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     

Pattern of projections of group I afferents from elbow muscles to motoneurones supplying wrist muscles in man

Summary The pattern of projections of low threshold afferents from triceps and biceps brachii muscles onto motoneurones innervating muscles acting at the wrist was assessed by a reflex and a poststimulus time histogram (psth) technique. Activation of low-threshold afferents originating from elbow flexors or extensors resulted in an early, short-lasting inhibition of wrist flexor motoneurones (flexor carpi radialis, flexor carpi ulnaris). An inhibition was also found in the extensor carpi radialis (ECR) motoneurones after stimulation of low-threshold afferents from triceps. Evidence is presented that Ia fibres contribute to these effects. The inhibitory effects were found in all subjects, but they were constant in only 57% of the reflex experimental sessions and in 25% of the explored motor units. Stimulation of biceps low-threshold afferents was always ineffective on ECR motoneurones. No early facilitation was ever seen in motor nuclei innervating wrist muscles following stimulation of low threshold afferents from biceps and triceps. The pattern of transjoint projections of group I afferents from proximal to distal muscles and from distal to proximal ones (Cavallari and Katz 1989) is discussed in relation to that described in the cat forelimb.     

Early and late stretch responses of human foot muscles induced by perturbation of stance

In eight subjects standing on a movable platform, surface EMG activity was recorded from the foot muscles extensor digitorum brevis (EDB) and flexor digitorum brevis (FDB) and from the leg muscles soleus (Sol) and tibialis anterior (TA) during perturbations of upright stance. Perturbations inducing foot dorsiflexion (upward tilt and backward translation) evoked a short-latency response (SLR) and a medium-latency response (MLR) to stretch in the physiological extensors FDB and Sol, and a long-latency response (LLR) in the physiological flexors EDB and TA. Perturbations inducing plantarflexion (downward tilt and forward translation) evoked the MLR in EDB and TA, and the LLR in FDB and Sol. The latency of the FDB and Sol SLR was compared to that of the H and T reflexes evoked in the same muscles by electrical or mechanical stimulation, respectively. In both muscles, the T reflex and the SLR followed the H reflex at delays accounted for by the different stimulation mode, indicating that the SLR induced in both muscles by upward tilt and backward translation was a true autogenetic stretch reflex from spindle primaries. The time interval between the onset of SLR and of MLR was significantly greater for the FDB than the Sol muscle, suggesting that MLR is a spinal reflex travelling through slower peripheral afferent pathways than SLR. From these latency differences and from the distance between the muscles, we calculated in four subjects the conduction velocity of the afferent fibres presumably responsible for the MLR in FDB. This was about 29 m/s. LLRs were evoked in TA and EDB during upward tilt and backward translation, and in Sol and FDB during downward tilt, but not forward translation. LLRs did not adhere to a proximal-to-distal pattern, since these could appear earlier in the foot than in the leg muscles. All responses were modulated by perturbation type (tilt vs translation) and body posture (normal stance vs forward leaning). Both the large amplitude of the foot muscle responses and their temporal pattern indicate that the muscles acting on the toes play a major role in stabilising posture. Their action increases in amplitude and extends in time the foot-ground reaction force, thereby improving the efficiency of the superimposed action of the leg muscle responses.     

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.     

Stretch reflexes in human abdominal muscles

Homonymous and heteronymous reflex connections of the abdominal muscles were investigated by the application of a tap to the muscle belly and observation of surface electromyographic responses. Reflex responses of the following abdominal muscles were investigated both ipsilateral and contralateral to the tap: rectus abdominis (RA), external oblique (EO) and internal oblique (IO). Reflexes were evoked in each of the homonymous muscles with latencies and estimated conduction velocities compatible with being evoked by Ia muscle afferents and having a monosynaptic component. Short latency heteronymous excitatory reflex connections were also observed in muscles on both ipsilateral and contralateral sides in response to the same stimulus. The latencies of the crossed responses were only marginally longer than responses evoked in the respective ipsilateral muscle. Moreover, the reflexes evoked in the IO muscle from ipsilateral and contralateral IO muscle afferents were of comparable amplitude, as were those reflexes evoked in ipsilateral and contralateral EO and RA muscles when tapping IO. These similarities in the reflex characteristics on the ipsilateral and contralateral sides suggest that abdominal muscle afferents activate similar pathways to muscles on both sides of the body. It follows that if the homonymous stretch reflex of abdominal muscles have a monosynaptic component, then a similar monosynaptic pathway activates synergistic motoneurones, not only ipsilaterally but also contralaterally.     

Characterisation of the quadriceps stretch reflex during the transition from swing to stance phase of human walking

The main objective of this study was to characterize the stretch reflex response of the human thigh muscles to an unexpected knee flexion at the transition from stance to swing during walking. Eleven healthy subjects walked on a treadmill at their preferred speed. Reliable and constant knee flexions (6–12° amplitude, 230–350°/s velocity, 220 ms duration) were applied during the late swing and early stance phase of human walking by rotating the knee joint with a specifically designed portable stretch apparatus affixed to the left knee. Responses from rectus femoris (RF), vastus lateralis (VL), vastus medialis (VM), biceps femoris (BF), medial hamstrings (MH) and medial gastrocnemius (GM) were recorded via bipolar surface electromyograms (EMG). The onset of the response in the RF, VL and VM, remained stable and independent of the time in the step cycle when the stretch was applied. Across all subjects the response onset (mean ± SD) occurred at 23±1, 24±1 and 23±1 ms for RF, VL and VM, respectively. The duration of the initial response was 90–110 ms, at which time the EMG signal returned towards baseline levels. Three reflex response windows, labelled the short latency reflex (SLR), the medium latency reflex (MLR) and the late latency reflex response (LLR), were analysed. The medium and late reflex responses of all knee extensors increased significantly (p=0.008) as the gait cycle progressed from swing to stance. This was not related to the background EMG activity. In contrast, during standing at extensor EMG levels similar to those attained during walking the reflex responses were dependent on background EMG. During walking, LLR amplitudes expressed as a function of the background activity were on average two to three times greater than SLR and MLR reflex amplitudes. Distinct differences in SLR and LLR amplitude were observed for RF, VL and VM but not in the MLR amplitude. This may be related to the different pathways mediating the SLR, MLR and LLR components of the stretch response. As for the knee extensor antagonists, they exhibited a response to the stretch of the quadriceps at latencies short enough to be monosynaptic. This is in agreement with the suggestion by Eccles and Lundberg (1958) that there may be functional excitatory connections between the knee extensors and flexors in mammals.     

Reflex control of ipsilateral and contralateral paraspinal muscles

Homonymous and heteronymous reflex connections of the paraspinal muscles were investigated by the application of a tap to the muscle bellies of the lumbar multifidus and iliocostalis lumborum muscles and observation of surface electromyographic responses in the same muscles on both sides of the trunk. Reflexes were evoked in each of the homonymous muscles with latencies and estimated conduction velocities compatible with being evoked by Ia muscle afferents and having a monosynaptic component. Short latency heteronymous excitatory reflex connections were observed in muscles on the ipsilateral side, whilst reflex responses in the contralateral muscles were inhibitory in response to the same stimulus. The latencies of the crossed responses were on average 9.1 ms longer than the ipsilateral excitatory responses. These results are in contrast to the crossed excitatory responses observed between the abdominal muscles and trapezius muscles on the opposite aspect of the trunk. Such a difference in the reflex pathways between these two groups of trunk muscles compliments the different anatomical arrangement of the muscle groups and suggests a contribution to their commonly observed activation patterns.     

The actualization of heteronomous monosynaptic connections of IA afferents in the muscle motor centers of the human hand

Reflex excitability of spinal centers of hand muscles was examined in normal subjects and patients with traumatic lesion of forearm nerves. Central and peripheral muscle responses were evoked by stimulation of homonymous and heteronymous nerves. Patients with median and/or ulnar nerve lesion showed the heteronymous abductor pollicis brevis and abductor digit minimi H-responses. The cause of appearance of heteronymous responses is supposed to be actualization of latent heteronymous monosynaptic connections following traumatic lesion and afferent deprivation. It seems obvious that activation of heteronymous pathways corresponds to the process in afferent system known as "revitalization" of median-to-ulnar anastomoses after damage.     

Pattern of monosynaptic heteronymous Ia connections in the human lower limb

Changes in the firing probability of single motor units in response to electrical stimulation of muscle nerves were used to derive the projections of muscle spindle Ia afferents to the motoneurones of various leg and thigh muscles. Discharges of units in soleus, gastrocnemius medialis, peroneus brevis, tibialis anterior, quadriceps, biceps femoris and semitendinosus were investigated after stimulation of inferior soleus, gastrocnemius medialis, superficial peroneal, deep peroneal and femoral nerves. Homonymous facilitation, occurring at the same latency as the H reflex and therefore attributed to monosynaptic Ia EPSPs, was found in virtually all the sampled units. In many motor nuclei an early facilitation was also evoked by heteronymous low-threshold afferents. The heteronymous facilitation was considered to be mediated through a monosynaptic pathway when the difference between the central latencies of heteronymous and homonymous peaks was not more than 0.2 ms. The heteronymous Ia connections were widely distributed. In particular, monosynaptic coupling between muscles operating at different joints appears to be the rule in humans, though it is rare between ankle and knee muscles in the cat and the baboon.     

Pattern of projections of group I afferents from forearm muscles to motoneurones supplying biceps and triceps muscles in man

Summary (1) Two independent methods were used, in man, to assess the modifications of the excitability of biceps and triceps brachii motoneurone pools following the stimulation of group I afferents coming from muscles acting at the wrist: (a) the modifications of the excitability of a motoneuronal population were studied using a reflex technique, (b) the modifications of the excitability of an isolated motor unit were estimated using a post-stimulus time histogram (p.s.t.h.) method. (2) The activation of group I afferents contained in the median nerve, originating from wrist flexors and pronators, resulted in a strong, short-latency facilitation of the biceps brachii motoneurones. A similar effect was also evoked by stimulation of group I afferents in the radial nerve, distally to the branch supplying the brachio-radialis muscle. The latency of both median and radial-induced facilitations is compatible with a monosynaptic linkage. (3) The stimulation of group I afferents in the median or the radial nerves produced inhibition of triceps motoneurones, with a latency compatible with a disynaptic linkage. (4) The prolonged vibration of the tendon of the flexor carpi radialis (FCR) or of the extensor carpi radialis (ECR) raised the threshold for both the facilitation of biceps and the inhibition of triceps motoneurones. The same pattern of excitatory and inhibitory convergence could also be obtained when the electrical conditioning stimulus to the median or radial nerves was replaced by a tap applied to the tendons of FCR or ECR respectively. Both results suggest that the conditioning fibres were Ia fibres. (5) The pattern of distribution of Ia afferents from muscles acting at the wrist onto motoneurones of muscles acting at the elbow has been compared to that described in the cat and monkey. A comparison has also been made between Ia connections of muscles acting at different joints in the upper and lower limb in man. The differences are discussed in relation to the manipulating capacity of the hand.     

Facilitation and suppression of wrist and digit muscles from single rubromotoneuronal cells in the awake monkey.

1. The output effects of 214 cells in the magnocellular red nuclei of two rhesus monkeys (Macaca mulatta) were tested with spike-triggered averaging of electromyogram (EMG) activity from six forearm extensor and six flexor muscles. The monkeys performed an alternating wrist movement task (auxotonic paradigm) or generated wrist torque trajectories alternating between flexion and extension (isometric paradigm). 2. Sixty-five cells (30%) were identified as rubromotoneuronal (RM) cells on the basis of their postpike effects on forearm flexor and extensor muscles. Three major types of RM cell output organization were identified: 1) pure facilitation (28 cells), 2) reciprocal (18 cells), and 3) cofacilitation (16 cells). 3. RM cell output showed a strong preference for facilitation of extensor forearm muscles. This preference was reflected in the fact that 69% (43 of 62) of RM cells facilitated extensors exclusively or most strongly; 27% facilitated flexors exclusively or most strongly; and 5% facilitated flexors and extensors equally. Postspike facilitation (PSpF) was observed in 45% of the extensor muscles and 20% of the flexors tested. In contrast, postpike suppression (PSpS) was observed in 3% of the extensors and 7% of the flexors. 4. The mean number of extensors facilitated per RM cell was 3.1 (53% of tested) compared with 2.8 (51% of tested) flexors facilitated per cell. The extensor and flexor PSpS muscle field sizes were both 2.0 (35% of extensors and 36% of flexors tested). The mean number of muscles facilitated by cofacilitation cells was 5.8 (48%) per cell. No clear preference was found for facilitation of particular combinations of synergist muscles. 5. PSpF magnitude was assessed by measuring both the percent change of facilitation or suppression from baseline and the signal-to-noise ratio of effects. The overall average magnitudes of RM PSpF and PSpS were 4.1 +/- 2.0 and 4.0 +/- 2.3% change from baseline, respectively. The average magnitude of PSpF in flexors was not significantly different from that of extensors; neither was there a difference in the average magnitude of PSpS in flexors and extensors. 6. The mean onset latency of RM cell PSpS was greater than PSpF (9.2 +/- 3.0 vs. 5.7 +/- 1.8 ms; P less than or equal to 0.05). This can be attributed to an underlying minimal disynaptic linkage to motoneurons for suppression effects, whereas most PSpFs are probably mediated by underlying monosynaptic connections. The mean onset latency of flexor PSpFs was greater than that of extensors (6.4 +/- 2.3 vs. 5.4 +/- 1.5 ms; P less than or equal to 0.05).(ABSTRACT TRUNCATED AT 400 WORDS)     

Muscle activity during forelimb stepping in decerebrate cats.

In decerebrate cats with the lower thoracic cord transected, electromyographic activities were analyzed in up to 41 forelimb muscles, almost all muscles involved in forelimb stepping (intrinsic hand muscles were not included). From the active period in the step cycle, muscles were classified into three groups: extensors, of which activity is like that of elbow extensors; flexors, activity like that of elbow flexors; others, including dorsiflexors of the wrist, pronators, and supinator. The results were well consistent with those from conscious animals as well as efferent pattern of fictive locomotion in elbow and distal muscles. Nevertheless, in some proximal muscles discrepancies were noted, suggesting their changeability depending on environmental conditions. Recording from almost all muscles allowed to estimate rhythmic change of the overall output of the forelimb central pattern generator.     

Excitability changes in human hand motor area induced by voluntary teeth clenching are dependent on muscle properties

To investigate whether the early effects of voluntary teeth clenching (VTC) among the first dorsal interosseous (FDI), abductor digiti minimi (ADM), and abductor pollicis brevis (APB) muscles are differently modulated depending on their muscle properties, we examined the responses of motor evoked potentials (MEPs) elicited by transcranial magnetic stimulation with selected current directions and by brainstem magnetic stimulation (BMS). Although MEP responses with anterior-medially current direction (preferentially elicited I1-waves) were facilitated in all three muscles, those responses with posterior-laterally current direction (preferentially elicited I3-waves) were different among FDI, ADM, and APB muscles. That is, MEP responses in FDI and APB muscles were significantly reduced, whereas those responses in ADM muscle were not significantly reduced. Further, inhibitory effects of VTC in FDI muscle were more potent than those in ADM or APB muscles. On the other hand, the responses to BMS were unchanged by VTC in all three muscles, suggesting that the modulations of MEP were attributed to the cortical origin. On the basis of our previous findings that the inhibitory connections in FDI muscle are more potent than those in ADM muscle (Takahashi et al. in Clin Neurophysiol 116:2757–2764, 2005), the cortical effects of VTC among three hand muscles are differently modulated, depending on muscle properties, presumably the extents of inhibitory connections to corticospinal tract neurons. Considering that the functional capacity in FDI muscle is higher than that in ADM or APB muscles, the cortical inhibitory effect of VTC might contribute to the sophisticated regulation of the motor outputs even during VTC.     

Neuronal pathways for spinal reflexes activated by group I and group II muscle afferents in the spinal segment (Co1) innervating the tail in the low spinalized cat

 We studied neuronal pathways for spinal reflexes activated by group-I and group-II muscle afferents in the spinal segments innervating the tail in unanesthetized and spinalized (L1) cats. Experiments were performed on 25 adult cats of both sexes. The effects of stimulating nerves innervating six tail muscles on both sides were recorded from tail motoneurons in the first coccygeal spinal segment (Co1) using glass microelectrodes. Stable recordings were obtained from 150 tail motoneurons. Stimulation of group-I muscle afferents (stimulus intensity <1.8 T) often produced EPSPs (82/150) after stimulating nerves innervating neighboring tail muscles. Motoneurons innervating the long-tendoned muscles, M. extensor caudae lateralis and M. flexor caudae longus (ECL and FCL), received heteronymous monosynaptic connections from group-I muscle afferents innervating the ipsilateral tail muscles. The motoneurons innervating segmental muscles, M. extensor caudae medialis and M. flexor caudae brevis (ECM and FCB), received heteronymous monosynaptic connections from group-I muscle afferents innervating tail muscles on both sides. The motoneurons innervating tail muscles originated from the Ossa coxae, M. abductor caudae externus and M. abductor caudae internus (ACE and ACI), received monosynaptic connection from group-I muscle afferents innervating most of the tail muscles on both sides. Crossed disynaptic inhibitory pathways activated by primary muscle afferent inputs were observed in ECM, ACE, FCL, and FCB motoneurons. The effects of group-II afferent inputs were not dependent on the kind of motoneuron, and alternative excitatory and inhibitory pathways were not clearly observed in the tail motoneuron pool. It is suggested that variability of the neuronal pathways from group-I and -II muscle afferents to tail motoneurons corresponds to functional relationships among tail muscles, depending on the tail movements. Received: 9 April 1997 / Accepted: 8 August 1998     

Proportional myoelectric control of a virtual object to investigate human efferent control

We used proportional myoelectric control of a one-dimensional virtual object to investigate differences in efferent control between the proximal and distal muscles of the upper limbs. Eleven subjects placed one of their upper limbs in a brace that restricted movement while we recorded electromyography (EMG) signals from elbow flexors/extensors or wrist flexors/extensors during isometric contractions. By activating their muscles, subjects applied virtual forces to a virtual object using a real-time computer interface. The magnitudes of these forces were proportional to EMG amplitudes. Subjects used this proportional EMG control to move the virtual object through two tracking tasks, one with a static target and one with a moving target (i.e., a sine wave). We hypothesized that subjects would have better control over the virtual object using their distal muscles rather than using their proximal muscles because humans typically use more distal joints to perform fine motor tasks. The results indicated that there was no difference in subjects ability to control virtual object movements when using either upper arm muscles or forearm muscles. These results suggest that differences in control accuracy between elbow joint movements and wrist joint movements are more likely to be a result of motor practice, proprioceptive feedback or joint mechanics rather than inherent differences in efferent control.     

Changes in Corticospinal Excitability in the Reactions of Forearm Muscles in Humans to Vibration

Transcranial magnetic stimulation was used to study changes in corticospinal excitability during vibration of the flexor and extensor muscles of the wrist in healthy humans. The ratios of muscle stimulation responses to activity levels in these muscles on contraction associated with vibration (the tonic vibratory reflex, TVR) and after vibration of antagonist muscles in isometric conditions (the antagonist vibratory reflex, AVR) were analyzed. The normalized muscle response in the wrist flexors was found to increase by 66% compared with threshold values in the direct vibratory response (TVR), by 75% in the relayed vibratory response (AVR), and by 18% on voluntary contraction. However, increases in the motor response in vibratory responses as compared with those on voluntary contraction did not reach significance, which contrasted with the responses in the forearm flexors. These results are discussed from the point of view that the motor cortex plays different roles in vibratory responses in the distal and proximal muscles.     

Properties of primary motor cortex output to forelimb muscles in rhesus macaques

Stimulus-triggered averaging (StTA) of electromyographic (EMG) activity from 24 simultaneously recorded forelimb muscles was used to investigate properties of primary motor cortex (M1) output in the macaque monkey. Two monkeys were trained to perform a reach-to-grasp task requiring multijoint coordination of the forelimb. EMG activity was recorded from 24 forelimb muscles including 5 shoulder, 7 elbow, 5 wrist, 5 digit, and 2 intrinsic hand muscles. Microstimulation (15 microA at 15 Hz) was delivered throughout the movement task. From 297 stimulation sites in M1, a total of 2,079 poststimulus effects (PStE) were obtained including 1,398 poststimulus facilitation (PStF) effects and 681 poststimulus suppression (PStS) effects. Of the PStF effects, 60% were in distal and 40% in proximal muscles; 43% were of extensors and 47% flexors. For PStS, the corresponding numbers were 55 and 45% and 36 and 55%, respectively. M1 output effects showed extensive cofacilitation of proximal and distal muscles (96 sites, 42%) including 47 sites that facilitated at least one shoulder, elbow, and distal muscle, 45 sites that facilitated an elbow muscle and a distal muscle, and 22 sites that facilitated at least one muscle at all joints. The muscle synergies represented by outputs from these sites may serve an important role in the production of coordinated, multijoint movements. M1 output effects showed many similarities with red nucleus output although red nucleus effects were generally weaker and showed a strong bias toward facilitation of extensor muscles and a greater tendency to facilitate synergies involving muscles at noncontiguous joints.     

Reflex Responses in the Lower Leg following Landing Impact on an Inverting and Non-Inverting Platform

In the lower leg, landing after a jump induces reflexes, the role of which is not well understood. This is even more so for reflexes following landing on inverting surfaces. The latter condition is of special interest since ankle inversion traumata are one of the most common injuries during sport. Most studies have investigated ankle inversions during a static standing condition. However, ankle injuries occur during more dynamic activities such as jumping. Therefore, the present study aimed at reproducing these situations but in a completely safe setting. EMG responses were recorded after landing on an inverting surface, which caused a mild ankle inversion of 25 deg of rotation (in a range sufficient to elicit reflexes but safe enough to exclude sprains). The results are compared with data from landing on a non-inverting surface to understand the effect of the inversion. In general, landing on the platform resulted in short and long latency responses (SLR and LLR) in triceps surae (soleus, gastrocnemius medialis and lateralis) and peroneal muscles (long and short peroneal) but not in the tibialis anterior muscle. Landing on the inverting platform caused significant LLRs in the peroneal muscles (which underwent the largest stretch) but not in the triceps muscles. Conversely, landing on a non-inverting platform induced larger SLRs in triceps than in the peroneal muscles. Although the peroneal LLRs thus appeared to be selectively recruited in an inverting perturbation, their role during such perturbations should be limited since the latency of these responses was about 90 ms while the inversion lasts only 42 ms. The SLRs, if present, had an onset latency of around 44 ms. In the period following the inversion, however, the responses may be important in preventing further stretch of these muscles.     

Physiological changes underlying bilateral isometric arm voluntary contractions in healthy humans

Many bilateral motor tasks engage simultaneous activation of distal and proximal arm muscles, but little is known about their physiological interactions. Here, we used transcranial magnetic stimulation to examine motor-evoked potentials (MEPs), interhemispheric inhibition at a conditioning-test interval of 10 (IHI(10)) and 40 ms (IHI(40)), and short-interval intracortical inhibition (SICI) in the left first dorsal interosseous (FDI) muscle during isometric index finger abduction. The right side remained at rest or performed isometric voluntary contraction with the FDI, biceps or triceps brachii, or the tibialis anterior. Left FDI MEPs were suppressed to a similar extent during contraction of the right FDI and biceps and triceps brachii but remained unchanged during contraction of the right tibialis anterior. IHI(10) and IHI(40) were decreased during contraction of the right biceps and triceps brachii compared with contraction of the right FDI. SICI was increased during activation of the right biceps and triceps brachii and decreased during activation of the right FDI. The present results indicate that an isometric voluntary contraction with either a distal or a proximal arm muscle, but not a foot dorsiflexor, decreases corticospinal output in a contralateral active finger muscle. Transcallosal inhibitory effects were strong during bilateral activation of distal hand muscles and weak during simultaneous activation of a distal and a proximal arm muscle, whereas GABAergic intracortical activity was modulated in the opposite manner. These findings suggest that in intact humans crossed interactions at the level of the motor cortex involved different physiological mechanisms when bilateral distal hand muscles are active and when a distal and a proximal arm muscle are simultaneously active.     

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

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