Glycine immunoreactivity of multipolar neurons in the ventral cochlear nucleus which project to the dorsal cochlear nucleus.

Certain distinct populations of neurons in the dorsal cochlear nucleus are inhibited by a neural source that is responsive to a wide range of acoustic frequencies. In this study, we examined the glycine immunoreactivity of two types of ventral cochlear nucleus neurons (planar and radiate) in the rat which project to the dorsal cochlear nucleus (DCN) and thus, might be responsible for this inhibition. Previously, we proposed that planar neurons provided a tonotopic and narrowly tuned input to the DCN, whereas radiate neurons provided a broadly tuned input and thus, were strong candidates as the source of broadband inhibition (Doucet and Ryugo [1997] J. Comp. Neurol. 385:245-264). We tested this idea by combining retrograde labeling and glycine immunohistochemical protocols. Planar and radiate neurons were first retrogradely labeled by injecting biotinylated dextran amine into a restricted region of the dorsal cochlear nucleus. The labeled cells were visualized using streptavidin conjugated to indocarbocyanine (Cy3), a fluorescent marker. Sections that contained planar or radiate neurons were then processed for glycine immunocytochemistry using diaminobenzidine as the chromogen. Immunostaining of planar neurons was light, comparable to that of excitatory neurons (pyramidal neurons in the DCN), whereas immunostaining of radiate neurons was dark, comparable to that of glycinergic neurons (cartwheel cells in the dorsal cochlear nucleus and principal cells in the medial nucleus of the trapezoid body). These results are consistent with the hypothesis that radiate neurons in the ventral cochlear nucleus subserve the wideband inhibition observed in the dorsal cochlear nucleus.     

Projections from the ventral cochlear nucleus to the dorsal cochlear nucleus in rats

Local circuit interactions between the dorsal and ventral divisions of the cochlear nucleus are known to influence the evoked responses of the resident neurons to sound. In the present study, we examined the projections of neurons in the ventral cochlear nucleus to the dorsal cochlear nucleus by using retrograde transport of biotinylated dextran amine injected into restricted but different regions of the dorsal cochlear nucleus. In all cases, we found retrogradely labeled granule, unipolar brush, and chestnut cells in the granule cell domain, and retrogradely labeled multipolar cells in the magnocellular core of the ventral cochlear nucleus. A small number of the labeled multipolar cells were found along the margins of the ventral cochlear nucleus, usually near the boundaries of the granule cell domain. Spherical bushy, globular bushy, and octopus cells were not labeled. Retrogradely-labeled auditory nerve fibers and the majority of labeled multipolar neurons formed a narrow sheet extending across the medial-to-lateral extent of the ventral cochlear nucleus whose dorsoventral position was topographically related to the injection site. Labeled multipolar cells within the core of the ventral cochlear nucleus could be divided into at least two distinct groups. Planar neurons were most numerous, their somata found within the associated band of labeled fibers, and their dendrites oriented within this band. This arrangement mimics the organization of isofrequency contours and implies that planar neurons respond best to a narrow range of frequencies. In contrast, radiate neurons were infrequent, found scattered throughout the ventral cochlear nucleus, and had long dendrites oriented perpendicular to the isofrequency contours. This dendritic orientation suggests that radiate neurons are sensitive to a broad range of frequencies. These structural differences between planar and radiate neurons suggest that they subserve separate functions in acoustic processing. J. Comp. Neurol. 385:245–264, 1997. © 1997 Wiley-Liss, Inc.     

Origins and targets of commissural connections between the cochlear nuclei in guinea pigs

Our objective was to identify the origins and targets of axons that project from one cochlear nucleus to the other. First, retrograde tracers were injected into one cochlear nucleus to label commissural cells in the opposite nucleus. In the dorsal cochlear nucleus, a few cells in the deep layers were labeled; they were not further classified according to type. In the ventral cochlear nucleus, all commissural cells that could be classified were multipolar cells. Second, an anterograde tracer was injected into one cochlear nucleus, and the distribution of boutons in the opposite cochlear nucleus was examined. Labeled boutons were present throughout the ventral cochlear nucleus, where they appeared to contact multipolar cells, spherical and globular bushy cells, and octopus cells. In the dorsal cochlear nucleus, labeled boutons were present in the fusiform cell and deep layers and appeared to contact fusiform cells and cells of unknown type. Many labeled terminals were also present in the granule cell regions. Injections into regions associated with high or low frequencies labeled boutons in corresponding regions in the contralateral ventral cochlear nucleus. Third, multiple tracers were used to determine whether cells that project to the inferior colliculus are contacted by commissural axons. Boutons labeled by anterograde transport of one tracer placed in the cochlear nucleus were frequently observed to be apposed to cells that were labeled by retrograde transport of a different tracer placed in the contralateral inferior colliculus. We conclude that commissural projections originate from multipolar cells throughout the ventral cochlear nucleus (and from a small number of cells in the dorsal cochlear nucleus) and make contact with all major cell types of the cochlear nuclei, including at least some of those that project to the inferior colliculus. © 1996 Wiley-Liss, Inc.     

Origins of axons in the cat's acoustic striae determined by injection of horseradish peroxidase into severed tracts.

Origins and terminations of fibers of the dorsal and intermediate acoustic striae were studied by surgically severing these tracts and injecting HRP into the incision. This procedure results in filling the severed axons with HRP. Filled axons were traced to cell groups of origin and to some terminations of the acoustic striae. HRP-labeled terminals were found in the cochlear nuclei as well as in periolivary cell groups. Filling of cells with HRP RANged from being complete, resulting in Golgi-like images, to being barely detectable. Labeled cells were abundant in the dorsal and posteroventral cochlear nucleus adjacent to the injection as well as scattered throughout the periolivary cell groups of both sides, being highest in concentration around the ipsilateral lateral superior olive. On the side contralateral to the injection, labeled cells were found along the medial border of the dorsal cochlear nucleus, in the interstitial nucleus of the stria of Held, and sparsely throughout the ventral cochlear nucleus. The distribution of labeled cells was similar following HRP injections of the dorsal cochlear nucleus, except that these injections revealed additional descending projections from the inferior colliculi and from the ventral nucleus of the trapezoid body of both sides. These additional projections were interpreted as entering the CN by a ventral route. Findings of this study are in accord with physiological recordings made from fibers of the acoustic striae.     

Planar multipolar cells in the cochlear nucleus project to medial olivocochlear neurons in mouse

Medial olivocochlear (MOC) neurons originate in the superior olivary complex and project to the cochlea, where they act to reduce the effects of noise masking and protect the cochlea from damage. MOC neurons respond to sound via a reflex pathway; however, in this pathway the cochlear nucleus cell type that provides input to MOC neurons is not known. We investigated whether multipolar cells of the ventral cochlear nucleus have projections to MOC neurons by labeling them with injections into the dorsal cochlear nucleus. The projections of one type of labeled multipolar cell, planar neurons, were traced into the ventral nucleus of the trapezoid body, where they were observed terminating on MOC neurons (labeled in some cases by a second cochlear injection of FluoroGold). These terminations formed what appear to be excitatory synapses, i.e., containing small, round vesicles and prominent postsynaptic densities. These data suggest that cochlear nucleus planar multipolar neurons drive the MOC neuron's response to sound.     

Axonal pathways to the lateral superior olive labeled with biotinylated dextran amine injections in the dorsal cochlear nucleus of rats

The lateral superior olive (LSO) contains cells that are sensitive to intensity differences between the two ears, a feature used by the brain to localize sounds in space. This report describes a source of input to the LSO that complements bushy cell projections from the ventral cochlear nucleus (VCN). Injections of biotinylated dextran amine (BDA) into the dorsal cochlear nucleus (DCN) of the rat label axons and swellings in several brainstem structures, including the ipsilateral LSO. Labeling in the ipsilateral LSO was confined to a thin band that extended throughout the length of the structure such that it resembled an LSO isofrequency lamina. The source of this labeled pathway was not obvious, because DCN neurons do not project to the LSO, and VCN bushy cells were not filled by these injections. Filled neurons in several brainstem structures emerged as possible sources. Three observations suggest that most of the axonal labeling in the LSO derives from a single source. First, the number of labeled VCN planar multipolar cells and the amount of labeling in the LSO were consistent and robust across animals. In contrast, the number of labeled cells in most other structures was small and highly variable. Second, the locations of planar cells and filled axons in the LSO were related topographically to the position of the DCN injection site. Third, labeled terminal arborizations in the LSO arose from collaterals of axons in the trapezoid body (output tract of planar cells). We infer that planar multipolar cells, in addition to bushy cells, are a source of ascending input from the cochlear nucleus to the LSO.     

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.     

Unmyelinated axons of the auditory nerve in cats.

This paper describes some central terminations of type II spiral ganglion neurons as labeled by extracellular injections of horseradish peroxidase (HRP) into the auditory nerve of cats. After histological processing with diaminobenzidine, both thick (2-4 microns) and thin (0.5 microns) fibers of the auditory nerve were stained. Whenever traced, thick fibers always originated from type I spiral ganglion neurons and thin fibers always from type II ganglion neurons. Because the labeling of type II axons faded as fibers projected into the cochlear nucleus, this report is limited to regions of the ventral cochlear nucleus near the auditory nerve root. The central axons of type II neurons are unmyelinated, have simple yet variable branching patterns in the cochlear nucleus, and form both en passant and terminal swellings. Under the light microscope, most swellings are located in the neuropil but they are also found in the vicinity of cell bodies, nodes of Ranvier of type I axons, and blood vessels. Eighteen en passant swellings in the neuropil were located by light microscopy and resectioned for electron microscopy; two of these swellings exhibited ultrastructural features characteristic of chemical synapses. The data indicate that inputs from outer hair cells might be able to influence auditory processing in the cochlear nucleus through type II primary neurons.     

Dorsal cochlear nucleus projections to the inferior colliculus in the cat: A light and electron microscopic study

In the Present report retrograde and anterograde labeling techniques are used to study the Projections of the dorsal cochlear nucleus (DCN) to the inferior colliculus in the cat. Horseradish peroxidase (HRP) or wheat germ agglutinin (WGA-HRP) injections into the inferior colliculus produce large numbers of labeled neurons in the DCN on the opposite side. Labeled cells with projections to the colliculus are identified as fusiform and giant cells and are organized into rostrocaudal bands. The axons of these DCN neurons are labeled by anterograde transport of ³H-leucine and/or proline and studied in light and electron microscopic autoradiographs. Axons from the DCN terminate within the central nucleus of the inferior colliculus in densely labeled, rostrocaudally oriented bands. Less heavily labeled extensions of these bands are found in the deepest layer of the dorsal cortex, and light labeling is found adjacent to the bands in the central nucleus and in the ventrolateral nucleus. Cells in the dorsomedial DCN project to the most ventromedial part of the central nucleus while progressively more ventrolateral cells in the DCN project to more dorsolateral parts of the central nucleus. This present evidence suggests that the DCN sends afferents to only two of the four subdivisions of the central nucleus. Within these regions, the axons from the DCN form terminal boutons or boutons de passage characterized by medium-sized, round synaptic vesicles. The labeled endings nearly always make asymmetric synaptic contacts on the dendrites of disc-shaped and stellate cells in the central nucleus. A few axosomatic contacts are found on one particular cell type, possibly the stellate variety. The results support the hypothesis that each subdivision of the central nucleus receives afferents from a different set of cell types in the auditory nuclei of the lower brainstem. The banding patterns of the efferent cells in the cochlear nucleus and the axons within the central nucleus suggest that these inputs are congruent to the fibrodendritic layers of the central nucleus and may contribute to tonotopic organization in the central nucleus. Finally, the results suggest that each of the two major classes of cells in the central nucleus receives different patterns of inputs from the DCN. These morphological differences could contribute to different electrophysiological responses to the sound stimuli by these cells.     

Afferent connections to the amygdaloid complex of the rat and cat: II. Afferents from the hypothalamus and the basal telencephalon

The projections from the basal telencephalon and hypothalamus to each nucleus of the amygdaloid complex of the rat, and to the central amygdala of the cat, were investigated by the use of retrograde transport of horseradish peroxidase (HRP). The enzyme was injected stereotaxically by microiontophoresis, using three different approaches. The ventral pallidum (Heimer, '78) and ventral part of the globus pallidus were found to project to the lateral and basolateral nuclei of the amygdala. The substantia innominata projects diffusely to the entire amygdaloid complex, except to the lateral nucleus and the caudal part of the medial nucleus. The anterior amygdaloid area shows a similar projection field, the only difference being that this structure does not project to any parts of the medial nucleus. The dorsal subdivision of the nucleus of the lateral olfactory tract sends fibers to the ipsilateral as well as the contralateral basolateral nucleus, and possibly to the ipsilateral basomedial and cortical amygdala. The ventral subdivision of the nucleus of the lateral olfactory tract was massively labeled after an injection in the ipsilateral central nucleus, but this injection affected the commissural component of the stria terminalis. The nucleus of the horizontal limb of the diagonal band of Broca connects with the medial, central, and anterior cortical nuclei, whereas the bed nucleus of stria terminalis and medial preoptic area are related to the medial nucleus predominantly. The lateral preoptic area is only weakly labeled after intra-amygdaloid HRP injections. The hypothalamo-amygdaloid projections terminate preponderantly in the medial part of the amygdaloid complex. Thus, axons from neurons in the area dorsal and medial to the paraventricular nucleus of the hypothalamus distribute to the medial nucleus and intra-amygdaloid part of the bed nucleus of stria terminalis. Most of the amygdalopetal fibers from the ventromedial, ventral premammillary, and arcuate nuclei of the hypothalamus end in the medial nucleus, but some extend into the central nucleus. A few fibers from the ventromedial nucleus of the hypothalamus reach the basolateral nucleus. The lateral hypothalamic area projects heavily to the central nucleus, and more sparsely to the medial and basolateral nuclei. The dorsal hypothalamic area and supramammillary nucleus show restricted projections to the central and basolateral nuclei, respectively. There are only a modest number of crossed hypothalamo-amygdaloid fibers. Most of these originate in the ventromedial nucleus of the hypothalamus and terminate in the contralateral medial nucleus. The projections from the basal telencephalon and hypothalamus to the central nucleus of the amygdala of the cat are similar to the corresponding projections in the rat.     

Ventral nucleus of the lateral lemniscus in guinea pigs: Cytoarchitecture and inputs from the cochlear nucleus

Cytoarchitectonic criteria were used to distinguish three subdivisions of the ventral nucleus of the lateral lemniscus in guinea pigs. Axonal tracing techniques were used to examine the projections from the cochlear nucleus to each subdivision. Based on the cell types they contain and their patterns of input, we distinguished ventral, dorsal, and anterior subdivisions of the ventral nucleus of the lateral lemniscus. All three subdivisions receive bilateral inputs from the cochlear nucleus, with contralateral inputs greatly outnumbering ipsilateral inputs. However, the relative density of the inputs varies: the ventral subdivision receives the densest projection, whereas the anterior subdivision receives the sparsest projection. Further differences are apparent in the morphology of the afferent axons. Following an injection of Phaseolus vulgaris-leucoagglutinin into the ventral cochlear nucleus, most of the axons on the contralateral side and all of the axons on the ipsilateral side are thin. Thick axons are present only in the ventral subdivision contralateral to the injection site. The evidence from both anterograde and retrograde tracing studies suggests that the thick axons originate from octopus cells, whereas the thin axons arise from multipolar cells and spherical bushy cells. The differences in constituent cell types and in patterns of inputs suggest that each of the three subdivisions of the ventral nucleus of the lateral lemniscus makes a distinct contribution to the analysis of acoustic signals. J. Comp. Neurol. 379:363–385, 1997. © 1997 Wiley-Liss, Inc.     

Descending projections from the superior olivary complex to the cochlear nucleus of the cat

Subdivisions of the cochlear nuclear complex give rise to a number of discrete projections to certain cell groups of the superior olivary complex and also received substantial descending projections from the periolivary nuclei. In the present study, we sought to determine by means of retrograde transport of horseradish peroxidase (HRP), and anterograde transport of radiolabeled protein, if the periolivary nuclei give rise to discrete projections to the various subdivisions of the cochlear nuclear complex. Following medium to large injections of HRP into the cochlear nucleus, irrespective of location, labeled cells were found in all periolivary nuclei bilaterally. In every case more than 40% of the labeled cells were found in the lateral nucleus of the trapezoid body on the same side and the ventral nucleus of the trapezoid body of both sides. Other periolivary nuclei contributing more than 5% of the total number of cells in individual cases were the contralateral lateral nucleus of the trapezoid body and the ipsilateral anterolateral and dorsal periolivary nuclei. Injections of tritiated leucine into periolivary nuclei gave rise to axonal labeling to the trapezoid body and the dorsal acoustic stria, usually bilaterally, and to terminal labeling that was widely distributed within the cochlear nuclear complex. In several cases with small injections, particularly in the lateral nucleus of the trapezoid body, the projections from the periolivary nuclei to the anteroventral and dorsal cochlear nuclei connected areas described as having similar best-frequency representation. The autoradiographic data corroborated the main results from the HRP experiments and provided additional information permitting these conclusions: the projections from the periolivary nuclei to the cochlear nuclear complex are organized tonotopically, at least in part; each periolivary nucleus (and perhaps individual cells), projects widely throughout the cochlear nuclear complex; the pattern of termination of projections from different periolivary nuclei to a given region of the cochlear nuclear complex are similar, as seen in autoradiograms, and the lateral and dorsal periolivary nuclei project mainly ipsilaterally, while the medial periolivary nuclei project bilaterally with a contralateral bias. The magnitude of these projections and their widespread distribution within the cochlear nuclear complex would suggest an important role for the descending projections in the normal functioning of the cochlear nucleus.     

Transneuronal labeling of cochlear nucleus neurons by HRP-labeled auditory nerve fibers and olivocochlear branches in mice.

Auditory nerve fibers were labeled by extracellular injections of horseradish peroxidase into the spiral ganglion in mice. The labeled fibers were traced in an anterograde direction through the auditory nerve into the cochlear nucleus. In almost half of the injections, the labeled endings of auditory nerve fibers contacted cochlear nucleus neurons that were also labeled with horseradish peroxidase and were presumably transneuronally labeled. Only darkly labeled endings were associated with transneuronally labeled neurons, but not all darkly labeled endings had targets that were transneuronally labeled. Transneuronal labeling occurred almost exclusively in the ventral cochlear nucleus, often between endbulbs and bushy cells. Both "modified" endbulbs and the larger endbulbs of Held transneuronally labeled the bushy cells that they contacted. At the ultrastructural level, transneuronal labeling was evident as a darkening of ribosomes and the membrane surfaces of mitochondria, endoplasmic reticulum, and the nucleus. Transneuronal labeling occurred rarely in octopus, small, and stellate cells, and in neurons of the dorsal cochlear nucleus. Spiral ganglion injections also label olivocochlear fibers, efferent fibers that pass through the ganglion en route to the hair cells. These fibers give off branches to the cochlear nucleus that were rarely associated with transneuronal labeling. In eight instances, the targets of olivocochlear branches were stellate cells or small cells. We suggest that in our mouse preparation, horseradish peroxidase is effective as a transneuronal marker because the short distance from injection site to the cochlear nucleus results in a high concentration of horseradish peroxidase in the endings of the auditory nerve fibers.     

Uncrossed and crossed projections from the upper cervical spinal cord to the cerebellar nuclei in the rat, studied by anterograde axonal tracing

In the upper cervical spinal segments, neurons in the medial part of lamina VI give rise to uncrossed spinocerebellar axons, whereas the central cervical nucleus (CCN) and neurons in laminae VII and VIII give rise to crossed spinocerebellar axons. Using anterograde labeling with biotinylated dextran in the rat, we examined the projections of these neuronal groups to the cerebellar nuclei. Uncrossed and crossed projections were distinguished by cerebellar lesions placed on the side contralateral or ipsilateral to the tracer injections confined to the second and third cervical spinal segments (C2 and C3, respectively). Labeled terminals of uncrossed projections were seen in the middle, dorsal, and ventrolateral parts of the middle subdivision and in the ventral part of the caudomedial subdivision of the medial nucleus. In the anterior interpositus nucleus, terminals were seen in the middle of the mediolateral extent, whereas, in the posterior interpositus nucleus, they were seen in lateral and caudal parts. The terminals of crossed projections from the CCN were distributed ventrally in medial to ventrolateral parts of the middle subdivision of the medial nucleus. Some terminals were seen in the caudomedial subdivision of the medial nucleus. In the anterior interpositus nucleus, labeled terminals were seen mainly in rostromedial parts, whereas, in the posterior interpositus nucleus, they were seen in caudal and dorsal parts of the medial half. The present study suggests that the medial lamina VI group and the CCN in the upper cervical segments project to the different areas of the cerebellar nuclei and are concerned with different functions.     

Tonotopic projection from the dorsal to the anteroventral cochlear nucleus of mice

To understand how auditory information is processed in the cochlear nuclei, it is crucial to know what circuitry exists and how it functions. In slice preparations, horseradish peroxidase (HRP) injections into the anteroventral cochlear nucleus (AVCN) reveal two circuits: a connection between the dorsal cochlear nucleus (DCN) and AVCN and a local circuit confined to the AVCN. Extracellular injection in the AVCN labels a band of cells in the DCN. The labeled cells in the DCN lie within a band of auditory nerve fiber terminals that are labeled by the same injection, showing that the connection from the DCN to the AVCN is frequency specific. The injections into the AVCN also labeled a cluster of neurons in the AVCN dorsal to the injection site. These cells may be interneurons that relay information from areas encoding higher frequencies to areas encoding lower frequencies within the AVCN. In the parasagittal plane, the AVCN is organized along two orthogonal axes that are indicated with HRP labeling of fibers and cell bodies. The tonotopic axis runs approximately dorsoventrally; the isofrequency axis runs approximately rostrocaudally. The axons of labeled DCN neurons and the cluster lie along the tonotopic axis, whereas the labeled auditory nerve fibers define the isofrequency axis. Where they cross is where HRP is taken up by the fibers. The area of uptake is small and lies in the middle of the darkly stained injection site.     

Development of ventral cochlear nucleus projections to the superior olivary complex in gerbil

The postnatal development of the projection from the ventral cochlear nucleus to the principal nuclei in the superior olivary complex in gerbil (Meriones unguiculatus) was studied in an age-graded series of pups ranging from 0 to 18 days old. Small crystals of 1, 1′-dioctadecyl3, 3, 3′, 3′-tetramethylindocarbocyanine perchlorate (DiI) were inserted into the ventral cochlear nucleus of aldehyde-fixed brains, and the labeled projections were examined with epifluorescence microscopy. Selected sections were photooxidized in a solution of diaminobenzidine and subsequently processed for electron microscopy to examine the development of labeled synapses in the target nuclei. Horseradish peroxidase was injected into the ventral cochlear nucleus of adult gerbils to assess the form and persistence of projections observed in the neonatal animals. In addition, electrophysiological responses to acoustic stimuli of single units in the adult auditory brainstem were analyzed to confirm the functionality of the novel projection from the ventral cochlear nucleus to the contralateral lateral superior olive. By the day of birth (PO), developing axons from the ventral cochlear nucleus have already established highly ordered pathways to the three primary nuclei of the superior olivary complex: the ipsilateral lateral superior olive, the contralateral medial nucleus of the trapezoid body, and at the lateral and medial dendrites of the ipsilateral and contralateral medial superior olive, respectively. Developing axons from the ventral cochlear nucleus that innervated the contralateral medial nucleus of the trapezoid body lacked the terminal morphology characteristic of the calyx of Held, but began to adopt a more characteristic form on P5. The mature calyx appeared around P14–16. Exuberant developmental projections to topographically inappropriate areas of the superior olivary complex were not observed at the postnatal ages studied. In addition to the projections of the ventral cochlear nucleus to the superior olivary complex described in other species, we observed the development and maintenance of a major direct projection from the ventral cochlear nucleus to the contralateral lateral superior olive. On PO, ventral cochlear nucleus axons decussate in the dorsal trapezoid body, form a plexus at the dorsal edge of the contralateral medial superior olive, and enter the ventrolateral limb of the contralateral lateral superior olive. Over the next 2 weeks, fascicles of fibers form on the hilar and ventral aspects of the ventrolateral limb. Fibers arising from these fascicles form converging, but nonoverlapping, arborizations within the ventrolateral limb at right angles to the curvature of the nucleus. The medial region was devoid of labeled axons. The direct innervation of the contralateral lateral superior olive was confirmed in the adult gerbil with anterograde horseradish peroxidase histochemistry and by the recording of excitatory responses in the innervated region to acoustic stimulation of the contralateral ear. © 1995 Wiley-Liss, Inc.     

Forebrain projections to the rostral nucleus of the solitary tract in the hamster

The rostral nucleus of the solitary tract (NST) is the first central site of taste information processing. Specific anatomical subdivisions of the NST receive taste afferent input and contain interneurons and projection neurons that engage ascending or premotor taste pathways. The forebrain projects to the NST and can influence taste responses, but the anatomical relationship between forebrain inputs and the subdivisions of the NST and their cellular elements is not understood. To evaluate this, in this study, we used cholera toxin B (CTb) as a retrograde and anterograde marker. CTb was injected into the rostral NST to label, by retrograde transport, the sources of forebrain inputs. Cells were labeled bilaterally in the lateral and paraventricular hypothalamic nuclei, bed nucleus of the stria terminalis, central nuclei of the amygdala, and the agranular and dysgranular divisions of insular cortex. Within the medulla, labeled cells were located in the parvicellular reticular formation and spinal trigeminal nuclei. In addition, labeled cells and anterograde axonal labeling were present in the rostral NST contralateral to the injections. Injections of CTb centered in the dysgranular insular cortex, the site of most forebrain-NST cells, labeled axon endings confined to the rostral NST. These endings were concentrated in the rostral central and ventral subdivisions. Corticofugal endings in the rostral central subdivision are positioned to influence microcircuits that include taste afferent synapses, presumed inhibitory interneurons, and neurons that project to the parabrachial nucleus. The many corticofugal endings in the ventral subdivision synapse among premotor neurons that ultimately influence salivatory and oromotor outflow. Intramedullary CTb labeling after NST injection indicates that the rostral central subdivision also receives projections from the contralateral rostral NST.     

Synaptic distribution of afferents from reticular nucleus in ventroposterior nucleus of cat thalamus

This study was aimed at determining the synaptic circuitry that contributes to the alterations in thalamic function that accompany changes in behavioral states. The somatosensory sector of the thalamic reticular nucleus (RTN) was identified by microelectrode recording in cats and injected with Phaseolus vulgaris-leucoagglutinin (PHA-L). The axons of labeled RTN cells gave rise to collaterals within the RTN and continued into the dorsal thalamus where they terminated predominately in the ventral posterior lateral nucleus (VPL). After small injections in the upper limb representation of RTN, most labeled terminations in VPL were confined to its medial part, suggesting the presence of a topographic organization in the projection. Terminations were concentrated in localized, focal aggregations of boutons. Combined electron microscopic immunocytochemistry, using immunogold labeling for γ-aminobutyric acid (GABA), showed that the PHA-L labeled boutons were GABA-positive terminals that ended in symmetrical synapses. Eighty-two percent of these synapses were on dendrites of relay neurons, 8.5% on dendrites of interneurons, and 9.3% on somata. The terminals of RTN axons form the majority of axon terminals ending in symmetrical synapses in VPL. Their concentration on relay neurons probably underlies the capacity of the RTN projection to reduce background activity of VPL relay neurons in the awake state and to maintain oscillatory behavior of these neurons in drowsiness and early phases of Sleep. © 1995 Wiley-Liss, Inc.