Medial Part Of Lamina VII Research Articles

Premotor nitric oxide synthase immunoreactive pathway connecting lumbar segments with the ventral motor nucleus of the cervical enlargement in the dog

In this study we investigate the occurrence and origin of punctate nitric oxide synthase immunoreactivity in the neuropil of the ventral motor nucleus in C7-Th1 segments of the dog spine, which are supposed to be the terminal field of an ascending premotor propriospinal nitric oxide synthase-immunoreactive pathway. As the first step, nitric oxide synthase immunohistochemistry was used to distinguish nitric oxide synthase-immunoreactive staining of the ventral motor nucleus. Dense, punctate nitric oxide synthase immunoreactivity was found on control sections in the neuropil of the ventral motor nucleus. After hemisection at Th10–11, axotomy-induced retrograde changes consisting in a strong upregulation of nitric oxide synthase-containing neurons were found mostly unilaterally in lamina VIII, the medial part of lamina VII and in the pericentral region in all segments of the lumbosacral enlargement. Concurrently, a strong depletion of the punctate nitric oxide synthase immunopositivity in the neuropil of the ventral motor nucleus ipsilaterally with the hemisection was detected, thus revealing that an uncrossed ascending premotor propriospinal pathway containing a fairly high number of nitric oxide synthase-immunoreactive fibers terminates in the ventral motor nucleus. Application of the retrograde fluorescent tracer Fluorogold injected into the ventral motor nucleus and analysis of alternate sections processed for nitric oxide synthase immunocytochemistry revealed the presence of Fluorogold-labeled and nitric oxide synthase-immunoreactive axons in the ventrolateral funiculus and in the lateral and medial portions of the ventral column throughout the thoracic and upper lumbar segments. A noticeable number of Fluorogold-labeled and nitric oxide synthase-immunoreactive somata detected on consecutive sections were found in the lumbosacral enlargement, mainly in laminae VIII–IX, the medial part of lamina VII and in the pericentral region (lamina X), ipsilaterally with the injection of Fluorogold into the ventral motor nucleus. In summary, the present study provides evidence for a hitherto unknown ascending premotor propriospinal nitric oxide synthase-immunoreactive pathway connecting the lumbosacral enlargement with the motoneurons of the ventral motor nucleus in the dog.

Distribution of the high-affinity choline transporter in the human and macaque monkey spinal cord

The distribution of the high-affinity choline transporter (CHT) was determined in the human and macaque monkey spinal cord using in situ hybridization histochemistry and immunohistochemistry. Signals for CHT mRNA were observed in somatic motor neurons, sympathetic preganglionic neurons, and neurons in the medial part of lamina VII. The mRNA for CHT was co-localized in single neurons with the mRNAs for vesicular acetylcholine transporter and cholineacetyltransferase. These same cholinergic neuronal groups were labeled by immunohistochemistry for human CHT. Of somatic motor neurons, smaller cell bodies of γ-motor neurons were labeled very intensely, whereas larger cell bodies of α-motor neurons showed various degrees of labeling from weak to moderately intense. Human CHT is thus a novel cholinergic marker, which not only labels cholinergic neurons, but also reveals their heterogeneity.

Projections from the lowest lumbar and sacral‐caudal segments to the cerebellar nuclei in the rat, studied by anterograde axonal tracing

Spinocerebellar fibers originate from neurons in the medial part of lamina VII, including Stilling's sacral nucleus, the lateral part of lamina V, and the ventrolateral part of the ventral horn in the lowest lumbar (L6) and sacral-caudal segments. By using the anterograde transport of biotinylated dextran in the rat, this study examined whether these spinocerebellar fibers project to the cerebellar nuclei. Anterogradely labeled axons that originated from the L6-caudal segments ascended contralaterally through the superficial layer of the lateral cord and entered the cerebellum through the superior cerebellar peduncle and the restiform body. Labeled terminals were bilaterally seen in the medial nucleus and the anterior and posterior interpositus nucleus. They were most numerous contralaterally in the anterior interpositus nucleus. In the anterior interpositus nucleus, they were distributed mainly in the most ventral part at rostral levels and in small numbers in the medial part. In the medial nucleus, labeled terminals were seen in limited rostromedial parts of the middle and the caudomedial subdivision. Only a small number of labeled terminals were seen in the caudomedial part of the posterior interpositus nucleus. The present study shows that the spinocerebellar fibers originating from the lowest lumbar and sacral-caudal segments project to specific areas of the anterior interpositus and the medial nucleus. This finding suggests that the neurons of origin convey information from muscles, joints, and integument to the cerebellar cortex and nuclei, in relation to movement of the hindlimbs. J. Comp. Neurol. 404:21–32, 1999. © 1999 Wiley-Liss, Inc.

Projections from the lowest lumbar and sacral-caudal segments to the cerebellar nuclei in the rat, studied by anterograde axonal tracing

Spinocerebellar fibers originate from neurons in the medial part of lamina VII, including Stilling's sacral nucleus, the lateral part of lamina V, and the ventrolateral part of the ventral horn in the lowest lumbar (L6) and sacral-caudal segments. By using the anterograde transport of biotinylated dextran in the rat, this study examined whether these spinocerebellar fibers project to the cerebellar nuclei. Anterogradely labeled axons that originated from the L6-caudal segments ascended contralaterally through the superficial layer of the lateral cord and entered the cerebellum through the superior cerebellar peduncle and the restiform body. Labeled terminals were bilaterally seen in the medial nucleus and the anterior and posterior interpositus nucleus. They were most numerous contralaterally in the anterior interpositus nucleus. In the anterior interpositus nucleus, they were distributed mainly in the most ventral part at rostral levels and in small numbers in the medial part. In the medial nucleus, labeled terminals were seen in limited rostromedial parts of the middle and the caudomedial subdivision. Only a small number of labeled terminals were seen in the caudomedial part of the posterior interpositus nucleus. The present study shows that the spinocerebellar fibers originating from the lowest lumbar and sacral-caudal segments project to specific areas of the anterior interpositus and the medial nucleus. This finding suggests that the neurons of origin convey information from muscles, joints, and integument to the cerebellar cortex and nuclei, in relation to movement of the hindlimbs.

Dual projections of the ventromedial lamina VI and the medial lamina VII neurones in the second sacral spinal cord segment to the thalamus and the cerebellum in the cat

Neurons of origin of the crossed spinocerebellar and/or spinothalamic tracts giving off axon collaterals at supraspinal levels were antidromically identified and labeled with horseradish peroxidase in lower sacral spinal cord segments of cats. Their cell bodies were located mainly in the medial part of lamina VII and rarely in the ventromedial aspect of lamina VI. In the S2 segment 77 neurons with axons in the opposite dorsolateral funiculus were recorded; 36 of them were invaded from only the thoracic level, 21 from both the thoracic level and the restiform body, 3 from both the thoracic level and the thalamus, and 17 from both the thoracic level, the restiform body and the thalamus on the contralateral side. These termination areas suggest that the S2 neurons under study transmit peripheral signals of proprioceptive, exteroceptive and nociceptive sensations.

The periaqueductal gray in the cat projects to lamina VIII and the medial part of lamina VII throughout the length of the spinal cord.

The periaqueductal gray (PAG) plays an important role in analgesia as well as in motor activities, such as vocalization, cardiovascular changes, and movements of the neck, back, and hind limbs. Although the anatomical pathways for vocalization and cardiovascular control are rather well understood, this is not the case for the pathways controlling the neck, back, and hind limb movements. This led us to study the direct projections from the PAG to the spinal cord in the cat. In a retrograde tracing study horseradish peroxidase (HRP) was injected into different spinal levels, which resulted in large HRP-labeled neurons in the lateral and ventrolateral PAG and the adjacent mesencephalic tegmentum. Even after an injection in the S2 spinal segment a few of these large neurons were found in the PAG. Wheat germ agglutinin-conjugated HRP injections in the ventrolateral and lateral PAG resulted in anterogradely labeled fibers descending through the ventromedial, ventral, and lateral funiculi. These fibers terminated in lamina VIII and the medial part of lamina VII of the caudal cervical, thoracic, lumbar, and sacral spinal cord. Interneurons in these laminae have been demonstrated to project to axial and proximal muscle motoneurons. The strongest PAG-spinal projections were to the upper cervical cord, where the fibers terminated in the lateral parts of the intermediate zone (laminae V, VII, and the dorsal part of lamina VIII). These laminae contain the premotor interneurons of the neck muscles. This distribution pattern suggests that the PAG-spinal pathway is involved in the control of neck and back movements. Comparing the location of the PAG-spinal neurons with the results of stimulation experiments leads to the supposition that the PAG-spinal neurons play a role in the control of the axial musculature during threat display.

Transneuronal labeling of spinal interneurons and sympathetic preganglionic neurons after pseudorabies virus injections in the rat medial gastrocnemius muscle

The distribution of retrogradely and transneuronally labeled neurons was studied in CNS of rats 4 days after injections of the Bartha strain of pseudorabies virus (PRV) into the medial gastrocnemius (MG) muscle. Tissue sections were processed for immunohistochemical detection of PRV. Retrogradely labeled cells were identified in the ipsilateral MG motor colum in the caudal L4 and the L5 spinal segments. In order to evaluate the efficacy of PRV retrograde cell body labeling, the number of PRV retrogradely labeled neurons in the MG motor column was compared to the number labeled with two conventional retrograde cell body markers — Fluoro-Gold and cholera toxin-HRP. A ratio of 1:3 representing medium-sized (< 30 μm) versus large neurons (>30 μm) was found in the Fluoro-Gold dye experiments; a 1:2 ratio was seen in the PRV experiments. In contrast, when cholera toxin-HRP was used as a retrograde marker, mainly large neurons were labeled; the medium-to-large cell body ratio was 1:10 suggesting cholera toxin-HRP may have a greater affinity for the terminals of α-motoneurons as opposed to γ-motoneurons. Transneuronally labeled cells were identified in the L1–L6 spinal gray matter, intermediolateral cell column (T11-L2), lateral spinal nucleus and medial part of lamina VII in C4 and C5 spinal segments, brainstem (caudal raphe nuclei, rostral ventrolateral medulla, A5 cell group, paralemniscal nucleus, locus coeruleus, subcoeruleus nucleus, red nucleus) and paraventricular hypothalamic nucleus. In the L5 spinal cord, transneuronally labeled neurons were seen in the ipsilateral spinal laminae I and II and bilaterally in spinal laminae IV–VIII, and X. Similar results were obtained in rats that had chronic unilateral L3–L6 dorsal rhizotomies indicating most of the labeling was due to retrograde transneuronal cell body labeling. In order to determine whether PRV was transported into the spinal cord by the dorsal root axons, the ipsilateral dorsal root ganglia (DRGs) were examined for PRV immunoreactivity; none was found. However, using the polymerase chain reaction, viral DNA was shown to be present in the ipsilateral DRGs indicating that some of spinal cord cell body labeling may have resulted from anterograde transneuronal labeling, as well.

Projections of the tectospinal tract to the upper cervical spinal cord of the cat: A study with the anterograde tracer PHA‐L

The goal of the present experiments was to re-examine the spinal projections of neurons in the superior colliculus (SC) of the cat by taking advantage of the high sensitivity of the anterograde tracer, phaseolus vulgaris leucoagglutinin (PHA-L). In seven experiments, multiple injections of PHA-L into different regions of the SC labelled a total of 172 axons in the predorsal bundle; yet only 11 tectospinal tract (TST) axons were found in the upper cervical spinal cord. Collaterals emerging from these axons were rare and arose exclusively from TST axons with a diameter of less than 1 micron. Individual collaterals had different termination zones: some terminated in the lateral part of lamina V and VI after taking a dorsolateral course through lamina VII and VIII; others terminated in the medial part of lamina VII. One collateral terminated within lamina IX and the ventral part of lamina VIII. The combined termination of all collaterals was densest in lamina VII and dorsal lamina VIII. A small number of boutons were also found in the lateral parts of laminae V and VI, and in lamina IX and immediately adjacent regions in lamina VIII. Compared to axons belonging to other spinal descending systems, individual TST axons give rise to much simpler intraspinal collaterals with relatively few boutons. This feature, together with the relative paucity of TST axons, suggests that direct connections from the SC to neurons in the upper cervical spinal cord are sparse. Furthermore, our results are consistent with electrophysiological studies that show that few, if any, neck motoneurons receive monosynaptic connections from TST neurons. Projections to neck motoneurons must therefore involve a relay, either through other descending pathways, such as the reticulospinal system, or via local segmental interneurons.

Inputs to spinocerebellar tract neurones located in Stilling's nucleus in the sacral segments of the rat spinal cord

Spinocerebellar neurones located in the sacral segments of the rat spinal cord have been identified electrophysiologically. The neurones studied were located 0.7-1.1 mm deep to the cord dorsum, lateral and dorsal to the central canal in the medial part of lamina VII. Neurones were identified as spinocerebellar by antidromic activation from the cerebellar surface, the lowest threshold stimulation sites being near the midline on the posterior lobe vermis (lobule VIII). Estimates of conduction velocities of the axons ranged from 15-32 m/s (mean 22.8 m/s) and are directly comparable to velocities of presumed ventral spinocerebellar tract neurones recorded in the same animals. In intact animals, activity was most strongly influenced by passive movement of the tail. Activation by proprioceptors was confirmed with nerve stimulation: all of the neurones studied were discharged by stimulation of nerves which innervate ipsilateral tail muscles. In many cases responses appeared close to the threshold of the nerve, indicating that the largest, fastest conducting afferents (group Ia muscle spindle primary afferents) were responsible for them. Latencies of EPSPs or spikes were brief and in many cases indicative of a monosynaptic connection. We conclude that this group of neurones is powerfully and monosynaptically excited by group I muscle afferents and thus resemble the cells of Clarke's column and cells of the central cervical nucleus, both of which occupy a similar location in the grey matter of more rostral segments.

Routes of entry into the cerebellum of spinocerebellar axons from the lower part of the spinal cord

Among the newly discovered spinocerebellar cell groups, those at lumbar and more caudal levels of the cat's spinal cord were studied with regard to which of the two cerebellar peduncles, the restiform body or the superior cerebellar peduncle, is used by their axons. Bilateral injections with horseradish peroxidase were made into either of the anterior lobe or the posterior cerebellar termination area for spinocerebellar fibers, following unilateral transections of either the superior cerebellar peduncle or the restiform body, combined with low contralateral transections of the lateral and ventral funiculi. Following transection of the superior cerebellar peduncle, labeled neurons were found ipsilateral to the transection in the column of Clarke and in laminae IV-VI at L 3-L 7. Contralaterally, labeled neurons were found in the ventromedial nucleus and lamina VIII of the ventral horn in the sacro-coccygeal segments and in the medial part of lamina VII at L 6 and more caudal levels. All these neurons were regarded as sending their axons through the restiform body. Following transection of the restiform body, labeled neurons were found in the following areas contralateral to the transection: the dorsolateral nucleus of the L 3-L 6 segments, the lateral part of lamina VII at L 3-L 5/6, the medial part of lamina VII in L 6 and more caudal segments, and the ventrolateral nucleus of L 4-L 5. Ipsilaterally, labeled neurons were found in lamina VIII at L 4-L 6. All these neurons were regarded as sending their axons through the superior cerebellar peduncle. In addition to new information about the peduncular routes of spinocerebellar neurons, the study has given confirming evidence as to the crossing conditions for different spinocerebellar cell groups. The findings should be useful in future studies on the organization of the spinocerebellar systems.

Collateral projections of neurons from the lower part of the spinal cord to anterior and posterior cerebellar termination areas

The collateral projections of spinocerebellar neurons located in the L 2 to Ca 1 spinal segments in the cat were investigated by retrograde fluorescent double labeling technique. Rhodamine labeled latex microspheres (Rm) and Fast Blue (FB) were used for injections into the cerebellum in 8 cats. Two additional cats, with injections of Fluoro-Gold (FG) combined with Rm were excluded because lipofuchsin autofluorescence obscured the labeling. After injections with one tracer unilaterally in the paramedian lobule and another tracer bilaterally in the anterior lobe, double labeled neurons were found on the side of the paramedian lobule injection in the column of Clarke at L 2-L 4, laminae IV-VI at L 2-L 5 and the dorsolateral nucleus at L 2-L 6. After bilateral injections of one tracer in lobule VIII B and another in the anterior lobe, double labeled neurons were found bilaterally in the column of Clarke at L 2-L 4, laminae IV-VI at L 2-L 5, the medial part of lamina VII at L 6-L 7 and in certain cell groups at sacro-coccygeal levels. Neurons in the lateral part of lamina VII at L 3-L 5 and the ventrolateral nucleus of L 4-L 5 were labeled exclusively from injections in the anterior lobe. The findings indicate that spinocerebellar neurons at lumbar and more caudal levels of the cat spinal cord have different projection patterns in the cerebellum. A certain number of neurons which project to the anterior lobe have divergent axon collaterals supplying also the posterior vermis and/or the paramedian lobule. Other neurons project to the anterior or to the posterior lobe only.

Chapter 2 Vestibular projections to the spinal cord: the morphology of single vestibulospinal axons

The three-dimensional distribution of LVST and MVST axons was examined in the cat cervical spinal cord using an intra-axonal staining method. LVST and MVST axons were electrophysiologically identified by their responses to stimulation of the vestibular nucleus, bilateral vestibular primary afferents, the LVST and the MVST were stained with injection of HRP. The axonal trajectory was reconstructed from serial histological sections. LVST axons were found to have multiple axon collaterals in the cervical cord. The maximum number of the identified collaterals for one neuron was 7. These collaterals were observed in either LVST axons terminating at the cervical cord or those projecting below Th2. The rostro-caudal extension of terminals for each collateral was very restricted (mean = 760 μm) and much narrower than intercollateral intervals (mean = 1470 μm). In the gray matter, collaterals ramified successively, pursued a delta-like path, and terminated mainly in lamina VIII and in the medial part of lamina VII and many boutons made apparent contact with the cell bodies and the proximal dendrites of motoneurons in the ventromedial nucleus. Some terminals were also distributed to the ventrolateral part of lamina VII adjacent to lamina IX. One group of LVST axons projected to lamina IX in the lateral ventral horn and terminated on large neurons there, probably motoneurons of forelimb muscles. MVST axons had one to seven axon collaterals at C1–C3 within the range of the stained axon. Stem axons ran in the ventromedial funiculus and primary collaterals arose from them at right angles. Each collateral had a very nar-row rostrocaudal spread as in LVST axons. Terminals were distributed in laminae VIII and IX, including the ventromedial nucleus, the spinal accessory nucleus, and the commissural nucleus. Many terminals seemed to make contact with retrogradely labelled motoneurons of neck muscles. Both crossed and uncrossed MVST axons had these characteristics.

Retrograde HRP study of neurons in the cervical enlargement projecting to the cerebellum in the cat.

After cerebellar HRP injections in kittens labeled neurons were found in laminae V-VIII in the cervical enlargement. Most of the labeled neurons were localized in two groups, one in laminae V-VI, the other centrally in lamina VII. Labeled neurons were also observed in the medial part of lamina VII of C5 and T1 and a few in lamina VIII. Neurons in the cervical enlargement seem to terminate largely in cerebellar lobules IV-V of the anterior lobe. Some neurons in laminae V-VI terminated in the ipsilateral paramedian lobule. Neurons in laminae V-VI and central lamina VII of C5-T1 had uncrossed axons. Neurons in medial lamina VII of C5, in lamina VIII and neighbouring parts of lamina VII of C6-T1 had crossed axons. The ramifications of proximal dendrites and axons of the labeled neurons are described using the tetramethylbenzidine (TMB) method for HRP histochemistry. The neurons in the various laminae differed in their characteristic morphology. In conclusion, the findings of Matsushita et al. (1979) concerning the localization and axonal course of cerebellar projecting neurons in the cervical enlargement are confirmed. In addition new data concerning the morphology of the labeled neurons are presented.

Spinocerebellar projections to lobules III to V of the anterior lobe in the cat, as studied by retrograde transport of horseradish peroxidase.

Spinocerebellar tract (SCT) neurons projecting to lobules III to V of the cerebellar anterior lobe were identified by the retrograde horseradish peroxidase technique. SCT neurons projecting to lobule III with crossed ascending axons were located mainly in the central cervical nucleus (CCN), the medial part of lamina VII of L6 to the caudal segments, and the dorsal horn (lamina V) and ventral horn (lamina VIII) of the sacral-caudal segments. Spinal border cells with crossed ascending axons also projected to lobule III. SCT neurons projecting to this lobule with uncrossed ascending axons were located in the medial part of lamina VI of the cervical segments and the middle part of lamina VII of C6 to T1, lamina V of the lower cervical, thoracic and the lumbar segments, Clarke's column including marginal neurons, and the medial part of lamina VI of L5 and L6. These neuronal groups also projected to lobule IV, except for those present caudal to L6 (in the medial part of lamina VII, and laminae V and VIII of the sacral-caudal segments). A far smaller number of similar neurons projected to lobule V. Injections of HRP restricted to the vermal region labeled mainly neurons in the CCN and Clarke's column while restricted injections to the intermediate-lateral regions labeled ipsilaterally spinal border cells, lamina V neurons, and Clarke column neurons, especially of the lumbar segments as well as marginal neurons of this column.

The location of cerebellar-projecting neurons within the lumbosacral spinal cord in the cat. An anatomical study with HRP and retrograde chromatolysis.

The locations of lumbosacral spinocerebellar neurons were examined by two anatomical methods in kittens. In one group of animals chromatolytic changes were provoked by cerebellar lesions. In another group horseradish peroxidase (HRP) was injected into the cerebellum. Projection laterality was investigated by making unilateral spinal lesions prior to the cerebellar HRP injections. Diaminobenzidine (DAB) or tetramethylbenzidine (TMB) was used as substrate fo HRP. The morphological characteristics of HRP-labeled neurons in the TMB-processed material were examined. Neurons marked by the two methods were located within the same regions. A greater number of cells were marked with the HRP method, however, than with the retrograde chromatolysis method. Spinocerebellar neurons were found in laminae IV-IX with large differences with regard to specific locations depending on segmental level. Numerous marked neurons were found in the following areas: laminae IV-Vi in L3-L7, the column of Clarke in L3-L4, the medial part of lamina VII in L6-L7, the lateral part of lamina VII in L3-L4, the dorsolateral nucleus of lamina IX in L3-L6, the ventrolateral nucleus of lamina IX in L4-L5, and the ventromedial nucleus of lamina IX in S3 (and Ca1). Dorsally located neurons were in general more likely to project ipsilaterally than ventrally located neurons. Marked structural differences were frequently observed between spinocerebellar neurons in different locations. These results provide additional information on the anatomical complexity of the spinocerebellar pathways from the lumbosacral region in the cat. Together with results from some other recent anatomical studies on spinocerebellar tracts, they also form a basis for further anatomical and physiological investigations which could contribute to a better understanding of the organization of the spinocerebellar tracts.

Spinocerebellar projections to lobules I and II of the anterior lobe in the cat, as studied by retrograde transport of horseradish peroxidase.

Spinocerebellar tract (SCT) neurons projecting to lobules I and II of the cerebellar anterior lobe were identified by the retrograde horseradish peroxidase technique in the cat. Instead of a conventional stereotaxic approach, we removed ventral parts of the vermis of the posterior lobe and approached the posterior aspect of lobule I through the fourth ventricle. Under direct visual guidance, discrete injections were made into lobule I or II with a glass micropipette. Neurons projecting to lobule I were located mainly in the central cervical nucleus (CCN), the medial part of lamina VII of L6 to the causal segments, and in lamina VIII of S2 to the caudal segments (with crossed ascending axons). The latter two groups correspond to medial lamina VII group of the lumbar to the caudal segments and the ventral horn group of the sacral-caudal segments of our previous studies. A small number of Clarke column neurons (with uncrossed ascending axons) also projected to lobule I. All of these neuronal groups projected to lobule II. In addition, large neurons in lamina V and the border between laminae IV and V from S2 to the caudal segments projected to sublobule IIA, and more numerously to sublobule IIB (with crossed ascending axons). They belong to the dorsal horn group of the sacral-caudal segments of our previous studies. Spinal border cells (with crossed ascending axons) projected to sublobule IIB, and a small number, to sublobule IIA. It was suggested that the CCN neurons project more densely to the median region whereas Clark column neurons project to the lateral part of these lobules.

Spinocerebellar projections to the vermis of the posterior lobe and the paramedian lobule in the cat, as studied by retrograde transport of horseradish peroxidase.

Spinal neurons projecting to the posterior lobe of the cerebellum were identified with the retrograde horseradish peroxidase technique in the cat. In four cases with the injections, which were preceded by hemisections at cervical or thoracic levels, it was determined whether in the spinal cord the identified neurons give rise to crossed ascending axons or uncrossed ascending ones. The main groups of neurons projecting to sublobule VIIIB were located in the central cervical nucleus (with crossed ascending axons), Clarke's column (with uncrossed ascending axons), and the medial part of lamina VII of L6 to the caudal segments (with crossed ascending axons). Additional labeled neurons were found in the medial part of lamina VI between C2 and T1 and L5 and L6 (with uncrossed ascending axons), and in the ventral as well as dorsal horns of the sacral-caudal segments (with crossed ascending axons). On the other hand, neurons projecting to sublobule C (the copular part) of the paramedian lobule, which appeared always ipsilaterally to the side of the injections, were located in lamina V of C8 to L4 (with uncrossed ascending axons). Marginal neurons of Clarke's column (with uncrossed ascending axons) and spinal border cells (with crossed ascending axons that recross in the cerebellum) projected specifically to this part. At L1 and L2 or L2 and L3 labeled large and medium-sized neurons were also found within Clarke's column. The present study suggests that there are segregated projections of spinal neurons to the cortex of the cerebellar posterior lobe.

Cells of origin of the spinocerebellar tract in the rat, studied with the method of retrograde transport of horseradish peroxidase

Following injections of horseradish peroxidase into the cerebellum, the distribution of labeled neurons was studied in the whole length of the spinal cord of the rat. To find the ascending side of the axons, injections were made following hemisections at C1 or between C1 and C2. Labeled spinocerebellar tract neurons were classified into two groups according to the axonal course in the spinal cord; one is composed of neurons with uncrossed ascending axons and the other, neurons with crossed ascending axons. Neurons of origin of the uncrossed tracts were located in the medial part of lamina VI of C2 to C8, the central part of lamina VII of C4 to C8, lamina V of C7 to L3 and Clarke's column. Neurons of origin of the crossed tracts were found in the central cervical nucleus of C1 to C3, the intermediate zone and the ventral horn of the lower thoracic and the lumbar segments (T11 to L3), and in the dorsal horn, the medial part of lamina VII and the ventrolateral part of the ventral horn of the sacral and caudal spinal cord. In comparison with our previous results in the cat, it was suggested that the spinocerebellar system in the rat is organized in the same fashion as in the cat, in terms of the location and the intraspinal axonal course of the cells of origin.

Cells of origin of propriospinal fibers and of fibers ascending to supraspinal levels. A HRP study in cat and rhesus monkey.

In the spinal cord of cat and rhesus monkey the cells of origin of long and short propriospinal fibers and those of the spinal fibers ascending to supraspinal levels were identified by means of retrograde HRP labeling after large unilateral HRP-injections, i.e. in the spinal white and grey matter at different levels, in the pons and in the dorsal column nuclei. The findings indicate the existence of the following arrangement. Long ascending supraspinal fibers arise mainly from neurons in the dorsal grey (laminae I-IV and the medial parts of laminae V and VI) as well as from neurons in the medial part of the ventral grey (lamina VIII), in Clarke's column and in the spinal border cell area. Some neurons in the dorsal grey projects to the dorsal column nuclei, which in turn distribute fibers back to the spinal cord. Long propriospinal fibers mainly derived from neurons in the medial part of the ventral grey (lamina VIII), while short propriospinal fibers are characteristically derived from neurons in the intermediate zone (lateral halves of laminae V and VI and lamina VII). The neurons located laterally in laminae V-VII distribute fibers mainly ipsilaterally, while those located medially in lamina VII distribute them to some degree bilaterally. The findings in cats with transections of either the dorsal or the ventral halves of the spinal white matter (both above and below the injected segment), show that the fibers from the dorsal grey and the lateral parts of laminae V-VII travel mainly through the dorsal half of the white matter, while those from the medial part of lamina VII and from lamina VIII travel mainly through the ventral half.