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Neural correlates of temporal integration in the cochlear nucleus of the chinchilla.

Single unit thresholds were measured as a function of stimulus duration for Primary-like and Chopper units in the anteroventral cochlear nucleus (AVCN) of the chinchilla to examine the neural correlates of temporal integration. Thresholds were measured with a two-alternative, forced-choice (2AFC) adaptive tracking procedure. The time constants and the slopes of the threshold-duration functions were estimated by fitting the threshold data with an exponential function and a power law function. The results showed that Primary-like units exhibited greater threshold improvement and a longer time constant than Chopper units. Units with low characteristic frequencies (CF) showed a larger decrease in threshold with increasing duration and a longer time constant than mid-CF or high-CF units. Units with low spontaneous rates (SR) showed a smaller threshold decrease with increasing duration and a shorter time constant than mid-SR or high-SR units. The single unit time constants and the rate of threshold improvement are similar to those measured psychophysically in the chinchilla.

Acoustic Stimulation↗

Deafferentation increases the intracellular calcium of cochlear nucleus neurons in the embryonic chick.

1. Ratiometric fura-2 imaging was used to measure the intracellular calcium concentration ([Ca2+]i) of neurons in the embryonic avian cochlear nucleus, nucleus magnocellularis (NM), after an in ovo unilateral cochlea removal (deafferentation). 2. The mean [Ca2+]i of NM neurons receiving normal input was 113 nM. 3. Deafferentation increased the mean [Ca2+]i of NM neurons to 247, 311, 339, and 314 nM at 1, 3, 6, and 12 h after cochlear removal, respectively. These values did not differ significantly. 4. The percent frequency distribution of deafferented NM neuron [Ca2+]i shifts away from normative levels toward higher [Ca2+]i at 1 and 3 h after cochlear removal, but shifts back toward normative levels at 6 and 12 h after cochlear removal. 5. This increased [Ca2+]i following cochlear removal temporally coincides with well-characterized changes in NM neurons following activity deprivation. 6. These data suggest that deregulation of [Ca2+]i homeostasis plays a key role in NM neuron degeneration and death following activity deprivation.

Animals↗

Modulation of spontaneous activity by acetylcholine receptors in the rat dorsal cochlear nucleus in vivo.

In vitro studies have implicated muscarinic acetylcholine receptors (mAChRs) in the modulation of spontaneous activity (SA) of neurons in the rat dorsal cochlear nucleus (DCN) (Chen et al., 1994,1998). Early studies suggest that cholinergic pathways also modulate SA in vivo, but these effects have not been investigated pharmacologically. The purpose of the present study was to determine whether multiunit SA can be modulated in vivo by application of cholinergic agents to the surface of the DCN. Sprague Dawley rats were used in the current experiment. The influence of cholinergic activation on SA was tested by applying carbachol (5-500 microM) to the DCN surface while recording multiunit SA at a depth of 250 microm. Out of a total of 32 sites tested, all but 2 (94%) showed well-defined responses to carbachol, characterized by suppression, activation or a combination of both (two-component responses). The most common responses were pure suppression and suppression accompanied by transient activation. Both the proportion of sites showing suppressive responses and the magnitude of suppression averaged across sites increased with dose. Although the proportion of sites showing pure activation in response to carbachol decreased with dose, there was no clear trend in the magnitude of activation with dose. The suppressive responses to high doses of carbachol were blocked by pre-application of atropine. These results extend previous work by suggesting that muscarinic receptors play an important role in the modulation of SA in vivo.

Action Potentials↗

Multisensory integration in the dorsal cochlear nucleus: unit responses to acoustic and trigeminal ganglion stimulation.

A necessary requirement for multisensory integration is the convergence of pathways from different senses. The dorsal cochlear nucleus (DCN) receives auditory input directly via the VIIIth nerve and somatosensory input indirectly from the Vth nerve via granule cells. Multisensory integration may occur in DCN cells that receive both trigeminal and auditory nerve input, such as the fusiform cell. We investigated trigeminal system influences on guinea pig DCN cells by stimulating the trigeminal ganglion while recording spontaneous and sound-driven activity from DCN neurons. A bipolar stimulating electrode was placed into the trigeminal ganglion of anesthetized guinea pigs using stereotaxic co-ordinates. Electrical stimuli were applied as bipolar pulses (100 micros per phase) with amplitudes ranging from 10 to 100 microA. Responses from DCN units were obtained using a 16-channel, four-shank electrode. Current pulses were presented alone or preceding 100- or 200-ms broadband noise (BBN) bursts. Thirty percent of DCN units showed either excitatory, inhibitory or excitatory-inhibitory responses to trigeminal ganglion stimulation. When paired with BBN stimulation, trigeminal stimulation suppressed or facilitated the firing rate in response to BBN in 78% of units, reflecting multisensory integration. Pulses preceding the acoustic stimuli by as much as 95 ms were able to alter responses to BBN. Bimodal suppression may play a role in attenuating body-generated sounds, such as vocalization or respiration, whereas bimodal enhancement may serve to direct attention in low signal-to-noise environments.

Acoustic Stimulation↗

Postnatal morphologic changes and glial fibrillary acidic protein immunoreactivity in the anteroventral cochlear nucleus of the acoustically-deprived gerbil.

This study investigated the morphological changes and glial fibrillary acidic protein immunoreactivity (GFAP-IR) in the anteroventral cochlear nucleus (AVCN) of acoustically-deprived gerbils during postnatal development. The mongolian gerbil, Meriones unguiculatus, had been acoustically deprived on the right side or left side by a surgical ligation of the external auditory canal at postnatal day 12-14. No discernible microcysts were located in the ipsilateral AVCN at one, three, six and nine months after monaural ligation. Also, no discernible microcysts were located in the contralateral AVCN at one and three months after monaural ligation. Numerous microcysts were located in the contralateral AVCN at six months after monaural ligation and were slightly reduced in number at nine months after monaural ligation. Some of the microcysts closely apposed to and connected with the blood vessels through a leakage route or channel. A foamy region was found in the superficial granule cell cap of the AVCN. The foamy region became evident in the ipsilateral AVCN at three months after monaural ligation. However, the foamy region became evident in the contralateral AVCN at three and nine months after monaural ligation. Vacuoles were mainly found in the neuronal cells at the junction of the superficial and deep layers in the AVCN. These vacuoles were found in the contralateral AVCN at one, three, six, and nine months after monaural ligation. However, vacuoles were found in the ipsilateral AVCN only at three months after monaural ligation. Morphological changes of the myelin sheath were found to be more severe in the contralateral AVCN than in the ipsilateral. GFAP-IR was located in the superficial layer of the contralateral AVCN at three and nine months after monaural ligation. However, GFAP-IR was found in the superficial and deep layers of the ipsilateral AVCN at three and nine months after monaural ligation. GFAP-IR was also found in the superficial layers of the ipsilateral AVCN at six months after monoaural ligation. Microcysts are presumably derived from the detachment of the myelin sheath from the retracted axons, protrusion of the myelin sheath, and disruption of the myelin sheath. The major conclusions were that (1) microcysts were greatly reduced following acoustical ligation during postnatal development, and (2) blood vessels and GFAP-immunoreactive astrocytes may be involved in the depletion of microcysts for maintaining the homeostasis of the microenvironment in the cochlear nuclei.

Animals↗

A computational model for signal processing by the dorsal cochlear nucleus. I. Responses to pure tones.

Much information is available on the anatomical organization and neurophysiological properties of the major cell types in the dorsal cochlear nucleus (DCN). The complicated response properties of individual cells and units in the DCN indicate that substantial information processing already occurs at the level of the DCN. A large number of connectional hypotheses have been put forward to explain various aspects of the response characteristics of DCN cells, but many of the consequences of these hypotheses have not been investigated quantitatively. In this paper, we investigate these hypotheses by constructing and testing mathematical and computational models and compare our results to those of previous modeling studies. The simplest versions of our models include auditory nerve (AN) fibers, type II cells (inhibitory interneurons) and type IV cells (fusiform and giant cells). The model response maps, i.e., the pattern of output of model type IV cells, generated by the simplest model have some but not all the features of the experimental response maps of type IV neurons. In particular, the excitatory region which occurs at best frequency is not isolated and the excitatory region at low frequencies and high amplitude is narrower than observed. Since experimental evidence exists that some of the connections between these cell types are divergent or convergent across adjacent isofrequency sheets, the effect of such convergence and divergence was then investigated. Response maps so obtained reproduce many of the qualitative features of the experimental maps. Enlargement of the model by including inhibitory interneurons (e.g., stellate cells) that receive convergent input from AN fibers and which inhibit type II cells results in the generation of response maps that, with some variations in connectional patterns and physiological properties of neurons, match most of the essential features seen in the large variety of experimental maps obtained from the cat DCN.

Animals↗

Dynamic properties of excitation and inhibition in the cochlear nucleus.

The dynamic properties of inhibition and excitation of single units in the cochlear nucleus were studied using tones that were amplitude modulated either sinusoidally or with pseudorandom noise. The cross-correlation analysis of the unit discharge rate and the pseudorandom noise modulation of the stimuli showed that the dynamic properties determined from the responses to a single modulated excitatory tone (at CF), to a modulated excitatory tone together with an unmodulated inhibitory tone (above CF), and to a modulated inhibitory tone together with an an unmodulated excitatory tone were almost identical with regard to latency as well as to delay of the peak of the cross-covariance function. On the basis hereof it is inferred that the inhibition is either a cochlear phenomenon or that it is transmitted over pathways with identical temporal properties as those of the excitation. When 2 tones were presented simultaneously the modulation of the excitatory tone usually gave a higher degree of modulation of the discharge rate than did modulation of an inhibitory tone. Addition of an unmodulated inhibitory tone to a modulated tone at CF resulted in an extension of the range of intensities over which the maximal gain was relatively constant. In many units this range extended from about 10dB above the unit's threshold to 60 dB or more above.

Acoustic Stimulation↗

Dendritic Ca2+ transients evoked by action potentials in rat dorsal cochlear nucleus pyramidal and cartwheel neurons.

Simultaneous fluorescence imaging and electrophysiologic recordings were used to investigate the Ca(2+) influx initiated by action potentials (APs) into dorsal cochlear nucleus (DCN) pyramidal cell (PC) and cartwheel cell (CWC) dendrites. Local application of Cd(2+) blocked Ca(2+) transients in PC and CWC dendrites, demonstrating that the Ca(2+) influx was initiated by dendritic Ca(2+) channels. In PCs, TTX eliminated the dendritic Ca(2+) transients when APs were completely blocked. However, the Ca(2+) influx could be partially recovered during an incomplete block of APs or when a large depolarization was substituted for the blocked APs. In CWCs, dendritic Ca(2+) transients evoked by individual APs, or simple spikes, were blocked by TTX and could be recovered during an incomplete block of APs or by a large depolarization. In contrast, dendritic Ca(2+) transients evoked by complex spikes, a burst of APs superimposed on a slow depolarization, were not blocked by TTX, despite eliminating the APs superimposed on the slow depolarization. These results suggest two different mechanisms for the retrograde activation of dendritic Ca(2+) channels: the first requires fast Na(+) channel-mediated APs or a large somatic depolarization, whereas the second is independent of Na(+) channel activation, requiring only the slow depolarization underlying complex spikes.

Action Potentials↗

Increases in spontaneous neural activity in the hamster dorsal cochlear nucleus following cisplatin treatment: a possible basis for cisplatin-induced tinnitus.

Recent investigations in the hamster have implicated increased spontaneous activity (SA) in the dorsal cochlear nucleus (DCN) as a contributing factor in the etiology of tinnitus induced by intense sound exposure. It might therefore be expected that increased SA would also develop in the DCN of hamsters treated with cisplatin, another cause of tinnitus. We tested this hypothesis by measuring the effects of cisplatin on SA in the DCN. Adult hamsters were divided into three groups, each receiving five injections of cisplatin at one of the following doses: 3 mg/kg, 2.25 mg/kg, or 1.5 mg/kg. Each group had corresponding controls receiving injections of isotonic saline. The effects of cisplatin were studied electrophysiologically 1 month after treatment by recording multiunit SA on the surface of the DCN. Measurements of SA were obtained in three rows of 13-15 locations spaced roughly 100 microm apart and spanning the length of the DCN along the tonotopic axis. Effects of cisplatin were evaluated by comparing plots of mean SA vs. tonotopic locus for cisplatin-treated groups with those of their corresponding untreated control groups. The results demonstrated a consistently higher level of SA in cisplatin-treated groups than in untreated controls. Whereas the highest rates of mean SA in control groups were between 10 and 15 events/s, the highest mean spontaneous rates in cisplatin-treated groups were between 25 and 38 events/s. The cisplatin-induced hyperactivity was greatest in the medial half of the DCN corresponding to the high frequency portion of the tonotopic range. These results suggest that cisplatin treatment is an effective inducer of hyperactivity in the DCN. This hyperactivity may be an important neural correlate of cisplatin-induced tinnitus.

Animals↗

GABA can improve acoustic contrast in the rat ventral cochlear nucleus.

The effect of microiontophoretically applied gamma-aminobutyric acid (GABA) and its agonists and antagonists on the response pattern of single units in the ventral cochlear nucleus (VCN) of the rat was examined in order to study GABA's physiological function in auditory processing. The effects of the drugs were judged by changes of spontaneous and sound-evoked activity in peristimulus-time histograms (PSTHs) of at least 20 consecutive presentations of acoustic stimuli. GABA inhibited the discharge activity of the majority of neurons. All response types found in the VCN except onset-I responders were sensitive to GABA. The GABAergic inhibition is most probably mediated by GABAA receptors, since the GABAA-receptor agonist muscimol, but not the GABAB-receptor agonist baclofen, mimicked the effect of GABA. The GABAA-receptor antagonists, bicuculline and picrotoxin, had an excitatory effect on the neurons' spontaneous activity, suggesting a tonic endogeneous release of GABA which exerts a permanent inhibition on VCN neurons. Although inhibitory, iontophoresis of GABA emphasized the response to stimulus onset in the PSTHs by means of a stronger inhibition of spontaneous activity. When using iontophoretical currents which did not suppress the neuronal activity completely, a strong inhibition of spontaneous activity was accompanied by only a small inhibition of tone-evoked activity. Under these conditions, the response to tone onset was frequently not inhibited at all. Therefore, GABA's physiological function is possibly to improve the contrast between transient acoustic signals and ongoing background activity. In order to test this hypothesis, the test tone was masked by continuous background noise. Indeed, GABA reduced the noise-evoked discharge more than the tone-evoked discharge, leaving the onset peak in the PSTHs almost unchanged. Thus, GABAergic input improves the signal-to-noise ratio for acoustic transients in VCN neurons. Our data suggest that a functional role of GABA in the VCN is to act as a transmitter within a descending inhibitory feedback loop of the auditory brainstem which serves to improve the transmission of relevant acoustic signals in constant background noise.

Animals↗

Coding of envelope modulation in the auditory nerve and anteroventral cochlear nucleus.

We have investigated responses of the auditory nerve fibres (ANFS) and anteroventral cochlear nucleus (AVCN) units to narrowband 'single-formant' stimuli (SFSS). We found that low and medium spontaneous rate (SR) ANFS maintain greater amplitude modulation (AM) in their responses at high sound levels than do high SR units when sound level is considered in dB SPL. However, this partitioning of high and low SR units disappears if sound level is considered in dB relative to unit threshold. Stimuli with carrier frequencies away from unit best frequency (BF) were found to generate higher AM in responses at high sound levels than that observed even in most low and medium SR units for stimuli with carrier frequencies near BF. AVCN units were shown to have increased modulation depth in their responses when compared with high SR ANFS with similar BFS and to have increased or comparable modulation depth when compared with low SR ANFS. At sound levels where AM almost completely disappears in high SR ANFS, most AVCN units we studied still show significant AM in their responses. Using a dendritic model, we investigated possible mechanisms of enhanced AM in AVCN units, including the convergence of inputs from different SR groups of ANFS and a postsynaptic threshold mechanism in the soma.

Acoustic Stimulation↗

The representations of the steady-state vowel sound /e/ in the discharge patterns of cat anteroventral cochlear nucleus neurons.

1. We have recorded the responses of neurons in the anteroventral cochlear nucleus (AVCN) of barbiturate-anesthetized cats to the synthetic, steady-state-vowel sound /e/, presented over a range of stimulus intensities. 2. The responses of (putative) spherical bushy cells [primary-like (Pri) units] to the vowel resemble those of auditory-nerve fibers (ANFs) in terms of both rate and temporal encoding at low and moderate stimulus levels. It was not possible to study the responses of most Pri units at the highest stimulus level because of the large neurophonic component present in recordings from most primarylike units at higher stimulus levels. 3. The responses of many (putative) globular bushy cells [primarylike with notch (PN) units] to the vowel resemble those of ANFs; however, there appears to be greater heterogeneity in the responses of units in the PN population than in the Pri population in terms of both temporal and rate encoding. 4. Populations of stellate cells (chopper units) have degraded representations of the temporal information in ANF population discharge patterns in response to the vowel; this is consistent with the responses of these units to pure tones. Both regular (ChS) and irregular (ChT) chopper subpopulations, however, maintain better rate-place representations of the vowel spectrum than does the population of ANFs as a whole. The rate-place representations of the vowel spectrum by both chopper populations closely resemble those of low and medium spontaneous rate ANFs at most stimulus levels. 5. The data presented in this paper suggest that a functional partition of the AVCN chopper population could yield two distinct rate representations in response to a complex stimulus: one that is graded with stimulus level (over a 30 to 40 dB range) and that, even at rate saturation, maintains a "low contrast" stimulus representation; and a second that maintains a robust, "high contrast" stimulus representation at all levels but that confers less information about stimulus level.

Acoustic Stimulation↗

Regularity analysis in a compartmental model of chopper units in the anteroventral cochlear nucleus.

1. We investigate the discharge patterns of chopper units in the anteroventral cochlear nucleus (AVCN) by developing an equivalent cylinder compartmental model of AVCN stellate cells, which are the sources of the chopper response pattern. The model consists of a passive dendritic tree connected to somatic and axonal compartments with voltage-sensitive channels. Synaptic inputs to the model are simulated auditory nerve fiber responses to best-frequency tones. 2. We adjust the anatomic and electrical parameters of the model to agree with available intracellular data from stellate cells in the AVCN of the mouse and the cat and compare the response of the model to injected current with responses recorded in vitro. The model shows approximately linear current-voltage characteristics for small hyperpolarizing currents. The model's input resistance and the time course of its response to hyperpolarizing current applied at the soma are comparable with those measured from stellate cells in vitro. In response to sustained depolarizing current, the model fires repetitively with nearly perfect regularity, a property also observed in vitro. 3. Auditory nerve inputs to the cell are modeled as deadtime-modified Poisson processes with a multiexponential adaptation in the Poisson rate. We are able to adjust the number, rate, and location of excitatory and inhibitory inputs to the model and succeed in simulating chopper response patterns seen in vivo. 4. Chopper units exhibit a variety of regularity and adaptation patterns in response to tone stimuli. Physiological data from brain slice experiments and experiments in vivo imply that this heterogeneity is primarily due to differences in input configurations. By systematically varying the number and position of excitatory and inhibitory inputs, we can simulate a range of chopper response patterns. 5. We quantify the regularity of the model's response using the coefficient of variation (CV) of the interspike interval. We find that the CV decreases, i.e., the regularity increases, as the number of converging inputs or their distance from the soma increases. The regularity of the output is more sensitive to the number of converging inputs than to their location on the dendritic tree. The statistics of the first spike latency (FSL) are also sensitive to the configuration of excitatory inputs. The mean and minimum FSL are more sensitive to the electrotonic distance of the inputs from the soma than to the number of inputs, whereas the standard deviation of the FSL is highly dependent on the number of converging inputs and is nearly independent of their location.(ABSTRACT TRUNCATED AT 400 WORDS)

Axons↗

Sodium pentobarbital abolishes bursting spontaneous activity of dorsal cochlear nucleus in rat brain slices.

There is evidence that pentobarbital, a commonly used anesthetic, can affect neuronal activity, but its effects on particular neurons of the dorsal cochlear nucleus (DCN) are not well known. Bursting (complex spiking) spontaneous activity has been observed in the DCN in brain slice preparations and in recordings from unanesthetized decerebrate animals, but seldom in experiments with anesthetized animals. This study investigated the effects of pentobarbital on spontaneous activity in the DCN in brain slices. Most extracellularly recorded bursting neurons decreased firing rates and reversibly changed their firing to simple spiking with irregular intervals during pentobarbital. Some reversibly stopped firing after the change to an irregular pattern. Most neurons with regular spontaneous activity (simple spiking) showed decreased firing rates and more irregular intervals during pentobarbital. The results also suggest some involvement of gamma-aminobutyric acid type A receptors in the pentobarbital effects.

Action Potentials↗

Expression of Shal potassium channel subunits in the adult and developing cochlear nucleus of the mouse.

The pattern of expression of potassium (K(+)) channel subunits is thought to contribute to the establishment of the unique discharge characteristics exhibited by cochlear nucleus (CN) neurons. This study describes the developmental distribution of mRNA for the three Shal channel subunits Kv4.1, Kv4.2 and Kv4.3 within the mouse CN, as assessed with in situ hybridization and RT-PCR techniques. Kv4.1 was not present in CN at any age. Kv4.2 mRNA was detectable as early as postnatal day 2 (P2) in all CN subdivisions, and continued to be constitutively expressed throughout development. Kv4.2 was abundantly expressed in a variety of CN cell types, including all of the major projection neuron classes (i.e., octopus, bushy, stellate, fusiform, and giant cells). In contrast, Kv4.3 was expressed at lower levels and by fewer cell types. Kv4.3-labeled cells were more prevalent in ventral subdivisions than in the dorsal CN. Kv4.3 expression was significantly delayed developmentally in comparison to Kv4.2, as it was detectable only after P14. Although the techniques employed in this study detect mRNA and not protein, it can be inferred from the differential distribution of Kv4 transcripts that CN neurons selectively regulate the expression of Shal K(+) channels among individual neurons throughout development.

Animals↗

Octopus cells of the mammalian ventral cochlear nucleus sense the rate of depolarization.

Whole cell patch recordings in slices show that the probability of firing of action potentials in octopus cells of the ventral cochlear nucleus depends on the dynamic properties of depolarization. Octopus cells fired only when the rate of rise of a depolarization exceeded a threshold value that varied between 5 and 15 mV/ms among cells. The threshold rate of rise was independent of whether depolarizations were evoked synaptically or by the intracellular injection of current. Previous work showed that octopus cells are contacted by many auditory nerve fibers, each providing less than 1-mV depolarization. Summation of synaptic input from multiple fibers is required for an octopus cell to reach threshold. In firing only when synaptic depolarization exceeds a threshold rate, octopus cells fire selectively when synaptic input is sufficiently large and synchronized for the small, brief unitary excitatory postsynaptic potentials (EPSPs) to sum to produce a rapidly rising depolarization. The sensitivity to rate of depolarization is governed by a low-threshold, alpha-dendrotoxin-sensitive potassium conductance (g(KL)). This conductance also shapes the peaks of action potentials, contributing to the precision in their timing. Firing in neighboring T stellate cells depends much less strongly on the rate of rise. They lack strong alpha-dendrotoxin-sensitive conductances. Octopus cells appear to be specialized to detect synchronization in the activation of groups of auditory nerve fibers, a common pattern in responses to natural sounds, and convey its occurrence with temporal precision.

Action Potentials↗

Neuronal nitric oxide synthase expression in the cochlear nucleus in a salicylate model of tinnitus.

Although a number of studies suggest that the development of tinnitus is associated with hyperactive neuronal discharges in the brainstem cochlear nucleus (CN), there is relatively little evidence to indicate the neurochemical basis of this phenomenon. While some studies suggest that it may be partly due to a decrease in GABAergic inhibition, it is also possible that increased excitability is a contributing factor. In the current study, we investigated whether the salicylate animal model of tinnitus is associated with changes in the number of CN neurons expressing neuronal nitric oxide synthase (nNOS), one of the NOS isoforms that results in the production of the neurotransmitter, nitric oxide. We used a behavioral conditioning paradigm to confirm that animals receiving salicylate injections experienced tinnitus, and used immunohistochemistry with stereology to quantify the number of nNOS-expressing neurons in the dorsal and ventral CN (DCN and VCN, respectively) in salicylate- and vehicle control-treated animals. We also employed Western blotting to quantify the amount of nNOS protein expression in the total CN (i.e., the DCN and VCN together). We found a significant increase (of approximately 70%) in the number of nNOS-expressing principal neurons in the VCN of salicylate-treated animals compared to controls, with no significant differences in the DCN; nor did we find any significant difference in the overall level of nNOS protein in the total CN using Western blotting. These results suggest that changes in the number of neurons in the VCN expressing nNOS may be implicated in the mechanisms of tinnitus.

Animals↗

Spontaneous and sound-evoked discharge characteristics of complex-spiking neurons in the dorsal cochlear nucleus of the unanesthetized decerebrate cat.

1. We examined the spontaneous and sound-evoked discharge characteristics of 20 complex-spiking units recorded in the dorsal cochlear nucleus (DCN) of 15 unanesthetized, decerebrate cats. 2. The extracellularly recorded complex spikes consisted of bursts of two to five action potentials whose size gradually decreased during the burst. Complex spikes were observed both in the spontaneous and sound-evoked activity of the units in our sample. 3. The spontaneous rates (SRs) of DCN complex-spiking units ranged from 0 to 30 spikes/s. Spontaneous activity consisted of complex and simple (i.e., the common single neuronal action potential) spikes. Comparison of the SR distributions of the DCN complex-spiking units with that of a total sample of 194 DCN units (from 9 cats) suggests that the complex-spiking units tended to be in the lower half of the DCN SR distribution. 4. Sound-evoked discharges could consist of both complex and simple spikes. On the basis of their sound-driven responses, we divided the DCN complex-spiking units into two groups. The majority (15 of 20, 75%) were weakly driven by pure tones and inhibited by broadband noise. They tended to have broad response areas. Their response latencies to pure tone and noise stimuli were relatively long (10-20 ms). The recording depths of these units tended to be superficial (i.e., 10 of 15 units were located within 400 microns of the dorsal surface of the DCN). A minority (5 of 20, 25%) of the complex-spiking units were strongly driven by pure tone and broadband noise stimuli. These units had more clearly defined excitatory regions of response areas than the weakly driven units. Their response latencies to pure tone and noise stimuli were short (< 10 ms). The recording depths of these units tended to be deeper (i.e., 4 of 5 units were located at 400-700 microns) than those of the weakly driven units. 5. Intracellular recording and labeling studies of in vitro DCN slice preparations have correlated complex spikes with the superficially located cartwheel cells. Given the complex spikes of the units, many of which were located superficially, we suggest that our sample, particularly the weakly driven group of neurons, corresponds to the cartwheel cells. 6. Cartwheel cells are putative inhibitory interneurons whose axons primarily contact on the main projection neurons of DCN, the fusiform cells. The present finding of sound-evoked discharges by the superficially located complex-spiking units suggests that cartwheel cells should play a role in modifying the sound-evoked responses of the fusiform cells.

Acoustic Stimulation↗