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 from the vestibulospinal tract on transmission from primary afferents to ventral spino-cerebellar tract neurones.

Convergence of vestibulospinal and segmental effects onto spinal interneurones which project to the ventral spino-cerebellar tract (VSCT) neurones has been studied by intracellular recording in VSCT cells. The disynaptic Ia IPSPs evoked in a group of VSCT neurones from the quadriceps nerve are monosynaptically facilitated by the vestibulospinal tract while there was no facilitation of Ia IPSP evoked from a flexor nerve. These results support the view that Ia inhibition to VSCT cells and motoneurones is mediated by common interneurones. The disynaptic inhibition evoked in other VSCT cells from the vestibulospinal tract is facilitated by volleys in the contralateral flexor reflex afferents (FRA) or bilaterally from the FRA. It is postulated that these actions are mediated by collaterals of the interneurones responsible for the analogous effects in motoneurones. Findings are reported suggesting that the monosynaptic vestibulospinal EPSP in VSCT cells in most cases is collateral to the excitatory input to the last order interneurones of reflex pathways from the FRA to motoneurones and only exceptionally to the corresponding input to Ia inhibitory interneurones. In many VSCT cells the vestibulospinal tract evoked disynaptic EPSPs which are facilitated from the FRA; the functional significance of this action is uncertain. The results are consistent with the hypothesis that VSCT neurones signal information on interneuronal transmission to motoneurones.     

Convergence on interneurones mediating the reciprocal Ia inhibition of motoneurones. III. Effects from supraspinal pathways.

Supraspinal effects were investigated in interneurones identified as mediating the disynaptic reciprocal Ia inhibition of motoneurones (referred to as Ia inhibitory interneurones). It was revealed that volleys in the vestibulospinal tract may evoke mono- and disynaptic EPSPs in interneurones monosynaptically excited from extensor muscles, i.e. extensor coupled Ia inhibitory interneurones. Flexor coupled interneurones instead received disynaptic inhibition. Volleys in the rubrospinal tract evoked a dominating polysynaptic excitation, usually mixed with inhibition, in flexor as well as extensor coupled interneurones. Disynaptic rubrospinal EPSPs and IPSPs were also revealed. The pyramidal tract also gives rise to a dominating polysynaptic excitation, usually mixed with inhibition, in flexor as well as extensor coupled Ia inhibitory interneurones. Rubrospinal and pyramidal volleys were shown to facilitate transmission in various segmental reflex pathways to the Ia inhibitory interneurones. A detailed comparison reveals a striking parallelism of segmental and supraspinal effects on alpha-motoneurones and Ia inhibitory interneurones connected to the same muscles. This considerably strengthens the hypothesis of an "alpha-gamma-linkage in the reciprocal inhibition".     

Convergence on Interneurones Mediating the Reciprocal Ia Inhibition of Motoneurones III. Effects from supraspinal pathways

Supraspinal effects were investigated in interneurones identified as mediating the disynaptic reciprocal Ia inhibition of motoneurones (referred to as “Ia inhibitory interneurones”). It was revealed that volleys in the vestibulospinal tract may evoke mono- and disynaptic EPSPs in interneurones monosynaptically excited from extensor muscles, i.e. extensor coupled Ia inhibitory interneurones. Flexor coupled interneurones instead received disynaptic inhibition. Volleys in the rubrospinal tract evoked a dominating polysynaptic excitation, usually mixed with inhibition, in flexor as well as extensor coupled interneurones. Disynaptic rubrospinal EPSPs and IPSPs were also revealed. The pyramidal tract also gives rise to a dominating polysynaptic excitation, usually mixed with inhibition, in flexor as well as extensor coupled Ia inhibitory interneurones. Rubrospinal and pyramidal volleys were shown to facilitate transmission in various segmental reflex pathways to the Ia inhibitory interneurones. A detailed comparison reveals a striking parallelism of segmental and supraspinal effects on α-motoneurones and Ia inhibitory interneurones connected to the same muscles. This considerably strengthens the hypothesis of an “α–γ-linkage in the reciprocal inhibition”.     

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.     

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.     

Integration in descending motor pathways controlling the forelimb in the cat. 4. Corticospinal inhibition of forelimb motoneurones mediated by short propriospinal neurones

Summary A previously described inhibitory trisynaptic cortico-motoneuronal pathway (Illert et al., 1976a) was analysed in order to identify the two relay stations. Intracellular recording was made from motoneurones to elbow muscles. Corticospinal fibres were stimulated in the contralateral medullary pyramid.Pyramidal IPSPs were abolished by a transection of the Corticospinal tract in C2 but remained after a corresponding lesion in C5.After a C5 lesion pyramidal volleys facilitated transmission in the Ia inhibitory pathway with a time course suggesting disynaptic excitatory action on the Ia inhibitory interneurones.The trisynaptic pyramidal IPSPs were depressed by volleys in the appropriate recurrent motor axon collaterals as would be expected if these IPSPs were mediated by Ia inhibitory interneurones.It is concluded that trisynaptic cortico-motoneuronal inhibition is evoked by consecutive activation of propriospinal neurones in C3-C4 and segmental Ia inhibitory interneurones.Supported by the Deutsche ForschungsgemeinschaftIBRO/UNESCO Fellow     

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.     

Integration in descending motor pathways controlling the forelimb in the cat

Summary With intracellular recording from forelimb motoneurones the spatial facilitation technique has been used to investigate interaction between descending pathways and forelimb afferents.As previously shown for the hindlimb, pyramidal volleys effectively facilitate interneuronal transmission in reflex pathways from different primary afferents. Evidence is presented suggesting disynaptic excitation from corticospinal fibres of interneurones in the reciprocal Ia inhibitory pathway. Interneurones of other reflex pathways from group I muscle afferents receive monosynaptic pyramidal excitation. During pyramidal facilitation volleys in cutaneous afferents may evoke PSPs in motoneurones after a central delay of 1.3 ms suggesting that the minimal linkage is disynaptic.Information regarding convergence on the neurones intercalated in the disynaptic cortico-motoneuronal pathway was obtained by investigating the effect from primary afferents and from other descending pathways on the disynaptic pyramidal EPSPs. Volleys in cutaneous and group I muscle afferents facilitate transmission in the disynaptic cortico-motoneuronal pathway with a time course showing oligosynaptic (probably monosynaptic) action on the intercalated neurone. Rubrospinal volleys likewise effectively facilitate disynaptic cortico-motoneuronal transmission with a time course showing monosynaptic action on the intercalated neurone. Spatial facilitation experiments involving three tests revealed that those intercalated neurones which receive convergent monosynaptic excitation from corticospinal and rubrospinal fibres are excited also from cutaneous forelimb afferents.Disynaptic cortico-motoneuronal transmission was also monosynaptically facilitated by stimuli in the dorsal mesencephalic tegmentum probably activating tectospinal fibres. Disynaptic, presumed tectospinal EPSPs were facilitated from cutaneous forelimb afferents.The convergence onto the neurones intercalated in the disynaptic excitatory cortico-motoneuronal pathway suggests that these neurones integrate the activity in different descending pathways and primary forelimb afferents.Supported by the Deutsche ForschungsgemeinschaftIBRO/UNESCO Fellow     

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.     

Effects of Volleys in Cortico-spinal Tract Fibres on Ventral Spino-cerebellar Tract Cells in the Cat

Both excitation and inhibition has been found in cells of origin of the ventral spino-cerebellar tract (VSCT) to be evoked by volleys in cortico-spinal fibres. The earliest EPSPs and IPSPs had features of disynaptically evoked postsynaptic potentials; these were, however, found only in a small proportion of cells and polysynaptic EPSPs and IPSPs were dominating. Postsynaptic potentials evoked in VSCT cells from primary afferents were effectively facilitated by cortico-spinal volleys. The cortico-spinal effects on VSCT cells may thus well be mediated by the same interneurones which mediate their excitation or inhibition from the periphery and which could evoke similar postsynaptic potentials in motoneurones. Generally all the observations are in keeping with the hypothesis (Lundberg 1971) that VSCT cells monitor transmission through interneurones interposed in various reflex paths to motoneurones.     

Recurrent Control from Motor Axon Collaterals of la Inhibitory Pathways to Ventral Spinocerebellar Tract Neurones

The effects of impulses in recurrent motor axon collaterals on transmission in different inhibitory pathways to ventral spinocerebellar tract (VSCT) neurones were investigated in the cat by conditioning of IPSPs evoked in intracellularly recorded VSCT cells. Disynaptic IPSPs from large muscle spindle (la) afferenls were depressed in many but not all VSCT cells following an antidromic stimulation of ventral roots. The effect was found in VSCT neurones which themselves did not receive recurrent inhibition from motor axon collaterals. In cells with affected la IPSPs also some polysynaptic IPSPs evoked from ipsi- and contralateral group II muscle afferents and low threshold cutaneous afferents were depressed by a ventral root volley as well as disynaptic IPSPs from fibres descending on the ipsilateral side of the spinal cord. In unanesthetized preparations recurrent facilitatory potentials similar to those in motoneurones were evoked in VSCT neurones with la IPSPs. The findings indicate that some VSCT neurones receive collateral connexions from the interneurones which mediate la recipiocal inhibition to motoneurones and support the hypothesis that the VSCT conveys information about transmission in inhibitory reflex pathways to motoneurones (Lundberg 1971).     

Facilitation from contralateral primary afferents of interneuronal transmission in the Ia inhibitory pathway to motoneurones.

The action of volleys in contralateral primary afferents on transmission in the Ia inhibitory pathways to motoneurones was investigated with intracellular recording from motoneurones. Ia IPSPs in flexor as well as most extensor motoneurones were regularly facilitated by volleys in contralateral high threshold muscle, cutaneous and joint afferents in spinal cats under chloralose anaesthesia. In decerebrate cats with a low pontine lesion transmission in Ia inhibitory pathways was not facilitated but rather depressed by volleys in these afferents. The recurrent effects from motor axon collaterals were investigated on inhibitory transmission from different contralateral afferents to motoneurones. Previous investigations have shown that the interneurones mediating the reciprocal Ia inhibition receive recurrent inhibition via motor axon collaterals and Renshaw cells. Now a strong positive correlation was revealed between recurrent depression of IPSPs evoked from different contralateral afferents and facilitation of Ia IPSPs by the same afferent volleys. These results suggest that the recurrent depression of IPSPs from different contralateral primary afferents depends on their excitatory convergence onto the Ia inhibitory interneurones, which then partly mediate the IPSP evoked in the motoneurone from these afferents.     

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.     

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)     

Facilitation from Contralateral Primary Afferents of Interneuronal Transmission in the la Inhibitory Pathway to Motoneurones

The action of volleys in contralateral primary afferents on transmission in the la inhibitory pathways to motoneurones was investigated with intracellular recording from motoneurones. Ia IPSPs in flexor as well as most extensor motoneurones were regularly facilitated by volleys in contralateral high threshold muscle, cutaneous and joint afferents in spinal cats under chloralose anaesthesia. In decerebrate cats with a low pontine lesion transmission in la inhibitory pathways was not facilitated but rather depressed by volleys in these afferents. The recurrent effects from motor axon collaterals were investigated on inhibitory transmission from different contralateral afferents to motoneurones. Previous investigations have shown that the interneurones mediating the reciprocal la inhibition receive recurrent inhibition via motor axon collaterals and Renshaw cells. Now a strong positive correlation was revealed between recurrent depression of IPSPs evoked from different contralateral afferents and facilitation of la IPSPs by the same afferent volleys. These results suggest that the recurrent depression of IPSPs from different contralateral primary afferents depends on their excitatory convergence onto the la inhibitory interneurones, which then partly mediate the IPSP evoked in the motoneurone from these afferents.     

Reflex pathways from group II muscle afferents

The convergence of group II muscle afferents on interneurones in reflex pathways has been elucidated by investigating interaction in transmission to motoneurones. Recording was also made from interneurones activated from group II afferents. Maximal group II EPSPs evoked in motoneurones from different muscles (extensors or flexors and extensors) did not summate linearly but with a deficit of 35-40%. The corresponding deficit in summation with Ia EPSPs was 7%. It is suggested that the difference in deficit is caused largely by occlusion due to shared interneuronal discharge zones and that it gives an approximate minimal measure of the convergence of group II afferents from different muscles on the interneurones. Tests with weak group II volleys from different muscles gave no or little evidence for spatial facilitation in the disynaptic excitatory pathway to flexor motoneurones, and there was no or little temporal facilitation of transmission in this pathway. It is suggested that group II excitation of the interneurones in this pathway depends on few afferents giving large unitary EPSPs. Convergence of cutaneous afferents and joint afferents on the interneurones was evidenced by spatial facilitation from these afferents of group II transmission to motoneurones. Convergence on interneurones in the trisynaptic inhibitory pathway from group II afferents to extensor motoneurones was also investigated with the spatial facilitation technique. There was convergence on common interneurones of group II afferents from different muscles (extensors or flexors and extensors) and from cutaneous afferents as well as joint afferents. Trisynaptic group II IPSPs, including those depending on spatial facilitation from different muscles were resistant to recurrent depression from motor axon collaterals and are therefore not mediated by the reciprocal Ia inhibitory pathway. Interneurones with monosynaptic group II EPSPs were recorded from in the dorsal horn and intermediate region. Graded stimulation revealed large unitary EPSPs from few group II afferents. The EPSP evoked by a single group II afferent may produce firing (extracellular recording). Convergence of monosynaptic group II EPSPs from different muscles was rather limited but could be from flexors and extensors. Extensive multisensory convergence onto some of these interneurones was indicated by di- or polysynaptic EPSPs from group II and III muscle afferents, from joint afferents and from cutaneous afferents.(ABSTRACT TRUNCATED AT 400 WORDS)     

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.     

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