Acoustic Stria Research Articles

Development of excitatory synaptic transmission to the superior paraolivary and lateral superior olivary nuclei optimizes differential decoding strategies

The superior paraolivary nucleus (SPON) is a prominent structure in the mammalian auditory brainstem with a proposed role in encoding transient broadband sounds such as vocalized utterances. Currently, the source of excitatory pathways that project to the SPON and how these inputs contribute to SPON function are poorly understood. To shed light on the nature of these inputs, we measured evoked excitatory postsynaptic currents (EPSCs) in the SPON originating from the intermediate acoustic stria and compared them with the properties of EPSCs in the lateral superior olive (LSO) originating from the ventral acoustic stria during auditory development from postnatal day 5 to 22 in mice. Before hearing onset, EPSCs in the SPON and LSO are very similar in size and kinetics. After the onset of hearing, SPON excitation is refined to extremely few (2:1) fibers, with each strengthened by an increase in release probability, yielding fast and strong EPSCs. LSO excitation is recruited from more fibers (5:1), resulting in strong EPSCs with a comparatively broader stimulus–response range after hearing onset. Evoked SPON excitation is comparatively weaker than evoked LSO excitation, likely due to a larger fraction of postsynaptic GluR2-containing Ca2+-impermeable AMPA receptors after hearing onset. Taken together, SPON excitation develops synaptic properties that are suited for transmitting single events with high temporal reliability and the strong, dynamic LSO excitation is compatible with high rate-level sensitivity. Thus, the excitatory input pathways to the SPON and LSO mature to support different decoding strategies of respective coarse temporal and sound intensity information at the brainstem level.

A novel method for selectively labelling olivocochlear collaterals in the rat

Axons of olivocochlear neurons originate from the brainstem and project to the cochlea. A subpopulation, medial olivocochlear (MOC) neurons, also projects collateral branches to the cochlear nucleus. The precise targets of these collaterals are as yet unknown. Previous methods for labelling these collaterals include firstly, cochlear injections of retrograde tracers, but this is technically demanding and can also label afferent projections or secondly, labelling by injecting tracers into the nuclei of origin of MOC neurons. However, this latter method is non-specific because it also labels non-MOC projections. A technique was used to specifically label MOC collaterals, which involved injections of the tracer biocytin at the floor of the fourth ventricle and fixation 3 hours later. Biocytin injections resulted in labelled neurons in the ventral nucleus of the trapezoid body and rostral periolivary nucleus, confirming MOC axonal labelling. Labelled neurons in dorsal cochlear nucleus indicated labelling of the dorsal acoustic stria and these injections were discarded. After selective MOC labelling, collateral branches were found to innervate granule cell regions, medial edge and core of the ventral cochlear nucleus, as well as the dorsal cochlear nucleus, in agreement with previous data. Therefore we conclude that injections at the floor of the fourth ventricle provide a simple, rapid and specific technique for labelling the majority of MOC axons and their collaterals and this technique may assist in defining the precise neuronal targets of olivocochlear collaterals in cochlear nucleus.

Commissural axons of the mouse cochlear nucleus

The axons of commissural neurons that project from one cochlear nucleus to the other were studied after labeling with anterograde tracer. Injections were made into the dorsal subdivision of the cochlear nucleus in order to restrict labeling only to the group of commissural neurons that gave off collaterals to, or were located in, this subdivision. The number of labeled commissural axons in each injection was correlated with the number of labeled radiate multipolar neurons, suggesting radiate neurons as the predominant origin of the axons. The radiate commissural axons are thick and myelinated, and they exit the dorsal acoustic stria of the injected cochlear nucleus to cross the brainstem in the dorsal half, near the crossing position of the olivocochlear bundle. They enter the opposite cochlear nucleus via the dorsal and ventral acoustic stria and at its medial border. Reconstructions of single axons demonstrate that terminations are mostly in the core and typically within a single subdivision of the cochlear nucleus. Extents of termination range from narrow to broad along both the dorsoventral (i.e., tonotopic) and the rostrocaudal dimensions. In the electron microscope, labeled swellings form synapses that are symmetric (in that there is little postsynaptic density), a characteristic of inhibitory synapses. Our labeled axons do not appear to include excitatory commissural axons that end in edge regions of the nucleus. Radiate commissural axons could mediate the broadband inhibition observed in responses to contralateral sound, and they may balance input from the two ears with a quick time course.

Ventral cochlear nucleus responses to contralateral sound are mediated by commissural and olivocochlear pathways.

In the normal guinea pig, contralateral sound inhibits more than a third of ventral cochlear nucleus (VCN) neurons but excites <4% of these neurons. However, unilateral conductive hearing loss (CHL) and cochlear ablation (CA) result in a major enhancement of contralateral excitation. The response properties of the contralateral excitation produced by CHL and CA are similar, suggesting similar pathways are involved for both types of hearing loss. Here we used the neurotoxin melittin to test the hypothesis that this "compensatory" contralateral excitation is mediated either by direct glutamatergic CN-commissural projections or by cholinergic neurons of the olivocochlear bundle (OCB) that send collaterals to the VCN. Unit responses were recorded from the left VCN of anesthetized, unilaterally deafened guinea pigs (CHL via ossicular disruption, or CA via mechanical destruction). Neural responses were obtained with 16-channel electrodes to enable simultaneous data collection from a large number of single- and multiunits in response to ipsi- and contralateral tone burst and noise stimuli. Lesions of each pathway had differential effects on the contralateral excitation. We conclude that contralateral excitation has a fast and a slow component. The fast excitation is likely mediated by glutamatergic neurons located in medial regions of VCN that send their commissural axons to the other CN via the dorsal/intermediate acoustic striae. The slow component is likely mediated by the OCB collateral projections to the CN. Commissural neurons that leave the CN via the trapezoid body are an additional source of fast, contralateral excitation.

Preferential localization of neural cell recognition molecule NB‐2 in developing glutamatergic neurons in the rat auditory brainstem

NB-2 is a neuronal cell recognition molecule that is preferentially expressed in auditory pathways. Mice deficient in the NB-2 gene exhibit aberrant responses to acoustic stimuli. Here we examined the expression and localization of NB-2 in the auditory brainstem during development in the rat. NB-2 was strongly expressed in the ventral cochlear nucleus (VCN), ventral acoustic stria, lateral and medial superior olivary complex (SOC), superior paraolivary nucleus, medial nucleus of the trapezoid body (MNTB), ventrolateral lemniscus, and central nucleus of the inferior colliculus (CIC). In the VCN and CIC, NB-2 was expressed in the regions that are known to respond to high frequencies. In situ hybridization combined with immunohistochemistry suggested that NB-2 is expressed only in neurons. NB-2 was colocalized with glutamatergic elements in the neuropil and the calyces of Held but not with glycinergic or GABAergic elements. NB-2 expression in the SOC was first detected at embryonic day (E)19, reached a maximum level at postnatal day (P)7, and declined thereafter. Immunolabeling with VGLUT1 and VGLUT2, markers for mature and premature glutamatergic synapses, respectively, in combination with NB-2 immunolabeling revealed that NB-2 is expressed at glutamatergic synapses. Collectively, our findings suggest that NB-2 plays a key role in maturation of glutamatergic synapses in the brainstem during the final stages of auditory development.

Molecular guidance cues necessary for axon pathfinding from the ventral cochlear nucleus

During development, multiple guidance cues direct the formation of appropriate synaptic connections. Factors that guide developing axons are known for various pathways throughout the mammalian brain; however, signals necessary to establish auditory connections are largely unknown. In the auditory brainstem the neurons whose axons traverse the midline in the ventral acoustic stria (VAS) are primarily located in the ventral cochlear nucleus (VCN) and project bilaterally to the superior olivary complex (SOC). The circumferential trajectory taken by developing VCN axons is similar to that of growing axons of spinal commissural neurons. Therefore, we reasoned that netrin-DCC and slit-robo signaling systems function in the guidance of VCN axons. VCN neurons express the transcription factor, mafB, as early as embryonic day (E) 13.5, thereby identifying the embryonic VCN for these studies. VCN axons extend toward the midline as early as E13, with many axons crossing by E14.5. During this time, netrin-1 and slit-1 RNAs are expressed at the brainstem midline. Additionally, neurons within the VCN express RNA for DCC, robo-1, and robo-2, and axons in the VAS are immunoreactive for DCC. VCN axons do not reach the midline of the brainstem in mice mutant for either the netrin-1 or DCC gene. VCN axons extend in pups lacking netrin-1, but most DCC-mutant samples lack VCN axonal outgrowth. Stereological cell estimates indicate only a modest reduction of VCN neurons in DCC-mutant mice. Taken together, these data show that a functional netrin-DCC signaling system is required for establishing proper VCN axonal projections in the auditory brainstem.

Differentiation of simple spike waveforms in the hamster dorsal cochlear nucleus

The dorsal cochlear nucleus (DCN) consists of many cell types with different morphologies and properties. DCN cells belonging to different morphological classes are distinguished by differences in their physiological characteristics such as their spectral and temporal response patterns, their levels of spontaneous activity, and certain biophysical properties. Recent studies suggest that they may also exhibit different action potentials, such as simple and complex spikes. In the present study, we systematically examined the spike waveforms of spontaneously active DCN neurons using extracellular recording methods. Neurons were found to exhibit simple spikes consisting of trains of individual action potentials. Spikes fell into two discrete groups of opposite polarity, those with M-shaped and those with W-shaped waveforms. The shapes of these waveforms recorded from a given unit remained constant, despite large changes in amplitude that occurred as the electrode was moved along its axis of penetration. A quantitative analysis of the fine details of the waveforms demonstrated that, although the durations of W- and M-shaped spikes exhibited considerable variation, the variants within each category fell along a continuous gradient rather than into discrete subgroups. Both M- and W-shaped waveforms were found predominantly in the fusiform cell and deep layers, with smaller numbers found in the dorsal acoustic stria. Consideration of their depths of occurrence, their response properties, and levels of spontaneous activity of the recorded neurons suggests that W-shaped waveforms probably are associated with fusiform cells, whereas M-shaped spikes likely originate from more than one cell type.

Effect of unilateral noise exposure on the tonotopic distribution of spontaneous activity in the cochlear nucleus and inferior colliculus in the cortically intact and decorticate rat

Effects of unilateral noise exposure on spontaneous activity (SA) in the anteroventral and dorsal cochlear nuclei (AVCN and DCN) and the central nucleus of the inferior colliculus (ICc) were studied in cortically intact and decorticate rats. SA was measured 1 week following exposure using uptake of 14C-labeled 2-deoxyglucose (2DG) in quiet. Optical density (OD) measurements were obtained in low- and high-frequency (LF and HF) areas of each nucleus. We refer to the ipsilateral AVCN and DCN (side of the noise-exposed ear) and the contralateral ICc as direct nuclei and to their opposite side counterparts as indirect nuclei. Noise exposure altered the tonotopic profile of SA in the direct pathway by causing a decrease in the ratio of HF OD to LF OD (HF/LF ratio). In intact animals, the decreased HF/LF ratio was due to decreased HF OD. In decorticate animals, it was due to decreased HF OD and increased LF OD, the latter occurring mainly in the DCN and ICc. Decorticate-intact differences may reflect corticofugal feedback inhibition. Lesion of the dorsal acoustic stria caused a substantial decrement of SA in the contralateral ICc. Furthermore, strong positive correlations between HF/LF ratios in the DCN, AVCN, and contralateral ICc suggest that the cochlear nucleus is a major contributor to SA in the ICc. Noise exposure had opposite and weaker effects on 2DG uptake in the indirect pathway that were attributed to crossed inhibition. Noise-induced changes in the tonotopic profile of SA may represent a neural correlate of tinnitus.

The effect of dorsal cochlear nucleus ablation on tinnitus in rats

A growing body of evidence implies that the dorsal cochlear nucleus (DCN) plays an important role in tinnitus. To test the hypothesis that the rostral output of the DCN is necessary for the experience of chronic tinnitus, the dorsal DCN and the dorsal acoustic stria of rats with psychophysical evidence of tinnitus was ablated. If the DCN plays a necessary role in the generation of chronic tinnitus, ablating the DCN should decrease the evidence of tinnitus in subjects previously shown to have tinnitus. Contrary to prediction, bilateral dorsal DCN ablation did not significantly ( n = 11, p = 0.707) affect the psychophysical evidence of tinnitus, and ipsilateral dorsal DCN ablation appeared to increase the evidence of tinnitus ( n = 9, p = 0.018) compared to pre-ablation performance. It was concluded that the DCN does not act as a simple feed-forward source of chronic tinnitus. Alternative hypotheses were considered, among them that elevated DCN activity following acoustic trauma triggers persistent pathological changes distributed across more than one level of the auditory system. In addition to serving as a trigger, the DCN may also modify the experience of tinnitus, since the evidence of tinnitus was enhanced by ipsilateral DCN ablation.

Acoustic stria: Anatomy of physiologically characterized cells and their axonal projection patterns

The mammalian cochlear nucleus (CN) has been a model structure to study the relationship between physiological and morphological cell classes. Several issues remain, in particular with regard to the projection patterns and physiology of neurons that exit the CN dorsally via the dorsal (DAS), intermediate (IAS), and commissural stria. We studied these neurons physiologically and anatomically using the intra-axonal labeling method. Multipolar cells with onset chopper (O(C)) responses innervated the ipsilateral ventral and dorsal CN before exiting the CN via the commissural stria. Upon reaching the midline they turned caudally to innervate the opposite CN. No collaterals were seen innervating any olivary complex nuclei. Octopus cells typically showed onset responses with little or no sustained activity. The main axon used the IAS and followed one of two routes occasionally giving off olivary complex collaterals on their way to the contralateral ventral nucleus of the lateral lemniscus (VNLL). Here they can have elaborate terminal arbors that surround VNLL cells. Fusiform and giant cells have overlapping but not identical physiology. Fusiform but not giant cells typically show pauser or buildup responses. Axons of both cells exit via the DAS and take the same course to reach the contralateral IC without giving off any collaterals en route.

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 100μm 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.

Ultrastructure and immunocytochemical characteristics of cells in the octopus cell area of the rat cochlear nucleus: comparison with multipolar cells

Cells in the octopus cell area of the rat ventral cochlear nucleus have been connected to the monaural interpretation of spectral patterns of sound such as those derived from speech. This is possible by their fast onset of firing after each octopus cell and its dendrites have been contacted by many auditory fibres carrying different frequencies. The cytological characteristics that make these large cells able to perform such a function have been studied with ultrastructural immunocytochemistry for glycine, GABA and glutamate, and compared to that of other multipolar neurons of other regions of the ventral cochlear nucleus. Cells in the octopus cell area have an ultrastructure similar to large-giant D-multipolar neurons present in other areas of the cochlear nucleus, from which they differ by the presence of a larger excitatory axo-somatic synaptic input and larger mitochondria. Octopus cells are glycine and GABA negative, and glutamate positive with different degree. Large octopus cells receive more axo-somatic boutons than smaller octopus cells. Fusiform octopus cells are found sparsely within the intermediate acoustic striae. These cells are large to giant excitatory neurons (23-35 microm) with 62-85% of their irregular perimeter covered with large axo-somatic synaptic boutons. Most boutons contain round vesicles and are glycine and GABA negative but glutamate positive. The latter excitatory boutons represent about 70% of the input to octopus cells. Glycine positive boutons with flat and pleomorphic vesicles account for 9-10% of the input while GABA-ergic boutons with pleomorphic vesicles represent about 20% of the synaptic input. Other few, multipolar cells within the rat octopus cell area are surrounded by more inhibitory than excitatory terminals which contain flat and pleomorphic vesicles, a feature distinctive from that of true octopus cells. The latter resemble multipolar cells seen outside the octopus cell area that project to the contralateral inferior colliculus and cochlear nucleus. Based on this study, two types of large multipolar cells are present in the octopus cell area: 1) those that receive about 70% of axo-somatic R boutons and stain more intensely for glutamate may correspond to pure onset neurons (Oi); 2) those with less than 33% of R axosomatic boutons, with less immunoreactivity to glutamate and sometimes glycine positive may represent the onset chopper neurons (Oc). In the octopus cell area the first type appears more prevalent. The present study suggests that octopus cells are a special type of excitatory D-multipolar neuron confined to the octopus cell area and mainly innervated by glutamatergic cochlear nerve terminals.

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.

Role of the dorsal cochlear nucleus in the sound localization behavior of cats

The role of the dorsal cochlear nucleus (DCN) in directional hearing was evaluated by measuring sound localization behaviors before and after cats received lesions of the dorsal and intermediate acoustic striae (DAS/IAS). These lesions are presumed to disrupt spectral processing in the DCN without affecting binaural time and level difference cues that exit the cochlear nucleus via the ventral acoustic stria. Prior to DAS/IAS lesions, cats made accurate head orientation responses toward sound sources in the frontal sound field. After a unilateral DAS/IAS lesion, subjects showed increased errors in the azimuth and elevation of their responses; in addition, the final orientation of head movements tended to be more variable. Largest deficits in response elevation were observed in the hemifield that was ipsilateral to the lesion. When a second lesion was placed in the opposite DAS/IAS, increased orientation errors were observed throughout the frontal field. Nonetheless, bilaterally lesioned cats showed normal discrimination of changes in sound source location when tested with a spatial acuity task. These findings support previous interpretations that the DCN contributes to sound orientation behavior, and further suggest that the identification of absolute sound source locations and the discrimination between spatial locations involve independent auditory processing mechanisms.

Connections of Purkinje cell axons of lobule X with vestibulospinal neurons projecting to the cervical cord in the rat.

Connections of Purkinje cell axons of lobule X (nodulus vermis) with vestibulospinal neurons have been demonstrated in the rat, by anterograde labeling of axons with biotinylated dextran (BD) injected into sublobule Xa and by retrograde labeling of neurons with cholera toxin subunit B (CTB) injected into cervical segments. Labeled terminals of Purkinje cell axons were numerous in the superior vestibular nucleus, the parvocellular (MVpc) and the caudal part (MVc) of the medial vestibular nucleus (MV), and group y. A limited number of labeled terminals were seen in the caudal part of the descending vestibular nucleus (DV). Occasional labeled terminals were seen in the lateral part of the lateral vestibular nucleus (LV) whereas few labeled terminals were seen in the magnocellular part of the MV (MVmc). Vestibulospinal neurons labeled from the C2 and C3 segments were seen bilaterally in the MVmc, MVpc, MVc, and DV, and ipsilaterally in the LV. CTB-labeled vestibulospinal neurons in contact with BD-labeled terminals of Purkinje cell axons were identified in the lateral part of the MVpc, near the border between the MVpc and MVmc, or close to the dorsal acoustic stria, and in the middle part of the MVc at its rostral level. The present study suggests that Purkinje cells of lobule X regulate the output of cervical-projecting vestibulospinal neurons in the MVpc and MVc.

Temporal and Binaural Properties in Dorsal Cochlear Nucleus and Its Output Tract

The dorsal cochlear nucleus (DCN) is one of three nuclei at the terminal zone of the auditory nerve. Axons of its projection neurons course via the dorsal acoustic stria (DAS) to the inferior colliculus (IC), where their signals are integrated with inputs from various other sources. The DCN presumably conveys sensitivity to spectral features, and it has been hypothesized that it plays a role in sound localization based on pinna cues. To account for its remarkable spectral properties, a DCN circuit scheme was developed in which three inputs converge onto projection neurons: auditory nerve fibers, inhibitory interneurons, and wide-band inhibitors, which possibly consist of Onset-chopper (Oc) cells. We studied temporal and binaural properties in DCN and DAS and examined whether the temporal properties are consistent with the model circuit. Interneurons (type II) and projection (types III and IV) neurons differed from Oc cells by their longer latencies and temporally nonlinear responses to amplitude-modulated tones. They also showed evidence of early inhibition to clicks. All projection neurons examined were inhibited by stimulation of the contralateral ear, particularly by broadband noise, and this inhibition also had short latency. Because Oc cells had short-latency responses and were well driven by broadband stimuli, we propose that they provide short-latency inhibition to DCN for both ipsilateral and contralateral stimuli. These results indicate more complex temporal behavior in DCN than has previously been emphasized, but they are consistent with the recently described nonlinear behavior to spectral manipulations and with the connectivity scheme deduced from such manipulations.

Role of acoustic striae in hearing: Reflexive responses to elevated sound-sources

This report is the fourth in a series describing the results of ablation-behavior experiments directed to the ascending output of the cochlear nuclei as it is conducted centrally within the acoustic striae. This fourth report focuses on the unique physiology of the fusiform or ‘output’ cells of the dorsal cochlear nucleus whose axons course through the dorsal acoustic stria (DAS). Because electrophysiological studies have shown that the cues for sensing the elevation of a sound source would seem to be best analyzed by the dorsal cochlear nucleus and projected centrally via its DAS, we tested normal cats and cats deprived of DAS for their ability to orient to elevated sources of broad-band noise. For behavioral testing, we made use of reflexive or unconditioned orienting responses to elevated sound sources using a similar method to one we have used previously for azimuth testing (Thompson GC, Masterton RB. Brainstem auditory pathways involved in reflexive head orientation to sound. J Neurophysiol 1978;41:1183–1202). The results show that cats deprived of their DAS do indeed have a marked deficit in their ability to orient to an elevated sound source. Further behavioral testing indicated that this deficit is not the secondary result of an attentional or peripheral motor deficit. Although the present results do not prove that the reflexive deficit is strictly auditory in nature, the deficit is notable in that it is the only one yet known to result from a lesion of the dorsal cochlear nucleus or its central projections.

Role of acoustic striae in hearing: discrimination of sound-source elevation

After years of systematic experimentation, we finally uncovered one thing the dorsal system contributes to hearing which the ventral system may not – the mechanism for orienting to an elevated sound source [Sutherland, D.P., Masterton, R.B., Glendenning, K.K. (1998) Behav. Brain Res. in press]. This paper follows up this one positive result on a historical background of uniformly negative results. The focus of this report is on the fusiform cells of the dorsal cochlear nucleus whose axons course through the dorsal acoustic stria (DAS). Because electrophysiological studies have shown that the cues for sensing the elevation of a sound source would seem to be best analyzed by the dorsal cochlear nucleus, we tested, behaviorally, normal cats and cats deprived of their DAS or intermediate acoustic stria, bilaterally or ipsilaterally (with or without their contralateral ear deafened), for their ability to orient to elevated sources of broad-band noise. For behavioral testing, we made use of a conventional shock-avoidance procedure. The results lead to the conclusion that DCN and DAS may play no role in learned elevation discriminations. This result builds on that of another of our papers which suggests that a deficit in reflexive discrimination of elevation is strictly auditory in nature [Sutherland, D.P., Masterton, R.B., Glendenning, K.K. (1998) Behav. Brain Res. in press].

Response Classes in the Dorsal Cochlear Nucleus and Its Output Tract in the Chloralose-Anesthetized Cat

Neurons in the dorsal cochlear nucleus (DCN) can be classified into three major physiological classes on the basis of responses to pure tone and broadband noise stimuli. A circuit diagram that associates these classes with different cell types has been proposed. According to this proposal, type II cells are inhibitory interneurons that respond well to tones and poorly to broadband noise, type IV cells are projection neurons with the opposite behavior, and type III cells are an inhomogeneous class with intermediate properties. To test the associations proposed, I compared the response type distribution in the DCN with its output tract, the dorsal acoustic stria (DAS), in chloralose-anesthetized cats. Axonal recordings in the DAS showed type III and IV responses as in DCN, but no type II responses. Compared with reports in decerebrate animals, fewer type IV neurons were encountered having sustained inhibition that generated strongly nonmonotonic responses to tones in both DCN and DAS. The presence of type II responses in the nucleus, but not in the output tract, offers strong support for the proposed association with DCN interneurons. On the other hand, the distinction between type III and IV responses needs refinement because the differences are only graded and because both types of responses occur in DAS, which shows that they are both associated with projection neurons.

Electron microscopic histochemistry for acetylcholinesterase in the dorsal cochlear nucleus of rat

Acetylcholinesterase (AChE), the enzyme which catalyzes hydrolysis of the neurotransmitter acetylcholine, is located in cholinergic neurons, cholinoceptive neurons, and extracellularly in the synaptic clefts of cholinergic synapses. In the rat cochlear nucleus, high activities of AChE have been found in locations rich in granule cells. In the rat dorsal cochlear nucleus (DCN), light microscopic histochemistry for AChE is characterized by stained dots and some patches, especially in its middle, fusiform soma layer, which contains a dense population of granule cells. The fine structure of AChE staining in the DCN has not previously been described. Electron microscopic histochemistry for AChE was performed on perfusion-fixed rat brains. Preliminary results indicate that AChE is distributed differentially across DCN layers. In the molecular layer, AChE staining was found in the extracellular space between nerve fibers and inside the cytoplasm of some dendrites. In deeper layers, extracellularly located AChE was less common, but staining was found inside some of the larger dendrites, some large neuron somata (probably fusiform cells), and some glomeruluslike structures. There appeared to be some labeled fibers coursing in the acoustic striae. [Supported by NIH Grant No. DC00172.]