Reciprocal Ia Inhibition Research Articles

Reflex connexions of motoneurones of muscles involved in head movement in the cat.

1. The reflex connexions from muscle afferents and ventral root fibres to the motoneurones of the muscles biventer-cervicis, complexus, sternocleidomastoid, trapezius and splenius, the principal muscles involved in head movement in the cat, were studied with the technique of intracellular recording. 2. Electrical stimulation of homonymous muscle afferents of biventer-cervicis and complexus, sternocleidomastoid and trapezius, at strengths below 1.6 times threshold of the dorsal root afferent volley, produced monosynaptic e.p.s.p.s in the corresponding motoneurones. Recruitment of higher threshold muscle afferents produced additional p.s.p.s with longer central delays. 3. Stimulation of low-threshold muscle afferents did not produce any p.s.p.s in the motoneurones of the ipsilateral antagonist. Stimulation of higher threshold afferents evoked i.p.s.p.s with central delays longer than 1.6 msec, or mixed e.p.s.p.-i.p.s.p.s in the ipsilateral antagonist. 4. Mixed e.p.s.p.-i.p.s.p.s or i.p.s.p.s with central delays longer than 1.5 msec were evoked in trapezius motoneurones upon stimulation of high threshold afferents from biventer-cervicis and complexus, while stimulation of low-threshold biventercervicis and complexus afferents evoked no p.s.p.s in trapezius motoneurones. 5. Stimulation of contralateral low-threshold biventer-cervicis and complexus afferents evoked a sequence of i.p.s.p. disinhibition in sternocleidomastoid motoneurones, and vice versa, with central delays longer than 1.7 msec. 6. Stimulation of the deafferented biventer-cervicis, complexus, splenius, sternocleidomastoid and trapezius muscle nerves frequently activated interneurones in the ventral horn at monosynaptic central delays. Activation of homoynmous ventral root fibres rarely evoked p.s.p.s in biventer-cervicis, complexius, splenius or sternocleidomastoid motoneurones, while it produced disynaptic i.p.s.p.s in 80% of trapezius motoneurones. 7. It is concluded that Ia reciprocal inhibition and recurrent inhibition, two reflex circuits which are so prominent in limb segments of the spinal cord, do not play a major role in the generation of head movement. Rather, head movement may be primarily controlled from supraspinal centres.

Reciprocal inhibition during the tonic stretch reflex in the decerebrate cat.

1. The aim of this study was to investigate post-synaptic reciprocal Ia inhibition during the stretch reflex; particularly the extent to which an increased Ia excitation of the Ia inhibitory interneurones will be counteracted by recurrent inhibition from motor axon collaterals. For this purpose we investigated depression of monosynaptic test reflexes antagonist flexors (reciprocal inhibition) during static stretch of quadriceps or triceps surae in unanaesthetized decerebrate cats. 3. With increasing stretch of the extensor muscle there was first a linear augmentation of reciprocal inhibition, but along with the stretch reflex in the extensor a plateau appeared in the inhibition of the flexors, although the extensor stretch reflex (judged by the e.m.g.) increased with further stretching. Within the range of stretching of triceps surae which gave increased stretch reflexes the plateau in the reciprocal inhibition was usually maintained, while during stretching of quadriceps a second phase of augmenting reciprocal inhibition often appeared. Stretch beyond the level which increased the stretch reflex activity gave augmenting reciprocal inhibition both in case of quadriceps and triceps surae. 3. Excitability measurements from central terminals of Ia afferents revealed that the increasing reciprocal inhibition during increasing stretch reflex activity in quadriceps was associated with a primary afferent depolarization in knee flexor Ia afferents; there was no corresponding effect in ankle flexor Ia afferents during stretch reflexes in triceps surae. 4. The primary afferent depolarization evoked in knee flexor Ia afferents by electrical nerve stimulation was then compared with the presynaptic inhibition of knee flexor monosynaptic test reflexes produced by the same stimuli. The results suggest that the second phase of increasing reciprocal inhibition in knee flexors is due to presynaptic inhibition and accordingly that the depth of post-synaptic reciprocal inhibition remains constant at different degrees of stretch reflex activity in both knee and ankle extensors. 5. It is postulated that during increasing stretch reflex activity the increment in Ia excitation and recurrent inhibitio; on to the Ia inhibitory interneurones almost exactly balance each other. It is suggested that recurrent inhibition of Ia inhibitory interneurones may serve as a segmental autoregulatory mechanism to keep 'alpha-gamma-linked reciprocal inhibition' at a constant depth during different levels of agonist activity.

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. II. Effects from segmental flexor reflex pathways.

Interneurones identified as mediating the disynaptic reciprocal Ia inhibition of motoneurones (referred to as "Ia inhibitory interneurones") were recorded in the lumbar spinal cord of the cat. Volleys in ipsilateral and contralateral high threshold muscle afferents, cutaneous and high threshold joint afferents evoked a mixture of polysynaptic excitation and inhibition. These effects were ascribed to pathways activated by flexor reflex afferents (FRA) and in addition a specific ipsilateral low threshold cutaneous pathway. Ia inhibitory interneurones excited monosynaptically from flexor nerves received stronger net excitation by volleys in ipsilateral FRA than did extensor coupled interneurones, while the opposite pattern was seen from the contralateral FRA. These patterns are similar to those found in flexor and extensor motoneurones respectivey. The FRA inhibition in Ia inhibitory interneurones was partly mediated by "opposite" Ia inhibitory interneurones, i.e. those which are mediating the Ia inhibition of Ia inhibitory interneurones. The extent to which the FRA inhibition is transmitted by Ia inhibitory interneurones was roughly estimated by its susceptibility to recurrent depression by antidromic ventral root stimulation. The main conclusion is that most segmental pathways seem to evoke their effects in parallel to motoneurones and Ia inhibitory interneurones which are monosynaptically linked to the same muscle. The functional importance of this conclusion is discussed in a following report.

Convergence on interneurones mediating the reciprocal Ia inhibition of motoneurones. I. Disynaptic Ia inhibition of Ia inhibitory interneurones.

Interneurones identified as mediating the disynaptic reciprocal Ia inhibition of motoneurones (referred to as "Ia inhibitory interneurones") were recorded in the lumbar spinal cord of the cat. It was revealed that the Ia inhibitory interneurones themselves receive disynaptic Ia inhibition. The muscles from which this inhibition is evoked are strictly antagonistic to those supplying their Ia excitation. Similar to the Ia inhibition in motoneurones the Ia inhibition in the Ia inhibitory interneurones is decreased when preceded by an antidromic stimulation of ventral roots. Furthermore, transmission of Ia inhibition to the Ia inhibitory interneurones is facilitated from ipsilateral and contralateral primary afferents as well as several supraspinal pathways analogous to earlier findings for the Ia inhibition of motoneurones. The pattern and control of the Ia inhibition of motoneurones and of Ia inhibitory interneurones display so striking similarities that it is suggested that identical interneurones are responsible. The conclusion thus emerges that "opposite" Ia inhibitory interneurones (i.e. interneurones monosynaptically connected to antagonistic muscles) are mutually inhibiting each other. The functional significance of this organization is discussed.

Morphology of interneurones mediating Ia reciprocal inhibition of motoneurones in the spinal cord of the cat.

1. Interneurones identified by physiological criteria (Hultborn, Jankowska & Lindström, 1971b) to mediate Ia reciprocal inhibition of motoneurones in the spinal cord of the cat were stained by intracellular injection of a fluorescent dye (Procion Yellow).2. The somas of the stained cells were found in Rexed's lamina VII, just dorsal or dorsomedial to the motor nuclei. Their size was about 30 x 20 mum. The cells had four to five slender, weakly branching dendrites. The total extension of their dendritic trees was about 600 mum dorsoventrally, 400 mum mediolaterally and 300 mum rostrocaudally.3. The axons originated from a separate axon hillock or from the base of a dendrite. They were myelinated with external diameter of about 6-14 mum and projected to either the ipsilateral ventral or lateral funiculi. Early collaterals were found only exceptionally. Some axons bifurcated into an ascending and a descending branch within the funiculi.4. The possibility of identifying the Ia inhibitory interneurones on purely morphological grounds is discussed.

Synaptic actions of single interneurones mediating reciprocal Ia inhibition of motoneurones.

1. The investigation was aimed at defining the function of the interneurones which, according to indirect evidence, mediate the reciprocal Ia inhibition of motoneurones (Hultborn, Jankowska & Lindstrom, 1971 b) by studying their direct synaptic actions. These actions were tested by recording post-synaptic potentials in motoneurones following spike activity of single interneurones activated by iontophoretic application of glutamate. The interneurones were found to produce unitary monosynaptic IPSPs in those motoneurones in which disynaptic IPSPs are evoked by the group Ia afferents which monosynaptically excite the interneurones.2. Unitary IPSPs were found in more than 80% of the motoneurones impaled in the immediate vicinity of the axonal branches of the investigated Q interneurones in the PBSt motor nucleus. It is estimated that each interneurone might inhibit about every fifth PBSt motoneurone. The amplitudes of the unitary IPSPs ranged between 8 and 220 muV and were 10-200 times smaller than the maximal Ia IPSPs evoked in the same motoneurones.3. The synaptic delay in the generation of unitary IPSPs was measured in relation to the spike potentials recorded from the terminal branches of interneurones in the immediate vicinity of the impaled motoneurones. The synaptic delays ranged between 0.28 and 0.42 msec.4. From chloride reversal tests and an analysis of the time course of the unitary IPSPs it was concluded that the terminals of the investigated interneurones make synaptic contact predominantly on the soma and/or on the proximal parts of the dendrites of the motoneurones, their distribution being, however, not quite uniform.

An electrophysiological demonstration of the axonal projections of single spinal interneurones in the cat.

1. Single interneurones excited from group Ia afferents and located in the ventral horn of the spinal cord in the cat were activated antidromically by stimulation of their axons with one micro-electrode while recording extracellularly close to their somas with a second micro-electrode. The interneurones studied were those which, according to previous indirect evidence, should mediate the reciprocal Ia inhibition of motoneurones.2. Two subgroups of these interneurones were studied: those excited from group Ia afferents in the quadriceps (Q) and posterior bicepssemitendinosus (PBSt) nerves. Most of them could be activated from a number of separate loci in the PBSt and Q motor nuclei respectively and from the ventral or lateral funiculi.3. The location of the axonal branches and the extent of their branching in the motor nuclei were reconstructed by comparing the latencies of the responses and the thresholds (0.1-5 muA) for antidromic activation of single interneurones from different electrode positions in a number of tracks, having previously established the relation between the threshold and the distance from the stimulated fibres. For the main branches in the white matter the conduction velocity was found to be about 70 m/sec.4. The axonal projections of the investigated interneurones were found to be fully consistent with the hypothesis that they mediate reciprocal inhibition of motoneurones.

Reciprocal group I inhibition on triceps surae motoneurons in man.

Reciprocal group I inhibition on triceps surae motoneurons in man.

Neuronal pathway of the recurrent facilitation of motoneurones.

1. The recurrent facilitation of motoneurones is a disinhibition, i.e. a release of the motoneurones from a sustained hyperpolarization evoked by tonically active inhibitory interneurones. Only two groups of interneurones are known to receive recurrent inhibition from motor axon collaterals via Renshaw cells; the interneurones mediating the reciprocal Ia inhibition and the Renshaw cells themselves. The properties of these two groups of neurones were studied to determine if they could produce the tonic inhibition of motoneurones removed during recurrent facilitation.2. It was found that the tonic firing of Ia inhibitory interneurones is sensitive to anaesthetics to the same degree as is recurrent facilitation. The range of frequencies of tonic discharges of Renshaw cells appeared to be similarly low in unanaesthetized and anaesthetized preparations although in individual cells the discharge rates were decreased by anaesthesia.3. The recurrent inhibition of Ia interneurones inhibiting a given group of motoneurones and the recurrent facilitation of the same group of motoneurones were, as a rule, evoked from the same nerves, although in some cats the origin of the recurrent facilitation was somewhat wider. In contrast no evidence could be found that the Renshaw cells which inhibit a functional group of motoneurones are inhibited by volleys in the nerves from which recurrent facilitation is regularly evoked.4. It was concluded that the recurrent facilitation is caused mainly by inhibition of the tonic activity of Ia inhibitory interneurones and that it is thus a manifestation of the recurrent control of Ia reciprocal inhibition of motoneurones.

Recurrent inhibition of interneurones monosynaptically activated from group Ia afferents.

1. Interneurones monosynaptically excited from large muscle spindle (Ia) afferents and inhibited from motor axon collaterals were searched for in the lumbar spinal cord of the cat.2. Monosynaptic Ia excitation was found in sixty-seven of sixty-nine interneurones inhibited by antidromic volleys. These interneurones were excited from Ia afferents from one or a few muscles (mainly close synergists). Volleys in high threshold muscle and skin afferents (FRA) evoked polysynaptic excitation or inhibition. Weak inhibition from Ia afferents (from antagonists to those giving Ia excitation) was seen in a few cells. Monosynaptic excitation was evoked from the ventral quadrant of the spinal cord and polysynaptic excitation from the dorsal quadrant.3. Inhibition from motor axon collaterals was evoked with a latency (1.2-2.0 msec) suggesting a disynaptic linkage and had the same time course as in motoneurones. It prevented synaptic activation of 60% of interneurones and decreased the firing index and delayed generation of spikes in the remaining.4. The interneurones with convergence of monosynaptic Ia excitation and inhibition from motor axon collaterals were found in the ventral horn dorsomedial to motor nuclei. No inhibition by antidromic volleys could be detected in interneurones located in intermediate nucleus and activated monosynaptically from Ia, Ib, group I or cutaneous afferents.5. It was concluded that the ventral Ia interneurones inhibited by volleys in recurrent motor axon collaterals mediate the reciprocal Ia inhibition to motoneurones.

Central pathway for direct inhibitory action of impulses in largest afferent nerve fibres to muscle.

Central pathway for direct inhibitory action of impulses in largest afferent nerve fibres to muscle.

FACILITATION AND INHIBITION OF SPINAL MOTONEURONS

FACILITATION AND INHIBITION OF SPINAL MOTONEURONS

INFLUENCE OF DISCHARGE OF MOTONEURONS UPON EXCITATION OF NEIGHBORING MOTONEURONS

INFLUENCE OF DISCHARGE OF MOTONEURONS UPON EXCITATION OF NEIGHBORING MOTONEURONS