Maps of interaural delay in the owl's nucleus laminaris
Catherine Emily Carr2015, Journal of Neurophysiology
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Abstract
Axons from the nucleus magnocellularis form a presynaptic map of interaural time differences (ITDs) in the nucleus laminaris (NL). These inputs generate a field potential that varies systematically with recording position and can be used to measure the map of ITDs. In the barn owl, the representation of best ITD shifts with mediolateral position in NL, so as to form continuous, smoothly overlapping maps of ITD with iso-ITD contours that are not parallel to the NL border. Frontal space (0°) is, however, represented throughout and thus overrepresented with respect to the periphery. Measurements of presynaptic conduction delay, combined with a model of delay line conduction velocity, reveal that conduction delays can account for the mediolateral shifts in the map of ITD.
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Interaural time difference is an important cue for sound localization. In the barn owl (Tyto alba) neuronal sensitivity to this disparity originates in the brainstem nucleus laminaris. Afferents from the ipsilateral and contralateral magnocellular cochlear nuclei enter the nucleus laminaris through its dorsal and ventral surfaces, respectively, and interdigitate in the nucleus. Intracellular recordings from these afferents show orderly changes in conduction delay with depth in the nucleus. These changes are comparable to the range of interaural time differences available to the owl. Thus, these afferent axons act as delay lines and provide anatomical and physiological bases for a neuronal map of interaural time differences in the nucleus laminaris.
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Journal of neurophysiology, 2014
Inputs from the two sides of the brain interact to create maps of interaural time difference (ITD) in the nucleus laminaris of birds. How inputs from each side are matched with high temporal precision in ITD-sensitive circuits is unknown, given the differences in input path lengths from each side. To understand this problem in birds, we modeled the geometry of the input axons and their corresponding conduction velocities and latencies. Consistent with existing physiological data, we assumed a common latency up to the border of nucleus laminaris. We analyzed two biological implementations of the model, the single ITD map in chickens and the multiple maps of ITD in barn owls. For binaural inputs, since ipsi- and contralateral initial common latencies were very similar, we could restrict adaptive regulation of conduction velocity to within the nucleus. Other model applications include the simultaneous derivation of multiple conduction velocities from one set of measurements and the dem...
Delay Line Models of Sound Localization in the Barn Owl
American Zoologist, 1993
SYNOPSIS. The detection of interaural time differences underlies azimuthal sound localization in the barn owl. Sensitivity to these time differences arises in the brainstem nucleus laminaris. Auditory information reaches the nucleus laminaris via bilateral projections from the cochlear nucleus magnocellularis. The magnocellular inputs to the nucleus laminaris act as delay lines to create maps of interaural time differences. These delay lines are tapped by postsynaptic coincidence detectors that encode interaural time differences. The entire circuit, from the auditory nerve to the nucleus magnocellularis to the nucleus laminaris, is specialized for the encoding and preservation of temporal information. A mathematical model of this circuit (Grun et al, 1990) provides useful predictions.
Functional delay of myelination of auditory delay lines in the nucleus laminaris of the barn owl
Developmental Neurobiology, 2007
In the barn owl, maps of interaural time difference (ITD) are created in the nucleus laminaris (NL) by interdigitating axons that act as delay lines. Adult delay line axons are myelinated, and this myelination is timely, coinciding with the attainment of adult head size, and stable ITD cues. The proximal portions of the axons become myelinated in late embryonic life, but the delay line portions of the axon in NL remain unmyelinated until the first postnatal week. Myelination of the delay lines peaks at the third week posthatch, and myelinating oligodendrocyte density approaches adult levels by one month, when the head reaches its adult width. Migration of oligodendrocyte progenitors into NL and the subsequent onset of myelination may be restricted by a glial barrier in late embryonic stages and the first posthatch week, since the loss of tenascin-C immunoreactivity in NL is correlated with oligodendrocyte progenitor migration into NL.
Sensitivity to spectral interaural intensity difference cues in space-specific neurons of the barn owl
Journal of Comparative Physiology A: Sensory, Neural, and Behavioral Physiology, 2004
Barn owls use interaural intensity differences to localize sounds in the vertical plane. At a given elevation the magnitude of the interaural intensity difference cue varies with frequency, creating an interaural intensity difference spectrum of cues which is characteristic of that direction. To test whether space-specific cells are sensitive to spectral interaural intensity difference cues, pure-tone interaural intensity difference tuning curves were taken at multiple different frequencies for single neurons in the external nucleus of the inferior colliculus. For a given neuron, the interaural intensity differences eliciting the maximum response (the best interaural intensity differences) changed with the frequency of the stimulus by an average maximal difference of 9.4±6.2 dB. The resulting spectral patterns of these neurally preferred interaural intensity differences exhibited a high degree of similarity to the acoustic interaural intensity difference spectra characteristic of restricted regions in space. Compared to stimuli whose interaural intensity difference spectra matched the preferred spectra, stimuli with inverted spectra elicited a smaller response, showing that space-specific neurons are sensitive to the shape of the spectrum. The underlying mechanism is an inhibition for frequency-specific interaural intensity differences which differ from the preferred spectral pattern. Collectively, these data show that space-specific neurons are sensitive to spectral interaural intensity difference cues and support the idea that behaving barn owls use such cues to precisely localize sounds. Keywords Binaural AE Interaural level difference AE Pattern recognition AE Spectral integration AE Template matching Abbreviations ABI average binaural intensity AE HRTF head-related transfer function AE ICx external nucleus of the inferior colliculus AE IID interaural intensity difference AE ITD interaural time difference AE OT optic tectum AE RMS root mean square AE VLVp nucleus ventralis lemnisci laterale, pars posterior
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