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Synaptic responses and electrical properties of cells in brain slices of the mouse anteroventral cochlear nucleus.

Intracellular recordings were made from cells in brain slices of the anteroventral cochlear nucleus. Responses to electrical stimulation of the stump of the auditory nerve were: (1) all-or-none, following the stimulus with no delay, and insensitive to the removal of extracellular Ca2+, probably representing the firing of directly stimulated auditory nerve fibers, or (2) graded, excitatory postsynaptic potentials, with synaptic delays of about 0.7 msec, which were often followed by graded, inhibitory synaptic potentials with synaptic delays of 1.2 msec or longer. The excitatory and inhibitory synaptic potentials were abolished by the removal of extracellular Ca2+. The result that delays of inhibitory synaptic responses were at least 2 times as long as those of excitatory ones indicates that probably an additional synapse was interposed. Responses to intracellularly injected current pulses show that cells in the anteroventral cochlear nucleus have one of two types of electrical characteristics. Type I properties are characterized by linear current-voltage relationships in the subthreshold voltage range and large, regularly firing action potentials in the suprathreshold range. Type II properties are characterized by nonlinear current-voltage relationships; suprathreshold depolarization elicits only one or two small action potentials. Type II characteristics are particularly well suited for maintaining the information contained in the timing and patterns of firing of the auditory nerve fibers.

Action Potentials↗

[Damage of the cochlea and of the cochlear nucleus after the application of aminoglycosid antibiotics--a comparative light-, transmission- and scanning electronmicroscopic study in the guinea pig (author's transl)].

Ototoxic alterations were studied in the guinea pig organ of Corti and in the cochlear nucleus after high administrations of aminoglycosid antibiotics (Gentamicin/Tobramycin, 150 mg/kg body weight/day or Amikacin 300 mg/kg body weight for 10 days). After survival times up to 22 days the animals were examined. By means of morphology degeneration was found in the cochlear nucleus before we could state it in the organ of Corti. After longer survival times, when we observed heavy degeneration in the organ of Corti we found only a few myelin figures of axons in the cochlear nerve. Cochlear nucleus: As an early sign of degeneration alterations in the mitochondria were found. Compared with the organ of Corti the damage was less widespread in the second order neurons of the afferent auditory pathways, their dendrites or nerve endings. Quantitative calculations however, were not performed. In axosomatic synapses we could observe a way of degeneration which was not reported before in damage caused by aminoglycosid antibiotics. Free postsynaptic densities were observed to be invaginated into the neuron as a probable way of sequestration. As well we could observe reoccupation of the synaptic sites by a probable sliding of neighbouring nerve endings into the free intracellular space. (Similar observations were reported first by Gentschev and Sotelo [1973] after ablation of the cochlear nerve in rats.

Aminoglycosides↗

Subcellular compartmentalization of a potassium channel (Kv1.4): preferential distribution in dendrites and dendritic spines of neurons in the dorsal cochlear nucleus.

Voltage-dependent ion channels have specific patterns of distribution along the neuronal plasma membrane of dendrites, cell bodies and axons, which need to be unravelled in order to understand their contribution to neuronal excitability and firing patterns. We have investigated the subcellular compartmentalization of Kv1.4, a transient, fast-inactivating potassium channel, in fusiform cells and related interneurons of the rat dorsal cochlear nucleus. A polyclonal antibody which binds to a region near the N-terminus domain of a Kv1.4 channel was raised in rabbits. Using a high-resolution combination of immunocytochemical methods, Kv1.4 was localized mainly in the apical dendritic trunks and cell bodies of fusiform cells, as well as in dendrites and cell bodies of interneurons of the dorsal cochlear nucleus, likely cartwheel cells. Quantitative immunogold immunocytochemistry revealed a pronounced distal to proximal gradient in the dendrosomatic distribution of Kv1. 4. In plasma membrane localizations, Kv1.4 was preferentially present in dendritic spines, either in the spine neck or in perisynaptic locations, always away from the postsynaptic density. These findings indicate that Kv1.4 is largely distributed in dendritic compartments of fusiform and cartwheel cells of the dorsal cochlear nucleus. Its preferential localization in dendritic spines, where granule cell axons make powerful excitatory synapses, suggests a role for this voltage-dependent ion channel in the regulation of dendritic excitability and excitatory inputs.

Amino Acid Sequence↗

Effects of unilateral cochlea ablation on the distribution of calretinin mRNA and immunoreactivity in the guinea pig ventral cochlear nucleus.

The predominantly neuronal, calcium-binding protein calretinin is highly expressed in the guinea pig auditory system. Within the ventral cochlear nucleus (VCN), calretinin-positive auditory nerve fibers terminate on many calretinin-containing bushy, octopus, and multipolar cells. The abundance of calretinin in the cochlear nucleus provides an ideal system for examining the effects of altered neuronal input on the expression of this calcium-binding protein. The present experiments examined the effects of unilateral cochlea ablation on calretinin immunoreactivity and mRNA levels in the VCN. Calretinin mRNA was labeled by in situ hybridization histochemistry using a radioactive oligonucleotide probe and was quantified by optical density measures on autoradiograms. Survival times of 1, 7, and 56 days postlesion were examined. The results revealed a consistent increase in calretinin mRNA in the rostral portion of the ipsilateral anterior VCN 1 day postlesion but no effect on calretinin mRNA in this region at 7 and 56 days postlesion. The intensity of immunohistochemical label was also increased at 1 and 7 days after surgery. In contrast, calretinin mRNA was not affected 1 day postlesion in the ipsilateral posterior VCN but was decreased at both 7 and 56 days postlesion. The decrease in calretinin mRNA in the posterior VCN at longer survival times was accompanied by decreased immunolabeling of fibers projecting from VCN cells to the superior olivary complex. These results suggest that calretinin gene expression is regulated in part by auditory nerve activity in some cochlear neurons but that additional factors related to the unique cellular milieu also control calretinin expression.

Analysis of Variance↗

Effects of neonatal deafening and chronic intracochlear electrical stimulation on the cochlear nucleus in cats.

Four newborn kittens were deafened by daily intramuscular injections of neomycin sulfate, beginning the day after birth and continuing for 14-16 days. At 10-16 weeks of age the deaf kittens were implanted unilaterally with a four wire intracochlear electrode array. The animals were stimulated daily (starting at 13-18 weeks of age), for a period of one hour, at 6 dB above the electrically evoked auditory brainstem response threshold. After 3 months of chronic intracochlear electrical stimulation, animals were studied in acute electrophysiological experiments and euthanized for histological studies. This study compares the stimulated and control cochlear nuclei (CN) of these deafened animals to the CN of four normal adult cats. Statistical comparisons of spherical cell densities in the anteroventral cochlear nucleus (AVCN), cross-sectional spherical cell areas, and volumes of the cochlear nucleus subdivisions were included in the analysis. The results indicate that, by all of these measures, the cochlear nuclei in neonatally deafened animals were significantly different from the cochlear nuclei of control animals. As a result of deafening, the density of spherical cells was decreased by 30%, the cross-sectional areas of spherical cells were reduced by 20%, and the volume of the cochlear nucleus was reduced by 25%. These changes were observed in both cochlear nuclei (ipsilateral to both stimulated and unstimulated ears) of the deafened animals. With the measures employed, no significant difference was demonstrated in comparisons between the deafened/unstimulated and the deafened/stimulated cochlear nuclei. That is, no reversal of the profound effects of deafening was observed in the cochlear nuclei as a consequence of chronic intracochlear electrical stimulation which was begun 11 to 16 weeks after deafening.

Animals↗

Quantitative autoradiography of 5-[3H]6-cyano-7-nitro-quinoxaline-2,3-dione and (+)-3-[3H]dizocilpine maleate binding in rat vestibular nuclear complex after unilateral deafferentation, with comparison to cochlear nucleus.

The distributions of non-N-methyl-D-aspartate and N-methyl-D-aspartate receptors in the rat vestibular nuclear complex were estimated by quantitative autoradiography of 5-[3H]6-cyano-7-nitro-quinoxaline-2,3-dione and (+)-3-[3H]dizocilpine maleate binding, respectively. The binding of 5-[3H]6-cyano-7-nitro-quinoxaline-2,3-dione in the vestibular nuclear complex was also compared with that in the cerebellar cortex and cochlear nucleus. Measurements were made in control rats and in rats with unilateral destruction of the inner ear and removal of the vestibular ganglion. Compared to the unlesioned side, 5-[3H]6-cyano-7-nitro-quinoxaline-2,3-dione binding in the lesioned-side vestibular nuclear complex was decreased significantly in all regions at two to four postoperative days. However, the bilateral asymmetry disappeared in most regions by 30 days. 5-[3H]6-Cyano-7-nitro-quinoxaline-2,3-dione binding increased in the molecular layer of the cerebellar cortex at 30 days after lesion, although there were no clear changes at two to seven days. 5-[3H]6-Cyano-7-nitro-quinoxaline-2,3-dione binding in the cochlear nucleus decreased on the lesioned side, compared to the unlesioned side, in regions receiving significant auditory nerve innervation, but increased in the molecular layer of the dorsal cochlear nucleus. (+)-3-[3H]Dizocilpine maleate binding in regions of the vestibular nuclear complex was reduced on the lesioned side, compared to the unlesioned side, after deafferentation, with the largest reductions usually at 30 postoperative days. It is suggested that: (i) non-N-methyl-D-aspartate receptors are involved in synaptic transmission for both vestibular and auditory nerve fibers, while the involvement of N-methyl-D-aspartate receptors is less certain; (ii) unilateral deafferentation of the vestibular nuclear complex can result in bilateral asymmetries for non-N-methyl-D-aspartate and N-methyl-D-aspartate receptors, which are most prominent at earlier and later survival times, respectively; and (iii) vestibular compensation may involve regulation of both non-N-methyl-D-aspartate and N-methyl-D-aspartate receptors in the vestibular nuclear complex and activation of non-N-methyl-D-aspartate receptor-related processes in cerebellar cortex.

6-Cyano-7-nitroquinoxaline-2,3-dione↗

Choline acetyltransferase, glutamate decarboxylase and tyrosine hydroxylase in the cochlea and cochlear nucleus of the guinea pig.

Activities of choline acetyltransferase (ChAC), glutamate decarboxylase (GAD) and tyrosine hydroxylase (TH), enzymes catalyzing the synthesis of acetylcholine (ACh), gamma-aminobutyric acid (GABA) and catecholamines, respectively, were measured in the cochlea and cochlear nucleus of the guinea pig. ChAc activity in the organ of Corti, third turn, was 1270 pmole ACh formed/min/mg protein (ChAc, 1270) and was higher than in turn 4 (ChAc, 543). ChAc activity was higher when the preparation included the inner hair cell region than when not. GAD activity in samples of turn 3 and 4 combined was low, 0.17 nmole GABA formed/min/mg protein (GAD, 0.17). All 3 enzymes were low in auditory nerve: ChAc, 1.7, GAD, 0.10 and TH, 1.0 pmole DOPA formed/min/mg protein. In the cochlear nucleus, the values were: ChAc, 129, GAD, 1.70 and TH, 2.7. The findings on the distribution of ChAc activity in the organ of Corti fit the hypothesis that the olivocochlear nerve fibers are cholinergic. Because of low GAD in the cochlea, GABA is unlikely to be transmitter in the organ of Corti. Similarly, it is unlikely that ACh, GABA or a catecholamine is a transmitter between the auditory nerve and the cochlear nucleus.

Acetyltransferases↗

Tonotopic organization of the anteroventral cochlear nucleus of the cat.

A quantitatively accurate map of the tonotopic organization of the anteroventral cochlear nucleus (AVCN) was derived from single unit recordings. Histologically localized single unit recordings from many animals were mapped onto a computerized atlas of the cochlear nucleus, and surfaces of constant characteristic frequency (CF) estimated with the aid of computer graphics. In anterior AVCN the surfaces of constant CF were found to be parallel planes, whereas in posterior AVCN they progressively deviated from this simple description. A further complication was noted in the most posterior portion of the AVCN where units with very different CF was found in close proximity. Comparison of the tonotopic map with descriptions of cellular organization shows conclusively that different CF ranges are dominant in the various cytoarchitectonic regions of the AVCN.

Animals↗

Response to acoustic stimuli increases in the ventral cochlear nucleus after stimulus pairing.

Recordings of activity in response to click and hiss were made from 364 units of the ventral cochlear nucleus of cats. The unit response to acoustic stimuli increased after forward or backward pairing of the stimuli with glabella tap and hypothalamic electrical stimulation. The results provide evidence against the widely held view that transmission through this initial brain stem relay of the auditory system is invariant, and suggest, instead, that the activity of the ventral cochlear nucleus changes to support increased attentiveness to acoustic signals after variably ordered pairing of conditioned and unconditioned stimuli.

Acoustic Stimulation↗

Age-related changes in GFAP-immunoreactive astrocytes in the rat ventral cochlear nucleus.

The age-related changes in the ventral cochlear nucleus (VCN) as revealed by glial fibrillary acid protein (GFAP) immunoreactivity were analyzed in the following age groups: 3-, 6-, 12-, 18-, and 24-month-old Sprague-Dawley rats. A cartographic and a quantitative analysis showed a significant increase in the number of GFAP positive astrocytes during the first year of life and a significant decrease in older rats. We also observed an age-induced modification in the spatial distribution of GFAP positive astrocyte. In the anterior part of the VCN of the 3- and 6-month-old rats, we observed a significant decrease in the rostro-caudal as well in the dorso-ventral axes. In the posterior part of the VCN, a significant decrease in the dorso-ventral axis could be also observed, but no significant difference in the spatial distribution was obtained in the rostro-caudal axis. In older rats, the distribution appeared homogeneous throughout the nucleus. Additionally, aging was associated with a significant increase in GFAP positive astrocyte sizes, except for immunolabelled astrocytes in the granule cell layer. The different levels of GFAP expression occurring in the VCN during normal aging could reflect a progressive decline of cellular activity in the VCN, without severe cell degeneration or synaptic loss.

Aging↗

Changes in dorsal cochlear nucleus blood flow during noise exposure.

Changes in dorsal cochlear nucleus (DCN) blood flow during noise exposure were assessed using both intravital microscopy (IVM) and laser Doppler flowmetry (LDF). Mature Syrian golden hamsters were anesthetized, tracheotomized and implanted with a carotid artery catheter for monitoring blood pressure and infusion of fluorescent dye. An occipital craniectomy was performed and the cerebellum partly aspirated for access to the DCN. Fluorescent dye was infused to enhance observations of the DCN surface using a customized IVM. Red blood cell velocity, vessel diameter and blood pressure were continuously monitored. Subjects were presented with 110 dB SPL broad-band noise for 15 min. A second group was presented with the same protocol for laser Doppler assessment of blood-flow changes. Control groups included animals not exposed to noise using both IVM and laser Doppler measures. Additional control measurements were obtained for noise-exposed and control groups with laser Doppler measures obtained from the obex, a brainstem structure with no known auditory function. Arterioles of the DCN showed a low-frequency oscillatory pattern of red blood cell velocity in control animals and in baseline conditions of the noise-exposure group. Presentation of noise abolished these velocity oscillations. Additionally, significant decreases in both red blood cell velocity and vessel diameter were measured during stimulation. These microvascular measures recovered slightly during the post-stimulus period. Laser Doppler measures of the overall blood flow in the nucleus were consistent with IVM findings. Measurements from DCN and obex in groups not exposed to noise, and animals exposed to noise and measured from the obex, showed stable oscillatory flow. These data show exposure to noise eliminates oscillatory patterns of blood flow and induces decreases in vascular perfusion. Furthermore, these changes appear specific to the auditory structure studied.

Acoustic Stimulation↗

Responses of anteroventral cochlear nucleus neurons of the unanesthetized decerebrate cat to click pairs as simulated echoes.

To elucidate the contribution of the anteroventral cochlear nucleus (AVCN) to 'echo' processing, this study documents the responses of AVCN neurons to simulated echoes and compares them to those of auditory nerve (AN) fibers. Single unit discharges were recorded from 121 units in the AVCN of 21 unanesthetized decerebrate cats in response to click pairs with inter-click intervals ranging from 1 to 32 ms between 45 and 105 dB SPL re 20 microPa. Units were classified according to the post-stimulus time histogram (PSTH) and excitatory-inhibitory response area (EI-area) schemes. Based on their spontaneous rates (SR), units were subdivided into low- ( < 20 spikes/s) and high- ( > 20 spikes/s) SR groups. A majority of the units exhibited second-click responses whose recovery time courses were similar to those of AN fibers. These units included primary-like, chopper and onset units in the PSTH scheme and Types I, I/III and III units in the EI-area scheme. A minority of the units exhibited responses that were distinct from those of AN fibers, in that they had second-click response recovery times that were either markedly reduced or prolonged. This group of units included those with primary-like, chopper and onset PSTHs and Type I/III and III EI-areas. No significant difference was found in the second-click response among various PSTH or EI-area types. High-SR AVCN units exhibited a decrease in the second-click response with increasing level. In contrast, low-SR AVCN units showed little level-dependent change in the second-click responses. This SR-based difference was similar to that previously found among AN fibers. The present results suggest that, although a majority of AVCN units exhibit similar time courses of second-click response recovery to those of AN fibers, there do exist mechanisms in the cochlear nucleus that can substantially alter this representation. Furthermore, the difference between the second-click response recovery functions of low- and high-SR AVCN units and the consistency of this finding between AVCN and AN suggest that SR represents an important dimension for signal representation in the AVCN neurons.

Acoustic Stimulation↗

A model of selective processing of auditory-nerve inputs by stellate cells of the antero-ventral cochlear nucleus.

Stellate cells in the cat antero-ventral cochlear nucleus (AVCN) maintain a robust rate-place representation of vowel spectra over a wide range of stimulus levels. This rate-place representation resembles that of low threshold, high spontaneous rate (SR) auditory nerve fibers (ANFs) at low stimulus levels, and that of high threshold, low-medium SR ANFs at high stimulus levels. One hypothesis accounting for this phenomenon is that AVCN stellate cells selectively process inputs from different SR population of ANFs in a level-dependent fashion. In this paper, we investigate a neural mechanism that can support selective processing of ANF inputs by stellate cells. We study a physiologically detailed compartmental model of stellate cells. The model reproduces PST histograms and rate-versus-level functions measured in real cells. These results indicate that simple and plausible distribution patterns of excitatory and inhibitory inputs within the stellate cell dendritic tree can support level dependent selective processing. Factors affecting selective processing are identified. This study thus represents a first step towards the development of a computational model of the AVCN stellate cell receptive field.

Animals↗

Cholinergic modulation of stellate cells in the mammalian ventral cochlear nucleus.

The main source of excitation to the ventral cochlear nucleus (VCN) is from glutamatergic auditory nerve afferents, but the VCN is also innervated by two groups of cholinergic efferents from the ventral nucleus of the trapezoid body. One arises from collaterals of medial olivocochlear efferents, and the other arises from neurons that project solely to the VCN. This study examines the action of cholinergic inputs on stellate cells in the VCN. T stellate cells, which form one of the ascending auditory pathways to the inferior colliculus, and D stellate cells, which inhibit T stellate cells, are distinguished electrophysiologically. Whole-cell recordings from stellate cells in slices of the VCN of mice demonstrate that most T stellate cells are excited by cholinergic agonists through three types of receptors, whereas all D stellate cells tested were insensitive to cholinergic agonists. Nicotinic excitation in T stellate cells has two components. The faster component was blocked by alpha-bungarotoxin and methyllycaconitine, suggesting that receptors contained alpha7 subunits; the slower component was insensitive to both. Muscarinic receptors excite T stellate cells by blocking a voltage-insensitive, "leak" potassium conductance. Our results suggest that cholinergic efferent innervation enhances excitation by sounds of T stellate cells, opposing the inhibitory action of cholinergic innervation in the cochlea that is conveyed indirectly through the glutamatergic afferents. The inhibitory action of D stellate cells on their targets is probably not affected by cholinergic inputs. Excitation of T stellate cells by cholinergic efferents would be expected to enhance the encoding of spectral peaks in noise.

Acetylcholine↗

Context-dependent synaptic action of glycinergic and GABAergic inputs in the dorsal cochlear nucleus.

Cartwheel cells are prominent interneurons in the dorsal cochlear nucleus (DCN) that bear considerable homology to cerebellar Purkinje cells. They contact other cartwheel cells as well as fusiform cells, the principal cells of the DCN. In fusiform cells, the inhibition from cartwheel cells interacts with excitation mediated by granule cells and auditory nerve fibers, and shapes the output of the DCN in its ascent to the inferior colliculi. With intracellular recordings from anatomically identified cells in slices, synaptic inputs to fusiform and cartwheel cells were analyzed pharmacologically. Shocks to the auditory nerve and granule cell domains evoked glutamatergic, glycinergic, and GABA(A)ergic postsynaptic potentials (PSPs) in both cartwheel and fusiform cells. The temporal patterns of spontaneous and evoked glycinergic PSPs in fusiform and cartwheel cells were similar and mirrored the pattern of firing of cartwheel cells, probably reflecting the anatomical connections between these cell types and supporting the conclusion that cartwheel cells are glycinergic. In fusiform cells, glycinergic and GABA(A)ergic IPSPs evoked with shocks reversed at -68 mV on average. In marked contrast, glycinergic and GABA(A)ergic PSPs in cartwheel cells, as well as responses to exogenous application of 50-100 mM glycine or 100 microns muscimol, were depolarizing. Reversal potentials of PSPs and responses to glycine and muscimol were similar and averaged -52 mV. Glycinergic and GABA(A)ergic PSPs could elicit firing from cartwheel cells at their resting potentials, but could also reduce rapid firing during strong depolarizations. Thus, the action of glycinergic and GABA(A)ergic inputs on cartwheel cells depends on the electrophysiological context in which they occur.

2-Amino-5-phosphonovalerate↗

Glutamate-stimulated phosphatidylinositol metabolism in the avian cochlear nucleus.

This study examined the ability of the excitatory amino acid glutamate and its analogs to stimulate phosphatidylinositol metabolism in isolated cochlear nucleus tissue from young chicks. In the presence of lithium chloride, glutamate and (+/-)-1-aminocyclopentyl-trans-1,3-dicarboxylate (ACPD) stimulated the formation of inositol phosphates to levels significantly above unstimulated control levels. Unexpectedly, quisqualate did not stimulate inositol phosphates formation. The N-methyl-D-aspartate (NMDA) receptor antagonist 2-amino-5-phosphonovalerate (APV), the ionotropic kainate/quisqualate receptor antagonist 6-cyano-7-nitroquinoxaline-2,3-dione (CNQX) and the putative metabotropic glutamate receptor antagonist 2-amino-3-phosphonopropionate (AP3) had no effect on the glutamate stimulated formation of inositol phosphates. We conclude that a metabotropic glutamate receptor is present on cochlear nucleus neurons of posthatch chicks and is able to stimulate formation of inositol phosphates.

2-Amino-5-phosphonovalerate↗

Computational model of response maps in the dorsal cochlear nucleus.

The neurons in the mammalian (gerbil, cat) dorsal cochlear nucleus (DCN) have responses to tones and noise that have been used to classify them into unit types. These types (I-V) are based on excitatory and inhibitory responses to tones organized into plots called response maps (RMs). Type I units show purely excitatory responses, while type V units are primarily inhibited. A computational model of the neural circuitry of the mammalian DCN, based on the MacGregor neuromime, was used to investigate RMs of the principal cells (P-cells) that represent the fusiform and giant cells. In gerbils, fusiform cells have been shown to have primarily type III unit response properties; however, fusiform cells in the cat DCN are thought to have type IV unit response properties. The DCN model is based on a previous computational model of the cat (Hancock and Voigt Ann Biomed Eng 27: 73-87, 1999) and gerbil (Zheng and Voigt Ann Biomed Eng 34: 697-708, 2006) DCN. The basic model for both species is architecturally the same, and to get either type III unit RMs or type IV unit RMs, connection parameters were adjusted. Interestingly, regardless of the RM type, these units in gerbils and cats show spectral notch sensitivity and are thought to play a role in sound localization in the median plane. In this study, further parameter adjustments were made to systematically explore their effect on P-cell RMs. Significantly, type I, type III, type III-i, type IV, type IV-T and type V unit RMs can be created for the modeled P-cells. Thus major RMs observed in the cat and gerbil DCN are recreated by the model. These results suggest that RMs of individual DCN projection neurons are the result of specific assortment of excitatory and inhibitory inputs to that neuron and that subtle differences in the complement of inputs can result in different RM types. Modulation of the efficacy of certain synapses suggests that RM type may change dynamically.

Animals↗

Neural encoding of single-formant stimuli in the cat. II. Responses of anteroventral cochlear nucleus units.

1. We have studied responses of anteroventral cochlear nucleus (AVCN) units to single-formant stimuli (SFS), in an effort to make quantitative comparisons with responses observed in auditory-nerve fibers (ANFs) to the same stimuli (Wang and Sachs 1993) and to reveal some of the signal processing mechanisms at the AVCN. Single-unit recordings and subsequent analyses were performed on each type of commonly recorded units, namely primarylike (Pri), primarylike with notch (PN), sustained chopper (ChS), transient chopper (ChT), and onset chopper (OnC), as well as a few onset (On) units, from the AVCN in anesthetized cats. The responses were obtained at a wide range of sound levels and at a frequency range of 1-10 kHz. Modulation in the envelopes of discharge patterns was quantified by a measure called modulation depth. 2. At moderate to high sound levels, most AVCN units were found to have enhanced modulation depth compared with that of ANFs, although the degree of enhancement varies among different types. All AVCN units, except Pri type, showed an enhancement in modulation depth over that of the highest of ANFs at moderate to high sound levels in the order of (from the highest to the lowest) On, OnC, ChT/PN, and ChS. Specifically, 1) modulation depth in Pri units was comparable to that of high spontaneous rate (SR) ANFs at low sound levels and to that of low/medium SR ANFs at high sound levels (in dB SPL). When sound level was normalized by unit threshold, Pri units, on average, exhibited only limited enhancement in envelope modulation at high sound levels (> 80 dB re threshold); 2) PN units showed substantially enhanced modulation depth over that of all SR groups of ANFs at moderate to high sound levels in dB SPL or dB re threshold scales; 3) significant enhancement in modulation depth was seen in both ChS and ChT units, with a slightly higher modulation depth in ChT type across sound levels (in dB SPL or dB re threshold); 4) modulation depth of OnC units was higher than those of primary-like (Pri and PN) and chopper (ChS and ChT) units at a wide range of sound levels; 5) responses from a limited sample of On units showed the highest modulation depth among all types of AVCN units. 3. Detailed analysis revealed that the enhanced modulation depth in the responses of AVCN units is the result of increased envelope peak height and decreased envelope minimum, relative to those of ANFs.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗