Dorsal Paraflocculus Research Articles

The entire trajectories of single pontocerebellar axons and their lobular and longitudinal terminal distribution patterns in multiple aldolase C-positive compartments of the rat cerebellar cortex.

The mammalian cerebellar cortex is compartmentalized, both anatomically and histochemically, into multiple parasagittal bands. To characterize the multiple zonal patterns of pontocerebellar mossy fiber projection, single neurons in the basilar pontine nucleus (BPN) were labeled by injecting biotinylated dextran amine into the BPN, and the entire axonal trajectory of single labeled neurons (n = 25) was reconstructed in relation to aldolase C compartments of Purkinje cells in rats. Single pontocerebellar axons, after passing through the contralateral middle cerebellar peduncle, ran transversely in the deep cerebellar white matter toward and often across the midline, and on their ways, gave rise to 2-10 primary collaterals at almost right angles in specific lobules only contralaterally or bilaterally with contralateral predominance. Each primary collateral further branched in a parasagittal plane to form a strip-shaped longitudinal termination zone with rosette-type swellings clustered in aldolase C-positive compartments in a single or multiple lobules, mainly in compartment 4+//5+, 5+//6+, and 6+//7+. Axons arising from the central, rostral, and lateral part of the BPN projected with multiple branches, mainly to simple lobule, crus II and paramedian lobule, to crus I and dorsal paraflocculus, and to ventral paraflocculus and lobule IXc, respectively. The results showed the pontocerebellar projection is closely related to lobular and compartmental organization of the cerebellum. A comparison of single axon morphologies of different mossy fiber systems indicates that the projection pattern of single pontocerebellar neurons with multiple collaterals innervating different longitudinal compartments arranged in a mediolateral direction represents a general feature of mossy fiber projection.

Compartmentation of the Cerebellar Cortex in the Naked Mole-Rat (Heterocephalus glaber)

Despite the apparent uniformity in cellular composition of the adult mammalian cerebellar cortex, it is actually highly compartmentalized into transverse zones and within each zone further subdivided into a reproducible array of parasagittal stripes. This basic cerebellar architecture is highly conserved in birds and mammals. However, different species have very different cerebellar morphologies, and it is unclear if cerebellar architecture reflects taxonomic relations or ecological niches. To explore this, we have examined the cerebellum of the naked mole-rat Heterocephalus glaber, a burrowing rodent with adaptations to a subterranean life that include only a rudimentary visual system. The cerebellum of H. glaber resembles that of other rodents with the remarkable exception that cerebellar regions that are prominent in the handling of visual information (the central zone, nodular zone, and dorsal paraflocculus) are greatly reduced or absent. In addition, there is a notable increase in size in the posterior zone, consistent with an expanded role for the trigeminal somatosensory system. These data suggest that cerebellar architecture may be substantially modified to serve a particular ecological niche.

Visuomotor cerebellum in human and nonhuman primates.

In this paper, we will review the anatomical components of the visuomotor cerebellum in human and, where possible, in non-human primates and discuss their function in relation to those of extracerebellar visuomotor regions with which they are connected. The floccular lobe, the dorsal paraflocculus, the oculomotor vermis, the uvula–nodulus, and the ansiform lobule are more or less independent components of the visuomotor cerebellum that are involved in different corticocerebellar and/or brain stem olivocerebellar loops. The floccular lobe and the oculomotor vermis share different mossy fiber inputs from the brain stem; the dorsal paraflocculus and the ansiform lobule receive corticopontine mossy fibers from postrolandic visual areas and the frontal eye fields, respectively. Of the visuomotor functions of the cerebellum, the vestibulo-ocular reflex is controlled by the floccular lobe; saccadic eye movements are controlled by the oculomotor vermis and ansiform lobule, while control of smooth pursuit involves all these cerebellar visuomotor regions. Functional imaging studies in humans further emphasize cerebellar involvement in visual reflexive eye movements and are discussed.

Mossy and climbing fiber collateral inputs in monkey cerebellar paraflocculus lobulus petrosus and hemispheric lobule VII and their relevance to oculomotor functions

Recent lesion studies on monkeys suggest that the cerebellar lobulus petrosus of the paraflocculus (LP) and crura I and II of hemispheric lobule VII (H-7) are involved in smooth pursuit eye movement control. To reveal the relationship between the LP and H-7, we studied mossy and climbing fiber collateral inputs to these areas in four cynamolgus monkeys. After unilateral injections of retrograde tracers into the LP, labeled mossy fibers were seen ipsilaterally in the crura I and II of H-7. A very small number of labeled mossy fiber collaterals were also seen in the dorsal paraflocculus (DP). Labeled climbing fibers were seen exclusively in the ipsilateral crus I. No labeled mossy/climbing fibers were seen in the flocculus, ventral paraflocculus and other cortical areas. Combined injections of fast blue in the LP and cholera toxin subunit B in the posterior crus I and crus II of H-7 resulted in a small number of the double-labeled pontine and principal olivary neurons. Combined injections in the LP and DP induced only a few double-labeled neurons in the pontine nuclei, and no double-labeled neurons in the olivary nuclei. These results suggest that the LP and crura I and II of H-7 may share some of their mossy and climbing fiber inputs and mediate similar functional roles involving smooth pursuit eye movement control.

Functional localization in the cerebellum

In this paper we review the basic anatomy and functional localization in the cerebellum. Experimental anatomical studies emphasize the predominance of skeletomotor and visuomotor connections. Parietal lobe visual inputs arise principally from the dorsal stream extrastriate visual areas, which are specialized for the visual control of movement. There are few or no inputs to the cerebellum from inferotemporal cortex. Much of the input from prefrontal cortex is from areas that control eye movements. Comparative anatomical studies of the hominoid dentate nucleus are consistent with the role of much of the cerebellar hemispheres in the visual guidance of movement. Although some mossy and climbing fibre afferents to the cerebellum are reciprocally organized, feeding back onto their original source, the reciprocity does not exist for the visuomotor division. The dorsal paraflocculus receives its mossy fibres from extrastriate areas of the dorsal visual stream, and projects to the frontal eye fields. Functional magnetic resonance imaging (fMRI) studies in human subjects confirm the presence of somatotopically organized anterior and posterior skeletomotor areas with eye movement activity centred in the oculomotor vermis. Oculomotor and skeletomotor representation extend into adjacent Crus I and II. Discrepancies between the results of imaging studies in human subjects and experimental data are discussed. Skeletomotor activity or eye movements may contaminate many fMRI studies of putative cognitive functions of the cerebellum.

Cerebellar Inputs to Intraparietal Cortex Areas LIP and MIP: Functional Frameworks for Adaptive Control of Eye Movements, Reaching, and Arm/Eye/Head Movement Coordination

Using retrograde transneuronal transfer of rabies virus in combination with a conventional tracer (cholera toxin B), we studied simultaneously direct (thalamocortical) and polysynaptic inputs to the ventral lateral intraparietal area (LIPv) and the medial intraparietal area (MIP) in nonhuman primates. We found that these areas receive major disynaptic inputs from specific portions of the cerebellar nuclei, the ventral dentate (D), and ventrolateral interpositus posterior (IP). Area LIPv receives inputs from oculomotor domains of the caudal D and IP. Area MIP is the target of projections from the ventral D (mainly middle third), and gaze- and arm-related domains of IP involved in reaching and arm/eye/head coordination. We also showed that cerebellar cortical "output channels" to MIP predominantly stem from posterior cerebellar areas (paramedian lobe/Crus II posterior, dorsal paraflocculus) that have the required connectivity for adaptive control of visual and proprioceptive guidance of reaching, arm/eye/head coordination, and prism adaptation. These findings provide important insight about the interplay between the posterior parietal cortex and the cerebellum regarding visuospatial adaptation mechanisms and visual and proprioceptive guidance of movement. They also have potential implications for clinical approaches to optic ataxia and neglect rehabilitation.

Independent roles for the dorsal paraflocculus and vermal lobule VII of the cerebellum in visuomotor coordination

Two distinct areas of cerebellar cortex, vermal lobule VII and the dorsal paraflocculus (DPFl) receive visual input. To help understand the visuomotor functions of these two regions, we compared their afferent and efferent connections using the tracers wheatgerm agglutinin horseradish peroxidase (WGA-HRP) and biotinilated dextran amine (BDA). The sources of both mossy fibre and climbing fibre input to the two areas are different. The main mossy fibre input to lobule VII is from the nucleus reticularis tegmenti pontis (NRTP), which relays visual information from the superior colliculus, while the main mossy fibre input to the DPFl is from the pontine nuclei, relaying information from cortical visual areas. The DPFl and lobule VII both also receive mossy fibre input from several common brainstem regions, but from different subsets of cells. These include visual input from the dorsolateral pons, and vestibular-oculomotor input from the medial vestibular nucleus (MVe) and the nucleus prepositus hypoglossi (Nph). The climbing fibre input to the two cerebellar regions is from different subdivisions of the inferior olivary nuclei. Climbing fibres from the caudal medial accessory olive (cMAO) project to lobule VII, while the rostral MAO (rMAO) and the principal olive (PO) project to the DPFl. The efferent projections from lobule VII and the DPF1 are to all of the recognised oculomotor and visual areas within the deep cerebellar nuclei, but to separate territories. Both regions play a role in eye movement control. The DPFl may also have a role in visually guided reaching.

Corticopontocerebellar pathway from the prearcuate region to hemispheric lobule VII of the cerebellum: an anterograde and retrograde tracing study in the monkey

The relationship between axons derived from the prearcuate region and pontine neurons projecting to contralateral cerebellar hemispheric lobule VII, lobulus petrosus (LP) of the paraflocculus or the dorsal paraflocculus (DPFl) was investigated in the monkey. The frontopontine axons were labeled with biotinylated dextran and pontocerebellar neurons with cholera toxin subunit B or fast blue. Labeled frontopontine axons and terminals were seen in the dorsal, medial, paramedian and dorsolateral parts of the pontine nuclei. The distribution of the labeled frontopontine axons overlapped that of labeled neurons projecting to hemispheric lobule VII but did not overlap that of labeled neurons projecting to LP or DPFl. The pathway from the prearcuate region to hemispheric lobule VII may provide an anatomical substrate for the involvement of hemispheric lobule VII in voluntary eye movements.

Cerebellar lesions and prism adaptation in macaque monkeys.

If a laterally displacing prism is placed in front of one eye of a person or monkey with the other eye occluded, they initially will point to one side of a target that is located directly in front of them. Normally, people and monkeys adapt easily to the displaced vision and correct their aim after a few trials. If the prism then is removed, there is a postadaptation shift in which the subject misses the target and points in the opposite direction for a few trials. We tested five Macaque monkeys for their ability to adapt to a laterally displacing prism and to show the expected postadaptation shift. When tested as normals, all five animals showed the typical pattern of adaptation and postadaptation shift. Like human subjects, the monkeys also showed complete interocular transfer of the adaptation but no transfer of the adaptation between the two arms. When preoperative training and testing was complete, we made lesions of various target areas on the cerebellar cortex. A cerebellar lesion that included the dorsal paraflocculus and uvula abolished completely the normal prism adaptation for the arm ipsilateral to the lesion in one of the five monkeys. The other four animals retained the ability to prism-adapt normally and showed the expected postadaptation shift. In the one case in which the lesion abolished prism adaptation, the damage included Crus I and II, paramedian lobule and the dorsal paraflocculus of the cerebellar hemispheres as well as lobule IX, of the vermis. Thus in this case, the lesion included virtually all the cerebellar cortex that receives mossy-fiber visual information relayed via the pontine nuclei from the cerebral cortex. The other four animals had damage to lobule V, the classical anterior lobe arm area and/or vermian lobules VI/VII, the oculomotor region. When tested postoperatively, some of these animals showed a degree of ataxia equivalent to that of the case in which prism adaptation was affected, but prism adaptation and the postadaptation shift remained normal. We conclude that in addition to its role in long-term motor learning and reflex adaptation, the region of the cerebellum that was ablated also may be a critical site for a short-term motor memory. Prism adaptation seems to involve a region of the cerebellum that receives a mossy-fiber visual error signal and probably a corollary discharge of the movement.

Differences in mossy and climbing afferent sources between flocculus and ventral and dorsal paraflocculus in the rat.

Sources of mossy and climbing fiber inputs to the flocculus (FL), ventral paraflocculus (VP) and/or dorsal paraflocculus (DP) were identified in the vestibular ganglion, medulla oblongata and pons of 19 Wistar rats after 26 local injections of horseradish peroxidase, wheat-germ agglutinin-conjugated horseradish peroxidase, fast blue or diamidino yellow into the FL, VP and/or DP. There were large differences in the sources of mossy fibers to the FL and VP/DP. Labeled neurons after injections into the FL were observed mainly in the ipsilateral vestibular ganglion, bilaterally in the vestibular and prepositus hypoglossal nuclei, and in the caudal part of the nucleus reticularis tegmenti pontis. Labeled neurons were rarely observed in the pontine nuclei after localized injections into the FL. By contrast, after injections into the VP and/or DP, numerous labeled neurons were observed in the pontine nuclei with a contralateral predominance and in the rostral part of the nucleus reticularis tegmenti pontis bilaterally, but not in the vestibular nuclei in either side. Sources of climbing fibers to the FL and paraflocculus were completely contralateral to the injection side. After injection into the FL, labeled neurons were observed in the caudal dorsal cap and ventrolateral outgrowth of the inferior olivary nucleus. After injections into the VP, labeled neurons were observed mainly in the rostral dorsal cap, ventral medial accessory olivary nucleus (MAO) and caudal half of the ventral leaf of the principal olivary nucleus. After injections into the DP, labeled neurons were observed in the ventral MAO and caudal half of the ventral leaf of the principal olivary nucleus. These differences in the sources of mossy and climbing fiber inputs may suggest functional differences between the FL and VP/DP. The present results are consistent with our previous observations in monkey that the FL and VP/DP exhibit quite different mossy fiber input organizations.

Anatomical compartments in the white matter of the rabbit flocculus

The white matter of the rabbit flocculus is subdivided into five compartments by narrow sheets of densely staining acetylcholinesterase-positive fibers. The most lateral compartment is continuous with the C2 compartment of the paraflocculus and contains the posterior interposed nucleus. The other four compartments are numbered from lateral to medial as floccular compartments 1, 2, 3, and 4 (FC1-4). FC1-3 continue across the posterolateral fissure into the adjacent folium (folium p) of the ventral paraflocculus. FC4 is present only in the rostral flocculus. In the caudal flocculus FC1 and FC3 abut dorsal to FC2. Fibers of FC1-4 can be traced into the lateral cerebellar nucleus and the floccular peduncle. The presence of acetylcholinesterase in the deep stratum of the molecular layer of the flocculus and ventral paraflocculus distinguishes them from the dorsal paraflocculus. The topographical relations to the flocculus and the floccular peduncle with group y and the cerebellar nuclei are discussed.

Zonal organization of the climbing fiber projection to the flocculus and nodulus of the rabbit: a combined axonal tracing and acetylcholinesterase histochemical study.

The localization and termination of olivocerebellar fibers in the flocculus and nodulus of the rabbit were studied with anterograde axonal transport methods [wheatgerm agglutinin-horseradish peroxidase (WGA-HRP) and tritiated leucine] and correlated with the compartments in the white matter of these lobules delineated with acetylcholinesterase histochemistry (Tan et al. J. Comp. Neurol., 1995, this issue). Olivocerebellar fibers originating from the caudal dorsal cap travel through floccular compartments FC2 and FC4 to terminate as climbing fibers in floccular zones FZII and FZIV. Fibers from the rostral dorsal cap and the ventrolateral outgrowth traverse compartments FC1 and FC3, which are interleaved with compartments FC2 and FC4, and terminate in zones FZI and FZIII. Fibers from the rostral pole of the medial accessory olive traverse the C2 compartment and terminate in the C2 zone. FZI-III extend into the adjoining folium (folium p) of the ventral paraflocculus. The C2 zone continues across folium p into other folia of the ventral paraflocculus and into the dorsal paraflocculus. Four compartments and five zones were distinguished in the nodulus. Medial compartment XC1 contains olivocerebellar fibers from the caudal dorsal cap and subnucleus beta that terminate in the XZI zone. Olivocerebellar fibers from the rostral dorsal cap and the ventrolateral outgrowth occupy XC2 and terminate in XZII. The XC4 compartment contains fibers from both the caudal dorsal cap and from the rostral dorsal cap and the ventrolateral outgrowth. The latter terminate in a central portion of the XZIV zone. The dorsomedial cell column projects to the XZIII zone, which is present only in the dorsal part of the nodulus. The rostral medial accessory olive projects to the XZV zone, which occupies the lateral border of the nodulus. These results confirm and extend the conclusions of Katayama and Nisimaru ([1988] Neurosci. Res. 5:424-438) on the zonal pattern in the olivo-nodular projection in the rabbit. Additional observations were made on the presence of a lateral A zone (Buisseret-Delmas [1988] Neurosci. Res. 5:475-493) in the hemisphere of lobules VI and VII. Retrograde labeling of the nucleo-olivary tract of Legendre and Courville ([1987] Neuroscience 21:877-891) was observed after WGA-HRP injections into the inferior olive including the rostral dorsal cap and the ventrolateral outgrowth. The anatomical and functional implications of these observations are discussed.

Visual pontocerebellar projections in the macaque

The cerebellum plays an important role in the visual guidance of movement. In order to understand the anatomical basis of visuomotor control, we studied the projection of pontine visual cells onto the cerebellar cortex of monkeys. Wheat germ agglutinin horseradish peroxidase was injected into the dorsolateral pons two monkeys. Retrogradely labelled cells were mapped in the cerebral cortex and superior colliculus, and orthogradely labelled fibers in the cerebellar cortex. The largest number of retrogradely labelled cells in the cerebral cortex was in a group of medial extrastriate visual areas. The major cerebellar target of these dorsolateral pontine cells is the dorsal paraflocculus. There is a weaker projection to the uvula, paramedian lobe, and Crus II, and a sparse but definite projection to the ventral paraflocculus. There are virtually no projections to the flocculus. There are sparse ipsilateral pontocerebellar projections to these same regions of cerebellar cortex. In nine monkeys, we made small injections of the tracer into the cerebellar cortex and studied the location of retrogradely filled cells in the pontine nuclei and inferior olive. Injections into the dorsal paraflocculus or rostral folia of the uvula retrogradely labelled large numbers of cells in the dorsolateral region of the contralateral pontine nuclei. Labelled cells were found ipsilaterally, but in reduced numbers. Injections outside of these areas in ventral paraflocculus or paramedian lobule labelled far fewer cells in this region of the pons. We conclude that the principal source of cerebral cortical visual information arises from a medial group of extrastriate visual areas and is relayed through cells in the dorsolateral pontine nuclei. The principal target of pontine visual cells is the dorsal paraflocculus.

The unipolar brush cells of the rat cerebellar cortex and cochlear nucleus are calretinin-positive: a study by light and electron microscopic immunocytochemistry.

Cell class-specific markers are powerful tools for the study of individual neuronal populations. The peculiar unipolar brush cells of the mammalian cerebellar cortex have only recently been definitively identified by means of the Golgi method, and we have explored markers of cerebellar neurons with the purpose of facilitating the analysis of this new cell population and, especially, its distribution and ultrastructural features. By light microscopic immunocytochemistry, we demonstrate that, in the rat, the unipolar brush cells are the cortical neurons that are most densely immunostained with antiserum to calretinin, a recently discovered calcium-binding protein. The unipolar brush cells are highly concentrated in the flocculo-nodular lobe, the ventral uvula and the ventral paraflocculus, occur at relatively high density in the lingula, at moderate-to-low density in other folia of the vermis and in the narrow intermediate cortex, and at low to very low density, with the exception of a few hot spots, in the lateral regions of the cerebellar hemispheres and in the dorsal paraflocculus. Unipolar brush cells are also found in the cochlear nucleus. In addition to the unipolar brush cells, calretinin antibody distinctly stains certain mossy fibers, and weakly to moderately stains other cerebellar elements, such as granule neurons and climbing fibers. In the lobules containing high densities of unipolar brush cells, the granule cell bodies and the parallel fibers are much less immunoreactive, and there are many more densely immunostained mossy fibers than in the lobules, where these cells are rare, which suggests some relationships between these elements. In the cerebellar nuclei, small neurons are densely immunostained, while large neurons are immunonegative. The unipolar brush cells reside nearly exclusively in the granular layer. They are small neurons, intermediate in size between granule cells and Golgi cells, and their features are remarkably similar across all lobules. They usually have a single, relatively thick dendrite of varying length that terminates in a brush-like tip consisting of several short branchlets. Utilizing a pre-embedding protocol, we have identified unipolar brush cells with the electron microscope. The cytoplasm of these cells is partially obscured by the electron dense product of calretinin immunoreaction in all regions of the soma and processes. The cells are often covered with non-synaptic appendages and contain a peculiar cytoplasmic inclusion consisting of ringlet subunits. Other characteristic components are numerous neurofilaments, mitochondria and large, dense-core vesicles. Individual brushes enter one or two glomeruli, where the dendritic branchlets establish an unusually extensive synapse with mossy fiber rosettes.(ABSTRACT TRUNCATED AT 400 WORDS)

Topographic organization of the corticonuclear projections from the paraflocculus in the albino rat: an autoradiographic orthograde tracing study.

The topographic organization of corticonuclear projections from the paraflocculus was studied in rats by an autoradiographic orthograde tracing method. The corticonuclear fibers from the paraflocculus terminated primarily in the caudoventral part of the lateral cerebellar nucleus and in the lateral part of the posterior interpositus nucleus. The rostrolateral part of the ventral paraflocculus projected only to the lateral nucleus, and the caudomedial part projected only to the posterior interpositus nucleus. Although differential projections from the dorsal and ventral paraflocculi were not clearly observed, the dorsal paraflocculus appeared to project to slightly more dorsal parts of the lateral and posterior interpositus nuclei than did the ventral paraflocculus. The lateral, medial, rostral, and caudal parts of each sublobule tended to project to the ventral, dorsal, rostral, and caudal regions within the caudoventral part of the lateral nucleus, respectively, despite massive overlap in the individual terminal fields. The corticonuclear projection from the dorsal and ventral paraflocculi to the posterior interpositus nucleus were organized in different directions; those from the dorsal paraflocculus were organized rostrocaudally, while those from the ventral paraflocculus were organized mediolaterally. These results suggest that in rats the corticonuclear projections from the paraflocculus are topographically organized in the mediolateral and rostrocaudal directions as well as in the dorsoventral direction.

Neuronal activity in the lateral cerebellum of trained monkeys, related to visual stimuli or to eye movements.

1. The responses of neurones in the lateral cerebellar cortex to visual stimuli and to eye movements were recorded in rhesus monkeys trained to perform visually guided arm and eye movements in a tracking task. 2. Twenty-two of 134 units recorded (16%) modulated their discharge in response to a bright Xenon flash. They were mainly located in the dorsal paraflocculus. Among those identified as Purkinje cells both simple spike and climbing fibre responses to the flash were seen. (72% of the units were related to arm movements; these were centred in the paramedian lobule, and have been described fully in Marple-Horvat & Stein (1987).) 3. The visual responsiveness of one of the units varied according to the phase of the monkey's task. Around the time that the target stepped, which was the monkey's cue to move, its sensitivity to other stimuli disappeared. 4. Only two neurones responded to the movements of the tracking target. These responses were conditional upon the monkey using visual signals to guide his movements; they did not respond to the target step if he moved before the target did. 5. Fourteen units (10%) located in crus I and II and lobulus simplex correlated strongly with the velocity of horizontal eye movements. Only one of these also responded to visual stimuli. 6. Thus most neurones were found to carry only visual, or eye movement, or limb movement information rather than combinations of these signals; they were located in different but overlapping regions of lateral cerebellar cortex. Visually responsive neurones are probably involved in planning the visual goal of movements, while eye and arm movement neurones probably help to create co-ordinative structures for executing voluntary eye and arm movements.

Organization of the cortico-ponto-cerebellar pathway to the dorsal paraflocculus. An experimental study with anterograde and retrograde transport of WGA-HRP in the cat.

To reveal the organization and relative magnitude of connections from various parts of the cerebral cortex to the dorsal paraflocculus via the pontine nuclei, WGA-HRP was injected in the dorsal paraflocculus in conjunction with injection of the same tracer in various parts of the cerebral cortex in 17 cats. Termination areas of cortical fibres (anterogradely labelled) and pontine neurons projecting to the dorsal paraflocculus (retrogradely labelled) were carefully plotted in serial transverse sections. As an average of countings in ten cats, 90% of the labelled cells were found in the pontine nuclei contralateral to the injection, and the majority (70%) were located in the rostral half of the nuclei. The highest degree of overlap between anterograde and retrograde labelling was found after injections of the parietal association cortex (areas 5 and 7). In an experiment with double anterograde tracing, it was shown that both area 5 and 7 contribute substantially to the cerebral inputs to the dorsal paraflocculus. High degree of overlap also occurred after injections of several visual cortical areas (areas 17, 18, 19, 20 and the posteromedial lateral suprasylvian visual area, PMLS). Cases with injections restricted to individual visual areas indicate that they all contribute to the parafloccular input. Considerably less overlap occurred after injections of the primary sensorimotor region (SI, MI) and second somatosensory area (SII), while the supplementary motor area, the auditory cortex and gyrus cinguli probably have no or very restricted access to the dorsal paraflocculus. It is concluded that the dorsal paraflocculus has its major cortical input from the parietal association cortex and the visual cortical areas. Since all the various cortical regions studied project to largely different parts of the pontine nuclei, and overlap with neurons projecting to the dorsal paraflocculus takes place at numerous places, it follows that the pontine neurons projecting to the dorsal paraflocculus must consist of many subgroups differing with regard to their cortical input.

Spinocerebellar projections from the upper lumbar segments in the cat, as studied by anterograde transport of wheat germ agglutinin-horseradish peroxidase.

The projection fields of the dorsal spinocerebellar tract (DSCT) arising from Clarke's column, marginal neurons of Clarke's column, and lamina V neurons in the upper lumbar segments were studied by the anterograde transport of wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP) in the cat. To label only these neuron groups with uncrossed ascending axons, the spinal cord was lesioned rostral and contralateral to the WGA-HRP injections. Following injections of WGA-HRP into the L1-L4 segments, labeled terminals were seen in sublobules Ia-VIc and VIIb-VIIIb, the simple lobule, the paramedian lobule, and the dorsal paraflocculus. About 70-80% and 20-30% of the total number of labeled terminals were in the anterior and the posterior lobe, respectively; the projections were predominantly ipsilateral to the cells of origin (about 87% or more labeled terminals of the total number in each of sublobules IIb-Va). The labeled terminals were abundant in sublobule IIb (6-11%), lobule III (12-27%), and sublobules IVa (14-17%) and IVb (14-21%). In the mediolateral extent of the lobules in the anterior lobe, the labeled terminals were most numerous between 1.1 and 3.0 mm lateral to the midline (45-75% of the total number of labeled terminals on the ipsilateral side). In the posterior lobe labeled terminals were numerous in sublobule VIIIb (13.6%) and sublobule C of the paramedian lobule (15-19%). The projection fields in the horizontal plane of the lobules were reconstructed from a series of cross sections through each lobule. In the anterior lobe the labeled terminals were distributed in eight major areas. In sublobules IIb-III, areas 1-3 were located within 1.0 mm of the midline in zone A of Voogd; areas 4-6, between 1.0 and 2.5 mm lateral to the midline in zones B-C1; and areas 7 and 8, lateral to 3.0 mm from the midline in zones C2 and C3. Areas 1-6 extended apicobasally in the middle part of the lobules. In sublobule VIIIb projections were confined to three longitudinal areas whereas in the paramedian lobule the projection areas were less distinct. The projection pattern of the lumbar DSCT was different from that of the thoracic DSCT reported previously. In the anterior lobe the thoracic DSCT projects to five areas in the medial (zone A) and the lateral part (zone B) of the vermis and to four areas in the intermediate region of the hemisphere (zones C1-C3).

Distribution of acetylcholinesterase in the granular layer of the cerebellum of the rhesus monkey (Macaca mulatta).

A nonuniform distribution of acetylcholinesterase (AChE) activity was identified in the granular layer of the cerebellum in rhesus monkeys. The distribution of darkly AChE-stained clumps in the granular layer was determined for each lobule of the vermis and the lateral cortex. The vermis contained a greater density of AChE reaction product than the lateral cortex. In the vermis, lobules IX and X had significantly the highest level of activity, followed by lobules VII and VIII, which were significantly higher than lobules II-VI. In the lateral cortex, the flocculus had the highest level of the AChE activity, followed by crus I and the dorsal paraflocculus, which had significantly higher levels than the remaining lobules. The high levels of AChE activity in the flocculonodular lobe and lobule IX may correspond to cholinergic mossy fiber transmission, but the high levels of AChE activity in other cerebellar regions probably involve noncholinergic functions. The significance of the nonuniform AChE distribution is not yet known, but may correspond to regional differences in neuronal or metabolic activity in the cerebellum that occur in conjunction with specific behaviors.

Demonstration of a Mamillo-Ponto-Cerebellar Pathway.

The pathway from the mamillary complex to the cerebellum via the pontine nuclei has been studied using several anterograde and retrograde tracing techniques in the cat. We have also compared the pontine terminal regions of fibres from the mamillary complex and from the cingulate gyrus. Implantations of crystalline horseradish peroxidase wheat germ agglutinin (HRP-WGA) in the mamillary complex and lesions of the cingulate gyrus were combined in the same animal with injections of HRP-WGA, rhodamine-B-isothiocyanate (RITC), and Fluoro-Gold in different parts of the cerebellar hemisphere. Fibres from both the mamillary complex and the cingulate gyrus terminate mainly within a transversely oriented, c-shaped band in the ipsilateral, rostral pontine nuclei. Within this band the terminal fields of fibres from the mamillary complex and the cingulate gyrus form a mosaic-like pattern of partly overlapping patches. Pontine regions receiving a mamillary input project mainly to the ventral paraflocculus, and to a lesser degree to the dorsal paraflocculus, but apparently not to the uvula or crus II. Judging from the literature it seems highly unlikely that other parts of the cerebellar hemispheres received projections from these pontine regions. Fibres from the ventral paraflocculus were shown to terminate in the parvicellular part of the lateral cerebellar nucleus only. The present findings would seem to imply that inputs from the mamillary complex and a related cortical region, the cingulate gyrus, are partly integrated, partly kept separate at the precerebellar level. This would ensure that small groups of cells in the rostral pontine nuclei receive a specific set of afferents. Conceivably, the information transmitted to the cerebellum by these groups of pontine cells might be related to functions of the mamillary complex, such as learning, motivation, and spatial memory.