Interneurones In Reflex Pathways Research Articles

A survey of spinal collateral actions of feline ventral spinocerebellar tract neurons

The aim of this study was to identify spinal target cells of spinocerebellar neurons, in particular the ventral spinocerebellar tract (VSCT) neurons, giving off axon collaterals terminating within the lumbosacral enlargement. Axons of spinocerebellar neurons were stimulated within the cerebellum while searching for most direct synaptic actions on intracellularly recorded hindlimb motoneurons and interneurons. In motoneurons the dominating effects were inhibitory [inhibitory postsynaptic potentials (IPSPs) in 67% and excitatory postsynaptic potentials (EPSPs) in 17% of motoneurons]. Latencies of most IPSPs indicated that they were evoked disynaptically and mutual facilitation between these IPSPs and disynaptic IPSPs evoked by group Ia afferents from antagonist muscles and group Ib and II afferents from synergists indicated that they were relayed by premotor interneurons in reflex pathways from muscle afferents. Monosynaptic EPSPs from the cerebellum were accordingly found in Ia inhibitory interneurons and intermediate zone interneurons with input from group I and II afferents but only oligosynaptic EPSPs in motoneurons. Monosynaptic EPSPs following cerebellar stimulation were also found in some VSCT neurons, indicating coupling between various spinocerebellar neurons. The results are in keeping with the previously demonstrated projections of VSCT neurons to the contralateral ventral horn, showing that VSCT neurons might contribute to motor control at a spinal level. They might thus play a role in modulating spinal activity in advance of any control exerted via the cerebellar loop.

Do premotor interneurons act in parallel on spinal motoneurons and on dorsal horn spinocerebellar and spinocervical tract neurons in the cat?

It has previously been established that ventral spinocerebellar tract (VSCT) neurons and dorsal spinocerebellar tract neurons located in Clarke's column (CC DSCT neurons) forward information on actions of premotor interneurons in reflex pathways from muscle afferents on α-motoneurons. Whether DSCT neurons located in the dorsal horn (dh DSCT neurons) and spinocervical tract (SCT) neurons are involved in forwarding similar feedback information has not yet been investigated. The aim of the present study was therefore to examine the input from premotor interneurons to these neurons. Electrical stimuli were applied within major hindlimb motor nuclei to activate axon-collaterals of interneurons projecting to these nuclei, and intracellular records were obtained from dh DSCT and SCT neurons. Direct actions of the stimulated interneurons were differentiated from indirect actions by latencies of postsynaptic potentials evoked by intraspinal stimuli and by the absence or presence of temporal facilitation. Direct actions of premotor interneurons were found in a smaller proportion of dh DSCT than of CC DSCT neurons. However, they were evoked by both excitatory and inhibitory interneurons, whereas only inhibitory premotor interneurons were previously found to affect CC DSCT neurons [as indicated by monosynaptic excitatory postsynaptic potentials (EPSPs) and inhibitory postsynaptic potentials (IPSPs) in dh DSCT and only IPSPs in CC DSCT neurons]. No effects of premotor interneurons were found in SCT neurons, since monosynaptic EPSPs or IPSPs were only evoked in them by stimuli applied outside motor nuclei. The study thus reveals a considerable differentiation of feedback information provided by different populations of ascending tract neurons.

Functional subdivision of feline spinal interneurons in reflex pathways from group Ib and II muscle afferents; an update

A first step towards understanding the operation of a neural network is identification of the populations of neurons that contribute to it. Our aim here is to reassess the basis for subdivision of adult mammalian spinal interneurons that mediate reflex actions from tendon organs (group Ib afferents) and muscle spindle secondary endings (group II afferents) into separate populations. Re-examining the existing experimental data, we find no compelling reasons to consider intermediate zone interneurons with input from group Ib afferents to be distinct from those co-excited by group II afferents. Similar patterns of distributed input have been found in subpopulations that project ipsilaterally, contralaterally or bilaterally, and in both excitatory and inhibitory interneurons; differences in input from group I and II afferents to individual interneurons showed intra- rather than inter-population variation. Patterns of reflex actions evoked from group Ib and II afferents and task-dependent changes in these actions, e.g. during locomotion, may likewise be compatible with mediation by premotor interneurons integrating information from both group I and II afferents. Pathological changes after injuries of the central nervous system in humans and the lineage of different subclasses of embryonic interneurons may therefore be analyzed without need to consider subdivision of adult intermediate zone interneurons into subpopulations with group Ib or group II input. We propose renaming these neurons 'group I/II interneurons'.

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.

Common interneurones in reflex pathways from cutaneous afferents innervating different foot regions in humans

Common interneurones in reflex pathways from cutaneous afferents innervating different foot regions in humans

Medullary reticulospinal tract mediating a generalized motor inhibition in cats: iii. functional organization of spinal interneurons in the lower lumbar segments

The previous report of intracellular recording of hindlimb motoneurons in decerebrate cats [ Neuroscience 103 (2001) 511] has suggested that the following mechanisms are involved in a generalized motor inhibition induced by stimulating the medullary reticular formation. First, the motor inhibition, which was prominent in the late latency (30–80 ms), can be ascribed to the inhibitory effects in parallel to motoneurons and to interneuronal transmission in reflex pathways. Second, both a group of interneurons receiving inhibition from flexor reflex afferents and a group of Ib interneurons mediate the late inhibitory effects upon the motoneurons. To substantiate the above mechanisms of motor inhibition we examined the medullary stimulus effects upon intracellular ( n=55) and extracellular ( n=136) activity of spinal interneurons recorded from the lower lumbar segments (L6–L7). Single pulses or stimulus trains (1–3) pulses, with a duration of 0.2 ms and intensity of 20–50 μA) applied to the medullary nucleus reticularis gigantocellularis evoked a mixture of excitatory and inhibitory effects with early (<20 ms) and late (>30 ms) latencies. The medullary stimulation excited 55 interneurons (28.8%) with a late latency. Thirty-nine of the cells, which included 10 Ib interneurons, were inhibited by volleys in flexor reflex afferents (FRAs). These cells were mainly located in lamina VII of Rexed. On the other hand, the late inhibitory effects were observed in 67 interneurons (35.1%), which included cells mediating reciprocal Ia inhibition, non-reciprocal group I (Ib) inhibition, recurrent inhibition and flexion reflexes. Intracellular recording revealed that the late inhibitory effects were due to inhibitory postsynaptic potentials with a peak latency of about 50 ms and a duration of 50–60 ms. The inhibitory effects were attenuated by volleys in FRAs. Neither excitatory nor inhibitory effects with a late latency were observed in 69 (36.1%) cells which were located in the intermediate region and dorsal horn. These results suggest the presence of a functional organization of the spinal cord with respect to the production of the generalized motor inhibition. Lamina VII interneurons that receive inhibition from volleys in FRAs possibly mediate the postsynaptic inhibition from the medullary reticular formation in parallel to motoneurons and to interneurons in reflex pathways.

Medullary reticulospinal tract mediating the generalized motor inhibition in cats: Parallel inhibitory mechanisms acting on motoneurons and on interneuronal transmission in reflex pathways

The present study was designed to elucidate the spinal interneuronal mechanisms of motor inhibition evoked by stimulating the medullary reticular formation. Two questions were addressed. First, whether there is a parallel motor inhibition to motoneurons and to interneurons in reflex pathways. Second, whether the inhibition is mediated by interneurons interposed in known reflex pathways. We recorded the intracellular activity of hindlimb motoneurons in decerebrate cats and examined the effects of medullary stimulation on these neurons and on interneuronal transmission in reflex pathways to them. Stimuli (three pulses at 10–60 μA and 1–10 ms intervals) delivered to the nucleus reticularis gigantocellularis evoked inhibitory postsynaptic potentials in α-motoneurons ( n=147) and γ-motoneurons ( n=5) with both early and late latencies. The early inhibitory postsynaptic potentials were observed in 66.4% of the motoneurons and had a latency of 4.0–5.5 ms with a segmental delay of more than 1.4 ms. The late inhibitory postsynaptic potentials were observed in 98.0% of the motoneurons and had a latency of 30–35 ms, with a peak latency of 50–60 ms. Both types of inhibitory postsynaptic potentials were evoked through fibers descending in the ventrolateral quadrant. The inhibitory postsynaptic potentials were not influenced by recurrent inhibitory pathways, but both types were greatly attenuated by volleys in flexor reflex afferents. Conditioning medullary stimulation, which was subthreshold to evoke inhibitory postsynaptic potentials in the motoneurons, neither evoked primary afferent depolarization of dorsal roots nor reduced the input resistance of the motoneurons. However, the conditioning stimulation often facilitated non-reciprocal group I inhibitory pathways (Ib inhibitory pathways) to the motoneurons in early (<20 ms) and late (30–80 ms) periods. In contrast, it attenuated test postsynaptic potentials evoked through reciprocal Ia inhibitory pathways, and excitatory and inhibitory pathways from flexor reflex afferent and recurrent inhibitory pathways. The inhibitory effects were observed in both early and late periods. The present results provide new information about a parallel inhibitory process from the medullary reticular formation that produces a generalized motor inhibition by acting on α- and γ-motoneurons, and on interneurons in reflex pathways. Interneurons receiving inhibition from flexor reflex afferents and a group of Ib interneurons may mediate the inhibitory effects upon motoneurons.

Nociceptive input to spinal interneurones in reflex pathways from group II muscle afferents in cats.

Effects of noxious stimulation of the skin by radiant heat were tested on responses of first order interneurones in reflex pathways from group II muscle afferents in mid-lumbar, lower-lumbar and sacral segments of the spinal cord. In mid- and lower-lumbar segments both background discharges and monosynaptically evoked responses of intermediate zone interneurones were facilitated. Those of mid-lumbar dorsal horn interneurones were also facilitated suggesting that both these interneuronal populations contribute to the facilitation of flexion reflexes by nociceptors. In contrast, the dominating effects of noxious heat on sacral dorsal horn group II interneurones were inhibitory. The effects evoked by selective activation of C fibres, after A-delta fibres had been blocked by TTX, were similar to those obtained before TTX application.

The medullary reticular formation (MRF) produces parallel inhibitory effects on motoneurons (MNs) and on interneurons in reflex pathways in cats

The medullary reticular formation (MRF) produces parallel inhibitory effects on motoneurons (MNs) and on interneurons in reflex pathways in cats

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)

Segmental actions of afferents of the interosseous nerve in the cat.

Electrical stimulation of the interosseous nerve evokes oligosynaptic inhibition of extensor motoneurones and excitation of flexor motoneurones. Lowest-threshold, shortest-latency post-synaptic potentials evoked at group I strength are attributed to the action of group Ib afferents. Post-synaptic potentials evoked at slightly higher stimulus strengths (within the higher group I and the group II range) and at longer latency are attributed to the action of afferents of Pacinian corpuscles. Facilitation of post-synaptic potentials evoked from afferents in the interosseous nerve by group I muscle afferents and by joint afferents is taken to indicate convergence of these afferents onto common interneurones in reflex pathways to motoneurones. Evidence is presented that afferents of Pacinian corpuscles project to the interneurones mediating group I (non-reciprocal) reflex actions to motoneurones. Unitary monosynaptic excitatory post-synaptic potentials (e.p.s.p.s) evoked from the interosseous nerve are taken to indicate that only a very small number of muscle spindle Ia afferents course through the interosseous nerve. Dorsal root potentials evoked by low-strength electrical stimulation of the interosseous nerve are largely attributable to the action of afferents of Pacinian corpuscles.

Inhibitory interactions between interneurones in reflex pathways from group Ia and group Ib afferents in the cat.

A hypothesis has been verified that laminae V-VI interneurones which mediate non-reciprocal inhibition of motoneurones from group I muscle afferents have collateral actions on other laminae V-VI interneurones. Stimulation within the areas of projection of these inhibitory interneurones in motor nuclei and in Clarke's column would be expected to give rise to monosynaptic i.p.s.p.s in interneurones with disynaptic i.p.s.p.s from group I afferents if the hypothesis were correct. Intracellular records were made from eighty-five laminae V-VI interneurones with input from group Ia muscle spindle and/or group Ib tendon organ afferents. Weak intraspinal stimuli applied in motor nuclei in L7 and S1 segments, or in the lateral funiculus just caudal to Clarke's column in L4, were found to evoke monosynaptic i.p.s.p.s in seventy-two interneurones. These i.p.s.p.s were systematically correlated with disynaptic inhibition from group Ia or Ib afferents but not from other fibres. Such monosynaptic i.p.s.p.s evoked by intraspinal stimuli were seen in forty-two interneurones which themselves projected to the level of Clarke's column and therefore (on the basis of previous evidence) should mediate inhibition of motoneurones. For seven of these interneurones it was also shown directly that they projected to motor nuclei. The inhibition of such interneurones demonstrates mutual interactions between those interneurones which are interposed in inhibitory pathways from group I afferents. Only indirect indications have been obtained for inhibition of interneurones in the excitatory pathways.

Facilitatory interaction between cutaneous afferents from low threshold mechanoreceptors and nociceptors in segmental reflex pathways to α-motoneurons

EPSPs evoked by air puff stimulation of the hairy skin of the hindlimb or by low strength stimulation of the lateral sural nerve were facilitated by radiant heat of 50–53°C but not by radiant heat of 42–43°C. The results indicate that afferents from low threshold mechanoreceptors and from nociceptors of the skin converge onto common interneurons in reflex pathways to α-motoneurons. This would allow a functionally useful cooperation of these two types of afferents in spinal motor control.

Convergence in segmental reflex pathways from fine muscle afferents and cutaneous or group II muscle afferents to α-motoneurones

Synaptic responses to hindlimb motoneurones, which were evoked by cutaneous or group II muscle afferents, were distinctly facilitated by selectively chemically activated group III and IV afferents of the gastrocnemius-soleus muscle. The results indicate a convergence from these different afferents onto common interneurones in reflex pathways to a-motoneurones. This convergence furnishes further evidence for a multisensory cooperation in movement control.

Common interneurones in reflex pathways from group 1a and 1b afferents of knee flexors and extensors in the cat.

1. Input from group I afferents of knee flexors and extensors to interneurones in Rexed's laminae V-VI in the cat spinal cord was analysed by use of intracellular recording and electrical stimulation of the nerves to differentiate between group Ia and Ib synaptic actions. The aim was to find out if these interneurones may mediate synaptic actions of both group Ia and Ib afferents. 2. 28% of the forty-nine neurones analysed were excited by both group Ia and group Ib afferents; 32% were inhibited by both and 35% were excited by the one and inhibited by the other. Taking into account all of these actions, input from both subgroups of group I afferents was found in nearly 60% of neurones. Most were also excited and/or inhibited by group I afferents of ankle and toe extensors. 3. Selective (excitatory and/or inhibitory) input from Ia afferents was found in 18% and from Ib afferents in 22% of the neurones. 4. Excitation was evoked from Ia afferents of either knee flexors or extensors but not from both. In several of the neurones Ia i.p.s.p.s were, however, evoked from both posterior biceps-semitendinosus and quadriceps. 5. Intracellular staining with horseradish peroxidase revealed axonal projections of laminae V-VI interneurones to motor nuclei as well as to the intermediate zone, ipsilateral as well as contralateral. No correlation was found between patterns of input from group I afferents and axonal projections, and interneurones co-excited by Ia and Ib afferents were among these with different axonal projections.

Common interneurones in reflex pathways from group 1a and 1b afferents of ankle extensors in the cat.

1. Input from group I afferents of ankle and toe extensors, other muscles, skin nerves and descending tracts to interneurones of Rexed's laminae V-VI in the cat spinal cord was analysed using intracellular recording from these interneurones. Adequate stimuli (muscle stretches) were used to activate selectively group Ia muscle spindle afferents of triceps surae and plantaris while other fibre systems were excited electrically. 2. Ia and Ib afferents of ankle and toe extensors were found to co-excite, co-inhibit or exert opposite synaptic actions in 41, 33, and 50% of the analysed interneurones, respectively. Taking into account both excitatory and inhibitory input from these two groups of afferents, 64% of the interneurones appeared to be used in common in reflex pathways from muscle spindles and tendon organs of ankle and toe extensors. 3. Selective input from Ib afferents of triceps surae and plantaris (excitation and/or inhibition) was found in 36% of the interneurones; there was evidence for a similarly selective input from Ia afferents. 4. A great majority (over 90%) of the interneurones excited by group I afferents were also inhibited by group I afferents, from either the same or other muscles. 5. Both monosynaptic and disynaptic e.p.s.p.s from Ia and/or Ib afferents from other muscles and from fibres in the ipsilateral funiculi were found in a great proportion of the same interneurones, together with disynaptic e.p.s.p.s from low threshold cutaneous afferents. 6. Intracellular staining with horseradish peroxidase revealed four different patterns of axonal projections of the analysed interneurones: (i) projections to motor nuclei and the intermediate region, (ii and III) projections only to the intermediate region, locally or combined with projections to different rostro-caudal levels, and (iv) projections to the opposite side of the spinal cord. 7. A large proportion of interneurones projecting to motor nuclei displayed input from both Ia and Ib afferents although such an input was a feature of interneurones with other projections as well. No systematic differences in the input from group I afferents were found for interneurones with different axonal projections. In contrast disynaptic e.p.s.p.s of cutaneous origin and monosynaptic e.p.s.p.s upon stimulation of ipsilateral spinal tracts appeared predominantly in interneurones projecting to motor nuclei.

Multisensory Control of Spinal Reflex Pathways

This chapter reviews the convergence from different primary afferents on interneurones in reflex pathways. It summarizes results showing that interneurones of the reciprocal Ia inhibitory pathway receive a wide convergence resembling that found in agonist motoneurones. Interneurones in reflex pathways from Ib afferents are excited from cutaneous and joint afferents and also from Ia afferents. The chapter describes reflexes from the FRA (flexor reflex afferents)—that is, reflex actions from high threshold muscle afferents, joint afferents and cutaneous afferents that are mediated by common interneurones. A distinction is made between short-latency reflexes in the acute spinal and long-latency reflexes in acute spinal cats after DOPA i.v.; the half-centre organisation of the interneuronal network mediating the latter reflexes is recapitulated and discussed in relation to spinal generation of alternating activity in flexors and extensors. An important role of reflex pathways from the FRA in motor control is indicated by findings showing that their interneurones are excited from different higher motor centres and that many ascending pathways, spinocerebellar in particular, signal information regarding their interneuronal activity.

NEURON PATTERNS CONTROLLING TRANSMISSION OF IPSILATERAL HIND LIMB REFLEXES IN CAT

NEURON PATTERNS CONTROLLING TRANSMISSION OF IPSILATERAL HIND LIMB REFLEXES IN CAT