Plasticity in the development of afferent patterns in the inferior colliculus of the rat after unilateral cochlear ablation.
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The central nucleus of the inferior colliculus (IC) is the site of convergence for nearly all ascending monaural and binaural projections. Several of these inputs, including inhibitory connections from the dorsal nucleus of the lateral lemniscus (DNLL), are highly ordered and organized into series of afferent bands or patches. Although inputs to the IC from the contralateral DNLL are present in the rat by birth [postnatal day 0 (P0)], the earliest indications of band formation are not evident until P4. Subsequently, the initially diffuse projection segregates into a pattern of bands and interband spaces, and by P12 adult-like, afferent-dense patches are established (Gabriele et al., 2000). To determine the role of the auditory periphery in the development of bands and patches before the onset of hearing (P12/P13), unilateral cochlear ablations were performed at P2 (before any evidence of banding). Rat pups were reared to P12, at which time glass pins coated with 1, 1'-dioctodecyl-3,3,3',3'-tetramethylindocarbocyanine perchlorate were placed in fixed tissue in the commissure of Probst where DNLL fibers cross the midline. The results indicate that a unilateral cochlear ablation disrupts the normal development of afferent patches in the IC. Although the crossed DNLL projections labeled via commissural dye placement always mirrored each other in P12 controls, ablation cases exhibited a consistent, bilateral asymmetry in pattern formation and relative density of the labeled projections. Possible developmental mechanisms likely to be involved in the establishment of afferent bands and patches before the onset of hearing are discussed. (+info)
Detection of synchrony in the activity of auditory nerve fibers by octopus cells of the mammalian cochlear nucleus.
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The anatomical and biophysical specializations of octopus cells allow them to detect the coincident firing of groups of auditory nerve fibers and to convey the precise timing of that coincidence to their targets. Octopus cells occupy a sharply defined region of the most caudal and dorsal part of the mammalian ventral cochlear nucleus. The dendrites of octopus cells cross the bundle of auditory nerve fibers just proximal to where the fibers leave the ventral and enter the dorsal cochlear nucleus, each octopus cell spanning about one-third of the tonotopic array. Octopus cells are excited by auditory nerve fibers through the activation of rapid, calcium-permeable, alpha-amino-3-hydroxy-5-methyl-4-isoxazole-propionate receptors. Synaptic responses are shaped by the unusual biophysical characteristics of octopus cells. Octopus cells have very low input resistances (about 7 M Omega), and short time constants (about 200 microsec) as a consequence of the activation at rest of a hyperpolarization-activated mixed-cation conductance and a low-threshold, depolarization-activated potassium conductance. The low input resistance causes rapid synaptic currents to generate rapid and small synaptic potentials. Summation of small synaptic potentials from many fibers is required to bring an octopus cell to threshold. Not only does the low input resistance make individual excitatory postsynaptic potentials brief so that they must be generated within 1 msec to sum but also the voltage-sensitive conductances of octopus cells prevent firing if the activation of auditory nerve inputs is not sufficiently synchronous and depolarization is not sufficiently rapid. In vivo in cats, octopus cells can fire rapidly and respond with exceptionally well-timed action potentials to periodic, broadband sounds such as clicks. Thus both the anatomical specializations and the biophysical specializations make octopus cells detectors of the coincident firing of their auditory nerve fiber inputs. (+info)
Linear and nonlinear pathways of spectral information transmission in the cochlear nucleus.
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At the level of the cochlear nucleus (CN), the auditory pathway divides into several parallel circuits, each of which provides a different representation of the acoustic signal. Here, the representation of the power spectrum of an acoustic signal is analyzed for two CN principal cells-chopper neurons of the ventral CN and type IV neurons of the dorsal CN. The analysis is based on a weighting function model that relates the discharge rate of a neuron to first- and second-order transformations of the power spectrum. In chopper neurons, the transformation of spectral level into rate is a linear (i.e., first-order) or nearly linear function. This transformation is a predominantly excitatory process involving multiple frequency components, centered in a narrow frequency range about best frequency, that usually are processed independently of each other. In contrast, type IV neurons encode spectral information linearly only near threshold. At higher stimulus levels, these neurons are strongly inhibited by spectral notches, a behavior that cannot be explained by level transformations of first- or second-order. Type IV weighting functions reveal complex excitatory and inhibitory interactions that involve frequency components spanning a wider range than that seen in choppers. These findings suggest that chopper and type IV neurons form parallel pathways of spectral information transmission that are governed by two different mechanisms. Although choppers use a predominantly linear mechanism to transmit tonotopic representations of spectra, type IV neurons use highly nonlinear processes to signal the presence of wide-band spectral features. (+info)
Ionic currents and spontaneous firing in neurons isolated from the cerebellar nuclei.
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Neurons of the cerebellar nuclei fire spontaneous action potentials both in vitro, with synaptic transmission blocked, and in vivo, in resting animals, despite ongoing inhibition from spontaneously active Purkinje neurons. We have studied the intrinsic currents of cerebellar nuclear neurons isolated from the mouse, with an interest in understanding how these currents generate spontaneous activity in the absence of synaptic input as well as how they allow firing to continue during basal levels of inhibition. Current-clamped isolated neurons fired regularly ( approximately 20 Hz), with shallow interspike hyperpolarizations (approximately -60 mV), much like neurons in more intact preparations. The spontaneous firing frequency lay in the middle of the dynamic range of the neurons and could be modulated up or down with small current injections. During step or action potential waveform voltage-clamp commands, the primary current active at interspike potentials was a tetrodotoxin-insensitive (TTX), cesium-insensitive, voltage-independent, cationic flux carried mainly by sodium ions. Although small, this cation current could depolarize neurons above threshold voltages. Voltage- and current-clamp recordings suggested a high level of inactivation of the TTX-sensitive transient sodium currents that supported action potentials. Blocking calcium currents terminated firing by preventing repolarization to normal interspike potentials, suggesting a significant role for K(Ca) currents. Potassium currents that flowed during action potential waveform voltage commands had high activation thresholds and were sensitive to 1 mm TEA. We propose that, after the decay of high-threshold potassium currents, the tonic cation current contributes strongly to the depolarization of neurons above threshold, thus maintaining the cycle of firing. (+info)
Expression of the Kv3.1 potassium channel in the avian auditory brainstem.
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The Shaw-like potassium channel Kv3.1, a delayed rectifier with a high threshold of activation, is expressed in the time coding nuclei of the bird auditory brainstem. In both barn owls and chickens, Kv3.1 mRNA was expressed in the cochlear nucleus magnocellularis (NM) and the nucleus laminaris (NL). Western blot analysis showed that an antibody raised against the synthetic peptide sequence of rat Kv3.1 (rKv3.1) specifically recognized the same 92 kDa protein bands in both rat and chicken synaptosomal preparations. Immunohistochemical analyses using this anti-rKv3.1 antibody revealed a prominent gradient in Kv3.1 immunoreactivity along the tonotopic axis of the barn owl NM and NL and a less prominent gradient in the chicken. The precise localization of the Kv3.1 immunoproduct was resolved by electron microscopy. In both the owl and the chicken, Kv3.1 was targeted postsynaptically in NM and NL. The major difference in localization of Kv3.1 protein between the two birds was the expression of Kv3.1 in the NM axons and terminals in the region of the barn owl NL. This location of Kv3.1 channels supports its postulated function in reducing the width of action potentials as they invade the presynaptic terminal. The presynaptic localization may be a specialization for enabling neurons in owl NM to transmit high-frequency temporal information with little jitter. (+info)
A physiologically based model of discharge pattern regulation by transient K+ currents in cochlear nucleus pyramidal cells.
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Pyramidal cells in the dorsal cochlear nucleus (DCN) show three characteristic discharge patterns in response tones: pauser, buildup, and regular firing. Experimental evidence suggests that a rapidly inactivating K+ current (I(KIF)) plays a critical role in generating these discharge patterns. To explore the role of I(KIF), we used a computational model based on the biophysical data. The model replicated the dependence of the discharge pattern on the magnitude and duration of hyperpolarizing prepulses, and I(KIF) was necessary to convey this dependence. Phase-plane and perturbation analyses show that responses to depolarization are critically controlled by the amount of inactivation of I(KIF). Experimentally, half-inactivation voltage and kinetics of I(KIF) show wide variability. Varying these parameters in the model revealed that half-inactivation voltage, and activation and inactivation rates, controls the voltage and time dependence of the model cell discharge. This suggests that pyramidal cells can adjust their sensitivity to different temporal patterns of inhibition and excitation by modulating the kinetics of I(KIF). Overall, I(KIF) is a critical conductance controlling the excitability of DCN pyramidal cells. (+info)
Temporal representation of iterated rippled noise as a function of delay and sound level in the ventral cochlear nucleus.
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The discharge patterns of single units in the ventral cochlear nucleus (VCN) of anesthetized guinea pigs were examined in response to iterated rippled noise (IRN) as a function of the IRN delay (which determines the IRN pitch) and the IRN sound level. Delays were varied over five octaves in half-octave steps, and sound levels were varied over a 30- or 50-dB range in steps of 5 dB. Neural responses were analyzed in terms of first-order and all-order inter-spike intervals (ISIs). The IRN quasi-periodicity was preserved in the all-order ISIs for most units independent of unit type or best frequency (BF). A deterioration of the temporal all-order code was found, however, when the neural response was influenced by inhibition. The IRN quasi-periodicity was also preserved in first-order ISIs for a limited range of IRN delays and levels. Sustained Chopper units (CS) in the VCN responded with very regular ISIs when the IRN delay corresponded to the unit's chopping period; i.e., the unit showed an increased proportion of intervals corresponding to the IRN delay (interval enhancement) relative to an equal-level, white-noise stimulation. This interval enhancement has a band-pass characteristic with a peak corresponding to the chopping period. Moreover, for CS units in rate saturation, the chopping period, and thus the interval enhancement to the IRN, did not vary with level. Units classified as onset-chopper also show a band-pass interval enhancement to the IRN stimuli; however, they show more level-dependent changes than CS units. Primary-like (PL) units also show level-dependent changes in their ability to code the IRN pitch in first-order intervals. The range of delays where PL units showed interval enhancement was broader and extended to shorter delays. Based on these findings, it is suggested that CS units may play an important role in pitch processing in that they transform a higher-order interval code into a first-order interval place code. Their limited dynamic range together with the preservation of the temporal stimulus features in saturation may serve as a physiological basis for the perceived level independence of pitch. (+info)
High-fidelity transmission acquired via a developmental decrease in NMDA receptor expression at an auditory synapse.
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Central auditory relay synapses in mature animals follow high-frequency inputs for computation of sound localization. In immature mice, however, transmission at the calyx of Held synapse in auditory brainstem was inaccurate for high-frequency inputs because the summed slow synaptic potential components caused aberrant firings or blocked action potentials. As the mice matured, synaptic potentials became shorter, with smaller and faster NMDA receptor components, thereby establishing the precise one-to-one transmission for high-frequency inputs. Developmental acquisition of this high-fidelity transmission could be mimicked experimentally in immature mice by blocking NMDA receptors with d(-)2-amino-5-phosphonovaleric acid (d-APV). Furthermore, bilateral cochlear ablations at postnatal day 7 (P7) attenuated the developmental decrease of NMDA receptor expression and prevented the acquisition of high-fidelity transmission. We suggest that auditory activity, which begins at P10-P12 in mice, downregulates the expression of postsynaptic NMDA receptors, thereby contributing to the establishment of high-fidelity synaptic transmission. (+info)