Reflex pathways from group II muscle afferents

The interneuronally mediated reflex actions evoked by electrical stimulation of group II muscle afferents in low spinal cats have been reinvestigated with intracellular recording with motoneurones to knee flexors and ankle extensors. The results of Eccles and Lundberg (1959) have been confirmed and extended. There was wide convergence from flexors and extensors of group II excitation to flexor and group II inhibition to extensor motoneurones. Some quantitative differences in the effect from the different nerves are described. Latency measurements suggest that the minimal linkage is disynaptic in the excitatory interneuronal pathways and trisynaptic in the inhibitory pathways. Disynaptic group II EPSPs were found in 14% of the ankle extensor motoneurones but were much more common in unanaesthetized high spinal cats (Wilson and Kato 1965). From these results and corresponding ones on flexors (Holmqvist and Lundberg 1961) it is postulated that secondary afferents in addition to the weak monosynaptic connexions (Kirkwood and Sears 1975) have disynaptic excitatory pathways and trisynaptic inhibitory pathways to both flexor and extensor motoneurones. It is proposed that the group II actions of the flexor reflex pattern characterizing the anaesthetized low spinal cat are due to suppression of the inhibitory pathway to flexor motoneurones and the excitatory pathway to extensor motoneurones. In some ankle extensor motoneurones the disynaptic group II EPSPs occurred in combination with IPSPs from the FRA (including group II and III muscle afferents). The possibility is considered that these group II EPSPs are mediated by an interneuronal group II pathway with little or no input from group III muscle afferents but probably from extramuscular receptors. In other ankle extensor motoneurones group II EPSPs were combined with EPSPs from group III muscle afferents, cutaneous afferents and joint afferents. It is postulated that these group II EPSPs are mediated by an interneuronal pathway from the FRA which also supply interneuronal pathways giving inhibition to extensor or/and flexor motoneurones and excitation to flexors as postulated by Eccles and Lundberg (1959) and Holmqvist and Lundberg (1961).     

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     

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.     

Interneurones in pathways from group II muscle afferents in sacral segments of the feline spinal cord.

1. Properties of dorsal horn interneurones that process information from group II muscle afferents in the sacral segments of the spinal cord have been investigated in the cat using both intracellular and extracellular recording. 2. The interneurones were excited by group II muscle afferents and cutaneous afferents but not by group I muscle afferents. They were most effectively excited by group II afferents of the posterior biceps, semitendinosus, triceps surae and quadriceps muscle nerves and by cutaneous afferents running in the cutaneous femoris, pudendal and sural nerves. The earliest synaptic actions were evoked monosynaptically and were very tightly locked to the stimuli. 3. EPSPs evoked monosynaptically by group II muscle afferents and cutaneous afferents of the most effective nerves were often cut short by disynaptic IPSPs. As a consequence of this negative feedback the EPSPs gave rise to single or double spike potentials and only a minority of interneurones responded with repetitive discharges. However, the neurones that did respond repetitively did so at a very high frequency of discharges (0.8-1.2 ms intervals between the first 2-3 spikes). 4. Sacral dorsal horn group II interneurones do not appear to act directly upon motoneurones because: (i) these interneurones are located outside the area within which last order interneurones have previously been found and (ii) the latencies of PSPs evoked in motoneurones by stimulation of the posterior biceps and semitendinosus, cutaneous femoris and pudendal nerves (i.e. the main nerves providing input to sacral interneurones) are compatible with a tri- but not with a disynaptic coupling. Spatial facilitation of EPSPs and IPSPs following synchronous stimulation of group II and cutaneous afferents of these nerves shows, however, that sacral interneurones may induce excitation or inhibition of motoneurones via other interneurones. 5. Comparison of the properties of group II interneurones in the sacral segments with those of previously studied group II interneurones in the midlumbar segments leads to the conclusion that these two populations of neurones are specialized for the processing of information from different muscles and skin areas. In addition, equivalents of only one of the two subpopulations of midlumbar interneurones have been found at the level of the pudendal nucleus: neurones with input from group II but not from group I muscle afferents. Neurones integrating information from group I and II muscle afferents and in direct contact with motoneurones thus seem to be scarce in the sacral segments.(ABSTRACT TRUNCATED AT 400 WORDS)     

Both dorsal horn and lamina VIII interneurones contribute to crossed reflexes from feline group II muscle afferents

Previous studies have demonstrated that group II muscle afferents exert powerful actions on contralateral motoneurones and that these actions are mediated primarily via lamina VIII commissural interneurones. We examined whether dorsal horn interneurones also contribute to these actions, as they have been shown to contribute to the actions of group II afferents on ipsilateral motoneurones. We tested the susceptibility of IPSPs and EPSPs evoked from group II afferents in contralateral motoneurones to presynaptic inhibition as an indicator of the relative contribution of dorsal horn interneurones to these PSPs, since the monosynaptic activation of dorsal horn interneurones is more weakly and more briefly depressed by presynaptic inhibition than is the monosynaptic activation of lamina VIII and other intermediate zone and ventral horn interneurones. While the earliest components of IPSPs and EPSPs evoked by group II afferents were abolished by conditioning stimulation of group II afferents, consistent with them being evoked disynaptically by commissural interneurones, trisynaptic components of these PSPs were only partly reduced and are therefore attributed to dorsal horn interneurones. The same conditioning stimuli depressed the disynaptic excitation of lamina VIII commissural interneurones by group II afferents much less effectively than they depressed monosynaptic excitation, indicating that dorsal horn interneurones contribute to this disynaptic excitation. On the basis of these observations we conclude that that dorsal horn interneurones contribute to the late actions of group II muscle afferents on contralateral motoneurones through their disynaptic actions on commissural interneurones.     

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.     

Transmission in a locomotor-related group Ib pathway from hindlimb extensor muscles in the cat

It has been previously shown that phasic stimulation of group I afferents from ankle and knee extensor muscles may entrain and/or reset the intrinsic locomotor rhythm; these afferents are thus acting on motoneurones through the spinal rhythm generators. It was also concluded that the major part of these effects originates from Golgi tendon organ Ib afferents. Transmission in this pathway to lumbar motoneurones has now been investigated during fictive locomotion in spinal cats injected with nialamide and l-DOPA, and in decerebrate cats with stimulation of the mesencephalic locomotor region. In spinal cats injected with nialamide and l-DOPA, it was possible to evoke long-latency, long-lasting reflexes upon stimulation of high threshold afferents before spontaneous fictive locomotion commenced. During that period, stimulation of ankle and knee extensor group I afferents evoked oligosynaptic excitation of extensor motoneurones, rather than the classical Ib inhibition. Furthermore, a premotoneuronal convergence (spatial facilitation) between this group I excitation and the crossed extensor reflex was established. During fictive locomotion, in both preparations, the transmission in these group I pathways was phasically modulated within the step cycle. During the flexor phase, the group I input cut the depolarised (active) phase in flexor motoneurones and evoked EPSPs in extensor motoneurones; during the extensor phase, the group I input evoked smaller EPSPs in extensor motoneurones and had virtually no effect on flexor motoneurones. The above results suggest that the group I input from extensor muscles is transmitted through the spinal rhythm generator and more particularly, through the extensor half-centre. The locomotor-related group I excitation had a central latency of 3.5–4.0 ms. The excitation from ankle extensors to ankle extensors remained after a spinal transection at the caudal part of L6 segment; the interneurones must therefore be located in the L7 and S1 spinal segments. Candidate interneurones for mediating these actions were recorded extracellularly in lamina VII of the 7th lumbar segment. Responses to different peripheral nerve stimulation (high threshold afferents and group I afferents bilaterally) were in concordance with the convergence studies in motoneurones. The interneurones were rhythmically active in the appropriate phases of the fictive locomotor cycle, as predicted by their response patterns. The synaptic input to, and the projection of these candidate interneurones must be fully identified before their possible role as components of the spinal locomotor network can be evaluated.     

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     

Reflex pathways from group II muscle afferents

Summary A hypothesis is forwarded regarding the role of secondary spindle afferents and the FRA (flexor reflex afferents) in motor control. The hypothesis is based on evidence (cf. Lundberg et al. 1987a, b) summarized in 9 introductory paragraphs. Group II excitation. It is postulated that subsets of excitatory group II interneurones (transmitting disynaptic group II excitation to motoneurones) may be used by the brain to mediate motor commands. It is assumed that the brain selects subsets of interneurones with convergence of secondary afferents from muscles whose activity is required for the movement. During movements depending on coactivation of static -motoneurones impulses in secondary afferents may servo-control transmission to -motoneurones at an interneuronal level. The large group II unitary EPSPs in interneurones are taken to indicate that, given an adequate interneuronal excitability, impulses in single secondary afferents may fire the interneurone and produce EPSPs in motoneurones; interneuronal transmission would then be equivalent to that in a monosynaptic pathway but with impulses from different muscles combining into one line. It is postulated that impulses in the FRA are evoked by the active movements and that the role of the multisensory convergence from the FRA onto the group II interneurones is to provide the high background excitability which allows the secondary spindle afferents to operate as outlined above. The working hypothesis is put forward that a movement governed by the excitatory group II interneurones is initiated by descending activation of these interneurones, but is maintained in a later phase by the combined effect of FRA activity evoked by the movement and by spindle secondaries activated by descending activation of static -motoneurones. As in the original follow up length servo hypothesis (Rossi 1927; Merton 1953), we assume that a movement at least in a certain phase can be governed from the brain solely or mainly via static -motoneurones. However, our hypothesis implies that the excitatory group II reflex connexions have a strength which does not allow transmission to motoneurones at rest and that the increase in the gain of transmission during an active movement is supplied by the movement itself. Group II inhibition. It is suggested that the inhibitory reflex pathways like the excitatory ones have subsets of interneurones with limited group II convergence. When higher centres utilize a subset of excitatory group II interneurones to evoke a given movement, they may mobilize inhibitory subsets to inhibit muscles not required in the movement. Inhibition may be reciprocal of extensors during flexor activation (the spinal pattern), of flexors during extensor activation or of flexors and extensors in more complex movements involving cocontraction of other flexors and extensors. It is postulated that group II inhibition depends on conjoint activation from spindle afferents and other sources (descending and/or the FRA) so that inhibition may be coupled to group II excitation of other motoneurones. Such a coupling would correspond to the --linkage in reciprocal Ia inhibition (Lundberg 1970) and is denoted --linkage in lateral group II inhibition. FRA and other reflex pathways. Results are summarized showing that the FRA evoke convergent excitation in interneurones not only in group II reflex pathways but also in other reflex pathways like the reciprocal Ia inhibitory, the nonreciprocal group I inhibitory and probably also in specialized reflex pathways from cutaneous afferents. It is inferred that facilitation of reflex transmission by impulses in the FRA evoked by the active movement may be a general principle. In this way reflex transmission to -motoneurones may be weak at rest and not disturb passive movements but have a high gain when the reflexes are required to regulate active movement.This work was supported by the Swedish Medical Research Council (project no. 94)     

The disynaptic group I inhibition between wrist flexor and extensor muscles revisited in humans

The present studies are designed to further characterise the interneuronal pathway mediating the disynaptic reciprocal group I inhibition between flexors and extensors at the wrist and the elbow levels in humans. In the first series of experiments, we compared the electrical threshold of the reciprocal group I inhibition at the wrist and the elbow level after a prolonged vibration aimed at raising the electrical threshold of the antagonistic activated Ia afferents. Prolonged vibration to the ‘conditioning’ tendon, which raised significantly the electrical threshold of the inhibition at the elbow level, did not alter it at the wrist level. These results suggest that the dominant input to the relevant interneurones is Ia in origin at the elbow level but Ib in origin at the wrist level. In the second series of experiments, using the spatial facilitation method, we compared the effects on the post-stimulus time histograms of single voluntarily activated motor units of two volleys delivered both separately and together to group I afferents in the nerves supplying the homonymous and antagonistic muscles. At the wrist, but not at the elbow level, the peak of homonymous monosynaptic group I excitation was reduced on combined stimulation, although the antagonistic IPSP was just at the threshold. Because the suppression did not involve the initial bins of the peak, it is argued that the suppression is not due to presynaptic inhibition of Ia terminals, but probably reflects convergence between the homonymous and antagonistic volleys onto the interneurones mediating the disynaptic inhibition. Taken together with the previously reported effects of recurrent inhibition on reciprocal inhibition, these results suggest that inhibition between flexors and extensors is differently organised at the elbow (reciprocal Ia inhibition) and the wrist (non-reciprocal group I inhibition) levels. It is argued that the particular connectivity at the wrist level might correspond to some functional requirements at this ball joint.     

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.     

Evidence for recurrent inhibition of reciprocal inhibition from soleus to tibialis anterior in Man

Reciprocal inhibition between ankle flexors and extensors has been the subject of numerous studies in Man. They have demonstrated that this reciprocal inhibition is in all likelihood caused by a disynaptic pathway at least partly fed by Ia afferents. It is thus generally agreed that this reciprocally organized inhibition between ankle flexors and extensors in Man is similar to the reciprocal Ia inhibition described in the cat. This conclusion has, however, been challenged, when Jankowska and McCrea described in the cat a non-reciprocal group I inhibition involving interneurones co-excited by Ia and Ib afferents and mediating inhibition to both antagonistic and non-antagonistic motoneurones. The only way to distinguish between reciprocal Ia inhibition and non-reciprocal group I inhibition is to test if the inhibition is blocked by recurrent inhibition, since only Ia interneurones are inhibited by recurrent inhibition. In the present study, reciprocal inhibition from soleus to tibialis anterior was thus investigated following activation of soleus-coupled Renshaw cells in normal human subjects. It was found that reciprocal inhibition induced in tibialis anterior motoneurones by the activation of soleus group I afferents is deeply depressed by activation of soleus-coupled Renshaw cells. This finding provides the missing data to identify disynaptic inhibition between antagonistic ankle muscles as a reciprocal Ia inhibition.     

Facilitatory interaction in spinal reflex pathways from nociceptive cutaneous afferents and identified secondary spindle afferents in the cat

Summary Using the technique of spike triggered averaging it could be demonstrated that di- or oligosynaptic EPSPs evoked from single identified secondary spindle afferents in lumbar motoneurones received spatial facilitation from nociceptive cutaneous afferents. This indicates that both types of afferents converge onto common interneurones in the segmental reflex pathways and confirms the assumption that secondary spindle afferents contribute to a multisensorial spinal system for motor control.Supported by the Deutsche Forschungsgemeinschaft (Scho 37/3-1)     

Cutaneous facilitation of transmission in reflex pathways from Ib afferents to motoneurones.

1. The effect of volleys in low threshold cutaneous afferents upon transmission of synaptic action from Ib afferents to motoneurones has been investigated with intracellular recording from alpha motoneurones to hind limb muscles. 2. There was facilitation from cutaneous afferents of transmission in excitatory and inhibitory reflex pathways from Ib afferents without any evidence for difference in effect on di- and trisynaptic pathways. It is postulated that volleys in cutaneous afferents evoke excitatory action in interneurones of these reflex pathways. 3. The time course of the facilitation suggest that cutaneous afferents have disynaptic excitatory connexions with the interneurones intercalated in the disynaptic Ib inhibitory pathways to motoneurones. 4. Some observations are reported suggesting that interneuronal transmission in Ib inhibitory pathways to motoneurones might be facilitated from Ia afferents. 5. The findings are discussed in relation to the presumed role of Ib reflex action in regulating muscle tension.     

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)     

Contribution of group III and IV muscle afferents to multisensorial spinal motor control in cats

The contribution of group III and IV muscle afferents to multisensorial segmental reflex pathways was investigated by testing for spatial facilitation between these afferents and non-nociceptive segmental afferents from skin, muscles and joints on postsynaptic potentials (PSPs) in alpha-motoneurones recorded in anaemically decapitated high spinal cats. Group III and IV muscle afferents were activated by intraarterial injection of potassium chloride (320 mM) or bradykinin triacetate (81 microM). Skin, joint and group I-II muscle afferents were stimulated by graded electrical stimulation of various nerves. Conditioning by stimulation of group III and IV muscle afferents spatially facilitated the transmission in segmental reflex pathways from low- to medium-threshold cutaneous and joint afferents as well as from lb and group II muscle afferents. Both excitatory and inhibitory pathways from these afferents were facilitated. Monosynaptic excitation and disynaptic antagonistic inhibition from group Ia afferents remained unaffected. It is concluded that the spatial facilitation observed between group III and IV muscle afferents and the other afferents indicate a convergence from group III and IV muscle afferents and the other afferents on common interneurones in segmental flexor reflex pathways. Under physiological conditions they thus contribute to the multisensorial feedback of the flexor reflex pathways. Pathophysiologically, the observed convergence may aggravate muscle weakness and atrophy of muscles induced by group III and IV muscle afferents.     

Integration in descending motor pathways controlling the forelimb in the cat

Summary A previously described disynaptic pathway from cortex to forelimb motoneurones whose intercalated neurones were excited both from other descending pathways and from forelimb afferents (Illert et al., 1976a, b) has been further analysed, mainly with respect to the location of the relay cells and their axons.Disynaptic EPSPs evoked in forelimb motoneurones by stimulation of the pyramid remained after complete transection of the corticospinal tract in C5 rostral to the forelimb segments but were abolished after a more rostral transection of the tract in the C2 segment. Corresponding findings were made with disynaptic rubral EPSPs after transection of the rubrospinal tract in these segments. It is concluded that disynaptic cortico-motoneuronal and rubro-motoneuronal excitation is relayed by propriospinal neurones originating in the C3–C4 segments. Other lesion experiments revealed that the axons of these propriospinal neurones descend to forelimb motoneurones in the ventrolateral part of the lateral funicle.Spatial facilitation of transmission from the corticospinal and rubrospinal tracts after transection of them in C5 occurred with a time course showing monosynaptic convergence from these pathways on common propriospinal neurones.Facilitation of disynaptic pyramidal EPSPs from the dorsal tegmentum remained after transection of the corticospinal tract at C5 but was abolished after a transection at C2. It is postulated that corticospinal and presumed tectospinal fibres converge onto common neurones in the propriospinal relay but evidence is also given for a more rostral relay (probably bulbar) with a similar convergence.The oligo- (probably mono-)synaptic facilitation of the disynaptic pyramidal EPSP evoked by volleys in cutaneous and group I muscle afferents from the forelimb likewise remained after a C5 transection of the corticospinal tract but was abolished after an additional C5 lesion in the dorsal column. It is concluded that propriospinal relay cells receive excitatory action from forelimb afferents ascending in the dorsal column. Spatial facilitation experiments using three tests revealed that propriospinal neurones monosynaptically excited from both corticospinal and rubrospinal fibres also receive excitation from cutaneous forelimb afferents.It is postulated that the propriospinal relay provides an important route for fast activation of forelimb motoneurones from the brain. The convergent monosynaptic excitation from several important motor centres in the brain is considered in relation to the general problem of the functional relationship between higher motor centres. The convergent action from forelimb afferents is taken to suggest that a descending command for a forelimb movement can be modified from the forelimb while on its way to the motoneurones.Supported by the Deutsche Forschungsgemeinschaft     

On organization of a neuronal network in pathways from group II muscle afferents in feline lumbar spinal segments

The aim of the study was to verify the hypothesis that trisynaptic actions of group II muscle afferents upon motoneurones are, at least in part, mediated by dorsal horn interneurones exciting the same intermediate zone interneurones that are interposed in disynaptic pathways from group II afferents. Population EPSPs (field potentials) and responses of individual interneurones evoked by group II afferents in the dorsal horn and in the intermediate zone were analysed in order to assess the possibility of a causal relationship between them. When direct actions of group II afferents in the intermediate zone were abolished by presynaptic inhibition, distinct later components of field potentials and delayed interneuronal responses were induced at latencies 0.5-1 ms longer than those seen originally. Both the latency and a marked temporal facilitation define these later group II actions as being evoked disynaptically. Under the same conditions, single stimuli activated more than one half of dorsal horn interneurones, and the second and third stimuli activated all of these interneurones. Responses of dorsal horn interneurones preceded disynaptically evoked responses of intermediate zone interneurones. The study indicates that intermediate zone interneurones may be activated by group II afferents both directly and via dorsal horn interneurones and that synaptic actions of group II afferents upon these interneurones, and their subsequent actions upon motoneurones, may be modulated in parallel at the level of intermediate zone and dorsal horn interneurones.     

Properties of human spinal interneurones: normal and dystonic control

The muscles that control wrist posture receive large inputs from reflexes driven by hand afferents. In several studies, we have investigated these reflexes by electrical stimulation of cutaneous (median nerve) and proprioceptive (ulnar nerve) afferents from the hand. Median stimulation produced short latency inhibition in all motor nuclei investigated, possibly through inhibitory propriospinal-like interneurones. Ulnar stimulation produced similar inhibition but only in wrist extensors. In the other motor nuclei, ulnar stimulation produced short latency excitation mediated by group I motoneuronal drive through both monosynaptic and non-monosynaptic pathways involving excitatory propriospinal-like interneurones. This was followed by late excitations mediated through spinal group II and trans-cortical group I pathways. These results show that these pathways are concerned with the integration of afferent inputs, proprioceptive and cutaneous, to control of wrist posture during hand movements. Patients with focal hand dystonia exhibit abnormal postures. To investigate whether these spinal pathways contribute to these conditions, the effects of ulnar stimulation on wrist muscle activity during voluntary tonic contraction were examined in patients who suffer writer's cramp. Ulnar-induced inhibition of the wrist extensors was reduced on the dystonic side of patients compared with their normal side and controls. In patients who exhibited abnormal wrist posture, group II excitation of the wrist flexors was also modified on the dystonic side. Cutaneous stimuli, by contrast, increased wrist flexor EMG on both sides and only in patients who exhibited normal posture. We conclude that spinal interneurones have a significant role in integrating afferent inputs from the hand to control wrist posture during hand movements and that altered function in these spinal networks is associated with the complex pathophysiology of writer's cramp.