The effect of transcranial magnetic stimulation on reciprocal inhibition in the human leg.

The effect of magnetic stimulation on reciprocal Ia inhibition of the human leg was investigated. Stimulation of the common peroneal nerve at the fibula head at the threshold of the alpha motoneuron axons resulted in inhibition of the soleus (SOL) H reflex at a conditioning-test interval of 2 ms. Magnetic stimulation over the contralateral motor cortex resulted in complex modulations of the SOL H reflex, including a short latency facilitation followed by inhibition. This inhibition may have been conveyed by Ia inhibitory interneurons projecting to SOL motoneurons. To test for convergence, whether or not the magnetic stimulation was capable of facilitating disynaptic reciprocal Ia inhibition of the SOL H reflex induced by stimulation of the peroneal nerve, the two stimuli were given together or separately. We observed the inhibition significantly increased when the two stimuli were given together than separately. These results suggest that the Ia inhibitory interneurons projecting to SOL motoneurons in humans might receive convergent input from the motor area of the brain and from Ia afferents of the tibialis anterior (TA) muscle in humans as well as in other animals.     

Modulation of transmission in the corticospinal and group Ia afferent pathways to soleus motoneurons during bicycling

Transmission in the corticospinal and Ia pathways to soleus motoneurons was investigated in healthy human subjects during bicycling. Soleus H reflexes and motor evoked potentials (MEPs) after transcranial magnetic stimulation (TMS) were modulated similarly during the crank cycle being large during downstroke [concomitant with soleus background electromyographic (EMG) activity] and small during upstroke. Tibialis anterior MEPs were in contrast large during upstroke and small during downstroke. The soleus H reflexes and MEPs were also recorded during tonic plantarflexion at a similar ankle joint position, corresponding ankle angle, and matched background EMG activity as during the different phases of bicycling. Relative to their size during tonic plantarflexion, the MEPs were found to be facilitated in the early part of downstroke during bicycling, whereas the H reflexes were depressed in the late part of downstroke. The intensity of TMS was decreased below MEP threshold and used to condition the soleus H reflex. At short intervals (conditioning-test intervals of -3 to -1 ms), TMS produced a facilitation of the H reflex that is in all likelihood caused by activation of the fast monosynaptic corticospinal pathway. This facilitation was significantly larger in the early part of downstroke during bicycling than during tonic plantarflexion. This suggests that the increased MEP during downstroke was caused by changes in transmission in the fast monosynaptic corticospinal pathway. To investigate whether the depression of H reflexes in the late part of downstroke was caused by increased presynaptic inhibition of Ia afferents, the soleus H reflex was conditioned by stimulation of the femoral nerve. At a short interval (conditioning-test interval: -7 to -5 ms), the femoral nerve stimulation produced a facilitation of the H reflex that is mediated by the heteronymous monosynaptic Ia pathway from the femoral nerve to soleus motoneurons. Within the initial 0.5 ms after its onset, the size of this facilitation depends on the level of presynaptic inhibition of the Ia afferents, which mediate the facilitation. The size of the facilitation was strongly depressed in the late part of downstroke, compared with the early part of downstroke, suggesting that increased presynaptic inhibition was indeed responsible for the depression of the H reflex. These findings suggest that there is a selectively increased transmission in the fast monosynaptic corticospinal pathway to soleus motoneurons in early downstroke during bicycling. It would seem likely that one cause of this is increased excitability of the involved cortical neurons. The increased presynaptic inhibition of Ia afferents in late downstroke may be of importance for depression of stretch reflex activity before and during upstroke.     

Recurrent and reciprocal inhibition of the human monosynaptic reflex shows opposite changes following intravenous administration of acetylcarnitine

A long-latency, long-lasting increase in the recurrent inhibitory effect on the soleus monosynaptic (Hoffmann, H) reflex was induced after intravenous administration of L-acetylcarnitine, a substance known to process central cholinergic activity. This effect was paralleled by disappearance of the H reflex inhibition (functionally disinhibition) induced by stimulation of Ia afferent fibres from the tibialis anterior (reciprocal inhibition) and gastrocnemius medialis muscle. Magnitude and time course of the L-acetylcarnitine-induced effects were significantly correlated. The data suggest that (1) the L-acetylcarnitine depression of the reciprocal inhibition is mediated by excitation of Renshaw cells impinging on Ia interneurones (INs), and (2) the inhibitory effect of GM Ia afferents onto Sol is mediated by INs subjected to Renshaw inhibition. The results point to the similarity in the wiring of the 'output stage' circuit between cat and humans, and provide a method for testing this network 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.     

Micturition-related neuronal firing in the periaqueductal gray area in cats

The midbrain periaqueductal gray (PAG) is the area promoting emotional motor responses, reproductive behaviors and analgesia. Recent studies suggest that neurons in the PAG may be crucial for regulating the micturition reflex in both experimental animals and humans. We examined single neuronal activities in the PAG and the adjacent area in response to isovolumetric spontaneous micturition reflexes in 20 supracollicular decerebrated cats. In total, 84 neurons were recorded in the PAG that were related to urinary storage/micturition cycles. Of the neurons recorded, the most common were tonic storage neurons (43%), followed by tonic micturition neurons (29%), phasic storage neurons (15%) and phasic micturition neurons (13%). In addition to the tonic/phasic as well as storage/micturition classification, the neurons showed diverse discharge patterns: augmenting, constant and decrementing, with the constant discharge pattern being most common. Of the 16 neurons located within the PAG that had similar discharge patterns to those just ventral to the PAG, the micturition neurons were distributed in a broader area, whereas the storage neurons seemed to be concentrated in the middle part of the PAG (P0-1, Horsley-Clarke coordinate). High-frequency stimulation (HFS; 0.2-ms duration, 100 Hz) applied in the PAG elicited inhibition of the micturition reflex. Effective amplitude of the electrical stimulation for evoking inhibitory responses was less than 50 microA. In conclusion, the results of the present study showed that HFS of the PAG inhibited the micturition reflex and there were micturition-related neuronal firings in the PAG in cats, suggesting that the PAG is involved in neural control of micturition.     

Convergence on Interneurones Mediating the Reciprocal Ia Inhibition of Motoneurones

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 afferents 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 respectively. 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 la 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.     

Influence of neck afferents on vestibulospinal neurons

Summary The effects of neck afferent stimulation on vestibulospinal neurons in Deiters' nucleus and in the descending nucleus, and the interaction of cervical and vestibular input, were examined extracellularly in decerebrate, decerebellate cats. Many of the vestibulospinal neurons were identified as having axons in the lateral or medial vestibulospinal tract (LVST or MVST) and as being driven antidromically from C₃ or C₆.Half of the spontaneously active neurons were excited with a latency of 2.5–5.5 ms (early excitation) by stimulation of the contralateral C₂ ganglion. In some neurons early excitation was followed by late excitation (latency > 6 ms), which was in other neurons the only effect seen. Early excitation was due to stimulation of proximal afferents because stimulation of the C₂ dorsal or ventral rami usually produced late excitation only. Early excitation was seen in LVST and MVST neurons terminating between C₃ and C₆ and in those projecting beyond C₆. Neurons with early excitation were scattered throughout Deiters' nucleus and the rostral part of the descending nucleus.In some neurons, mainly in the descending nucleus, the initial effect of contralateral C₂ ganglion stimulation was inhibition. Inhibition could be evoked by stimulation of the ganglion or dorsal rami bilaterally. The axons of all tested inhibited neurons were in the MVST.Thirty-five percent of the population studied received convergence of early excitation and short-latency input from the labyrinth, sometimes from the semicircular canals. There was also convergence between late excitation or inhibition and vestibular input.The influence of neck afferent input on vestibulospinal neurons provides one pathway for this input to the neck and limb segments of the spinal cord. This pathway may be part of the substrate of the tonic neck reflex. In addition, vestibulospinal neurons are one site of interaction between neck and vestibular reflexes.Supported by N.I.H. grant NS 02619     

The frequency transfer characteristics of cat soleus motoneurone during suppression of excitatory synaptic activity by “direct” and Ib-type inhibition

Summary The frequency transfer characteristics of the tonic stretch reflex have been studied using homonymous Ia afferent stimulation as input to the soleus motoneurone. By varying additionally the frequency of inhibitory volleys from either the Ia component of antagonist muscle nerve or the homonymous Ib afferents, the suppressive effect on the excitatory synaptic activity has been investigated for a wide variety of different phase relations.     

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.     

Sensitivity of monosynaptic test reflexes to facilitation and inhibition as a function of the test reflex size: a study in man and the cat

Summary In parallel experiments on humans and in the cat it was investigated how the sensitivity of monosynaptic test reflexes to facilitation and inhibition varies as a function of the size of the control test reflex itself. In man the monosynaptic reflex (the Hoffmann reflex) was evoked in either the soleus muscle (by stimulation of the tibial nerve) or the quadriceps muscle (by stimulation of the femoral nerve). In the decerebrate cat monosynaptic reflexes were recorded from the nerves to soleus and medial gastrocnemius muscles; they were evoked by stimulation of the proximal ends of the sectioned L7 and S1 dorsal roots. Various excitatory and inhibitory spinal reflex pathways were used for conditioning the test reflexes (e.g. monosynaptic Ia excitation, disynaptic reciprocal inhibition, cutaneous inhibition, recurrent inhibition, presynaptic inhibition of the Ia fibres mediating the test reflex). It was shown that the additional number of motoneurones recruited in a monosynaptic test reflex by a constant excitatory conditioning stimulus was very much dependent on the size of the test reflex itself. This dependency had the same characteristic pattern whatever the conditioning stimulus. With increasing size of the test reflex the number of additionally recruited motoneurones first increased, then reached a peak (or plateau) and finally decreased. A similar relation was also seen with inhibitory conditioning stimuli. The basic physiological factors responsible for these findings are discussed. Finally, the implications for the interpretation of experiments in man with the H-reflex technique are considered.     

Excitability of reciprocal and recurrent inhibitory pathways after voluntary muscle relaxation in man

Summary We studied the potential contribution of postsynaptic mechanisms to the depression of reflex excitability which occurs immediately after a voluntary release from tonic muscle contraction. The excitability of the Soleus (Sol) motor pool was tested at rest and after voluntary muscle relaxation. In both cases the Sol H-reflex was conditioned by 1. a single shock to the peroneal nerve, in order to activate the Ia interneurones (INs) mediating the reciprocal inhibition via a peripheral input, or by 2. a short-lasting voluntary contraction of the Tibialis Anterior (TA) muscle, to activate the Ia INs via a central command. Changes in excitability of Renshaw cells were also tested at rest and after release, to assess the role of recurrent inhibition in the release-induced inhibition of the Sol H-reflex. It was demonstrated that: 1. the excitability of the INs mediating the reciprocal inhibition was only slightly enhanced in comparison with resting conditions; 2. the H-reflex of the antagonist muscle (TA) evoked after Sol release was not consistently facilitated with respect to rest; 3. the command to contract the TA muscle reduced the H-reflex of the Sol muscle during rest but not after Sol release; 4. recurrent inhibition did not increase its effect in the post-release period. Such features suggest that recurrent and reciprocal post-synaptic inhibitions do not play a major role in reducing the reflex excitability of a relaxing muscle; rather, the command to release prevents the reciprocal inhibitory effect which accompanies the contraction of the antagonist muscle. The findings support the concept that release-induced reflex depression is mediated mainly by presynaptic inhibition of autogenetic spindle afferences (Schieppati and Crenna 1984).Supported by Italian M.P.I.     

Recurrent inhibition from motor axon collaterals of transmission in the Ia inhibitory pathway to motoneurones

1. The effects of impulses in recurrent motor axon collaterals on reflex transmission from different types of primary afferents to motoneurones were investigated in the cat by conditioning of PSPs evoked in motoneurones.

2. IPSPs evoked by volleys in large muscle spindle (Ia) afferents were effectively decreased when preceded by an antidromic stimulation of ventral roots. Some IPSPs from group II muscle afferents and low threshold cutaneous afferents were also slightly depressed, while other PSPs were unaffected.

3. The depression of the IPSPs could be evoked by antidromic volleys, which produced neither conductance changes in the motoneurones nor depolarization of Ia afferent terminals.

4. The effect on the Ia IPSPs is most likely due to post-synaptic inhibition of the Ia inhibitory interneurones, evoked through α-motor axon collaterals and Renshaw cells. The depression of some IPSPs from flexor reflex afferents is explained by a convergence of excitatory effects from these afferents on the Ia inhibitory interneurones.

5. The results indicate a selective recurrent control from motor axon collaterals of the interneurones in the reciprocal Ia inhibitory pathway to motoneurones.

     

Changes in the inhibitory control exerted by the antagonist Ia afferents on human wrist extensor motor units during an attention-demanding motor task

The aim of this study was to determine the extent to which an attention-demanding visuomotor task affects the strength of the inhibitory control exerted by the wrist flexor group Ia afferents on the wrist extensor motoneurons. Effects of median nerve stimulation on the tonic activity of wrist extensor single motor units were analyzed in terms of the interspike interval (ISI) lengthening. Results show that the inhibitory effects exerted by the antagonistic group Ia afferents were significantly enhanced when the wrist extensor motoneurons were involved in an attention-demanding task. Enhanced inhibition from antagonist afferents may reflect task-related changes in the excitability of the di- and/or polysynaptic pathways mediating reciprocal inhibition due to either the action of descending inputs and/or an increase in the efficiency of the Ia inputs to the premotoneuronal inhibitory interneurons. Modulation of the inhibition exerted by proprioceptive antagonist afferents may be one of the processes which contribute to the fine adjustment of the wrist muscle force output required in fine handling tasks.     

Characteristics of the output from the dentate nucleus to spinal neurons via pathways which do not involve the primary sensorimotor cortex

Summary Experiments were performed to determine the action of the dentate output on neurons in the spinal cord mediated by pathways which do not involve the primary sensorimotor and premotor cortices. The dentate nucleus was electrically stimulated by stereotaxically placed electrodes in Rhesus monkeys whose contralateral sensorimotor and premotor cortices were ablated. The resultant changes in excitability of lumbar alpha motoneurons activated by Ia afferents from nerves innervating femoral, hamstring, gastrocnemius-soleus and peroneal muscles were measured by intracellular recordings and by determining the percent change in the amplitude of the monosynaptic reflex recorded from ventral roots. The effect of stimulation of the dentate nucleus on proprioceptive reflexes was determined by recording the changes in postsynaptic potentials evoked by selective stimulation of Ia and Ib afferent fibers. The results demonstrated that the dentate nucleus exerts a significant action on the excitability of spinal neurons via pathways which do not include the sensorimotor and premotor cortices. Whether the dentate stimulus produced an increase or decrease in the excitability of these neurons was dependent upon the site within the dentate nucleus at which the stimulus was applied, demonstrating that, in the decorticate preparation, the output from this nucleus is quite heterogeneous. In addition, stimulation of the dentate nucleus in these monkeys did not affect the Ia reflex pathway but significantly changed the amplitude of the inhibitory postsynaptic potential evoked by Ib afferents in lumbar alpha motoneurons.     

Pelvic afferent reflex control of rectal motility and lumbar colonic efferent discharge mediated by the pontine sympatho-inhibitory region in guinea pigs

Rectal motility and the efferent discharge of lumbar colonic nerves (LCED) have previously been shown to be affected by reflex activity activated by rectal stimulation. The sensory limb of this reflex is represented by afferent fibers in pelvic nerves. The present study revealed that this reflex is modulated by supraspinal sympatho-inhibitory regions. Pelvic afferent stimulation led to rectal contraction through the withdrawal of a tonic inhibitory influence of lumbar colonic nerves. The supraspinal region responsible for this antagonism ofthe rectal-inhibitory colonic nerve activity was localized to the pons. Neither the intravenous administration of atropine nor that of guanethidine (and Eisai compound 865–123, another adrenergic neuron blocking agent) effected the ability of pelvic afferent stimulation to inhibit tonic discharge of lumbar colonic efferent nerves; nervertheless, both agents eliminated the mechanical response of the rectum to stimulation of pelvic afferents. These observations suggest that lumbar sympathetic nerves may tonically inhibit the release of acetylcholine from excitatory neurons in the rectal myenteric plexus. We conclude that descending fibers from the pons are activated as a result of pelvic afferent nerve stimulation. These descending pontine fibers in turn inhibit the firing of sympathetic lumbar colonic nerves. Removal of this tonic restraint leads to rectal contraction.     

Modulation of H reflex of pretibial muscles and reciprocal Ia inhibition of soleus muscle during voluntary teeth clenching in humans

A previous study has demonstrated that the soleus H reflex is facilitated in association with voluntary teeth clenching in proportion with biting force in humans. The present study tried to elucidate the functional significance of this facilitation of the soleus H reflex, by examining 1) whether the facilitation of the H reflex is reciprocal or nonreciprocal between the ankle extensors and flexors and 2) whether the reciprocal Ia inhibition of crural muscles is facilitated or depressed in association with voluntary teeth clenching. The H reflex of the pretibial muscles was evoked by stimulation of the common peroneal nerve in seven healthy subjects with no oral dysfunction. The pretibial H reflex was facilitated in association with voluntary teeth clenching in a force-dependent manner. The facilitation started preceding the onset of electromyographic activity of the masseter muscle. Stimulation of the common peroneal nerve at low intensities subthreshold for evoking the M wave of the pretibial muscles inhibited the soleus H reflex after a short latency corresponding with a disynaptic inhibition, indicating that the reciprocal Ia inhibition was depressed in association with voluntary teeth clenching. Thus, the present study has shown that voluntary teeth clenching evokes a nonreciprocal facilitation of ankle extensor and flexor muscles and attenuated reciprocal Ia inhibition from the pretibial muscles to the soleus muscle. It is concluded that voluntary teeth clenching contributes to improve stability of stance rather than smoothness of movements.     

Tonic neck reflex of the decerebrate cat: a role for propriospinal neurons

In decerebrate, acutely labyrinthectomized cats we used neck rotation to study the role of direct upper cervical afferents to the cervical enlargement and of cervical and lumbar propriospinal neurons in the tonic neck reflex. Interruption of the dorsal columns between C4 and C5 had no qualitative effect on the dynamics of the reflex although gain usually increased. Direct upper cervical afferents to the cervical enlargement therefore have no unique role in producing the reflex. Many medially located propriospinal neurons in C4 were modulated by neck rotation. About 40% had axons, mostly crossed, that terminated in the cervical enlargement. The others projected more caudally, some as far as L3-L4 or even the lumbar enlargement. For a population of C4 neurons, including propriospinal neurons, we measured the response vector with combinations of roll and pitch stimuli. These vectors ranged from pitch to roll. Many propriospinal neurons in L3-L4, projecting to the lumbosacral enlargement, were also modulated by neck rotation with a variety of response vectors. Some of these neurons had an ascending projection. As in previous experiments, C4 neurons were modulated by neck rotation after spinal transection rostral to the C1 dorsal root entry zone; a wide variety of response vectors was observed. In contrast, almost no modulated L3-L4 neurons were found in the same experiments. The results suggest a role for propriospinal neurons in the tonic neck reflex. They also demonstrate that responses of lumbar neurons to neck rotation are much more dependent on supraspinal pathways than are those of cervical neurons.     

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 10-60microA and 1-10ms intervals) delivered to the nucleus reticularis gigantocellularis evoked inhibitory postsynaptic potentials in alpha-motoneurons (n=147) and gamma-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 4.0-5.5ms with a segmental delay of more than 1.4ms. The late inhibitory postsynaptic potentials were observed in 98.0% of the motoneurons and had a latency of 30-35ms, with a peak latency of 50-60ms. 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 (<20ms) and late (30-80ms) 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 alpha- and gamma-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.     

Recurrent inhibition of individual Ia inhibitory interneurones and disinhibition of their target α-motoneurones during muscle stretches

The effects of ramp stretches applied to triceps surae muscle on the discharge patterns of single Ia inhibitory interneurones, monosynaptically invaded from various nerves, were studied in either decerebrate or anesthetized cats. Interneurones which received direct excitatory Ia input from the stretched muscle exhibited augmented activity both during the dynamic and static phase of stretch, which was, however, interrupted by a transient inhibitory influence during the dynamic phase of stretch. The influences on Ia inhibitory interneurones, monosynaptically invaded from hamstring or tibial nerve, were exclusively inhibitory. These stretch-induced inhibitions were better demonstrable in decerebrate than in anesthetized preparations. The timing of the discharge patterns of additionally recorded Renshaw cells during stretch, and the disappearance or reduction of the above described inhibitory effects after administration of DHE, strongly support the idea that these inhibitory actions are caused by Renshaw inhibition. In Ia inhibitory interneurones, monosynaptically activated from the antagonistic peroneal nerve, stretch induced also pronounced inhibitory effects, which were most probably caused by mutual inhibition between Ia inhibitory interneurones. The suppression of agonistic Ia inhibitory interneurone activity below the tonic resting activity corresponded to an enhancement of the monosynaptic reflex amplitude of the antagonistic motoneurone pool. The findings suggest that normal orthodromic activation of Renshaw cells, and consequently the recurrent inhibition of the Ia inhibitory interneurones, is predominantly linked with rapid phasic, rather than slow tonic, motoneuronal firing. The functional role of this mechanism for the performance of rapidly alternating movements and the damping of ballistic agonist contractions is discussed.