Dorsal Cap Research Articles

Neurons of the medial terminal accessory optic nucleus of the rat are poorly collateralized

The vast majority of neurons of the rat medial terminal nucleus (MTN) project to the nucleus of the optic tract (NOT), but the MTN also projects to a lesser degree upon a number of other brainstem nuclei controlling optokinetic nystagmus. Because of the diversity of targets of the MTN, it is possible that individual neurons have branched axons that project to two or more brainstem nuclei. The possibility that axons of MTN-NOT neurons collateralize to innervate other MTN targets is examined in the rat with the fluorescent, double-labeling, retrograde tracer technique. Fluoro-Gold was injected into the NOT while Fast Blue was simultaneously injected into each of five other known targets of the MTN: the supraoculomotor-periaqueductal gray; the dorsal cap of the inferior olive; the visual tegmental relay zone; the dorsolateral nucleus of the basal pons; and the superior/lateral vestibular nuclei. Brainstem sections were processed for fluorescence microscopy and the MTN was examined for single- and double-labeled neurons. Results show that virtually all neurons of the MTN (greater than 97.5%), together with neurons in the visual tegmental relay zone immediately surrounding the MTNd, are single-labeled in all paired injections involving the NOT and the other target nuclei. It was found that about 69% of MTN neurons project exclusively to the NOT, 5-10% project to each one of the other nuclei, and 3% of MTN neurons project to more than one target. Based upon cell counts from the fluorescent material, and previous analysis of Nissl-stained serial sections, the findings show that virtually all MTN neurons are projection neurons. It was concluded that the MTN is comprised of independent projection systems, possibly involved in different aspects of generating optokinetic nystagmus.

Olivary branching projections to the flocculus, nodulus and uvula in the rabbit. I. An electrophysiological study.

Olivocerebellar branching projections to the flocculus, nodulus and uvula were studied electrophysiologically in pigmented rabbits anesthetized with pentobarbital and halothane. Neurons in the dorsal cap of the inferior olive were antidromically activated by stimulation of the contralateral flocculus, nodulus and uvula. The antidromic responses in the dorsal cap from the flocculus, nodulus and uvula partially occluded with each other. Electrical stimulation of the nodulus (or uvula) evoked early and late climbing fiber responses in the flocculus with latencies of 4.0 ms and 10.0 ms, respectively. The early response was resistant to repetitive stimulation and not affected by interruption of the olivocerebellar fibers at the lower medulla, while the late response was abolished by these two procedures. This indicated that the early response was evoked through climbing fiber branches to the flocculus and nodulus (or uvula) as an axon reflex, while the late response was evoked via the inferior olive. Of 76 Purkinje cells in the flocculus, 35.6% showed climbing fiber activation through branches to the nodulus and/or uvula, 70% of which were orthodromically activated through climbing fibers from the contralateral optic tract. Electrical stimulation of the flocculus or uvula evoked the early and late climbing fiber responses in the nodulus. Of 60 Purkinje cells in the nodulus, 63.4% showed axon reflex activation from the flocculus and/or uvula, 42% of which were activated through climbing fibers from the contralateral optic tract. These findings demonstrated that the flocculus, nodulus and uvula receive branching climbing fiber projection from the inferior olive, a part of which conveys visual information.

Localized metabolic responses to optokinetic stimulation in the brain stem nuclei and the cerebellum investigated with the [ 14C]2-deoxyglucose method in rats

The localized metabolic effects of monocular optokinetic stimulation to the cerebellar flocculus and brain stem nuclei were measured in pigmented rats. Quantitative [ 14C]2-deoxyglucose autoradiography was performed on alert rats stimulated with a drum either rotated horizontally in the temporonasal direction (optokinetic group) or kept stationary (control group). The superior colliculus in both groups showed a higher amount of activity on the contralateral side to the stimulated eye than on the ipsilateral side. The dorsal cap of the inferior olive, the nucleus of the optic tract, and the lateral pontine nucleus showed a higher amount of activity on the contralateral side only in the optokinetic group. The nucleus reticularis tegmenti pontis and the ventromedial aspect of the cerebellar paraflocculus showed no lateralized activity in either group. Local glucose utilization rates of both flocculi were significantly enhanced in the optokinetic group. Only in the optokinetic group did the ipsilateral flocculus show a higher local glucose utilization rate than the contralateral flocculus. The most enhanced activity was localized in the middle aspect of the rostrocaudal extent of the ipsilateral flocculus. The activity was greater in the granular layer than in the molecular layer or in the white matter. The pattern of activation in the granular layer was characterized by a patchy appearance in the frontal sections. Serial reconstruction of these sections showed metabolic blobs appearing with intervals of several hundred micrometers.

Spatial organization of visual messages of the rabbit's cerebellar flocculus. I. Typology of inferior olive neurons of the dorsal cap of Kooy.

1. Single-unit responses to large-field visual stimuli were recorded extracellularly from neurons in the dorsal cap of Kooy of the inferior olive in anesthetized, paralyzed rabbits. The visually modulated neurons in the dorsal cap responded optimally to slow rotation of random dot stimuli, which were produced using handheld patterns or a planetarium projector. 2. Neurons had either monocular or binocular receptive fields. For binocular receptive field neurons, monocular stimulation of one eye, called the dominant eye, elicited greater modulation than did stimulation of the other eye. Rotation about a particular axis, called the preferred axis, produced both maximal excitation and inhibition. On the basis of differences in preferred axis orientation and in eye dominance, three neuron classes called Vertical Axis, Anterior (45 degrees) Axis, and Posterior (135 degrees) Axis were distinguished. 3. Vertical Axis neurons were modulated exclusively from the eye contralateral to the inferior olive recording site. This cell type responded strongly to large-field visual stimuli rotating about the vertical axis. Excitation resulted from movement in the temporal to nasal direction, and inhibition occurred during movement in the nasal to temporal direction. 4. Two subclasses of Anterior (45 degrees) Axis neurons were distinguished according to whether the receptive field was monocular or binocular. For both subclasses, the dominant eye was ipsilateral. The receptive field organization of the dominant eye was bipartite as described in the previous paper (51) for neurons in the midbrain visual tegmental relay zone. Anterior (45 degrees) Axis neurons were maximally excited when the stimulus pattern moved upward and posterior above the horizon in the anterior quadrant of the ipsilateral visual field, from 0 degrees (nose) to approximately 45 degrees azimuth. From 45 to 180 degrees azimuth (occiput) and above the horizon, these neurons were excited by downward and posterior movement. Inhibition occurred with oppositely directed movements. For rotating stimuli presented to the dominant eye, this class of neurons responded best to rotation of the visual world about an axis oriented near the horizontal plane and approximately 45 degrees azimuth. 5. The receptive field of Posterior (135 degrees) Axis neurons was always binocular, with the dominant eye contralateral. For the contralateral receptive field, from 0 degree (nose) to 135 degrees azimuth and above the horizon, excitation occurred during upward and posterior movement.(ABSTRACT TRUNCATED AT 400 WORDS)

The accessory optic system of rabbit. II. Spatial organization of direction selectivity.

1. To compare the spatial organization of the direction selectivity of neurons in the medial terminal nucleus (MTN) of the accessory optic system with that of neurons in the adjacent ventral tegmentum, extracellular single-unit recordings were made in the anesthetized rabbit. The ventral tegmental neurons were located in a region called the visual tegmental relay zone (VTRZ), which is defined by the ventral tegmental terminal field of contralaterally projecting MTN neurons. 2. Some of the present sample of MTN neurons (5 of 34) had monocular receptive fields composed of two parts distinguished by a marked difference in the orientation of their respective direction-selective tuning curves. For one part of the receptive field the preferred excitatory direction was "up," while for the other part it was "down." Such receptive fields for one eye were called bipartite, whereas the more usually encountered MTN receptive fields, which could be characterized by a single direction-selective tuning curve, were called uniform. 3. Of the 16 neurons recorded from the VTRZ, all but one were binocular. For these neurons, both uniform and bipartite receptive fields were found for each eye alone. The only monocular neuron encountered in the VTRZ had a contralateral, bipartite receptive field. 4. The spatial organization of the direction selectivity of bipartite receptive fields strongly suggests that they are suited to represent rotation of the visual field about a horizontal axis located in the vertical plane that divides the receptive field into two parts. 5. The boundary between the two parts of the bipartite receptive fields was found using handheld visual stimuli at one of two azimuthal locations, either close to 45 degrees or between 95 and 125 degrees (the 0 degree reference was rostral in the midsagittal plane). This particular structure of the bipartite receptive fields suggests that their preferred rotation axes have a close spatial relation to the best-response axes of the semicircular canals. 6. Seven VTRZ neurons were antidromically activated by electrical stimulation of the ipsilateral dorsal cap of the inferior olive. Since the receptive fields of VTRZ neurons have many of the structural features characteristic of the receptive fields of rostral dorsal cap neurons we conclude that the spatial organization of the receptive fields of dorsal cap neurons is, for the most part, synthesized prior to the inferior olive.(ABSTRACT TRUNCATED AT 400 WORDS)

The rat inferior olive as seen with immunostaining for glutamate decarboxylase

Boutons presumed to use gamma-aminobutyric acid as neurotransmitter (GABAergic boutons) were detected by glutamate decarboxylase (GAD) immunocytochemistry in all regions of the rat inferior olive. The remarkably high concentration of these boutons allowed a clear visualization of olivary subnuclei boundaries. Regional variations in GAD immunostaining intensity were observed within the nuclear complex and were graded both visually and photometrically. The regional staining variations, for the most part, followed subnuclei boundaries and olivary zonal compartments that have been delineated by the topography of climbing fibre projections. Some subnuclei were grouped by similar staining intensities. The beta nucleus and a medial region in the ventral fold of the dorsal accessory olive were most intensely immunostained, followed by the subnucleus c of the medial accessory olive. Lower staining intensities were observed in the dorsomedial cell column, the dorsal fold of the dorsal accessory olive and the dorsal cap. The lowest intensities were observed in the subnuclei a and b of the medial accessory olive, the ventrolateral outgrowth, the rostral lamella of the medial accessory olive, the principal olive, and the lateral part of the ventral fold of the dorsal accessory olive. The factors contributing to the variations in immunostaining intensity (bouton size and frequency of occurrence) were investigated. The largest boutons were observed in the beta nucleus. Intermediate sized boutons were observed in the dorsomedial cell column, dorsal cap and the dorsal fold of the dorsal accessory olive. The smallest boutons were present in the remaining regions of the inferior olive, including the principal olive, the rostral lamella of the medial accessory olive, and the ventral fold of the dorsal accessory olive. The medial region of the dorsal accessory olive ventral fold contained a higher density of GABAergic boutons than other regions. GABAergic bouton size and innervation density therefore largely accounted for the variations in GAD immunostaining intensity. This study provides a map of the rat inferior olive based on the distribution of GABAergic nerve terminals, and may serve as a basis for characterizing different GABAergic afferent systems in the inferior olive.

Yaw direction neurons in the cat inferior olive.

1. Single units that responded to yaw rotation were recorded extracellularly in the caudal inferior olive (IO) of barbiturate-anesthetized cats. Of 276 neurons, 55 responded reliably to yaw, and extensive quantitative data were recorded from 25. 2. No yaw-sensitive IO neuron responded to somatosensory or auditory stimuli but two responded, though unreliably, to flash. 3. Yaw-sensitive IO cells fired at low (1-4 spikes/s), irregular rates during one direction of rotation. Though cells responded reliably during yaw, firing rates varied considerably from cycle to cycle. Rotation speed and acceleration were not represented in any cell's firing rate. 4. Eighty five percent (47/55) of yaw-sensitive cells fired during contralateral rotation, 9% (5/55) during ipsilateral rotation, and 6% (3/55) fired from late in the ipsilateral phase of a sinusoidal oscillation to the middle of the contralateral phase. 5. Responses were tested to 0.1-Hz sinusoidal yaw oscillations with a range of peak angular velocities (1-200 degrees/s). Thresholds were not sharp because of the cycle to cycle variability in response rates but were estimated using averaged responses. The peak rate of the most sensitive cell was driven to criterion (2 SD above spontaneous rate) by an oscillation with a peak velocity of 1 degrees/s. Other cells reached criterion between 5 and 50 degrees/s. 6. Sinusoidal oscillation at all frequencies tested (0.01-0.5 Hz) elicited approximately the same firing rates. Even at 0.01 Hz cells responded well. Responses lagged acceleration by approximately 25 degrees at 0.01 Hz and shifted to later parts of the cycle as frequency increased so that firing lagged acceleration by approximately 200 degrees at 0.5 Hz. 7. Histological reconstruction showed that yaw-sensitive neurons were recorded in olivary subnucleus beta (N beta), the dorsal cap of Kooy (DC), the posterior medial region of the medial accessory division of the inferior olive (MAO), and in the medial-lateral center of the caudal MAO. 8. Yaw-sensitive neurons in the inferior olive provide a signal to the cerebellum that indicates the direction of passive rotation over a wide range of velocity and acceleration. The signal from individual neurons does not reliably encode either rotation velocity or acceleration. Yaw-sensitive IO neurons are therefore unlike other central vestibular neurons but are similar to somatosensory IO cells which signal the presence, but not the intensity of a stimulus.

Topographical and ultrastructural investigation of the habenulo-interpeduncular pathway in the rat: a wheat germ agglutinin-horseradish peroxidase anterograde study.

The topographical and ultrastructural organization of the habenular projection to the interpeduncular nucleus (IPN) of the rat was examined employing the anterogradely transported tracer wheat germ agglutinin-horseradish peroxidase (WGA-HRP) and the chromogen tetramethylbenzidine (TMB). Unilateral placements of WGA-HRP in the habenular complex resulted in heavy terminal labelling in the rostral, central, and intermediate subnuclei bilaterally, and in the lateral subnuclei ipsilaterally. The apical subnucleus possessed only a sparse amount of label. Placements confined to the medial habenula (mH) produced similar results to those observed when the entire habenula was filled, suggesting that the afferent contribution made by the lateral habenula (lH) to the IPN is small. Unilateral placements of WGA-HRP in the dorsal portion of the mH resulted in heavy, predominantly ipsilateral labelling in the lateral subnucleus and the dorsal cap of the rostral subnucleus. In the lateral subnucleus labelled habenular terminals consistently contacted single dendritic processes shared by one or more other boutons, possibly of nonhabenular origin. Labelled habenular terminals in the rostral subnucleus normally contacted one or two dendrites. Labelled terminals in both subnuclei possessed clear, spherical vesicles and a variable number of dense-core vesicles. Unilateral placements of WGA-HRP in the ventral portion of the mH resulted in heavy labelling in the rostral half of the rostral subnucleus with a slight ipsilateral predominance, and in the central and intermediate subnuclei bilaterally. Terminal labelling was observed in crest and S synapses in the intermediate and central subnuclei respectively. Crest synapses, which consist of two parallel habenular terminals contacting an attenuated dendritic process, normally possessed label in only one of the two boutons. In the central subnucleus labelled horizontal axons formed several en passant S synapses with dendritic processes of small and medium diameter. These synaptic specializations of habenular axons contained numerous clear, spherical vesicles. This study demonstrates that a major topographically organized projection to the IPN originates from two distinct subpopulations of habenular neurons which comprise a dorsal sector and a ventral sector of the mH. Ultrastructural examination demonstrated that axons originating from neurons in the ventral and dorsal mH form characteristic contacts in the various IPN subnuclei.

Parasagittal zonal pattern of olivo-nodular projections in rabbit cerebellum

Injection of horseradish peroxidase (HRP) into the nodulus of the rabbit retrogradely labeled cells in 5 subdivisions of the contralateral inferior olive. Localized HRP injections at each of the medial, intermediate and lateral parts of the nodulus revealed 6 longitudinal zones, two zones in each part, projected differentially from the 5 olivary subdivisions: (i) the medial-most zone projected from the β nucleus, (ii) the lateral zone of the medial part and (iii) the lateral zone of the intermediate part both from the dorsal cap, (iv) the medial zone of the intermediate part from the ventrolateral outgrowth, (v) the medial zone of the lateral part from the dorsomedial cell column, and (vi) the lateral-most zone from the rostrolateral part of the medial accessory olive. A complication is that the zones (v) and (vi) seemed to cover the dorsal lamina of the nodulus, but not the ventral lamina. The lateral part of the ventral nodulus seemed to be projected from the dorsal cap and may therefore be a lateral extension of zone (iv). There was an indication that the rostral-most area of the medial accessory olive also projects to the nodulus, but a specific receptive zone for this projection was unclear. The present results suggest that the small lateral area of the nodulus previously found to be involved in cardiovascular control is projected from the rostrolateral part of the medial accessory olive and/or the dorsomedial cell column.

Zonal organization of olivo-nodulus projections in albino rabbits

The organization of inferior olivary projections to the cerebellar nodulus in albino rabbits was assessed by autoradiographic, anterograde degeneration and retrograde transport techniques. These data indicate that the caudal aspect of the dorsal cap of Kooy projects to a band extending 0.5–1 mm lateral to the midline of the nodulus. The medial half of this region receives a projection from β nucleus over at least the dorsal surface of the nodulus; an extension onto the ventral surface, though, is consistent with the anterograde tracing data. The rostral aspect of the dorsal cap and ventrolateral outgrowth projects to an adjacent 0.5–1. mm wide band in the nodulus. A group of cells spanning the intermediocaudal dorsal cap and the adjacent, dorsomedial margin of the β nucleus appears to project laterally on the ventral surface of the nodulus. On the dorsal aspect of the nodulus and ventral surface of lobule IXd, though, comparisons of anterograde and retrograde tracing data suggest that this lateral field is innervated by the rostral aspect of the dorsomedial cell column and the rostromedial accessory olive. Finally, the regions of lobules X and IXd lining the posterolateral fissure represent a transition between lobule IX and ventral lobule X patterns of olivary projections. These data provide a basis for investigating the efferent projections of the nodulus to distinct olivo-vestibular terminal fields in the vestibular nuclei.

Analysis of the rat interpeduncular subnuclei by immunocytochemical double‐staining for enkephalin and substance P, with some reference to the coexistence of both peptides

The rat interpeduncular nucleus (IPN) was immunocytochemically double-stained for enkephalin (ENK) and substance P (SP) on the same sections. On the basis of both peptidergic distribution patterns and topographic relationship, the IPN was divided into nine subnuclei and one cap: the rostral subnucleus (IP-R), the central subnucleus (IP-C), the rostral-lateral subnucleus (IP-RL), the main lateral subnucleus (IP-L), the caudal-lateral subnucleus (IP-CL), the dorsal-lateral subnucleus (IP-DL), the dorsal-medial subnucleus (IP-DM), the apical subnucleus (IP-A), the intermediate subnucleus (IP-I), and the dorsal cap (IP-Cap). As the descriptions of the IP-RL, IP-L, and IP-CL were inconsistent with previous reports, they were reevaluated; the IP-RL was proposed as the region situated in the lateral portion at rostral levels and characterized by the lack of ENK and SP immunoreactive structures, the IP-L as the region situated throughout the rostrocaudal extent in the lateral portion of the IPN and containing the highest density of SP immunoreactive fibers but no ENK immunoreactive fibers, and the IP-CL as the region situated just laterocaudal to the IP-L in the caudal pole of the IPN and containing ENK immunoreactive cells and fibers but no SP immunoreactive structures. Our results also showed that some cells in the IP-R have both ENK and SP immunoreactivity. This coexistence was observed in some small spherical cells of the IP-R, but rarely in larger oval-shaped cells, which occasionally showed only ENK immunoreactivity. In addition, paired ENK immunoreactive fiber bundles entering the IP-R were found to run just rostral to the paired SP immunoreactive columns, both of which composed parts of the interpedunculotegmental tract. A three-dimensional model representing the subnuclear organization of the IPN was proposed on the basis of the present results.

Dynamic characteristics of optokinetically controlled eye movements following inferior olive lesions in the brown rat.

1. The inferior olive was destroyed by the drug 3-acetylpyridine in brown rats. Spontaneous and optokinetic eye movements in response to constant-velocity rotation (5-80 deg/s) or sinusoidal oscillations (0.05 and 0.1 Hz with 15 deg/s peak velocity and 0.3, 0.5, 1.0 and 2 Hz with 5 deg/s peak velocity) of the visual surround were recorded 4-6 days, 40-50 days and 3-4 months after the lesion using the magnetic search coil technique. 2. Persistent oculomotor deficits were observed in rats with a lesion of more than 97% of inferior olive neurones. In cases with a less complete lesion, no or only transient deficits were observed. In these latter cases the bulk of surviving neurones was located in the caudal half of the inferior olive, which includes the dorsal cap of Kooy. 3. Eye position holding after saccadic gaze shifts in the light was strongly deficient, showing pronounced postsaccadic centripetal drift for several hundred milliseconds. Similar deficits were observed in slow-phase components following quick phases of optokinetic nystagmus. In the dark, eye position holding was also deficient. 4. Closed-loop gains of optokinetic step responses obtained from rats with inferior olive lesions could be as good as those obtained from control animals. There was, however, a trend towards smaller gain values over the range of stimulus velocities tested. The duration of optokinetic after-nystagmus was not changed. 5. The initial fast rise of slow-phase velocity of optokinetic step responses was reduced by about 30-50%, showing no recovery in the follow-up experiments up to 3-4 months after the lesion. 6. Optokinetic responses to sinusoidal oscillations of the visual surround exhibited an increasing drop in gain for frequencies between 0.1 to 0.5 Hz. In the range of 0.5-2.0 Hz gain was only about 0.2 compared to 0.7-0.8 in control animals. Phase lag of sinusoidal responses was shifted to larger values by about 25-35 deg for frequencies increasing from 0.1 to 0.5 Hz. At 1.0 Hz phase shift was reduced to about 15 deg and at 2.0 Hz no significant change in phase was observed. Both gain and phase of sinusoidal responses showed some recovery when tested 3-4 months after inferior olive lesion. 7. The results suggest that inferior olive lesions impair velocity-to-position integration, mainly as a consequence of the missing climbing fibre input to the cerebellar flocculi.(ABSTRACT TRUNCATED AT 400 WORDS)

Anatomical evidence for interaction of ACTH1?39 immunostained fibers and hypothalamic paraventricular neurons that project to the dorsal vagal complex

Anatomical evidence is presented for an interaction of ACTH1-39 immunostained fibers and a specific population of hypothalamic paraventricular (PVN) neurons; these neurons project to the dorsal vagal complex (DVC) of brainstem medulla. Bilateral injection of 10% HRP-WGA into DVC is incorporated into nerve terminals and transported retrogradely to cell bodies in the parvocellular subdivision of PVN, as revealed by standard HRP-WGA histochemistry or antibody to wheatgerm agglutinin followed by immunocytochemical techniques. Labeled cells are localized predominantly in the ventral portion of the caudal medial parvocellular subdivision and ventrolaterally in the posterior subnucleus of PVN. Few labeled cells are seen in the anterior parvocellular PVN, rostrally in the medial parvocellular component and in the dorsal cap. HRP-WGA cells are rarely observed in the magnocellular divisions of PVN. Dual-staining immunocytochemical-retrograde tracing techniques in the same tissue section demonstrate ACTH1-39 fibers in intimate anatomical proximity to parvocellular PVN neurons that project to DVC. It is suggested that this interaction may partially account for the known cardiovascular effects of opiocortins and supports the role of the paraventricular nucleus in hypothalamic integration and modulation of cardiovascular control.

Distribution of inferior olivary projections to the vestibular nuclei of albino rabbits

This study analyses the course and topography of olivo-vestibular projections originating in the dorsal cap, ventrolateral outgrowth and beta nucleus of albino rabbits. Rabbits were given either single pressure-injections of [ 3H] l-leucine (20 μCi in 50 nl) or single or multiple injections of 3-acetylpyridine (0.2–0.25 μ1 of 27.5μg/μ1 in saline) into the medial aspect of the inferior olive. Brains from the former animals were processed for autoradiography after 2–3 days survival; brains from the latter animals were stained for degeneration with cupric-silver methods after a 16–24 h survival. In addition, four rabbits with kainic acid lesions of the flocculus were used to document flocculo-vestibular projections. Olivo-vestibular projections from the dorsal cap-ventrolateral outgrowth, beta nucleus and the medial accessory olive diverge from olivo-cerebellar projections at the caudal margin of the flocculus stalk, and course medially in a broad sheet. Fibers (1) ascend in the superior fascicle, with flocculo-vestibular projections, to the superior vestibular nucleus, (2) enter the medial fascicle, with flocculo-vestibular fibers, and course along the dorsolateral border of the 4th ventricle to innervate a distinct rostral subdivision of the medial vestibular nucleus, and (3) enter the lateral fascicle, with flocculo-vestibular fibers, to terminate in pars α and β of the lateral vestibular nucleus and the caudal subdivision of the medial vestibular nucleus. Comparison of different injection cases indicate that the caudal half to two-thirds of the dorsal cap contributes projections to the rostral medial vestibular nucleus, centrolateral and dorsomedial aspects of the superior vestibular nucleus, and a projection to both central and dorsal aspects of the caudal medial vestibular nucleus. By contrast, the rostral third to half of the dorsal cap-ventrolateral outgrowth projects sparsely to the rostral medial vestibular nucleus, contributing dense projections to the central aspect of the superior vestibular nucleus and dorsomedial and lateral regions in the caudal medial vestibular nucleus. Pars β of the lateral vestibular nucleus received densest projections from the intermediate third of the dorsal cap; the projection to pars α did not display a distinct topography. These data and results of previous retrograde tracing experiments [C.D. Balaban (1984) Neuroscience 12, 129–149] suggest that distinct groups of dorsal cap-ventrolateral outgrowth neurons project differentially to the vestibular nuclei in rabbits. Comparisons with previous electrophysiologic studies raise the testable hypothesis that they demarcate distinct pools of vestibular nuclear output neurons.

Thalamic projections to fields A, AI, P, and VP in the cat auditory cortex.

Thalamocortical projections to four tonotopic fields (A, AI, P, and VP) of the cat auditory cortex were studied by using combined microelectrode mapping and retrograde axonal transport techniques. Horseradish peroxidase (HRP) or HRP combined with either tritiated bovine serum albumin or nuclear yellow was injected into identified best-frequency sites of one or two different fields in the same brain. Arrays of labeled neurons were related to thalamic nuclei defined on the basis of their cytoarchitecture and physiology. In some cases, patterns of labeling were directly compared with thalamic best-frequency maps obtained in the same brain. We compared only patterns of labeling resulting from injections into similar parts of the frequency representation in different fields to insure that observed differences in patterns of labeling did not simply reflect differences in the frequency representation at the injection sites. The thalamic projection to the four fields is divided among seven nuclei, three tonotopic nuclei (ventral nucleus, V; lateral part of the posterior group of thalamic nuclei, Po; and dorsal cap nucleus, d) and four nontonotopic nuclei (caudodorsal nucleus, cd; ventrolateral nucleus, vl; and small, Ms; and medium-large, Mg, cell regions of the medial division). Projections to each field differ, and each field receives inputs from tonotopic and nontonotopic nuclei. Field A receives its major inputs from Po and Mg, and a minor input from V. Field AI receives its major inputs from V, Po, and Mg, although Po and Mg have heavier projections to field A. Field P receives its major inputs from V, d, and vl; and minor inputs from cd, Ms, Mg, and Po. Field VP receives major inputs from V, vl, and cd; and minor inputs from d, Ms, and Mg. There are segregated territories in V and Po in which most neurons projects to one cortical field (major projection), and a smaller proportion projects to one or more other fields (minor projections). Field VP receives a major projection from the caudal pole of V. Field P receives a major projection from the caudal half of V, and from a thin band along the dorsal border of rostral V. Field AI receives a major projection from most of the rostral one-half of V, and smaller areas in Po and the caudal half of V exclusive of its caudal pole. Field A receives a major projection from most of Po.(ABSTRACT TRUNCATED AT 400 WORDS)

The olivonodular projection: A re-examination based on folial cerebellar implants

In 5 cats implants of crystalline wheat germ agglutinin-horseradish peroxidase complex were placed in one or two folia of the nodulus with no contamination of adjacent cerebellar lobules. In the inferior olive only the dorsal cap with the adjacent ventrolateral outgrowth and the caudalmost part of nucleus β were found to project to the nodulus. Negative findings were consistently made in all other parts of the inferior olive, some of which have previously been described to contribute to the olivonodular projection.

Inferior olivary nuclear complex of the rat: Morphology and comments on the principles of organization within the olivocerebellar system

This study was undertaken (1) to provide a description of the normal morphology and anatomical interrelationships within the inferior olivary cell groups and (2) to determine the topographical organization of projections from this nuclear complex to the cerebellum. Conventional histological methods and the technique of retrograde transport of horseradish peroxidase (HRP) and WGA-HRP in conjunction with the sensitive chromogen tetramethyl benzidine were used in this series of experiments. In common with that of other animals, the inferior olivary nucleus of the rat consists of three subdivisions: the medial accessory olive (MAO), dorsal accessory olive (DAO), and principal olive (PO). The MAO is made of several subnuclei including a, b, c, and nucleus beta. On the basis of their common connections, the smaller subnuclei, dorsal cap, ventrolateral outgrowth, and dorsomedial cell column can be considered as parts of the MAO. The DAO is made of two subdivisions or lamellae--the dorsal and ventral folds--joined together laterally to form a twisted V-shaped structure. The principal olive consists of dorsal and ventral lamellae. A point-to-point projection was determined for four areas of the cerebellum: the anterior lobe, the posterior vermis, the intermediate and the lateral cerebellum. Analysis of the details of the projection system revealed that distinct groups of cells, here referred to as lamellae, project to sagittal zones of the cerebellum. (1) The medial accessory olive appears to be composed of three lamellae: horizontal, vertical, and rostral. The horizontal lamella (elsewhere denoted as groups a and b) projects to a sagittal zone in the vermal anterior lobe. The vertical lamella (groups c, beta, dorsal cap, ventrolateral outgrowth, and dorsomedial cell column) projects to a sagittal zone in the posterior vermis and the flocculus, and the rostral lamella projects to the lateral cerebellum. (2) The dorsal accessory olive is composed of two distinct lamellae which we have previously denoted as the dorsal and ventral folds. The dorsal fold projects to the vermal anterior lobe and receives afferents from the spinal cord, whereas the ventral fold projects to a sagittal zone in the intermediate cerebellum and receives afferents primarily from dorsal column nuclei. (3) The principal olive contains the two familiar lamellae: dorsal and ventral lamellae, each of which projects to a specific sagittal strip in the lateral cerebellum.(ABSTRACT TRUNCATED AT 400 WORDS)

Topographical organization of olivocerebellar and corticonuclear connections in the rat--an WGA-HRP study: I. Lobules IX, X, and the flocculus.

The organization of the olivocerebellar and corticonuclear relations for vermal lobules IX and X and the flocculus has been studied in the rat by using microinjections of wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP). This axonal tracer allowed us to study simultaneously the olivocortical connections (revealed by retrograde transport) and corticonuclear connections (revealed by anterograde transport) from a single injection in the cerebellar cortex. The results indicate that four modules can be distinguished, each of which consists of a region of cerebellar cortex receiving afferents from a single small region of the inferior olive (IO) and sending efferents to one or several portions of the cerebellar nuclei and/or vestibular nuclei. The first module includes a medial part of lobule X as well as all the flocculus. It receives afferents from the dorsal cap (dc) and sends efferents to the small cell (s) zone of the dentate nucleus as well as to the medial vestibular (VM) nucleus and subnucleus y. The second module includes a medial parasagittal region located in lobules IX and X. It receives afferents from the ventrolateral outgrowth (vlo) and/or beta nucleus (vlo + beta nucleus) and sends efferents principally to the ventrolateral part of fastigial nucleus and to the superior vestibular (VS), inferior vestibular (VI), and VM nuclei. The third module includes a lateral parasagittal region in lobules IX and X. It receives afferents from the dorsomedial cell column (dmcc) of IO and sends efferents principally to the interpositus nucleus and subnucleus y. The fourth module includes the most lateral part of lobules IX and Xa. It receives afferents from the principal olive (PO) and sends efferents to the s zone of the dentate nucleus. These results are comparable to those obtained in the cat although a few differences are discussed.

Autoradiographic study of descending pathways from the pontine reticular formation and the mesencephalic trigeminal nucleus in the rat.

Descending projections were studied in autoradiographically prepared material after injections of tritiated leucine in the pontine tegmentum of rats. Injections involving the medial pontine reticular formation resulted not only in labeling commissural fibers, the medial reticulospinal tract, and the dorsal cap of the inferior olive, but also, in two cases, in labeling a cerebellar projection that originated from a region near the midline and clearly dorsal to the nucleus reticularis tegmenti pontis. The labeled fibers passed ventral in the midline to the pontine gray, then laterally through the gray and into the middle cerebellar peduncle to terminate as mossy fibers primarily in the flocculus, lobulus simplex, and Crus I of the ansiform lobule. Injections involving the mesencephalic nucleus of the trigeminal nerve (Vmes), resulted in labeling of Probst's tract, which descends in the dorsolateral reticular formation. Probst's tract gave off extensive terminal branches to the lateral medullary reticular formation and weaker projections to restricted portions of the descending trigeminal nucleus, the solitary nucleus, and the hypoglossal nucleus. In one case, fibers could be traced into the dorsal horn of the upper cervical cord.

Ventroposterior thalamic regions projecting to cytoarchitectonic areas 3a and 3b in the cat.

The adequate stimulus and body site that excited neurons in cat cortical somatosensory areas 3a and 3b were recorded using low-impedance tungsten microelectrodes. Horizontal penetrations provided a good correlation between the electrophysiological and cytoarchitectonic data. Responses best driven by cutaneous stimuli were replaced with responses driven by manipulation of deep tissue at, or very near, the border between areas 3a and 3b. Following functional identification of these areas horseradish peroxidase was injected into one of them. Injections into area 3a labeled neurons in a rostral and dorsal cap of the ventroposterior thalamus. It was suggested that this region is a distinct nucleus termed the ventroposterior oralis nucleus (VPO). Injections into the forelimb portion of area 3b labeled neurons in the ventroposterior lateral nucleus (VPL). With both vertical and horizontal microelectrode trajectories through the ventroposterior thalamic nuclei, inputs from deep structures presumed to be muscles were consistently located in the VPO nucleus and cutaneous inputs activated neurons in the VPL. The existence of several functionally-distinct subdivisions within the somatosensory nuclei of the thalamus supports the hypothesis of parallel processing and relay of somatosensory information at this level of the pathway.