The lateral reticular nucleus in the cat

The afferent paths from the spinal cord and from trigeminal afferents to the lateral reticular nucleus (LRN) were investigated by intracellular recording from 204 LRN neurones in preparations with a spinal cord lesion at C3 that spared only the ipsilateral ventral quadrant. Stimulation of nerves in the limbs evoked EPSPs and JPSPs in 201 of 204 tested LRN neurones. The strongest input was from the ipsilateral forelimb (iF) which evoked EPSPs in 49% and IPSPs in 73% of the LRN neurones. Each of the other limbs evoked EPSPs in approximately 20% and IPSPs in approximately 25% of the neurones. Stimulation of the ipsilateral trigeminal nerve (iTrig) evoked EPSPs in 32% and IPSPs in 46% of the neurones. The shortest latencies of the EPSPs and IPSPs indicated a disynaptic connection between primary afferents in the iF and iTrig and the LRN. The most direct pathways for excitatory and inhibitory responses from the other limbs were trisynaptic. Stimulation of the ventral part of the ipsilateral funiculus (iVLF) at C3 (C3iVLF) evoked monosynaptic responses in 189 of 201 tested LRN neurones. Monosynaptic EPSPs were recorded in 104 neurones and monosynaptic IPSPs in 126 neurones. Monosynaptic EPSPs and IPSPs were encountered in all parts of the LRN. Stimulation of the iVLF at L1 (L1iVLF) evoked monosynaptic EPSPs and IPSPs in the ventrolateral part of the LRN. The termination areas of excitatory and inhibitory fibres appeared to be the same. LRN neurones without monosynaptic EPSPs or IPSPs from the L1iVLF were located mainly in the dorsal part of the magnocellular division. Stimulation of the dorsal funiculi (DF) at C2 and the ipsilateral trigeminal nerve (iTrig) evoked excitatory and inhibitory responses in the LRN. The shortest latencies of EPSPs and IPSPs indicated disynaptic connections.     

Organization of neurones in the cat cerebral cortex that are influenced from Group I muscle afferents

1. Neurones in the Group I projection area of the first sensori-motor cortex were investigated with extra- and intracellular technique.2. The majority of the neurones influenced by volleys in Group I afferents of contralateral forelimb nerves received exclusively excitation, but some received exclusively inhibition, and some mixed excitation and inhibition.3. The Group I influenced cells were usually found 500-1500 mu beneath the cortical surface.4. The EPSPs and IPSPs evoked from Group I afferents had a steep rising phase and a slow, approximately exponential decay. The duration was sometimes more than 50 msec. The EPSP evoked by a maximal Group I volley was often formed by a small number of large unitary EPSPs.5. Latency measurements indicate that the majority of the Group I activated neurones were monosynaptically excited from the thalamic fibres, and hence constitute the fourth-order neurones in the Group I projection system. The latencies of the IPSPs suggest a disynaptic linkage with thalamic fibres. Hence, the exclusively inhibited cells would constitute fifth-order neurones. It is suggested that most or all of the fourth-order neurones are inhibitory.6. The convergence of excitation and/or inhibition to individual cells was usually extensive and included effects not only from muscle groups working at different joints but also effects from antagonistic groups at the same joint. In addition cutaneous afferents contributed synaptic actions which had a longer latency than the synaptic actions from Group I afferents.7. The neurones influenced from Group I afferents were not antidromically activated on stimulation of the pyramidal tract.     

Rubrospinal effects on ventral spinocerebellar tract neurones.

Stimulation of the contralateral red nucleus evoked monosynaptic EPSPs in 14 of 82 ventral spinocerebellar tract neurones. In some of these cells the monosynaptic EPSP was followed by a disynaptic IPSP. The remaining cell population received di- or polysynaptic PSPs from the rubrospinal tract, either EPSPs or IPSPs or both. Convergence of the rubrospinal tract onto interneurones of the segmental pathways projecting to VSCT cells was demonstrated. Rubrospinal volleys facilitated disynaptic Ia IPSPs evoked in VSCT neurones from both flexors and extensors, as well as disynaptic Ib IPSPs. Facilitation of the Ia interneurones was disynaptic whereas facilitation of Ib interneurones was monosynaptic. Disynaptic rubrospinal EPSPs and IPSPs were facilitated by volleys in ipsi- as well as in contralateral cutaneous and high threshold muscle afferents. The complex pattern of projections from the rubrospinal tract onto VSCT neurones and the related reflex pathways gives further support to the hypothesis that these tract cells convey information on transmission through interneurones of the spinal segmental mechanisms.     

Rubrospinal Effects on Ventral Spinocerebellar Tract Neurones

Stimulation of the contralateral red nucleus evoked monosynaptic EPSPs in 14 of 82 ventral spinocerebellar tract neurones. In some of these cells the monosynaptic EPSP was followed by a disynaptic IPSP. The remaining cell population received di- or polysynaptic PSPs from the rubrospinal tract, either EPSPs or IPSPs or both. Convergence of the rubrospinal tract onto interneurones of the segmental pathways projecting to VSCT cells was demonstrated. Rubrospinal volleys facilitated disynaptic Ia IPSPs evoked in VSCT neurones from both flexors and extensors, as well as disynaptic Ib IPSPs. Facilitation of the Ia interneurones was disynaptic whereas facilitation of Ib interneurones was monosynaptic. Disynaptic rubrospinal EPSPs and IPSPs were facilitated by volleys in ipsi- as well as in contralateral cutaneous and high threshold muscle afferents. The complex pattern of projections from the rubrospinal tract onto VSCT neurones and the related reflex pathways gives further support to the hypothesis that these tract cells convey information on transmission through interneurones of the spinal segmental mechanisms.     

Effects on the ventral spinocerebellar tract neurones from deiters' nucleus and the medial longitudinal fascicle in the cat.

Effects from the vestibulospinal tract (VST) and from fibres descending in the medial longitudinal fascicle (MLF) on the cells of origin of the ventral spinocerebellar tract (VSCT) have been studied with intracellular recording. Out of 110 VSCT neurones, the VST evoked monosynaptic EPSPs in 27, di- or polysynaptic EPSPs in 56 and disynaptic IPSPs in 26. In 93 tested VSCT cells, MLF stimulation evoked monosynaptic EPSPs in 26, monosynaptic IPSPs in 2, di- or polysynaptic EPSPs in 25 and disynaptic IPSPs in 21. Convergence of monosynaptic EPSPs from VST and MLF was found in a small proportion of cells whereas the two descending pathways evoked reciprocal effects in another small group of neurones. Convergence of monosynaptic EPSPs from VST or MLF and from group I afferents was also modest. In 9 VSCT neurones there was convergence of monosynaptic excitation and disynaptic inhibition from the vestibulospinal tract and the same pattern from MLF was recorded in 9 neurones. The results are discussed in view of the hypothesis that VSCT neurones carry information on the interneuronal ttransmission in the spinal cord.     

Vestibular-evoked postsynaptic potentials in Deiters neurones

Summary Stimulation of the vestibular nerve induced EPSPs monosynaptically in 29% of cat's Deiters neurones sampled on the ipsilateral side. These EPSPs started with latencies of 0.6–1.0 msec, rose sharply with a summit time of 0.5 msec and decayed exponentially with a time constant of 0.9–1.7 msec. Then amplitudes were graded finely according to the intensity of the vestibular nerve stimulation, the maximal size being 5–10 mV. The unitary EPSPs, evoked by vestibular nerve stimulation at juxta-threshold intensity or appearing spontaneously, were as small as 0.2–0.3 mV in amplitude. Those neurones monosynaptically activated by vestibular nerve volleys were located in the ventral portion of the nucleus of Deiters, in agreement with histological data. The vestibular nerve impulses also produced delayed EPSPs with latencies of 1.0–1.8 msec, presumably disynaptically. They occurred in many Deiters neurones located not only ventrally but also dorsally. Even later EPSPs often were superposed on the monosynaptic EPSPs with latencies of 1.9–2.2 msec. There is evidence that they were caused by repetitive discharges in the vestibular nerve fibres which occur in response to single shock stimulation of the vestibular nerve. IPSPs were produced only polysynaptically in some Deiters neurones in association with the monosynaptic EPSPs.     

Corticofugal influences on transmission to the dorsal spinocerebellar tract from hindlimb primary afferents

Summary Effects from the cerebral cortex on neurones of the dorsal spinocerebellar tract (DSCT) were examined: I. In group I units (units receiving monosynaptic excitation from group I fibres) repetitive stimulation of the contralateral sensorimotor cortex usually inhibited impulse transmission from the primary afferents. The inhibition had a latency of 10–20 msec and lasted for 82-100 msec or more. Discharges induced by muscle stretch were also inhibited by the cortical stimulation. DSCT units belonging to extensors and flexors were both inhibited from the cortex. In a small percentage of group I units the inhibition was preceded by a shorter-lasting excitation. 2. FRA units (units receiving excitation from cutaneous and/or high threshold muscle afferents) were typically excited by the cortical stimulation. The excitation was often followed by a period of depression of transmission from the periphery. 3. It is suggested from the effective cortical area and experiments with lesions in the medullary pyramid and in the spinal cord that the inhibition in group I units and the excitation of FRA units are both mediated by the corticospinal tract.Experiments were also made to determine the level where the cell body of a given DSCT unit is located, and the results from 56 units are presented.     

Neuronal relays in double crossed pathways between feline motor cortex and ipsilateral hindlimb motoneurones

Coupling between pyramidal tract (PT) neurones and ipsilateral hindlimb motoneurones was investigated by recording from commissural interneurones interposed between them. Near maximal stimulation of either the left or right PT induced short latency EPSPs in more than 80% of 20 commissural interneurones that were monosynaptically excited by reticulospinal tract fibres in the medial longitudinal fascicle (MLF). The EPSPs were evoked at latencies that were only 1–2 ms longer than those of EPSPs evoked from the MLF, compatible with a disynaptic coupling between PT fibres and these commissural interneurones. EPSPs evoked by PT stimulation were frequently associated with IPSPs which either followed or preceded the EPSPs. The latencies of the IPSPs (on average about 1 ms longer than latencies of the earliest EPSPs) indicated that they were mediated via single additional inhibitory interneurones. Records from a sample of nine commissural interneurones from a different population (with monosynaptic input from group I and/or II muscle afferents, and disynaptically excited from the MLF) suggest that actions of PT fibres on such interneurones are weaker because only four of them were excited by PT stimuli and at longer latencies. By demonstrating disynaptic coupling between PT neurones and commissural interneurones via reticulospinal fibres, the results provide a direct demonstration of trisynaptic coupling in the most direct pathways between PT neurones and ipsilateral motoneurones, and thereby strengthen the proposal that the double crossed pathways between PT neurones and ipsilateral motoneurones might be used to replace crossed actions of damaged PT neurones.     

Uncrossed actions of feline corticospinal tract neurones on lumbar interneurones evoked via ipsilaterally descending pathways

Effects of stimulation of ipsilateral pyramidal tract (PT) fibres were analysed in interneurones in midlumbar segments of the cat spinal cord in search of interneurones mediating disynaptic actions of uncrossed PT fibres on hindlimb motoneurones. The sample included 44 intermediate zone and ventral horn interneurones, most with monosynaptic input from group I and/or group II muscle afferents and likely to be premotor interneurones. Monosynaptic EPSPs evoked by stimulation of the ipsilateral PT were found in 12 of the 44 (27%) interneurones, while disynaptic or trisynaptic EPSPs were evoked in more than 75%. Both appeared at latencies that were either longer or within the same range as those of disynaptic EPSPs and IPSPs evoked by PT stimuli in motoneurones, making it unlikely that premotor interneurones in pathways from group I and/or II afferents relay the earliest actions of uncrossed PT fibres on motoneurones. These interneurones might nevertheless contribute to PT actions at longer latencies. Uncrossed PT actions on interneurones were to a great extent relayed via reticulospinal neurones with axons in the ipsilateral medial longitudinal fascicle (MLF), as indicated by occlusion and mutual facilitation of actions evoked by PT and MLF stimulation. However, PT actions were also relayed by other supraspinal or spinal neurones, as some remained after MLF lesions. Mutual facilitation and occlusion of actions evoked from the ipsilateral and contralateral PTs lead to the conclusion that the same midlumbar interneurones in pathways from group I or II muscle afferents may relay uncrossed and crossed PT actions.     

post-synaptic excitation and inhibition from primary afferents in neurones of the spinocervical tract

1. Intra- and extracellular recordings were made from cells of the spinocervical tract in the lumbosacral spinal cord. A convergence of monosynaptic excitatory post-synaptic potentials (EPSPs) and disynaptic inhibitory post-synaptic potentials (IPSPs) was a general pattern of effects from the low threshold cutaneous fibres. Unitary IPSPs, probably mediated via the same disynaptic path, were evoked by light touch of hairs, which was also the adequate stimulus for exciting the cells. The receptive field for unitary IPSPs was closely related to the excitatory receptive field but was eccentric, not of a surround type.

2. EPSPs, IPSPs, or both, were evoked from the flexor reflex afferents in the great majority of neurones. Disynaptic IPSPs may be evoked from the interosseous nerve. No effects were produced by volleys in group I muscle afferents.

3. It is suggested, on the basis of the spatial organization of the excitatory and inhibitory receptive skin fields, that the spinocervical tract may give information regarding the direction of tactile stimuli.

     

Effects on the Ventral Spinocerebellar Tract Neurones from Deiters' Nucleus and the Medial Longitudinal Fascicle in the Cat

Effects from the vestibulospinal tract (VST) and from fibres descending in the medial longitudinal fascicle (MLF) on the cells of origin of the ventral spinocerebellar tract (VSCT) have been studied with intracellular recording. Out of 110 VSCT neurones, the VST evoked monosynaptic EPSPs in 27, di- or polysynaptic EPSPs in 56 and disynaptic lPSPs in 26. In 93 tested VSCT cells, MLF stimulation evoked monosynaptic EPSPs in 26, monosynaptic IPSPs in 2, di- or polysynaptic EPSPs in 25 and disynaptic IPSPs in 21, Convergence of monosynaptic EPSPs from VST and MLF was found in a small proportion of cells whereas the two descending pathways evoked reciprocal effects in another small group of neurones. Convergence of monosynaptic EPSPs from VST or MLF and from group 1 afferents was also modest. In 9 VSCT neurones there was convergence of monosynaptic excitation and disynaptic inhibition from the vestibulospinal tract and the same pattern from MLF was recorded in 9 neurones. The results are discussed in view of the hypothesis that VSCT neurones carry information on the interneuronal transmission in the spinal cord.     

Axon reflex activation of Deiters neurones from the cerebellar cortex through collaterals of the cerebellar afferents

Summary When recording intracellularly from cat's Deiters neurones, stimulation of the anterior lobe of the cerebellar cortex produced excitatory postsynaptic potentials (EPSPs) monosynaptically, in addition to the inhibitory ones (IPSPs) that were identified previously as being produced via Purkinje cell axons. The EPSPs were induced bilaterally from a wide area of the anterior and posterior lobes of the cerebellum, in contrast to the IPSPs that were evoked only ipsilaterally, mainly from the vermal cortex. The latency of the EPSPs was slightly, but significantly, shorter than that of the IPSPs. The presynaptic impulses responsible for these EPSPs were represented by the discrete field potentials and also by unit spikes of individual fibres. The pathway for these EPSPs and presynaptic impulses was pursued by testing their interference, in the manner of impulse collision and refractoriness, with those induced from various spots within or outside the cerebellum. It is found that the excitatory fibres for Deiters neurones extend transversely, and probably longitudinally too, over the culmen and pass out of the cerebellum through cerebellar peduncles. The major portion of them appears to originate from the medulla and a minority from the spinal cord. It is postulated that cerebellar afferents from these structures have synapses with Deiters neurones via their collateral branches, through which a kind of axon reflex occurs to Deiters neurones during stimulation of the cerebellar cortex.     

The lateral reticular nucleus in the cat

Intracellular recordings were obtained from 204 neurones in the lateral reticular nucleus (LRN). LRN neurones contacted by the bVFRT were identified by the responses evoked on stimulation of descending fibres in the contralateral ventral quadrant of the spinal cord (cVQ) at cervical (C5cVQ) and lumbar (L2cVQ) levels. Stimulation of the cVQ evoked excitatory or inhibitory responses in 124 of the 204 LRN neurones. EPSPs were evoked in 45, IPSPs in 52 and both EPSPs and IPSPs in 27 LRN neurones. The shortest latencies of the responses evoked from the cVQ indicated that both EPSPs and IPSPs were disynaptic. This finding was confirmed by direct stimulation of the ascending fibres in the ipsilateral ventrolateral funiculus at C3 (C3iVLF) or L1 (L1iVLF). In most LRN neurones activated or inhibited from the cVQ, stimulation of the iVLF evoked similar responses at a monosynaptic latency. These results indicate that the bVFRT consists of roughly equally large groups of excitatory and inhibitory neurones monosynaptically connected with the LRN. Excitatory and inhibitory bVFRT neurones had similar peripheral receptive fields and termination areas in the LRN. LRN neurones were divided into those contacted by cervical bVFRT neurones and lumbar bVFRT neurones. The former group consisted of LRN neurones responding to C5cVQ stimulation at latencies below 5 ms, whereas the latter group contained LRN neurones responding to stimulation of the L2cVQ. Cervical bVFRT neurones projected to most parts of the LRN whereas the projection of lumbar bVFRT neurones were confined to the ventrolateral part of the nucleus. Excitatory and inhibitory vVFRT neurones of each group had similar termination areas.(ABSTRACT TRUNCATED AT 250 WORDS)     

Neuronal activity in the lateral vestibular nucleus of the cat

The synaptic input to Deiters neurones evoked by stimulation of peripheral somatic nerves was measured by intracellular recordings. EPSPs with broad receptive fields and latencies which indicate polysynaptic connexions were commonly evoked from the FRA. In other cells, low threshold cutaneous afferents were effective at rather short latencies suggesting oligosynaptic connexions from fast ascending fibres. One example was found of EPSPs due to low threshold muscle afferents. IPSPs due to climging fibre activation of Purkinje cells as observed in most of the neurones were evoked by cutaneous volleys above 1.5-2.0T and muscle volleys above 5T (above 3-3.5T in case of Q). Often, IPSPs were evoked by stimulation of nerves, to the segmental level of which the the vestibulospinal neurone under investigation projected. A small proportion of cells received short latency IPSPs involving direct fast mossy fibre tracts, which were evoked from low threshold cutaneous afferents. IPSPs due to polysynaptic mossy fibre activation of Purkinje cells were evoked from the FRA bilaterally and from ipsilateral cutaneous afferents at 1.5-2.0T ("prolonged inhibition"). Prolonged excitatory/inhibitory events mediated by mossy fibre pathways may be involved in quadruped locomotion or other processes making use of a broad motor integration.     

Mutual inhibition between olivary cell groups projecting to different cerebellar microzones in the cat

Summary Intracellular recording was made in the C3-C4 segments from cell bodies of a previously described system of propriospinal neurones (PNs), which receive convergent monosynaptic excitation from different higher motor centres and mediate disynaptic excitation and inhibition from them to forelimb motoneurones. Inhibitory effects in these PNs have now been investigated with electrical stimulation of higher motor centres and forelimb nerves. Short-latency IPSPs were evoked by volleys in the cortico-, rubro- and tectospinal tracts and from the reticular formation. Latency measurements showed that those IPSPs which required temporal summation were disynaptically mediated. After transection of the corticospinal tract in C2, only small and infrequent disynaptic IPSPs were evoked from the pyramid. It is postulated that disynaptic pyramidal IPSPs only to a small extent are evoked by monosynaptic excitation of reticulospinal inhibitory neurones known to project directly to the PNs, and that they are mainly mediated by inhibitory interneurones in the C3-C4 segments. Tests with spatial facilitation revealed monosynaptic excitatory convergence from tecto-, rubro- and probably also from reticulospinal fibres on inhibitory interneurones monosynaptically excited from corticospinal fibres (interneuronal system I). Disynaptic IPSPs were also evoked in the great majority of the PNs by volleys in forelimb muscle and skin nerves. A short train of volleys was usually required to evoke these IPSPs from group I muscle afferents. In the case of cutaneous nerves and mixed nerves single volleys were often effective, and the lack of temporal facilitation of IPSPs produced by a train of volleys showed strong linkage from these nerves. The results obtained after transection of the dorsal column at different levels show that the relay is almost entirely rostral to the forelimb segments. Test with spatial facilitation revealed that interneurones monosynaptically activated from forelimb afferents receive convergent excitation from corticospinal but not or only weakly so from tecto- or rubrospinal fibres. There was also convergence from group I muscle afferents and low threshold cutaneous afferents on common interneurones. It is postulated that the disynaptic IPSPs from forelimb afferents are mediated by inhibitory interneurones (interneuronal system II) other than those receiving convergent descending excitation. Volleys in corticospinal fibres, in addition to the disynaptic IPSPs, evoke late IPSPs in the PNs. Similar late IPSPs were evoked from the ipsilateral forelimb by stimulation of the FRA. Monosynaptic IPSPs were evoked in the majority of the PNs on weak stimulation of the lateral reticular nucleus (LRN) and from regions dorsal to it. Results from threshold mapping suggest that these IPSPs are due to antidromic stimulation of ascending inhibitory neurones which also project to the C3-C4 PNs, and that the ascending collaterals terminate in the LRN or/and the base of the cuneate nuclei. Activity in the ascending collaterals may give higher centres information regarding inhibitory control of the PNs. It is postulated that interneuronal system I subserves descending feed-forward inhibition and interneuronal system II feed-back inhibition from the forelimb of transmission through the C3-C4 PNs to motoneurones.This work was supported by the Swedish Medical Research Council (project no. 94)     

Excitation of Group I activated thalamocortical relay neurones in the cat

1. Extracellular recordings have been made from 134 group I activated neurones in the ventrobasal thalamic complex. One hundred of the neurones were identified as thalamocortical relay neurones by antidromic activation from the projection areas for group I afferents.2. The discharge evoked by group I volleys showed characteristic fluctuations in latency and number of spikes. The mechanisms underlying these phenomena are discussed.3. Half of the relay neurones were activated from afferents in only one of six dissected forelimb nerves innervating muscle groups at the various forelimb joints. Two-thirds of the relay neurones were activated from at least two adjacent synergistic muscles.4. The pattern of group I convergence in the thalamic neurones is compared with that at the cuneate and cortical levels of the group I pathway to the cerebral cortex. It is suggested that integration of information from synergistic muscles occurs at the thalamic level and that integration of information from muscle groups of more unrelated function occurs at the cortical level.5. A few of the thalamocortical cells (15%) were activated by group II muscle afferents, often in the same nerve as that which provided group I excitation. Weakly linked excitation from cutaneous afferents was observed in 39% of the neurones.6. Stimulation of the group I projection area in the first sensorimotor cortex at moderately high stimulus frequencies produced a trans-synaptic excitation in most of the group I activated cells.     

Integration in descending motor pathways controlling the forelimb in the cat

A further analysis has been made of inhibitory pathways to motoneurones via C3-C4 propriospinal neurones (PNs). Intracellular recording was made from triceps brachi motoneurones and effects from higher centres and forelimb afferents on corticospinal IPSPs were investigated after transection of the corticospinal tract at the C5/C6 border. The shortest latencies of the IPSPs evoked by stimulation of the pyramid were as brief as those of the pyramidal EPSPs (Illert et al. 1977). It is postulated that the minimal linkage of the pyramidal IPSPs is disynaptic via inhibitory C3-C4 PNs projecting directly to motoneurones. It was confirmed that pyramidal IPSPs usually are depressed by volleys in forelimb motor axon collaterals (Illert and Tanaka 1978). A quantitative comparison was made of the recurrent depression of pyramidal IPSPs and of IPSPs caused by activation of the Ia inhibitory interneurones. The result support the hypothesis of two parallel inhibitory cortico-motoneuronal pathways via C3-C4 PNs, one disynaptic via the inhibitory PNs and the other trisynaptic via excitatory PNs and Ia inhibitory interneurones. Pyramidal volleys also evoked late IPSPs which in some cases were not depressed from forelimb motor axon collaterals. It is postulated that the late IPSPs are partly due to activation of inhibitory C3-C4 PNs. Disynaptic pyramidal IPSPs were effectively facilitated by volleys in rubro-, tecto- and reticulospinal fibres - but not from vestibulospinal fibres - showing a convergence from the former descending tracts on common inhibitory C3-C4 PNs. Projection from forelimb afferents and corticospinal fibres on common inhibitory C3-C4 PNs was revealed by strong facilitation of disynaptic pyramidal IPSPs from cutaneous forelimb afferents. No corresponding effect was evoked from C2 neck afferents. Stimulation in the lateral reticular nucleus (LRN) evoked monosynaptic IPSPs in some motoneurones. The results of threshold mapping in and around the LRN suggest that the IPSPs are caused by antidromic stimulation of ascending collaterals of inhibitory neurones also projecting to motoneurones, possibly the inhibitory C3-C4 PNs.     

Convergence of excitatory and inhibitory action on interneurones in the lumbosacral cord

Summary Intracellular recording has been made in spinal cats from more than 100 interneurones in the dorsal horn and intermediary region of the lumbosacral spinal cord. The majority of interneurones receive not only EPSPs but also IPSPs from primary afferents. The IPSPs are evoked from three different systems, group I muscle afferents (probably Ib), low threshold cutaneous afferents and the FRA. The shortest central latency of the IPSPs indicates a disynaptic linkage from primary afferents. Interneurones with monosynaptic EPSPs from group I muscle afferents may receive IPSPs from all the above mentioned afferent systems. Interneurones with monosynaptic EPSPs from cutaneous afferents receive their inhibition from the two latter afferent systems. Convergence of EPSPs and IPSPs from the FRA may occur on the same interneurone. The results are discussed mainly with respect to inhibitory interaction between spinal reflex pathways.This work was supported by the Swedish Medical Research Council (Project No 14X-94-02A).IBRO-Unesco fellow     

Neck muscle afferent input to spinocerebellar tract cells of the central cervical nucleus in the cat

Summary Extracellular and intracellular recordings were made from spinocerebellar tract neurones of the central cervical nucleus (CCN) in C1–C3 segments of the anaesthetized cat. These neurones were identified by antidromic activation from the cerebellar peduncle. Stimulation of the ipsilateral dorsal root elicited extracellular spikes or EPSPs with a monosynaptic latency in almost all CCN neurones in the same segment (segmental input). Late excitatory effects were also observed in about one third of CCN neurones. The monosynaptic EPSP was occasionally followed by an IPSP. The excitatory input from the dorsal root to CCN neurones was extended over several segments for some CCN neurons (extrasegmental input). Monosynaptic excitation was evoked in CCN neurones after stimulation of dorsal neck muscle nerves as well; i.e. splenius (SPL), biventer cervicis and complexus (BCC), rectus capitus dorsalis, and obliquus capitus caudalis. Thresholds for this excitation were near the threshold of the nerve, suggesting that it originated from group I fibres. The component of excitation added after strong stimulation of neck muscle nerves would be attributed to group II fibres. When a CCN neurone received excitatory input from the nerve of one muscle, it was generally not affected by stimulation of other nerves in the same segment. Such muscle specificity of segmental input was the principal pattern of connexion of neck muscle afferents with CCN neurones. In some cases, however, excitatory convergence from SPL and BCC nerves onto single CCN neurones or excitation from the SPL nerve and inhibition from the BCC nerve were also observed. Nearly half of the CCN neurones received input from one muscle nerve of the same segment and not from the afferent of the same muscle of different segments, indicating a segment specificity of input. In the remaining CCN neurones, weaker excitatory effects were induced from afferents of different segments as well. In such extrasegmental effects, inputs to CCN neurones from caudal segments predominated in frequency over those from rostral segments. The origin of extrasegmental input was generally confined to the same muscle. Low threshold muscle afferents from the SPL and BCC were intraaxonally stained with HRP. The collaterals of the stained fibre distributed branchlets and terminals to the CCN, laminae VII, VIII, and motor nuclei. Two fibres responding to local muscle prodding or stretch showed a similar morphology. The findings indicated that muscle spindle afferents from primary endings projected to the CCN.Supported by a Grant-in-Aid for Scientific Research from the Ministry of Education, Science and Culture of Japan     

Premotor interneurones contributing to actions of feline pyramidal tract neurones on ipsilateral hindlimb motoneurones

The aim of the study was to analyse the potential contribution of excitatory and inhibitory premotor interneurones in reflex pathways from muscle afferents to actions of pyramidal tract (PT) neurones on ipsilateral hindlimb motoneurones. Disynaptic EPSPs and IPSPs evoked in motoneurones in deeply anaesthetized cats by group Ia, Ib and II muscle afferents were found to be facilitated by stimulation of the ipsilateral, as well as of contralateral, PT. The ipsilateral actions were evoked by either uncrossed or double-crossed pathways. The results show that interneurones mediating reflex actions of muscle afferents may be activated strongly enough by PT stimulation to contribute to movements initiated by ipsilateral PT neurones and that PT actions relayed by them might be enhanced by muscle stretches and/or contractions. However, in some motoneurones disynaptic IPSPs and EPSPs evoked from group Ib or II afferents were depressed by PT stimulation. In order to analyse the basis of this depression, the transmitter content in terminals of 11 intracellularly labelled interneurones excited by PT stimulation was defined immunohistochemically and their axonal projections were reconstructed. The interneurones included 9 glycinergic and 2 glutamatergic neurones. All but one of these neurones were mono- or disynaptically excited by group I and/or II afferents. Several projected to motor nuclei and formed contacts with motoneurones. However, all had terminal projections to areas outside the motor nuclei. Therefore both inhibitory and excitatory interneurones could modulate responses of other premotor interneurones in parallel with direct actions on motoneurones.