Morphology of physiologically characterised ventral cochlear nucleus stellate cells

Stellate cells within the ventral cochlear nucleus (VCN) are a diverse cell group that have been classified according to their size and morphology. Some of these stellate cell types constitute major projection neurones into the brainstem and directly into the inferior colliculus, while others are implicated in more local processing. It is still not clear whether a specific physiological profile is uniquely associated with each distinct type of stellate cell. To investigate such associations, we have analysed 23 units with a battery of physiological stimuli in vivo and then examined their shape and outputs following juxtacellular labelling with biocytin. Five physiologically identified groups of cells were filled. These formed two major response classes: onset cells and chopper cells. The two classes could be separated purely on morphological grounds. The onset cells had large somata, large symmetrical dendritic trees and profuse axonal branches that were restricted to the cochlear nucleus on one (On-L) or both sides (On-C) of the brainstem. The chopper cells had smaller, asymmetric, dendritic trees, which were either planar or marginal, had smaller somata and an output axon that left via the trapezoid body. We have confirmed profuse projections into the dorsal cochlear nucleus from all onset cells, and more focal projections from some members of all three groups of chopper cells.     

Hyperpolarization-activated currents regulate excitability in stellate cells of the mammalian ventral cochlear nucleus

The differing biophysical properties of neurons the axons of which form the different pathways from the ventral cochlear nucleus (VCN) determine what acoustic information they can convey. T stellate cells, excitatory neurons the axons of which project locally and to the inferior colliculus, and D stellate cells, inhibitory neurons the axons of which project to the ipsi- and contralateral cochlear nuclei, fire tonically when they are depolarized, and, unlike other cell types in the VCN, their firing rates are sensitive to small changes in resting currents. In both types of neurons, the hyperpolarization-activated current (I(h)) reversed at -40 mV, was activated at voltages negative to -60 mV, and half-activated at approximately -88 mV; maximum hyperpolarization-activated conductances (g(h max)) were 19.1 +/- 2.3 nS in T and 30.3 +/- 2.6 nS in D stellate cells (means +/- SE). Activation and deactivation were slower in T than in D stellate cells. In both types of stellate cells, 50 microM 4(N-ethyl-N-phenylamino)1,2-dimethyl-6-(methylamino) pyridinium chloride (ZD7288) and 2 mM Cs(+) blocked a 6- to 10-fold greater conductance than the voltage-dependent g(h) determined from Boltzmann analyses at -62 mV. The voltage-insensitive, ZD7288-sensitive conductance was proportional to g(h max) and g(input). 8-Br-cAMP shifted the voltage dependence of I(h) in the depolarizing direction, increased the rate of activation, and slowed its deactivation in both T and D stellate cells. Reduction in temperature did not change the voltage dependence but reduced the maximal g(h) with a Q(10) of 1.3 and slowed the kinetics with a Q(10) of 3.3.     

Laminar inputs from dorsal cochlear nucleus and ventral cochlear nucleus to the central nucleus of the inferior colliculus: two patterns of convergence

The central nucleus of the inferior colliculus is a laminated structure composed of oriented dendrites and similarly oriented afferent fibers that provide a substrate for tonotopic organization. Although inputs from many sources converge in the inferior colliculus, how axons from these sources contribute to the laminar pattern has remained unclear. Here, we investigated the axons from the cochlear nuclei that terminate in the central nucleus of the cat and rat. After characterization of the best frequency of the neurons at the injection sites in the cochlear nucleus, the neurons were labeled with dextran in order to visualize their axons and synaptic boutons in the central nucleus. Quantitative methods were used to determine the size and distribution of the boutons within the laminar organization. Two components in the laminae were identified: (1) a narrow axonal lamina that included the largest fibers and largest boutons; (2) a wide axonal lamina, surrounding the narrow lamina, composed of thin fibers and only small boutons. The wide lamina was approximately 30-40% wider than the narrow lamina, and it often extended more than 100 microm beyond the larger boutons on each side. The presence of both thick and thin fibers within the acoustic striae following these injections suggests that large and small fibers/boutons within these bands may originate from different neuronal types in the dorsal and ventral cochlear nucleus. We conclude that the narrow laminae that contain large fibers and boutons originate from larger cell types in the cochlear nucleus. In contrast, the wide lamina composed exclusively of small boutons may represent an input from other, perhaps smaller neurons in the cochlear nucleus. Thus, two types of inferior colliculus laminar structures may originate from the cochlear nucleus, and the small boutons in the wide laminae may contribute a functionally distinct input to the neurons of the inferior colliculus.     

Development of the octopus cell area in the cat ventral cochlear nucleus

The octopus cell area (OCA) of the posteroventral cochlear nucleus was studied electron microscopically in kittens. The adult OCA, a region of morphologically homogenous neurons receiving heterotypic synapses from the cochlea, was used to define the mature state. The OCA reaches cytological maturity at three weeks postnatally, after progression through four stages, defined on the basis of octopus cell cytology (including relative numbers of somatic and dendritic filopodia and spines) and the frequency, ultrastructure and location of previously defined synaptic terminals. Octopus cell size was also studied in rapid Golgi impregnations. The OCA from birth through three postnatal days (stage 1) showed small neurons, few identifiable synaptic types, small, mostly unmyelinated axons, mitotic cells and undifferentiated glia. Between the fourth and seventh postnatal days (stage 2) distinct type 1 and type 2 endings appeared and dendrites thickened, expanded peripherally and developed mature spines. During stage 3 (8–19 days) loss of filopodia, increased somatic spicules, larger somas and clearer differentiation of type 1 and type 2 synapses occurred. After three postnatal weeks (stage 4) the OCA contained morphologically mature octopus cell somas, all three synaptic types ending upon somas and thick basal dendrites, and fascicles of myelinated fibers. Although cytologically mature, the OCA at this stage (about 20–35 days) is substantially smaller than the adult OCA. This smaller size will facilitate further study of OCA synaptic organization.     

Effects of OFF-BF tones on responses of chopper units in ventral cochlear nucleus. I. Regularity and temporal adaptation patterns.

1. We have recorded the responses of neurons in the anteroventral cochlear nucleus (AVCN) of barbiturate-anesthetized cats to pure tones [either at the unit's best frequency (BF) or at another frequency (OFF-BF)] and to two-tone combination stimuli. 2. The effects of OFF-BF input (either alone or presented simultaneously with a BF tone in a two-tone stimulus) on the response patterns of choppers may include not only rate inhibition but changes in the discharge regularity and the temporal adaptation properties of the spike trains. 3. In the majority of cases we studied (119 of 146 frequencies examined in 45 units), the discharge regularity of a response to an OFF-BF or two-tone stimulus is comparable with that of a "rate-matched" BF tone response. In a minority of cases (27 of 146 frequencies examined), however, OFF-BF input (either alone or in a two-tone stimulus format) changed the regularity compared with that of a rate-matched BF tone response. 4. In the majority of cases studied (139 of 171 frequencies examined in 53 units), the initial pattern of rate adaptation ["temporal adaptation pattern" (TAP)] was the same in response to a short tone burst at BF, to an OFF-BF tone burst, or to a pair of tones. The TAP can, however, be significantly altered by OFF-BF input, although this is a comparatively infrequent occurrence in our data sample (32 of 171 frequencies examined), from the response to BF tone to the response to the two-tone or OFF-BF stimulus, are as follows: sustained to slowly adapting; slowly adapting to transiently adapting, and transiently adapting to slowly adapting. Changes in the TAPs of chopper unit responses have been recorded from both regular and irregular choppers and cannot be accounted for on the basis of changes in sustained firing rate. These changes in the discharge regularity and TAP in the small minority of cases suggest that (at least in these cases) the inhibitory effect of OFF-BF input is not simply the result of two-tone suppression at the level of the auditory nerve fiber input. 5. We have observed that regular choppers may be transformed into irregular choppers by OFF-BF (rate inhibitory) input.(ABSTRACT TRUNCATED AT 400 WORDS)     

Dependence of discharge rate on sound pressure level in cochlear nerve fibers of the alligator lizard: implications for cochlear mechanisms

1. Rate-level functions for individual cochlear nerve fibers of the alligator lizard, Gerrhonotus multicarinatus, were generated by measuring a fiber's driven discharge rate (the difference between the average discharge rates in the presence and absence of a tone burst) as a function of sound pressure level. 2. When plotted in double logarithmic coordinates, the rate-level function approaches a straight line at low sound pressure levels and saturates at high levels. Thus the rate-level function is a saturating power function of sound pressure. We developed an algorithm to estimate the exponent of the straight-line portion of the function. When tested on simulated data with known parameters, the algorithm provided unbiased estimates of the exponent. 3. Nerve fibers innervating two distinct regions of the alligator lizard's auditory organ, the free-standing region and the tectorial region, have differing rate-level functions. 4. The mean exponent estimate of the rate-level functions of fibers innervating the free-standing region is approximately 2 at all frequencies. For stimulus frequencies at the characteristic frequency (CF), the mean value was 2.1 +/- 0.10 (SE, n = 131). For stimulus frequencies above and below CF, the mean exponent estimates were 2.1 +/- 0.13 (n = 49) and 2.1 +/- 0.11 (n = 34), respectively. A value of 2 is expected for a broad class of nonlinear systems. 5. The mean exponent estimates of the rate-level functions of fibers innervating the tectorial region were 3.0 +/- 0.30 (n = 32) for stimulus frequencies at CF, 2.5 +/- 0.33 (n = 3) for stimulus frequencies below CF, and 1.0 +/- 0.21 (n = 16) for stimulus frequencies above CF. Both the deviation from square-law behavior at CF and the frequency dependence of the exponent imply that nonlinear processing in the tectorial region differs intrinsically from that in the free-standing region. 6. For free-standing fibers, the saturation rate of the rate-level function (the maximum driven rate) is independent of stimulus frequency. This suggests that, in the free-standing region, 1) the alternating (AC) component of the receptor potential makes no significant contribution to the average rate of discharge and 2) neural saturation results from a process that occurs after the narrow-band frequency-selective process(es). 7. In tectorial fibers, the saturation rate is a bandpass function of sound frequency, with a broad peak between 150 and 300 Hz. This function appears to be common to all tectorial fibers.(ABSTRACT TRUNCATED AT 400 WORDS)     

Regularity and latency of units in ventral cochlear nucleus: implications for unit classification and generation of response properties

1. The responses of neurons in the ventral cochlear nucleus (VCN) of decerebrate cats are described with regard to their regularity of discharge and latency. Regularity is measured by estimating the mean and standard deviation of interspike intervals as a function of time during responses to short tone bursts (25 ms). This method extends the usual interspike-interval analysis based on interval histograms by allowing the study of temporal changes in regularity during transient responses. The coefficient of variation (CV), equal to the ratio of standard deviation to mean interspike interval, is used as a measure of irregularity. Latency is measured as the mean and standard deviation of the latency of the first spike in response to short tone bursts, with 1.6-ms rise times. 2. The regularity and latency properties of the usual PST histogram response types are shown. Five major PST response type classes are used: chopper, primary-like, onset, onset-C, and unusual. The presence of a prepotential in a unit's action potentials is also noted; a prepotential implies that the unit is recorded from a bushy cell. 3. Units with chopper PST histograms give the most regular discharge. Three varieties of choppers are found. Chop-S units (regular choppers) have CVs less than 0.35 that are approximately constant during the response; chop-S units show no adaptation of instantaneous rate, as measured by the inverse of the mean interspike interval. Chop-T units have CVs greater than 0.35, show an increase in irregularity during the response and show substantial rate adaptation. Chop-U units have CVs greater than 0.35, show a decrease in irregularity during the response, and show a variety of rate adaptation behaviors, including negative adaptation (an increase in rate during a short-tone response). Irregular choppers (chop-T and chop-U units) rarely have CVs greater than 0.5. Choppers have the longest latencies of VCN units; all three groups have mean latencies at least 1 ms longer than the shortest auditory nerve (AN) fiber mean latencies. 4. Chopper units are recorded from stellate cells in VCN (35, 42). Our results for chopper units suggest a model for stellate cells in which a regularly firing action potential generator is driven by the summation of the AN inputs to the cell, where the summation is low-pass filtered by the membrane capacitance of the cell.(ABSTRACT TRUNCATED AT 400 WORDS)     

Inferior colliculus responses to multichannel microstimulation of the ventral cochlear nucleus: implications for auditory brain stem implants

Multichannel techniques were used to assess the frequency specificity of activation in the central nucleus of the inferior colliculus (CIC) produced by electrical stimulation of localized regions within the ventral cochlear nucleus (VCN). Data were recorded in response to pure tones from 141 and 193 multiunit clusters in the rat VCN and the CIC, respectively. Of 141 VCN sites, 126 were individually stimulated while recording responses in the CIC. A variety of CIC response types were seen with an increase in both electrical and acoustic stimulation levels. The majority of sites exhibited monotonic rate-level types acoustically, whereas spike rate saturation was achieved predominantly with electrical stimulation. In 20.6% of the 364 characteristic frequency aligned VCN-CIC pairs, the CIC sites did not respond to stimulation. In 26% of the 193 CIC sites, a high correlation was observed between acoustic tuning and electrical tuning obtained through VCN stimulation. A high degree of frequency specificity was found in 58% of the 118 lowest threshold VCN-CIC pairs. This was dependent on electrode placement within the VCN because a higher degree of frequency specificity was achieved with stimulation of medial, central, and posterolateral VCN regions than more anterolateral regions. Broadness of acoustic tuning in the CIC played a role in frequency-specific activation. Narrowly tuned CIC sites showed the lowest degree of frequency specificity on stimulation of the anterolateral VCN regions. These data provide significant implications for auditory brain stem implant electrode placement, current localization, power requirements, and facilitation of information transfer to higher brain centers.     

Analysis of temporal discharge characteristics of dorsal cochlear nucleus neurons of unanesthetized decerebrate cats.

1. We examined the mean and standard deviation (SD) of interspike intervals (ISI) and the coefficient of variation (CV, the ratio of SD of ISI to mean ISI) of ISIs versus time to study discharge regularity of units in the dorsal cochlear nucleus (DCN) of decerebrate unanesthetized cats. The units were characterized by the use of both poststimulus time histograms (PSTH) and excitatory-inhibitory area (EI-area) schemes. We present results of a systematic examination of all of 87 DCN pause-build units recorded in this study. In addition, we present examples of chopper subtypes of the DCN. 2. A major finding of this study is that a majority of the pause-build units in the present sample exhibited regular discharges in response to short (50 ms) tone bursts at characteristic frequency (CF), as revealed by CVs less than 0.5. A predominant portion of pause-build units (80% of 44 units with sufficient number of spikes for CV analysis) exhibited mean CVs less than 0.5 in the 20- to 39.9-ms time window in response to 50-ms tone burst at CF at 60 dB SPL re 20 microPa; 39% of the 44 units exhibited highly regular discharges (mean CV less than 0.35). During the onset (2-14.9 ms) time window, 77% (of 53 units with sufficient number of spikes for CV analysis) of the units had mean CVs less than 0.5, and 55% were less than 0.35. 3. In our sample of 87 DCN pause-build units, 59% had spontaneous rates (SR) greater than 15 spikes/s. The pause-build units of the present sample were distributed across four different EI-area types: III (51%), I/III (25%), II (15%), or IV (9%). In the 20- to 39.9-ms time window, the pause-build units with mean CVs less than 0.35 were exclusively of types III and I/III. All of the above EI-area types were represented in the lowest CV group for the 2- to 14.9-ms window. 4. The mean ISIs of DCN pause-build units typically showed a decrease during the first 20-25 ms of the response to 50-ms CF tone bursts, and stable mean ISIs in the latter half, when off-discharges were absent. In the presence of off-discharges, the mean ISIs decreased further in the last 5-10 ms of the response.(ABSTRACT TRUNCATED AT 400 WORDS)     

Sensitivities of cells in anteroventral cochlear nucleus of cat to spatiotemporal discharge patterns across primary afferents

1. This study tested the hypothesis that a cell in the anteroventral cochlear nucleus (AVCN) that receives convergent input from auditory nerve (AN) fibers can be sensitive to the temporal pattern of discharges on the set of AN fibers providing its input. 2. The temporal discharge pattern across the population of low-frequency AN fibers was manipulated by varying the phase spectra of complex stimuli that had fixed, flat magnitude spectra. By introducing a phase shift with variable slope at a particular frequency, the relative times of discharge of phase-locked neurons with different characteristic frequencies (CFs) could be varied. In this manner the overall spatiotemporal discharge pattern across the array of AN fibers was systematically manipulated. 3. Some low-frequency cells in the AVCN were sensitive to changes in the slope of the phase transition of the complex stimulus. The cells that were sensitive came from several different cell types in the AVCN. Their responses were consistent with the hypothesis that these cells were sensitive to the temporal relationships between discharges on their primary inputs and that they received inputs with different CFs, because the phase shifts introduced relative time differences between different frequencies. 4. Other cells were not sensitive to the degree of phase shift of the stimulus. This insensitivity implied either that these cells received inputs of the same, or nearly the same, CF, or that they were not sensitive to the time differences introduced by these changes in the phase spectra, or both. 5. The cells that were sensitive to the manipulations of the phase spectrum were located in the posterior region of anterior AVCN and in the posterior region of AVCN and thus were presumably either globular bushy, small spherical bushy, or stellate cells. No sensitive cells were located in the most anterior region of the AVCN, where large spherical bushy cells are located. 6. Temporal discharge patterns across the AN population in response to complex stimuli change as a function of level. Accordingly, the sensitivity of neurons to changes in the phase transitions of the complex stimuli used in this study was often affected by the level of the stimulus. 7. The sensitivity to changes in the phase spectrum was a frequency-specific effect. That is, a cell was most sensitive to changes made in phase that were centered near its CF and less sensitive to changes in phase that were introduced at frequencies below or above CF.(ABSTRACT TRUNCATED AT 400 WORDS)     

Synaptic events and discharge patterns of cochlear nucleus cells. II. Frequency-modulated tones.

Responses of 99 cochlear nucleus cells and 24 cochlear nerve fibers were studied with FM signals; 14 cochlear nerve fibers and 57 cochlear nucleus cells were studied at four rates of modulation and several signal intensities. Classification of FM response patterns as symmetrical, asymmetrical, or unidirectional was based on the calculation of a symmetry factor (S), which compared the number of discharges evoked by the ascending and by the descending phases of the FM sweep. Certain FM response patterns could not adequately be described by the symmetry factor along and variables of modulation rate and signal intensity had significant influence. A correspondence was found between the four response classes evoked by a steady-frequency tone burst (primarylike, buildup, onset, and pause) and the FM response pattern. Cochlear nerve fibers showed symmetrical response patterns to FM stimulation. Primarylike units were similar to eighth nerve fibers and generally showed symmetrical FM responses. Occasional eighth nerve fibers and primarylike cells developed asymmetry at the fastest rate of modulation (50 sps). Buildup units showed a variety of response patterns to FM signals. Onset units generally showed asymmetrical response patterns with the greater response occurring to the ascending than to the descending phase of the FM sweep. Pause units showed a characteristic inhibition of activity at 5 sps (rate-dependent inhibition). Of the 57 cochlear nuclear cells studied in response to FM signals, 16 were symmetrical, another 16 were symmetrical except at the fastest modulation rate, 12 were asymmetrical, 3 were unidirectional, and 10 showed complex responses to certain signal rates or intensities. It is clear the the cat cochlear with its complex cytoarchitecture is involved in the recoding of acoustic information. Some units in cochlear nucleus demonstrate differential responses to the direction and to the rate of frequency movement. Other cochlear nucleus cells respond as eighth nerve fibers and may serve as simple "relays" in transmitting information from the cochlea to higher auditory centers.     

Synaptic events and discharge patterns of cochlear nucleus cells. I. Steady-frequency tone bursts.

Unitary discharge patterns (peristimulus time histograms or PSTH) and synaptic events were studies with intracellular recording techniques in 164 cat cochlear nucleus cells to steady-frequency tone bursts 250 ms in duration. There were four response types defined on the basis of the shape of the discharge patterns to tones at the characteristic or best frequency. Primarylike units resemble eighth nerve fibres and have a maximum discharge at tone onset, followed by a smooth decline to a steady level of activity. Buildup units have a transient response at tone onset, followed a period of little or not activity before gradually increasing their discharge rate for the remainder of the tone burst. Onset units have an initial burst of spikes at the onset, with little or no activity for the remainder of the tone burst. Pause units have a long latency (10-30 ms) between tone onset and the appearance of low levels of unit activity, which then gradually increase in rate for the remainder of the tone burst. Changes in signal frequency or intensity within the excitatory response area did not modify response patterns of primarylike and onset units, but could evoke primarylike patterns in buildup and pause units. Inhibition manifested by suppression of spontaneous activity and membrane hyperpolarization were of three kinds: 1) in response to signals at the edges of the excitatory response area (i.e., the inhibitory surround) and detected in onset buildup, and pause units but not in primarylike units; 2) occurring at the offset of tones in the excitatory response area and detected in all four types of cochlear nucleus cells; 3) during excitatory tone bursts in onset and buildup units associated with the periods of suppressed unit activity. Membrane hyperpolarization did not accompany the delay in unit activity after tone onset in pause units. Inhibitory events in cochlear nucleus cells provide mechanisms for producing diversity in the temporal pattern of discharges to acoustic signals which may underly the encoding of complex features of sounds.     

Fine structure and neurotransmitter cytochemistry of neurons in the rat ventral cochlear nucleus projecting to the ipsilateral dorsal cochlear nucleus

The neural tracer wheat germ agglutinin conjugated to horse radish peroxidase was injected into the rat dorsal cochlear nucleus and acoustic stria. Some labelled neurons in the ipsilateral ventral cochlear nucleus were found as a result. These neurons were studied at the ultrastructural level, and their axo-somatic synaptic profile and glycine immunoreactivity were determined. Most neurons were glycine negative and classified as type I multipolar neurons. The latter showed a different synaptic profile from that of neurons projecting to the contralateral inferior colliculus or cochlear nucleus. This suggests the presence of differing populations of multipolar cells based on their synaptic profile. Few labelled multipolar neurons of type II were found, which appeared glycine negative and, rarely, glycine positive. The latter show an ultrastructure and axo-somatic profile similar to that of glycinergic commissural neurons in the dorsal and ventral cochlear nucleus. In particular, about one-third of boutons contained round synaptic vesicles, which are believed to contain an excitatory neurotransmitter. The ultrastructural analysis of the synaptic boutons in the cochlear nucleus confirms the presence of numerous cases of colocalization of glycine and GABA where flat and pleomorphic synaptic vesicles are mixed. The present study is in accordance with previous tract-tracing light microscopic studies which have indicated that large glycinergic neurons in the ventral cochlear nucleus act as broad-band inhibitory neurons in microcircuits of the dorsal cochlear nucleus and contralateral cochlear nucleus.     

Differential expression of three distinct potassium currents in the ventral cochlear nucleus

In the ventral cochlear nucleus (VCN), neurons transform information from auditory nerve fibers into a set of parallel ascending pathways, each emphasizing different aspects of the acoustic environment. Previous studies have shown that VCN neurons differ in their intrinsic electrical properties, including the K+ currents they express. In this study, we examine these K+ currents in more detail using whole cell voltage-clamp techniques on isolated VCN cells from adult guinea pigs at 22 degrees C. Our results show a differential expression of three distinct K+ currents. Whereas some VCN cells express only a high-threshold delayed-rectifier-like current (IHT), others express IHT in combination with a fast inactivating current (IA) and/or a slow-inactivating low-threshold current (ILT). IHT, ILT, and IA, were partially blocked by 1 mM 4-aminopyridine. In contrast, only ILT was blocked by 10-100 nM dendrotoxin-I. A surprising finding was the wide range of levels of ILT, suggesting ILT is expressed as a continuum across cell types rather than modally in a particular cell type. IA, on the other hand, appears to be expressed only in cells that show little or no ILT, the Type I cells. Boltzmann analysis shows IHT activates with 164 +/- 12 (SE) nS peak conductance, -14.3 +/- 0.7 mV half-activation, and 7.0 +/- 0.5 mV slope factor. Similar analysis shows ILT activates with 171 +/- 22 nS peak conductance, -47.4 +/- 1.0 mV half-activation, and 5.8 +/- 0.3 mV slope factor.     

Structural and functional classes of multipolar cells in the ventral cochlear nucleus

Multipolar cells in the ventral cochlear nucleus (VCN) are a structurally and functionally diverse group of projection neurons. Understanding their role in the ascending pathway involves partitioning multipolar cells into distinct populations and determining where in the brain each sends its coded messages. In this study, we used retrograde labeling techniques in rats to identify multipolar neurons that project their axons to the ipsilateral dorsal cochlear nucleus (DCN), the contralateral CN, or both structures. Three rats received injections of biotinylated dextran amine in the ipsilateral DCN and diamidino yellow in the contralateral CN. Several radiate multipolar neurons (defined by their axonal projections to the ipsilateral DCN and their dendrites that traverse VCN isofrequency sheets) were double-labeled but over 70% were not. This result suggests two distinct populations: (1) radiate-commissural (RC) multipolar cells that project to the ipsilateral DCN and the contralateral CN, and (2) radiate multipolar cells that project exclusively (in this context) to the ipsilateral DCN. In a different group of animals, we retrogradely labeled multipolar neurons that project their axons to the contralateral CN and measured the size of their cell bodies. The mean size of this population (266 +/- 156 microm2) was significantly smaller than those of RC-multipolar cells (418 +/- 140 microm2). We conclude that the CN commissural pathway is composed of at least two components: (1) RC multipolar cells and (2) commissural multipolar cells that are small- and medium-sized neurons that project exclusively (in this context) to the contralateral CN. These results identify separate structural groups of multipolar cells that may correspond to physiological unit types described in the literature. They also provide protocols for isolating and studying different populations of multipolar cells to determine the neural mechanisms that govern their responses to sound.     

Encoding timing and intensity in the ventral cochlear nucleus of the cat

Physiological response properties of neurons in the ventral cochlear nucleus have a variety of features that are substantially different from the stereotypical auditory nerve responses that serve as the principal source of activation for these neurons. These emergent features are the result of the varying distribution of auditory nerve inputs on the soma and dendrites of the various cell types within the nucleus; the intrinsic membrane characteristics of the various cell types causing different responses to the same input in different cell types; and secondary excitatory and inhibitory inputs to different cell types. Well-isolated units were recorded with high-impedance glass microelectrodes, both intracellularly and extracellularly. Units were characterized by their temporal response to short tones, rate vs. intensity relation, and response areas. The principal response patterns were onset, chopper, and primary-like. Onset units are characterized by a well-timed first spike in response to tones at the characteristic frequency. For frequencies less than 1 kHz, onset units can entrain to the stimulus frequency with greater precision than their auditory nerve inputs. This implies that onset units receive converging inputs from a number of auditory nerve fibers. Onset units are divided into three subcategories, OC, OL, and OI. OC units have extraordinarily wide dynamic ranges and low-frequency selectivity. Some are capable of sustaining firing rates of 800 spikes/s at high intensities. They have the smallest standard deviation and coefficient of variation of the first spike latency of any cells in the cochlear nuclei. OC units are candidates for encoding intensity. OI and OL units differ from OC units in that they have dynamic ranges and frequency selectivity ranges much like those of auditory nerve fibers. They differ from one another in their steady-state firing rates; OI units fire mainly at the onset of a tone. OI units also differ from OL units in that they prefer frequency sweeps in the low to high direction. Primary-like-with-notch (PLN) units also respond to tones with a well-timed first spike. They differ from onset cells in that the onset peak is not always as precise as the spontaneous rate is higher. A comparison of spontaneous firing rate and saturation firing rate of PLN units with auditory nerve fibers suggest that PLN units receive one to four auditory nerve fiber inputs. Chopper units fire in a sustained regular manner when they are excited by sound.(ABSTRACT TRUNCATED AT 400 WORDS)