ch. biguttulus) as a model system, we investigated the possible contributions of temporal integration and parallel processing at the receptor level to the observed behavioural acuity. The precision of the animals' phonotactic orientation behaviour to stimuli of different durations was measured and compared with the spiking responses of auditory receptor cells, leading to the following results. A behavioural decision based on integrating the spikes from a pair of receptors (one cell in each ear) over a 1000-ms time window would substantially decrease the error probability, compared with evaluating a 250-ms period. The animal as a whole, however, responded as precisely to a 250 ms stimulus as to a stimulus of natural duration (c. 1000 ms). A phonotactic decision based on a 1.5 dB intensity difference between the ears corresponds to a spike count difference of approximately 1 spike per 100 ms in a pair of receptors. As these differences are in the order of the statistical spike count fluctuations, the error probabilities for an ideal observer of the spike count are substantially higher (13-18%) than the errors observed in the behavioural tests (only 5%). We conclude that the animals' nervous system has to sample information from a pool of 6-13 receptors to arrive at the observed behavioural precision.

Action Potentials↗