Effects of cortical stimulation on reciprocal inhibition in humans

We attempted to demonstrate convergence onto human spinal Ia inhibitory interneurons from Ia afferents and from fast conducting corticospinal axons. Stimulation of the common peroneal nerve at or below the threshold of the alpha motoneuron axons resulted in inhibition of the soleus H-reflex, attributed to reciprocal inhibition. Magnetic stimulation over the contralateral motor cortex resulted in complex modulations of the soleus H-reflex, including a short latency-inhibition. To test for convergence, the two stimuli were given together so that the two inhibitions coincided.When each stimulus alone produced clear inhibition, the inhibition produced by both stimuli was less than expected, implying an interaction between the two volleys, for example, occlusion occurring in interneurons or motoneurons.When the H-reflex was relatively unaffected by one or other conditioning volley, the inhibition produced by the combined stimulation was greater than expected, as might be expected with convergence onto a common pool of interneurons.     

Modulation of presynaptic inhibition of Ia afferents during voluntary wrist flexion and extension in man

Changes in presynaptic inhibition of Ia terminals directed to flexor carpi radialis (FCR) motoneurones (MNs) were investigated in normal human subjects at rest and during voluntary wrist flexion and extension. To that end, two independent methods were used: (1) the radial-induced D1 inhibition of the FCR H reflex, which assesses the excitability of PAD (primary afferent depolarisation) interneurones controlling presynaptic inhibition of Ia terminals mediating the afferent volley of the FCR H reflex; and (2) the heteronymous monosynaptic Ia facilitation induced in the FCR H reflex by intrinsic muscle Ia afferent stimulation, which assesses the ongoing presynaptic inhibition of Ia terminals. With respect to results at rest, it was found that at the onset of (and during tonic) voluntary wrist flexion, D1 inhibition was reduced and heteronymous monosynaptic Ia facilitation was increased. This suggests that, as in the lower limb, presynaptic inhibition is decreased on Ia terminals projecting to MNs involved in the voluntary contraction. In contrast with results observed in the lower limb, presynaptic inhibition of Ia terminals to FCR MNs was also found to be reduced at the onset of a voluntary contraction involving the antagonistic wrist extensors, suggesting that presynaptic inhibition of Ia terminals projecting to wrist flexors and extensors might be mediated through the same subsets of PAD interneurones. This is in keeping with other features showing that the organisation of reflex pathways between wrist flexors and extensors differs from that observed at other (elbow, ankle) joints.     

Modulation, probably presynaptic in origin, of monosynaptic Ia excitation during human gait

Modulation of presynaptic inhibition of Ia afferents projecting monosynaptically to soleus motoneurones was investigated during human gait. Changes in presynaptic inhibition of Ia afferents were deduced from alterations in the amount of heteronymous soleus H-reflex facilitation evoked by a constant femoral nerve stimulation. It has been shown that this facilitation is mediated through a monosynaptic Ia pathway and that during its first 0.5 ms it is still uncontaminated by any polysynaptic effect and can be used to assess ongoing presynaptic inhibition of Ia terminals to soleus motoneurones. During gait, heteronymous facilitation was reduced with respect to its control value (rest during sitting) and modulated during the step cycle: it reached its maximum at mid-stance and decreased to near zero by the end of stance. At the same time the H-reflex amplitude was to some extent similarly modulated. It is argued that this decrease in heteronymous Ia facilitation and in H-reflex amplitude reflects an increased, ongoing presynaptic inhibition of Ia terminals projecting onto soleus motoneurones, which could be from central and/or peripheral origin. D1 inhibition, i.e. the late and long-lasting inhibition of the soleus H-reflex evoked by a train of stimuli to the common peroneal nerve, was used as another method to assess presynaptic inhibition. This D1 inhibition was decreased during gait, and it is argued that this decrease might reflect an occlusion in presynaptic pathways or increased presynaptic inhibition of pathways mediating the conditioning volley.     

Ia afferents of the antagonist are inhibited presynaptically before the onset of a ballistic muscle contraction in man

Summary The aim of the present investigation was to study whether the pre-movement inhibition of the H reflex in the antagonist of a ballistic voluntary contraction was due to a reduced activity of the motoneuronal pool of the antagonist, or to a reduced excitatory effect of the afferent volley reaching these motoneurons. Human subjects performed visually conditioned ballistic dorsal flexions of the ankle. The inhibition of the H reflex in the antagonist (soleus muscle) was similar if the muscle was initially relaxed or if there was a preexisting level of motor discharge. Since the soleus muscle was inhibited before movement onset in parallel with the H reflex inhibition, the relation between the level of a background activity and the size of superimposed H reflexes was studied. The finding that H reflexes were only slightly reduced in size with decreasing steady EMG levels could not explain the pre-movement inhibition, and it was concluded than an increased presynaptic inhibition of Ia terminals was the source of the H reflex inhibition.     

Changes in presynaptic inhibition of Ia fibres to soleus motoneurones during voluntary dorsiflexion of the foot

Summary Variations of presynaptic inhibition in heteronymous Ia fibres projecting to soleus motoneurones were studied during the first 250 ms of phasic voluntary isometric contractions of the antagonist tibialis anterior muscle in human subjects. During the first 60–80 ms of TA e.m.g activity, presynaptic inhibition was often more marked than at rest, but not in all experimental sessions. After 60–80 ms, presynaptic inhibition was always increased compared to rest and to the onset of TA e.m.g. activity. A rebound in femoral nerve induced Ia facilitation was often observed between 90–150 ms. The early increase in presynaptic inhibition was widespread and non specific since it was observed at the onset of extensor carpi radialis contractions of maximal strength. The rebound in heteronymous Ia facilitation was interpreted as a relative decrease in presynaptic inhibition to which nonspecific suprasegmental and cutaneous effects contributed. The late increase in presynaptic inhibition in Ia fibers to soleus motoneurones was considered as reciprocally inhibiting the Sol H-reflex, thus counteracting the phasic stretch of the antagonist muscle during TA contraction.     

Evidence for presynaptic inhibition of muscle spindle Ia afferents in man

A new method for estimating presynaptic inhibition of soleus (Sol) Ia fibres in man is introduced based on the assessment of the amount of facilitation produced by a preceding homonymous Ia volley onto a Sol monosynaptic reflex. It was found in animal experiments that a reduction of such Ia facilitation reflects the amount of presynaptic inhibition exerted on these Ia fibres. Since, in man, a similar reduction of homonymous Ia facilitation was found after tibialis anterior vibration, this provides evidence for presynaptic inhibition of Sol Ia fibres, elicited here by the afferent discharge due to this flexor vibration.     

Selective cortical and segmental control of primary afferent depolarization of single muscle afferents in the cat spinal cord

 This study was primarily aimed at investigating the selectivity of the cortico-spinal actions exerted on the pathways mediating primary afferent depolarization (PAD) of muscle spindle and tendon organ afferents ending within the intermediate nucleus at the L6–L7 segmental level. To this end we analyzed, in the anesthetized cat, the effects produced by electrical stimulation of sensory nerves and of the cerebral cortex on (a) the intraspinal threshold of pairs of single group I afferent fibers belonging to the same or to different hindlimb muscles and (b) the intraspinal threshold of two collaterals of the same muscle afferent fiber. Afferent fibers were classified in three categories, according to the effects produced by stimulation of segmental nerves and of the cerebral cortex. Twenty-five of 40 fibers (62.5%) were depolarized by stimulation of group I posterior biceps and semitendinosus (PBSt) or tibialis (Tib) fibers, but not by stimulation of the cerebral cortex or of cutaneous and joint nerves, which instead inhibited the PBSt- or Tib-induced PAD (type A PAD pattern, usually seen in Ia fibers). The remaining 15 fibers (37.5%) were all depolarized by stimulation of the PBSt or Tib nerves and the cerebral cortex. Stimulation of cutaneous and joint nerves produced PAD in 10 of those 15 fibers (type B PAD pattern) and inhibited the PBSt- or Tib-induced PAD in the 5 remaining fibers (type C PAD pattern). Fibers with a type B or C PAD pattern are likely to be Ib. Not all sites in the cerebral cortex inhibited with the same effectiveness the segmentally induced PAD of group I fibers with a type A PAD pattern. With the weakest stimulation of the cortical surface, the most effective sites that inhibited the PAD of individual fibers were surrounded by less effective sites, scattered all along the motor cortex (area 4γ and 6) and sensory cortex (areas 3, 2 and 1), far beyond the area of projection of group I fibers from the hindlimb. With higher strengths of cortical stimulation, the magnitude of the inhibition was also increased, and previously ineffective or weakly effective sites became more effective. Maps obtained when using the weakest cortical stimuli have indicated that the most effective regions that produced PAD of group I fibers with a type B or type C PAD pattern were also scattered throughout the sensory-motor cortex, in the same general area as those that inhibited the PAD of group I afferents with a type A PAD pattern. In eight fibers with a type A PAD pattern it was possible to examine the intraspinal threshold of two collaterals of the same single afferent fiber ending within the intermediate nucleus at the L7 segmental level. In six fibers, stimulation of the PBSt nerve with trains of pulses between 1.5 and 1.86 times threshold (×T) produced a larger PAD in one collateral than in the other. In seven fibers, stimulation of the sensory-motor cortex and of cutaneous nerves produced a larger inhibition of the PBSt-induced PAD in one collateral than in the other. The ratio of the cortically induced inhibition of the PAD elicited in the two collaterals could be modified by changing the strength of cortical and of PBSt stimulation. In three fibers it was possible to inhibit almost completely the background PAD elicited in one collateral while having little or no effect on the PAD in the other collateral. Changes in the intraspinal threshold of pairs of collaterals following electrical stimulation of segmental nerves and of the somato-sensory cortex were examined in three fibers with a type B and two fibers with a type C PAD pattern. In four fibers the PAD elicited by stimulation of cutaneous (4–20×T) and muscle nerves (1.54–3.7×T), or by stimulation of the sensory-motor cortex, was of different magnitude in the two collaterals. In two experiments it was possible to find cortical sites in which weak surface stimulation produced PAD in one collateral only. The magnitude of the PAD elicited in pairs of collaterals of group I afferents with a type B or C PAD pattern, or the inhibition of the PAD in pairs of collaterals of fibers with a type A PAD pattern, appeared not to be topographically related to the site of spinal projection of the cutaneous and cortico-spinal fibers used for conditioning stimulation. The present demonstration of a differential control of the PAD exerted on two collaterals of the same afferent fiber suggests that the profuse intraspinal branching of muscle spindle and tendon organs is a potentially rich substrate for information transmission. By means of presynaptic control mechanisms, the terminal arborizations of the afferent fibers could function either as a simple unit or in a fractionated manner, allowing funneling of information to selected groups of central neurons. Received: 18 April 1996 / Accepted: 5 September 1996     

Distribution of heteronymous Ia facilitation and recurrent inhibition in the human deltoid motor nucleus

Summary Distribution of heteronymous Ia facilitation and of heteronymous recurrent inhibition in motoneurones innervating the anterior part of the deltoid muscle were investigated in normal human subjects following electrical stimulation of the nerves innervating the main muscles of the upper limb. Activation of group I afferents originating from deltoid, biceps, triceps and extensor carpi radialis (ECR) muscles resulted in an early increase in firing probability of voluntarily activated motor units belonging to the anterior part of the deltoid muscle whereas activation of motor axons supplying deltoid, triceps, ECR and flexor carpi radialis (FCR) muscles resulted in an early and long-lasting decrease in firing probability. No effect was seen following activation of group I afferents and motor axons contained in the ulnar nerve. The characteristics of the early facilitation suggest that it is at least partly due to heteronymous Ia monosynaptic connections while these of the long-lasting inhibition suggest that it is at least partly due to heteronymous recurrent inhibition. Their patterns of distribution are discussed with regards to the functional role of the human deltoid.     

Patterns of connectivity of spinal interneurons with single muscle afferents

 A technique was developed to measure, in the anesthetized and paralyzed cat under artificial ventilation, changes of excitability to intraspinal stimulation simultaneously in two different afferent fibers or in two collaterals of the same afferent fiber. Intraspinal stimulation reduced the threshold of single muscle afferent fibers ending in the intermediate nucleus. This effect was seen with strengths below those required to activate the afferent fiber tested (1.5–12 μA), occurred at a short latency (1.5–2.0 ms), reached a maximum between 15 and 30 ms, and lasted up to 100 ms. The effects produced by graded stimulation applied at the shortest conditioning-testing stimulus time intervals increased by fixed steps, suggesting recruitment of discrete elements, most likely of last-order interneurons mediating primary afferent depolarization (PAD). The short-latency increases in excitability produced by the weakest effective intraspinal stimuli were usually detected only in the collateral closest to the stimulating micropipette, indicating that the stimulated interneurons mediating PAD have spatially restricted actions. The short-latency PAD produced by intraspinal stimuli, as well as the PAD produced by stimulation of the posterior biceps and semitendinosus (PBSt) nerve or by stimulation of the bulbar reticular formation (RF), was depressed 19–30 min after the i.v. injection of 0.5 mg/kg of picrotoxin, suggesting that all these effects were mediated by GABAergic mechanisms. The PAD elicited by stimulation of muscle and/or cutaneous nerves was depressed following the i.v. injection of (–)-baclofen, whereas the PAD elicited in the same collateral by stimulation of the RF was baclofen-resistant. The short-latency PAD produced by intraspinal stimulation was not always depressed by i.v. injections of (–)-baclofen. Baclofen-sensitive and baclofen-resistant monosynaptic PADs could be produced in different collaterals of the same afferent fiber. The results suggest that the intraspinal terminals of single muscle afferents receive synapses from more than one PAD-mediating GABAergic interneuron and that a single last-order interneuron has synaptic connections with a restricted number of intraspinal terminals and/or collaterals of the same afferent fiber. In addition, they support the existence of separate subsets of last-order baclofen-sensitive and baclofen-resistant interneurons that respond predominantly to segmental and to descending inputs. It is suggested that the restricted nature of the PAD plays an important role in the central control of the synaptic effectiveness of group I muscle afferents. Received: 10 October 1996 / Accepted: 10 December 1996     

Synaptic connections from large afferents of wrist flexor and extensor muscles to synergistic motoneurones in man

 Short-latency excitatory Ia reflex connections were determined between pairs of human wrist flexor and extensor muscles. Spindle Ia afferents were stimulated by either tendon tap or electrical stimulation. The activity of voluntarily activated single motor units was recorded intramuscularly from pairs of wrist flexor or extensor muscles. Cross-correlation between stimuli and the discharge of the motor units provided a measure of the homonymous or heteronymous excitatory input to a motoneurone. Homonymous motoneurone facilitation was generally stronger than that of the heteronymous motoneurones. The principal wrist flexors, flexor carpi radialis (FCR) and flexor carpi ulnaris (FCU), were tightly connected through a bidirectional short-latency reflex pathway. In contrast, the extensor carpi ulnaris (ECU) and the extensor carpi radialis (ECR) did not have similar connections. ECU motoneurones received no short-latency excitatory Ia input from the ECR. ECR motoneurones did receive excitatory Ia input from ECU Ia afferents; however, its latency was delayed by several milliseconds compared with other heteronymous Ia excitatory effects observed. The wrist and finger extensors were linked through heteronymous Ia excitatory reflexes. The reflex connections observed in humans are largely similar to those observed in the cat, with the exception of heteronymous effects from the ECU to the ECR and from the extensor digitorum communis (EDC) to the ECU, which are present only in humans. The differences in the reflex organization of the wrist flexors versus the extensors probably reflects the importance of grasping. Received: 19 August 1996 / Accepted: 6 March 1997     

Evidence for mutual inhibition of opposite Ia interneurones in the human upper limb

Summary The H-reflex technique was used to collect indirect evidence for changes in excitability of the interneurones mediating reciprocal Ia inhibition between wrist extensors and flexors. Stimulating the radial nerve results in an inhibition of the flexor carpi radialis (FCR) H-reflex and evidence has previously been presented that the early phase of inhibition is mediated by extensor-coupled Ia interneurones (Ext Ia INs), i.e. by inhibitory interneurones fed by muscle spindle Ia afferents from wrist extensors. Variations in the level of this inhibition were used to assess changes in excitability of Ext Ia INs. Stimulation of group I fibres from flexors was shown to depress the reference Ia inhibition, i.e. to inhibit the Ext Ia INs. The central latency of this interneuronal inhibition was compatible with a disynaptic linkage between flexor Ia afferents and Ext Ia INs. Its threshold and time course profile could almost exactly be superimposed on those of reciprocal Ia inhibition from flexors to extensor carpi radialis (ECR) motoneurones (MNs). This suggests that the Ia inhibitions to extensor MNs and extensor Ia INs are collateral effects mediated by the same flexor-coupled Ia interneurones. In two subjects, in whom it was possible to elicit an H-reflex in the ECR, inhibition of flexor-coupled Ia interneurones by activation of extensor Ia interneurones could similarly be demonstrated.     

Effects of motor cortex stimulation on spinal interneurones in intact man

Summary The effect of a descending corticospinal volley on a spinal inhibitory pathway, has been studied in five intact human subjects. Approximately 63% inhibition of the H-reflex evoked in wrist and finger flexor muscles, was produced by motor threshold stimulation of the radial nerve. When a submotor threshold cortical shock was given 2 to 4 ms before the H-reflex, this inhibition was reduced to approximately 38%. The timing of this effect is compatible with either a monosynaptic or disynaptic corticospinal tract projection onto the spinal inhibitory interneurone.     

Changes in polysynaptic Ia excitation to quadriceps motoneurones during voluntary contraction in man

Summary Homonymous Ia facilitation of quadriceps (Q) motoneurones (MNs) was significantly larger during (and at the onset of) a very weak Q voluntary contraction than at rest. This increase in Ia facilitation only appeared with a conditioning-test interval within the narrow range of 5–9 ms, which fits the time course of the recently described polysynaptic Ia excitation to Q MNs. This suggests that interneurones mediating polysynaptic Ia excitatory effects to Q MNs receive a strong descending excitation during such a contraction. It is therefore argued that these interneurones might mediate part of the descending command to Q MNs during voluntary contraction.     

Reflex pathways from large muscle spindle afferents and recurrent axon collaterals to motoneurones of wrist and digit muscles: a comparison in cats, monkeys and humans

Whereas a large body of data is available on the control of hand motoneurones from the brain, not much is known about the contribution of the spinal interneuronal apparatus to the differentiated movement repertoire of the hand. This review summarises recent data on the excitatory Ia pathways and on recurrent inhibition for cats, monkeys and humans. The basic principles of organisation have, in general, been preserved in the different species. Thus the motoneurones to cat and human long and short digit muscles seem not to possess a recurrent axon collateral system. With regard to the Ia pathways, specialised connectivity patterns have developed in the long digit extensor and wrist extensor muscles. They allow the former group to support the differentiated movement repertoire of the digits, and the latter group to be part of a general extensor or flexor synergism. Modifications between the species are present, however, with regard to the proximodistal connectivity across the elbow. Whereas they are regularly present in the cat, they are less developed in the monkey and absent in man, which frees the human hand from the elbow position. Received: 28 September 1998 / Accepted: 22 January 1999     

Projections of group Ia afferents to motoneurons of thigh muscles in man

Summary Changes in the firing probability of single motor units in response to electrical stimulation of muscle nerves and to tendon taps were used to derive the projections of large muscle afferents to the motoneurons of various thigh muscles in man. Homonymous facilitation was demonstrated to virtually all of the sampled motor units of biceps (BI), semitendinosus (ST), vastus lateralis (VL) and vastus medialis (VM). Heteronymous facilitation was readily demonstrated between VM and VL but was never obtained from ST to BI and never unequivocally obtained from BI to ST. Reciprocal inhibition was demonstrated from femoral nerve afferents to all of the sampled units of BI, and ST but reciprocal inhibition of VM or VL was never obtained from BI afferents and infrequently from ST afferents. These projections of group I afferents in man show certain specific differences from those of the cat and baboon that may reflect the normal function of the limb.     

Effects of stimulation of group I afferents from flexor muscles on heterosynaptic facilitation of monosynaptic reflexes produced by Ia and descending inputs: a test for presynaptic inhibition

Summary 1. In the chloralose anesthetized cat, conditioning stimulation of group I flexor afferents depresses the monosynaptic potentials generated by Ia afferents in single spinal motoneurons or in populations of motoneurons without affecting the monosynaptic potentials produced by stimulation of descending fibers in the ipsilateral ventromedial fasciculus (VMF). 2. Heterosynaptic facilitation of monosynaptic reflexes was used to test changes in the presynaptic effectiveness of excitatory inputs with direct connections with motoneurons. We found that the heterosynaptic facilitation of Ia origin was reduced by conditioning stimulation of group I afferents from flexors, without affecting the heterosynaptic facilitation produced by stimulation of the VMF. 3. These results confirm and expand previous observations showing that the synaptic effectiveness of descending fibers synapsing with motoneurons is not subjected to a presynaptic control mechanism of the type acting on Ia fiber terminals, and provide further basis for the use of changes in heterosynaptic facilitation of monosynaptic reflexes of Ia origin as an estimate of changes in presynaptic inhibition of Ia fibers (Hultborn et al. 1987a).     

Segmental and supraspinal control of synaptic effectiveness of functionally identified muscle afferents in the cat

The present investigation documents the patterns of primary afferent depolarization (PAD) of single, functionally identified muscle afferents from the medial gastrocnemius nerve in the intact, anesthetized cat. Classification of the impaled muscle afferents as from muscle spindles or from tendon organs was made according to several criteria, which comprised measurement of conduction velocity and electrical threshold of the peripheral axons, and the maximal frequency followed by the afferent fibers during vibration, as well as the changes in discharge frequency during longitudinal stretch, the projection of the afferent fiber to the motor pool, and, in unparalyzed preparations, the changes in afferent activity during a muscle twitch. In confirmation of a previous study, we found that most muscle spindle afferents (46.1–66.6%, depending on the combination of criteria utilized for receptor classification) had a type A PAD pattern. That is, they were depolarized by stimulation of group I fibers of the posterior biceps and semitendinosus (PBSt) nerve, but not by stimulation of cutaneous nerves (sural and superficial peroneus) or the bulbar reticular formation (RF), which in many cases inhibited the PBSt-induced PAD. In addition, we found a significant fraction of muscle spindle primaries that were depolarized by stimulation of group I PBSt fibers and also by stimulation of the bulbar RF. Stimulation of cutaneous nerves produced PAD in 9.1–31.2% of these fibers (type B PAD pattern) and no PAD in 8.2–15.4% (type C PAD pattern). In contrast to muscle spindle afferents, only the 7.7–15.4% of fibers from tendon organs had a type A PAD pattern, 23–46.1% had a type B and 50–61.5% a type C PAD pattern. These observations suggest that the neuronal circuitry involved in the control of the synaptic effectiveness of muscle spindles and tendon organs is subjected to excitatory as well as to inhibitory influences from cutaneous and reticulospinal fibers. As shown in the accompanying paper, the balance between excitation and inhibition is not fixed, but can be changed by crushing the afferent axons in the peripheral nerve and allowing subsequent reconnection of these afferent fibers with muscle receptors.     

Tonic and phasic differential GABAergic inhibition of synaptic actions of joint afferents in the cat

The aim of this study was to examine the functional organization of the spinal neuronal networks activated by myelinated afferent fibers in the posterior articular nerve (PAN) of the anesthetized cat. Particular attention was given to the tonic and phasic GABAa inhibitory modulation of these networks. Changes in the synaptic effectiveness of the joint afferents were inferred from changes in the intraspinal focal potentials produced by electrical stimulation of the PAN. We found that conditioning stimulation of cutaneous nerves (sural, superficial peroneus and saphenous) and of the nucleus raphe magnus often inhibited, in a differential manner, the early and late components of the intraspinal focal potentials produced by stimulation of low and high threshold myelinated PAN afferents, respectively. The degree of the inhibition depended on the strength of both the conditioning and test stimuli and on the segmental level of recording. Conditioning stimulation of group I muscle afferents was less effective, but marked depression of the early and late focal potentials was produced by stimuli exceeding 5 xT. The i.v. injection of 1–2.5 mg/kg of picrotoxin, a GABAa blocker, had relatively minor effects on the early components of the PAN focal potentials, but was able to induce a significant increase of the late components. It also reduced the inhibitory effects of cutaneous and joint nerve conditioning on PAN focal responses. Conditioning autogenetic stimulation with high-frequency trains depressed the PAN focal potentials. The late components of the PAN responses remained depressed several minutes after discontinuing the conditioning train, even after picrotoxin administration. The present observations indicate that the neuronal networks activated by the low threshold PAN afferents show a relatively small post-activation depression and appear to be subjected to a minor tonic inhibitory GABAa control. In contrast, the pathways activated by stimulation of high threshold myelinated afferents have a strong post-activation depression and are subjected to a significant tonic GABAergic modulation. These contrasting features, together with the phasic differential GABAergic inhibition of the responses produced by stimulation of the different populations of joint afferents, may contribute to the preservation of the original information on joint position transmitted by large diameter joint afferents, in contrast with the tonic presynaptic inhibition exerted on the fine myelinated joint afferents, which may be involved in the adjustment of compensatory reactions to inflammation.     

Patterns of primary afferent depolarization of segmental and ascending intraspinal collaterals of single joint afferents in the cat

We have examined in the anesthetized cat the threshold changes produced by sensory and supraspinal stimuli on intraspinal collaterals of single afferents from the posterior articular nerve (PAN). Forty-eight fibers were tested in the L3 segment, in or close to Clarke’s column, and 70 fibers in the L6–L7 segments within the intermediate zone. Of these, 15 pairs of L3 and L6–L7 collaterals were from the same afferent. Antidromically activated fibers had conduction velocities between 23 and 74 m/s and peripheral thresholds between 1.1 and 4.7 times the threshold of the most excitable fibers (xT), most of them below 3 xT. PAN afferents were strongly depolarized by stimulation of muscle afferents and by cutaneous afferents, as well as by stimulation of the bulbar reticular formation and the midline raphe nuclei. Stimulation of muscle nerves (posterior biceps and semitendinosus, quadriceps) produced a larger PAD (primary afferent depolarization) in the L6–L7 than in the L3 terminations. Group II were more effective than group I muscle afferents. As with group I muscle afferents, the PAD elicited in PAN afferents by stimulation of muscle nerves could be inhibited by conditioning stimulation of cutaneous afferents. Stimulation of the cutaneous sural and superficial peroneal nerves increased the threshold of few terminations (i.e., produced primary afferent hyperpolarization, PAH) and reduced the threshold of many others, particularly of those tested in the L6–L7 segments. Yet, there was a substantial number of terminals where these conditioning stimuli had minor or no effects. Autogenetic stimulation of the PAN with trains of pulses increased the intraspinal threshold in 46% and reduced the threshold in 26% of fibers tested in the L6–L7 segments (no tests were made with trains of pulses on fibers ending in L3). These observations indicate that PAN afferents have a rather small autogenetic PAD, particularly if this is compared with the effects of heterogenetic stimulation. Therefore, the depression of the PAN intraspinal fields produced by autogenetic stimulation described by Rudomin et al. (Exp Brain Res DOI 10.1007/s00221-006-0600-x, 2006) may be ascribed to other mechanisms besides a GABAa PAD. It is suggested that the small or no autogenetic PAD displayed by the examined joint afferents prevents presynaptic filtering of their synaptic actions and preserves the original information generated in the periphery. This could be important for proper adjustment of limb position.     

Changes in reciprocal Ia inhibition from wrist extensors to wrist flexors during voluntary movement in man

Summary Transmission in the Ia inhibitory pathway from wrist extensor muscles onto flexor MNs was studied at various times after the onset of voluntary wrist extension or flexion. At the very onset of wrist movements Ia inhibition was not changed, as compared to at rest, whereas later it progressively increased during wrist extension and decreased during wrist flexion. These results are discussed in relation to the different inputs converging onto Ia interneurones and it is suggested that their inhibition by Renshaw cells might be responsible for the results found at the onset of contraction