Bidirectional neurones in the cervical enlargement of the cat spinal cord with axons descending to sacral segments and ascending to the cerebellum and the lateral reticular nucleus

Neurones located in cervical segments (C6/C7) of the spinal cord were investigated electrophysiologically in cats deeply anaesthetized with alpha-chloralose. Extracellular recordings of antidromic action potentials were performed in order to establish whether long descending propriospinal neurones projecting to sacral segments could have collateral axonal branches ascending to supraspinal centres. The effects of stimulation of the lateral reticular nucleus (LRN) and the inferior cerebellar peduncle (restiform body, RB), as well as the thirteenth thoracic (Th13) and sacral (S1/S2) segments of the spinal cord were tested in 93 cells. Two main groups of cells were identified: 54 % of the total sample were classified as purely propriospinal and 46 % as bidirectional neurones. Various patterns of projections, as well as the ipsi-, contra- or bilateral courses of axons in the lateral funiculi of the spinal cord, enabled several types of neurones to be distinguished within the above groups. Comparison between particular types showed no significant difference with respect to location in the grey matter (predominantly Rexed's laminae VII-VIII) and the conduction velocities of descending axons. However, the mean axonal conduction velocities of branches ascending to LRN and/or RB were significantly lower in comparison to those measured for spinal collaterals. The hypothetical function of the neurones examined is discussed. Since the same information can be conveyed simultaneously by these branching neurones to lower spinal segments and supraspinal centres, an integrative role in the system of motor control is suggested.     

Divergence of lamina VII and VIII neurones of S1 and S2 segments of the cat's spinal cord to the cerebellum and the reticular formation.

Cerebellar and reticular projections of neurones located in sacral segments of the spinal cord were electrophysiologically investigated in alpha-chloralose anaesthetized cats. Antidromic action potentials were recorded following stimulation of the contralateral restiform body (coRB), contralateral gigantocellular nucleus (coGRN) as well as ipsi- and contralateral lateral funiculus of the 13th thoracic segment (iTh13 and coTh13). Eighty-seven neurones were found in the medial lamina VII and lamina VIII of the gray matter of S1 and S2 segments. Their axons ascended in lateral funiculi on the contralateral side and in 46 cases also on the ipsilateral side of the spinal cord. A projection to coRB was found in 20 neurones, to coGRN in 10 and dual projections to both coRB and coGRN in 20 neurones. Axons of the remaining 37 cells were found to ascend to the level of Th13 only. Conduction velocities of neurones investigated were comprised in the range 35-83 m/s and no significant differences were found between particular groups. However, an evident decrease in conduction was observed in most neurones when comparing proximal to distal parts of their axons, suggesting the possibility of more extensive divergence than indicated in this study. The pattern of projections revealed that the information from the periphery is conveyed in parallel to various supraspinal and possibly also spinal centres.     

Branching neurones in the cervical spinal cord with axons that reach sacral segments and the lateral reticular nucleus. An electrophysiological study in the cat

Branching neurones in the cervical enlargement of the spinal cord were electrophysiologically studied in alpha-chloralose anaesthetized cats with the method of antidromic activation of axons. Stimulating electrodes were placed bilaterally at levels of lower thoracic and sacral segments and in the lateral reticular nucleus (LRN), ipsilaterally to the recording sites in C6/C7 segments. Thirty-nine out of a total one hundred neurones could be classified as bidirectional neurones with both descending and ascending collaterals. In the remaining cases only long descending projections to spinal segments were found. Comparison of conduction velocities measured in descending branches revealed no significant differences between individual neurones. On the other hand, descending collaterals of double direction neurones conducted impulses considerably faster than their axonal branches ascending to LRN. Our results suggest that parallel transmission of information to various, spinal or supraspinal centres of the nervous system is more common than reported before.     

Electrophysiological investigation of spino-olivary projections originating from sacral segments of the cat spinal cord

Ascending projections of sacral spinal cord neurones (S1-S2) to the dorsal accessory olivary nucleus (DAO) were electrophysiologically investigated in 3 adult cats under deep alpha-chloralose anaesthesia. Antidromic action potentials were recorded extracellularly from 19 cells following stimulation of their axons in both the contralateral dorsal accessory olivary nucleus (coDAO) and the contralateral lateral funiculus at the level of lower thoracic segments (Th13). Two groups of neurones were identified in the gray matter of S1-S2 segments: one distributed in the medial part of Rexed's laminae VI and VII (n = 5), the other located in the ventromedial part of lamina VIII (n = 14). Axonal conduction velocities of neurones investigated were comprised in the range 32-55 m/s. A significant decrease of conduction velocity was observed in each case when distal and proximal parts of the axon were compared. Our research confirmed anatomical data concerning spino-olivary neurones originating from sacral segments. However, we suggest that axons of this pathway give off collaterals to other spinal or supraspinal centres.     

Ascending tract neurones processing information from group II muscle afferents in sacral segments of the feline spinal cord.

1. Ascending tract neurones located in the dorsal horn of sacral segments of the spinal cord have been investigated by extracellular and intracellular recording in the anaesthetized cat. The aim was to determine whether information from group II afferents that terminate within the sacral segments is conveyed to supraspinal structures and which types of neurones are involved. 2. A considerable proportion of ascending tract neurones found in the dorsal horn in the same segments as the pudendal (Onuf's) motor nucleus were excited by group II muscle afferents. The great majority (93%) of these neurones had axons ascending in ipsilateral funiculi. Spinocervical tract neurones constituted the largest proportion (82%) of such neurones, while very few spinocerebellar tract and propriospinal neurones and no postsynaptic dorsal column neurones were found among them. 3. In addition to activation by group II muscle afferents all of the neurones were strongly excited by cutaneous afferents. The most potent excitation was evoked by afferents of the posterior biceps-semitendinosus and gastrocnemius muscle nerves and by afferents of the cutaneous femoris, sural and pudendal nerves. The latencies of intracellularly recorded excitatory potentials were indicative of a high incidence of monosynaptic coupling between the afferents and ascending tract neurones. 4. The highly effective monosynaptic excitation of spinocervical tract neurones in the sacral segments by group II afferents is in contrast to the weak disynaptically mediated actions of group II afferents on such neurones in the L6-L7 segments but comparable to the actions of group II afferents on ascending tract neurones in the midlumbar segments. 5. Both the patterns of peripheral input and the latencies of synaptic actions in ascending tract neurones were similar to those in interneurones at the same locations (accompanying report). Similar information is therefore likely to be processed by both categories of neurones. 6. The role of sacral spinocervical tract neurones as a system for transmitting information from group II muscle afferents to supraspinal centres and the potential contribution of this system to the perception of limb position are discussed.     

Projection from excitatory C3-C4 propriospinal neurones to lamina VII and VIII neurones in the C6-Th1 segments of the cat

Intracellular recording and injection of horseradish peroxidase (HRP) were made in neurones located medially in lamina VII and in lamina VIII of the forelimb segments (C6-Th1). The cells received disynaptic excitation from the contralateral pyramid after corticospinal transection in C5/C6 and monosynaptic excitation from the ipsilateral lateral reticular nucleus. The pyramidal excitation was facilitated by a conditioning volley evoked from the contralateral nucleus ruber, which suggests convergence of cortico- and rubrospinal fibres on the intercalated neurones. It is proposed that laminae VII and VIII neurones receive a collateral input from the same excitatory C3-C4 propriospinal neurones which project to motoneurones and/or Ia inhibitory interneurones. Reconstruction of HRP-stained lamina VII and VIII neurones revealed ipsi- and contralateral ascending and/or descending axonal projections and termination in laminae VII and VIII in the forelimb segments.     

Cells of origin of propriospinal and ascending supraspinal fibers in a lizard (Lacerta galloti)

The location of cells of origin of propriospinal and ascending supraspinal fibers has been determined by injecting horseradish peroxidase (HRP) unilaterally into various parts of the spinal cord of the lizard Lacerta galloti. The distribution of retrogradely labeled cells after unilateral high spinal cord injections suggests that ascending supraspinal fibers are derived from neurons in the following areas: in the cervical intumescence in most areas of the spinal gray, particularly ipsilaterally, in thoracic and lumbar segments, in deeper situated areas, especially contralaterally. The presence of cells of origin of long descending propriospinal pathways has been demonstrated following HRP-injections into the lumbar intumescence.     

Fluorescent double-labelling study of ascending and descending neurones in the feline lateral cervical nucleus

Summary The organization of ascending and descending neurones of the lateral cervical nucleus (LCN) was investigated in 10 adult cats after injections of the fluorescent tracers Fast Blue and Nuclear Yellow. Injections into the thalamus and tectum resulted in up to 3000 labelled cell profiles within the contralateral LCN. This corresponded to a calculated number of 4500 labelled LCN neurones. The greatest diameter of the labelled cell profiles was about 30 m. They were located throughout the nucleus, but were less numerous in its medial portion. Injections mainly into the dorsal horn of different pairs of cervical and lumbar segments of the spinal cord resulted in a calculated number of up to 305 labelled LCN cells. The diameter of these cell profiles was about 25 m and they were mainly situated in the rostro-ventral and medial parts of the LCN. Doublelabelled cells with ascending and descending projections were not encountered after injections into the thalamus-tectum and spinal segments C5-6. About 15% of the descending LCN cells were doublelabelled by pairs of spinal injections separated by intervals of one segment. It is concluded that the neurones descending down the spinal cord and ascending to the thalamus-tectum constitute different subpopulations of cells within the LCN and that a minor proportion of the descending cells seem to project to at least three adjacent segments of the spinal cord.     

Integration in descending motor pathways controlling the forelimb in the cat

Summary Recording was made in the C3-C4 segments from cell bodies of propriospinal neurones identified by their antidromic activation from more caudal segments. Monosynaptic excitatory effects from descending motor pathways and primary afferents were investigated by electrical stimulation of higher motor centres and peripheral nerves in the forelimb and neck.The cell bodies were located mainly laterally in Rexed's layer VII. Threshold mapping for single axons showed that they descend in the lateroventral part of the lateral funicle. Antidromic stimulation at different spinal cord levels showed that some neurones terminated in the forelimb segments, others in the thoracic cord or in the lumbar segments. Terminal slowing of the conduction velocity suggested axonal branching over some segments.Monosynaptic EPSPs were evoked in the neurones by stimulation of the contralateral pyramid, red nucleus and dorsal tegmentum-superior colliculus. It is concluded that corticospinal, rubrospinal and tectospinal fibres project directly to both short and long propriospinal neurones. There was marked frequency potentiation in tectospinal synapses. Convergence from two descending tracts was common and in half of the tested cells all three tracts contributed monosynaptic excitation. Experiments with collision of descending volleys and antidromic volleys from the brachial segments demonstrated that the corticospinal and rubrospinal monosynaptic projection to the propriospinal neurones is by collaterals from fibres continuing to the forelimb segments.Stimulation of cervical primary afferents in the dorsal column gave monosynaptic EPSPs in somewhat less than half of the tested propriospinal neurones. The further analysis with stimulation of forelimb nerves and C2-C3 dorsal rami showed that monosynaptic EPSPs may be evoked from low threshold cutaneous and group I muscle afferents in the forelimb and from C2-C3 neck afferents entering close to the spinal ganglia, possibly from joint receptors. Convergence from cervical afferents and at least two of the above descending tracts was common.It is postulated that the propriospinal neurones previously indirectly defined by their action on motoneurones as relaying disynaptic excitation from higher motor centres to forelimb motoneurones (Illert et al., 1977) belong to those neurones of the C3-C4 propriospinal systems which terminate in the cervical enlargement. The function of the neurones projecting beyond the upper thoracic segments is discussed.Supported by the Deutsche ForschungsgemeinschaftIBRO/UNESCO Fellow     

Neurochemical excitation of propriospinal neurons facilitates locomotor command signal transmission in the lesioned spinal cord

Previous studies of the in vitro neonatal rat brain stem-spinal cord showed that propriospinal relays contribute to descending transmission of a supraspinal command signal that is capable of activating locomotion. Using the same preparation, the present series examines whether enhanced excitation of thoracic propriospinal neurons facilitates propagation of the locomotor command signal in the lesioned spinal cord. First, we identified neurotransmitters contributing to normal endogenous propriospinal transmission of the locomotor command signal by testing the effect of receptor antagonists applied to cervicothoracic segments during brain stem-induced locomotor-like activity. Spinal cords were either intact or contained staggered bilateral hemisections located at right T1/T2 and left T10/T11 junctions designed to abolish direct long-projecting bulbospinal axons. Serotonergic, noradrenergic, dopaminergic, and glutamatergic, but not cholinergic, receptor antagonists blocked locomotor-like activity. Approximately 73% of preparations with staggered bilateral hemisections failed to generate locomotor-like activity in response to electrical stimulation of the brain stem alone; such preparations were used to test the effect of neuroactive substances applied to thoracic segments (bath barriers placed at T3 and T9) during brain stem stimulation. The percentage of preparations developing locomotor-like activity was as follows: 5-HT (43%), 5-HT/N-methyl-D-aspartate (NMDA; 33%), quipazine (42%), 8-hydroxy-2-(di-n-propylamino)tetralin (20%), methoxamine (45%), and elevated bath K(+) concentration (29%). Combined norepinephrine and dopamine increased the success rate (67%) compared with the use of either agent alone (4 and 7%, respectively). NMDA, Mg(2+) ion removal, clonidine, and acetylcholine were ineffective. The results provide proof of principle that artificial excitation of thoracic propriospinal neurons can improve supraspinal control over hindlimb locomotor networks in the lesioned spinal cord.     

Distribution of the sacral neurones of origin of the ascending spinal tracts with axons passing through the lateral funiculi of the lowermost thoracic segments: an experimental HRP study in the cat.

The locations of 249 cell bodies of the ascending tract neurones in the grey matter of S1-S3 segments of the spinal cord were reconstructed by histochemical staining, after their axons (or axonal collaterals) at the level of the Thl3 segment were injected with horseradish peroxidase (HRP). In three cats in which the injections of HRP were restricted to the lateral part of the lateral funiculi (llf), about 84% of 159 retrogradely labelled cells were found on the contralateral side, while about 16% were located ipsilaterally. They were the most numerous in S2, S3 and S1 segments, respectively, and the neurones were distributed mainly in the lateral laminae I-VII, medial laminae V, VI and lamina VIII. In three other animals in which the injections of the marker were limited to the dorsal part of the lateral funiculi (dlf), 84 of the 90 ascending tract neurones were found to be distributed in the S2 and S3 segments both ipsi- (lateral laminae III-V) and contralaterally, (lateral laminae IV and V as well as the medial laminae VII and VIII) in similar numbers. The remaining six of the 90 cells with only contralateral projections at the dorsolateral funiculus at Thl3 were scattered within the S1 segment. These data are consistent with the results of studies on sacral spinocerebellar, spinothalamic and spinoreticular projections, as well as the localization of sacral spinocervical and priopriospinal neurones. They may also imply the importance of the bilateral fiber course of the neurones of origin of ascending tracts in the S2 and S3 segments within the dorsolateral funiculus.     

Descending pathways to the cutaneous trunci muscle motoneuronal cell group in the cat

1. The cutaneus trunci muscle (CTM) is a thin broad sheet of skeletal muscle just beneath the skin. It does not contain muscle spindles and receives its afferents from the overlying skin. Contraction of the muscle can easily be triggered by pinching the skin or, in the cat, by gentle displacement of the fur (CTM reflex). The afferent information of this reflex is conveyed via the cutaneous nerves, which are segmentally organized. In the cat, the CTM motoneurons are located in a circumscribed cell group in the ventrolateral part of the ventral horn of the C8 and T1 spinal segments. The CTM motor nucleus corresponds with "nucleus X" of Giovanelli Barilari and Kuypers and with "ventral motor nucleus" of Matsushita and Ueyama. 2. Relatively long ascending propriospinal pathways, originating in the thoracolumbar cord, exist between the cutaneous afferents and the CTM motor nucleus. Such pathways have been described physiologically, as well as anatomically. Our results, based on anterograde autoradiographic experiments with [3H]leucine injections in the C1, C2, C6, and C8 segments, suggest that propriospinal pathways to the CTM motor nucleus originating in the cervical cord do not exist, although these propriospinal projections are very strong to all other motoneuronal cell groups surrounding the CTM motor nucleus. 3. The present results also demonstrate specific supraspinal projections to the CTM motor nucleus originating in 1) the contralateral nucleus retroambiguous (NRA) and 2) the ipsilateral dorsolateral pontine tegmentum. These projections suggest that the CTM motor nucleus is not only involved in spinal reflexes, but also in other functions such as abdominal straining.(ABSTRACT TRUNCATED AT 250 WORDS)     

Effects of groups of propriospinal interneurons on fictive swimming in the isolated spinal cord of the lamprey

Fictive swimming activity was induced in isolated spinal cords of adult lampreys Ichthyomyzon unicuspis and Petromyzon marinus by addition of D-glutamate or N-methyl-D,L-aspartate (NMA) to the bathing fluid. Propriospinal interneurons are defined as nerve cells within the spinal cord with projections longer than 1 segment. The hypothesis that propriospinal interneurons contribute to intersegmental coordination during fictive swimming was tested using electrical stimulation, extracellular recording, and separated compartments. Stimulation of the split caudal end of the spinal cord indirectly excited ascending propriospinal interneurons, which enhanced and entrained bursts in rostral contralateral ventral roots. Indirect electrical stimulation of descending propriospinal interneurons could delay and diminish bursts in caudal contralateral ventral roots. Extracellular recordings from the rostral and caudal split ends of the spinal cord sometimes showed spike activities in phase with contralateral or ipsilateral ventral roots. Inhibition of 1-3 segments by spot applications of glycine or gamma-aminobutyric acid (GABA) did not interrupt normal coordination or rostrocaudal phase lag. When a middle region of spinal cord was inhibited in a compartment with GABA or glycine, the caudal spinal cord could entrain the bursts in rostral ventral roots. In a few preparations the caudal region induced antiphasic bursts in previously silent rostral roots through the inhibited region. The maximum separation for caudal-upon-rostral antiphasic entrainment was approximately 20 segments in Ichthyomyzon and 36 segments in Petromyzon. Increased concentrations of an excitatory amino acid in a rostral compartment could produce descending entrainment of bursts in an adjacent caudal compartment at a higher frequency with rostrocaudal phase lag. The rostral-upon-caudal entrainment could still occur through spot applications of GABA or glycine but not through long inhibited regions. Two hypothetical groups of propriospinal interneurons are proposed for the coordination of swimming activities in the isolated spinal cords of adult lampreys. 1) Crossed, ascending interneurons may be excited in phase with nearby motoneurons and may excite and entrain rostral pattern generators on the opposite side. 2) Short, commissural interneurons may be excited in phase with nearby motoneurons and may inhibit contralateral generators.     

Propriospinal neurons with ascending collaterals to the dorsal medulla,the thalamus and the tectum: a retrograde fluorescent double-labeling study of the cervical cord of the rat

Summary Branching neurons with descending propriospinal collaterals and ascending collaterals to the dorsal medulla, the thalamus and the tectum were studied in the rat's cervical spinal cord (C1–C8), using the retrograde fluorescent double-labeling technique: Diamidino Yellow Dihydrochloride (DY) was injected in the cord at T2, True Blue (TB) was injected in the brain stem. DY-labeled descending propriospinal neurons were present in all laminae, except lamina IX. They were concentrated in lamina I, laminae IV to VIII, and in the lateral spinal nucleus, LSN. TB-labeled neurons projecting to the dorsal medulla were concentrated in lamina IV and the medial parts of laminae V and VI (probably representing postsynaptic dorsal column — PSDC — neurons), but were also present in lamina I, the LSN, the lateral dorsal horn, and in laminae VII and VIII. DY-TB double-labeled neurons giving rise to both a descending propriospinal collateral and an ascending collateral to the dorsal medulla were intermingled with the TB single-labeled neurons. About 4% of the descending propriospinal neurons gave rise to an ascending collateral to the dorsal column nuclei; these double-labeled cells constitute a sizable fraction (10%) of the PSDC neurons. TB-labeled spinothalamic and spinotectal neurons were located in lamina I, the lateral cervical nucleus (LCN), the LSN, the lateral lamina V, lamina VII and VIII, lamina X and in the spinal extensions of the dorsal column nuclei, predominantly contralateral to the TB injections. DY-TB double-labeled neurons were present throughout C1–C8 in the LSN, lateral lamina V, lamina VIII, ventromedial lamina VII, and lamina X. Only very few were observed in lamina I and the LCN, and none in the spinal extensions of the dorsal column nuclei. The double-labeled neurons constituted only a minor fraction of all labeled neurons; 3–5% of the spinothalamic neurons and about 1–7% of the spinotectal neurons were double-labeled. Conversely, only about 1% of the labeled descending propriospinal neurons gave rise to an ascending spinothalamic collateral, and even fewer (0.1 to 0.6%) to a collateral to the dorsal midbrain. The LSN displayed the highest relative content of branching neurons. Up to 20% of its ascending spinothalamic and spinotectal neurons and up to 8% of its descending propriospinal neurons were found to be branching neurons, indicating that the LSN constitutes an unique cell-group in the rat spinal cord.     

神经示踪定位SD大鼠长下行脊髓固有神经元胞体及其轴突投射的解剖位置

目的 利用神经示踪技术探讨SD大鼠长下行脊髓固有神经元及其轴突投射的解剖位置.方法 将荧光金(FG)注射入第1腰髓(L1)节段逆行标记大鼠下行脊髓固有神经元(DPNs)胞体;将顺行神经示踪剂生物素葡聚糖胺(BDA)注射到脊髓第3和第4颈髓处标记此处的DPNs胞体及其长下行脊髓固有束(LDPT).固定取材与切片染色后,检...     

Spinal distribution and collateral projections of rat spinomesencephalic tract cells.

The distribution of cells belonging to the rat spinomesencephalic tract was studied by means of the retrograde transport of fluorescent dyes. Bilateral midbrain injections of cytoplasmic and nuclear tracers were made in order to evaluate the location of ipsilateral, contralateral, or bilaterally projecting cells. Spinal neurons with ascending projections to midbrain and descending propriospinal projections were identified by midbrain and spinal injections of different cytoplasmic labels. The locations of spinomesencephalic tract cells included seven regions of the spinal gray matter: marginal zone, lateral neck of the dorsal horn, nucleus proprius, the region around the central canal, the lateral cervical and spinal nuclei and the ventral horn. Cells projecting to the ipsilateral or contralateral midbrain had similar distributions and were frequently found in clusters with overlapping dendritic fields. Approximately 75% of spinomesencephalic cells projected to the contralateral midbrain. The largest contribution to the spinomesencephalic tract cell population was found in cervical cord segments 1-4. Cells with bilateral projections accounted for nearly 2% of all labeled cells, whereas 5% had both ascending and descending projections. Spinomesencephalic cells were found to have varying dendritic fields and morphology, e.g. fusiform, pyramidal, round/oval, and multipolar. The results of the present study lend further support to the view that the spinomesencephalic tract is a multi-component pathway with varied origins and projection targets.     

Spinothalamic and propriospinal neurones in the upper cervical cord of the rat: terminations of primary afferent fibres on soma and primary dendrites

Summary Experiments were performed on rats to determine whether primary afferents from the upper cervical region terminate directly on Spinothalamic and propriospinal neurones. The central terminations of primary afferents from the upper cervical region were identified by diffusely filling their axons with horseradish peroxidase. Spinothalamic neurones or propriospinal neurones were identified in the same experimental animals by using retrograde transport of wheat germ agglutinin conjugated to horseradish peroxidase. Approximately 3–11 % of Spinothalamic cells in laminae 4–6 of spinal segments C2–4 received apparent synaptic contacts from primary afferents on the soma or primary dendrites. Approximately 18–36% of propriospinal neurones with axons descending to lower thoracic or lumbar levels received apparent synaptic contacts on the soma or primary dendrites. These data provide anatomical evidence that Spinothalamic and long propriospinal neurones in the upper cervical cord are excited directly by primary afferents. The data also help to clarify the neural circuitry underlying somatic sensation and reflex movements evoked by neck receptors.     

Integration in descending motor pathways controlling the forelimb in the cat

Summary The morphology of single C3–C4 propriospinal neurones (PNs) including the cell body, dendritic tree, axonal trajectory and the pattern of projection and termination of axonal collaterals in the C3–C4 segments was investigated by intra-somatic or intra-axonal injection of horseradish peroxidase. All the C3–C4 PNs could be antidromically activated from the lateral funicle in C6 and the lateral reticular nucleus but not from Th13. Another criterion was that they received monosynaptic excitation from corticospinal fibres in the contralateral pyramid. Twenty-four C3–C4 PNs were successfully stained. They were located in the lateral part of laminae VI–VIII except for two neurones which were located in lamina V and two in lamina IX. Five to eleven dendrites originated from the cell bodies and extended throughout laminae IV–VIII and even into the white matter in the transverse plane and up to 3 mm rostro-caudally. The axonal trajectory from the cell body was usually curved before reaching the lateral funicle. The bifurcation of the stem axon into a descending and an ascending branch was mostly observed in the white matter close to or at the border between the white and grey matter at the level of the cell body. The ascending and descending axonal branches maintained their location in the same part of the lateral funicle. Sixteen out of 24 stem axons gave off collaterals in the grey matter and/or in the white matter. One to five collaterals were given off from the axons in the grey matter. Boutons were found in a restricted region in the intermediate zone from lamina VI to the border between laminae VII and VIII, in the lateral part of laminae V–IX, in the middle and medial parts of laminae VI–VIII. The termination in the vicinity of large neurones in lamina VIII suggests that long PNs receive collateral projections from the C3–C4 PNs. The finding that some collaterals terminated laterally in lamina IX is in agreement with electrophysiological observations that spinal accessory motoneurones receive disynaptic pyramidal excitation which is mediated via C3–C4 PNs. The collateral projection from the C3–C4 PNs to lateral and medial regions in laminae VI-VII is discussed in relation to feed-forward and feed-back inhibitory control of the C3–C4 PNs.     

Spinocerebellar neurons and propriospinal neurons in the cervical spinal cord: a fluorescent double-labeling study in the rat and the cat

Summary In the cervical spinal cord of the rat and the cat, the distributions of spinocerebellar and of descending propriospinal neurons were investigated using the retrograde fluorescent double-labeling technique. Moreover, a search was made for the presence of neurons with both ascending spinocerebellar and descending propriospinal axoncollaterals. Diamidino Yellow Dihydrochloride (DY) was injected at T2, while True Blue (TB) (in rats) or Fast Blue (FB) (in cats) was injected in the cerebellum. The distributions of labeled neurons were very similar in the rat and the cat. DY-labeled propriospinal neurons, projecting to T2 or below, were most numerous in lamina I and laminae IV to VIII. In the rat, such neurons were also present in the lateral spinal nucleus (LSN). TB- or FB-labeled spinocerebellar neurons were concentrated in the central cervical nucleus (CCN) at C1-C4, in the central part of lamina VII at C5-T1, in the medial part of lamina VI and the adjoining dorsomedial part of lamina VII at C2/C3-T1, and in Clarke's column. They were also found in lamina V at C1 and C7-T1, and in lamina VIII at all levels. In both species only very few DYTB/FB double-labeled neurons, representing neurons with branching axons, were observed; in C1-T1, only about 0,5% of all TB/FB-labeled Spinocerebellar neurons and about 0,05% of all DY-labeled descending propriospinal neurons were double-labeled. The double-labeled neurons were all located centrally in lamina VII at C5-T1, but even in that area they constituted not more than 1,5% (rat) and 4% (cat) of the labeled spinocerebellar neurons. These findings indicate that, in the cervical cord of the rat and the cat, descending propriospinal neurons and spinocerebellar neurons are to a large extent separate populations.     

Projection from excitatory C3-C4 propriospinal neurones to spinocerebellar and spinoreticular neurones in the C6-Th1 segments of the cat

Extra- and intracellular recording was made from neurones in laminae VII and VIII of the C6-Th1 segments, which were disynaptically excited from the contralateral pyramid, nucleus ruber and monosynaptically from the ipsilateral lateral reticular nucleus. The results suggest collateral excitation from the C3-C4 propriospinal neurones which are excited monosynaptically from the former two inputs and antidromically from the latter nucleus. The cells were antidromically activated from the ipsilateral nucleus fastigus, and from the ipsilateral or contralateral reticular formation. Some of the spinocerebellar and spinoreticular neurones were also antidromatically activated from Th13. It is suggested that spinocerebellar, spinoreticular and bifurcating spinocerebellar and spinoreticular neurones receive collateral input from the same excitatory C3-C4 propriospinal neurones which project to motoneurones and/or Ia inhibitory interneurones.