Effect of clenching levels on heteronymous H-reflex in human temporalis muscle

 Selective stimulation of the masseteric nerve has been shown to elicit a heteronymous H-reflex in the ipsilateral temporalis muscle during voluntary clenching. However, the relation between the electromyographic (EMG) activity of the temporalis muscle and the amplitude of the H-reflex has not been previously described. In the present study, the hypothesis was tested that there would be a positive relationship between the level of EMG activity and the amplitude of the H-reflex. The direct motor response (M-response) in the masseter muscle and the heteronymous H-reflex in the anterior temporalis muscle were successfully elicited by electrical stimulation of the masseteric nerve in 12 of 13 subjects. A new automatic system was used to control the on-line EMG activity and to trigger the stimulus. In a random order, two series of 20 stimuli were delivered at each of four clenching levels (0, 25, 50, and 75% of maximal voluntary contraction). The analysis showed that both the masseteric M-response and the temporalis H-reflex were reproducible within and between series. The amplitude of the temporalis H-reflex increased significantly at higher clenching levels (ANOVA: P=0.003). Clenching at 50% and 75% of the maximal voluntary contraction caused significantly larger amplitudes of the H-reflex than clenching at 25% of the maximal voluntary contraction; at rest, no H-reflex could be recorded. There was a significant correlation between the background EMG activity in the ipsilateral temporalis muscle and the amplitude of the H-reflex (Pearson: r=0.313, P=0.008). These data indicate that the heteronymous H-reflex can be reliably elicited by means of an automatic system for stimulus delivery and that the amplitude of the H-reflex is dependent on the preceding activity of the motoneuron pool. Received: 27 November 1998 / Accepted: 16 February 1999     

Modulation of intracortical facilitatory circuits of the human primary motor cortex by digital nerve stimulation

We investigated the effect of electrical digit stimulation on two different intracortical facilitatory phenomena. Paired-pulse transcranial magnetic stimuli (TMS) with different conditioning stimulus (CS) intensities were applied over the primary motor cortex (M1). Electromyographic (EMG) recordings were made from the relaxed right abductor digiti minimi muscle (ADM). The effect of preceding sensory stimulation applied to the ipsilateral digit V on the conditioning magnetic stimulus was examined. Changing the CS intensity affected the influence of peripheral electrical stimulation on motor evoked potential (MEP) amplitudes evoked by paired pulse TMS. Inhibition induced by ipsilateral digit stimulation was strongest with the lowest CS intensity if MEP amplitudes were evoked by a subthreshold CS followed by a suprathreshold test stimulus (TS) at an interstimulus interval (ISI) of 10 ms. In contrast, inhibition induced by digit stimulation in a paired-pulse paradigm with a suprathreshold first and a subthreshold second stimulus at ISI of 1.5 ms was strongest with the highest CS intensity. These findings suggest that appropriately timed peripheral electrical stimuli differentially modulate facilitatory interactions in the primary motor cortex. They further support the hypothesis that intracortical facilitation (ICF) and short-interval intracortical facilitation (SICF) are evoked through different mechanisms. An erratum to this article can be found at     

Experimental skin pain and muscle pain induce distinct changes in human trigeminal motoneuronal excitability

Seeking information on the physiological properties of the trigeminal motoneuronal pool we investigated changes in the excitability of trigeminal motor system induced by two types of experimental pain (muscle and skin). In one session, we studied the effect of muscle pain induced by hypertonic saline infusion into the masseter muscle on the recovery cycle of the heteronymous H-reflex in the temporalis muscle and the homonymous silent period (SP) in the masseter muscle, both elicited by stimulation of the masseteric nerve in ten-healthy subjects. In another session, we studied the effect of laser stimuli applied to the perioral region, at conditioning intervals from 20 to 160 ms, on the temporalis H-reflex and masseter SP in nine healthy subjects. Whereas laser-induced skin pain significantly inhibited the temporalis H-reflex and facilitated the masseter SP (P < 0.01), muscle pain left the time course of the temporalis H-reflex and masseter SP unchanged (P > 0.05). The timing of temporalis H-reflex suppression and masseter-SP enhancement induced by laser stimuli indicates that facial skin nociceptors inhibit trigeminal motoneurones via multysynaptic reflex pathways. Hypertonic saline, a stimulus that predominantly activates group III and IV afferents, left both variables reflecting trigeminal motoneuron excitability unchanged. Due to the differences between the two experimental models, we cannot conclude that such inhibitory reflex pathway does not exist from muscle nociceptors to trigeminal motoneurones.     

Intracortical modulation of cortical-bulbar responses for the masseter muscle

Short interval intracortical inhibition (SICI) and intracortical facilitation (ICF) were evaluated in the masseter muscles of 12 subjects and the cortical silent period (SP) in nine subjects. Motor evoked potentials (MEPs) were recorded from contralateral (cMM) and ipsilateral (iMM) masseters, activated at 10% of maximal voluntary contraction (MVC). Interstimulus intervals (ISIs) were 2 and 3 ms for SICI, 10 and 15 ms for ICF. TMS of the left masseteric cortex induced MEPs that were larger in the cMM than the iMM; stimulation of right masseteric cortex produced a similar asymmetry in response amplitude. SICI was only observed using a CS intensity of 70% AMT and was equal in both cMM and iMM. SICI was stronger at higher TS intensities, was abolished by muscle activation greater than 10% MVC, and was unaffected by coil orientation changes. Control experiments confirmed that SICI was not contaminated by any inhibitory peripheral reflexes. However, ICF could not be obtained because it was masked by bilateral reflex depression of masseter EMG caused by auditory input from the coil discharge. The SP was bilateral and symmetric; its duration ranged from 35 to 70 ms depending on TS intensity and coil orientation. We conclude that SICI is present in the cortical representation of masseter muscles. The similarity of SICI in cMM and iMM suggests either that a single pool of inhibitory interneurons controls ipsi- and contralateral corticotrigeminal projections or that inhibition is directed to bilaterally projecting corticotrigeminal fibres. Finally, the corticotrigeminal projection seems to be weakly influenced by inhibitory interneurons mediating the cortical SP.     

Experimental jaw-muscle pain does not change heteronymous H-reflexes in the human temporalis muscle

Muscle pain generally has an inhibitory effect on voluntary orofacial motor function. However, it is not known whether muscle pain causes direct or indirect changes in motoneuron excitability. In this study a monopolar needle stimulation technique was used to evoke the direct motor response (M-response) in the left masseter muscle and the heteronymous H-reflex in the left temporalis muscle as an indirect measure of motoneuron excitability. Series of 20 repeated electrical stimuli were delivered at 50% of maximal voluntary contraction (MVC) before, during, and after periods with experimental jaw-muscle pain in 11 healthy subjects. Pain was induced by standardized infusion of hypertonic (5%) saline into the mid-portion of the right masseter muscle. The mean pain intensity rating on a 100-mm visual analog scale was 42±5 mm. The short-latency responses (less than 6 ms) could be evoked in all subjects. Analysis of the latency and amplitude of the temporal H-reflex indicated no significant effect of jaw-muscle pain. The amplitude of the masseteric M-response was significantly smaller in the postpain condition than in the pain conditions (ANOVA, P=0.018), but no differences were found between the prepain and postpain conditions. In nine subjects, poststimulus periods (mean offset latency, 69.6±8.6 ms) with significantly (more than 50%) suppressed EMG activity were detected in the ipsilateral masseter muscle following the M-response (mean offset latency, 5.5±0.2 ms). These reflex responses did not show a systematic change during the pain conditions. In conclusion, acute contralateral jaw-muscle pain does not seem to modulate the motoneuron excitability as measured by the heteronymous H-reflex. Received: 7 November 1997 / Accepted: 16 February 1998     

H-reflexes in masseter and temporalis muscles in man

In contrast with limb muscles, studies on H-reflexes in the trigeminal system are scarce. The present report aimed at reevaluating the responses obtained in the masseter and temporalis muscles after electrical stimulation of their nerves. Twenty-four subjects participated in the experiments. The reflexes were elicited in the masseter and temporal muscles by monopolar stimulation and recorded using surface electrodes. Stimulation of the masseteric nerve evoked an M-response in the masseter and an H-reflex in both the masseter and the temporal muscles. In contrast with the masseter muscle, where the homonymous H-reflex disappeared at higher stimulation intensities, the heteronymous temporal H-reflex remained and reached a plateau. Simultaneous stimulation of the masseteric and deep temporal nerves resulted in an M-response and an H-reflex in both the masseter and temporal muscles. Increasing stimulus intensitites led to disappearance of the H-reflex in both muscles. The results were compared with those obtained by others on limb muscles. As in these muscles, the presence of heteronymous H-reflexes in the jaw muscles can be used in future studies of motoneuronal excitability.     

Specific modulation of the Hoffmann reflex cutaneous facilitation during a reaction-time task

Summary The Hoffmann (H) reflex and its facilitation produced by electrical stimulation of the sural area were examined before a ballistic extension of the right foot. Modulations of the cutaneous facilitation of the H reflex (CFH) were used to assess the control exerted over the transmission of low threshold cutaneous afferents. The time-course of H and CFH changes were investigated at the end of the foreperiod and during the premotor period, i.e. between the response signal and the onset of the electromyogram (EMG) of the soleus muscle. Four stimulation conditions were set up depending on whether the H reflex was elicited on the contracting or non-contracting limb, and whether cutaneous stimulation was ipsilateral or contralateral to the reflex. During the 100 ms preceding the response signal, the inhibition of the H reflex was more marked in the contracting limb than in the noncontracting limb. At the end of the foreperiod, the CFHs had a symmetric time course: the CFHs evoked by conditioning stimulation of the contracting limb were facilitated just before the response signal, while those produced by conditioning stimulation to the non-contracting limb were depressed. It is suggested that these variations are related to postural adjustments taking place before the movement is performed. As previously reported, the H reflex of the contracting limb exhibited a marked increase in amplitude over the 50 ms preceding the EMG. Reflex facilitations showed specific variations according to the ankle stimulated and the soleus muscle tested. The CFHs produced by stimulation of the non-contracting limb regained, at the end of the premotor period, a value close to their reference level recorded in trials without movement. The CFHs, produced by conditioning stimulation of the contracting limb were modulated differently according to whether the tested soleus muscle was contracting or not: when the CFH was tested on the contracting muscle, it was found to be depressed throughout the premotor period; this contrasted significantly with the isolated depression recorded on the non-contracting muscle. Therefore, only the cutaneous afferents from the mobilized limb, modulating the H reflex of the same limb, were subject to a specific inhibition during the premotor period. Throughout the preparatory and premotor periods, negative correlations were observed between H and CFH amplitude, except just before the EMG onset in the condition where the H reflex was delivered to the contracting muscle and the cutaneous stimulation to the ankle of the non-contracting limb: in this case, CFH amplitude increase paralleled that of the reflex amplitude. The negative correlations between these two variables are discussed in terms of a central control regulating the balance between proprioceptive and cutaneous input.     

Effects of flexor reflex afferent stimulation on the soleus H reflex in patients with a complete spinal cord lesion: evidence for presynaptic inhibition of Ia transmission

Summary The effects of electrically stimulating the Flexor Reflex Afferent (FRA) on the soleus H reflexes were investigated in 34 paraplegic patients having a clinically complete spinal cord lesion. Conditioning stimuli (5–50 mA) were applied to the ipsilateral or contralateral sural nerve. The conditioning-test interval ranged from 20 to 1000 ms. A late ipsilateral flexor reflex (EMG) was found in all patients. A late contralateral extension reflex was sporadically observed in only 3 patients. The excitability curves usually showed two phases of ipsilateral H reflex inhibition and contralateral H reflex facilitation, one between 50 and 130 ms and the other after over 200 ms. These intervals correspond to early and late flexion reflexes. With high intensity stimulation the early and late ipsilateral inhibition fused. An early low threshold ipsilateral facilitation occured in 9 patients. The contralateral late facilitation was followed by prolonged inhibition in 10 patients. Changes in presynaptic inhibition were assessed by measuring the heteronymous monosynaptic Ia facilitation from quadriceps to soleus. For methodological reasons, it was only possible to investigate the effect of contralateral conditioning volleys which was performed in 5 patients. A significant and regular reduction of the heteronymous Ia facilitation was found in 4 patients. This reduction is taken to indicate that the FRA evokes presynaptic inhibition of Ia transmission to alpha motoneurones. Presynaptic inhibition was also indicated by the enhancement of a vibratory stimulus induced inhibition in 2 subjects. These results are consistent with the hypothesis that the reflex organization in patients with a spinal cord section is similar to that of the acute spinal cat injected with DOPA.     

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.     

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.     

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.     

Non-nociceptive upper limb afferents modulate masseter muscle EMG activity in man

Recent electrophysiological data obtained in anaesthetized rats evidenced jaw muscle excitatory responses to the electrical stimulation of type II limb somatosensory afferents. In the present work, we describe an inhibitory reflex evoked in human masseter muscles by stimulation of non-nociceptive fibres travelling in the median and radial nerves (MED and RAD, respectively). Eighteen healthy volunteers participated in the study. Subjects were seated on a comfortable chair, with the complex head-mandible-neck-trunk and the limbs securely fixed to the chair. Attempts were made to minimize possible interferences due to the activation of afferents other than the stimulated ones. The subjects were instructed to contract masseter muscles at a submaximal level and to maintain a stable level of muscle contraction during all trials. EMG voluntary activity was recorded from both masseter muscles by means of coaxial needle electrodes before and after the electrical stimulation of MED and/or RAD at intensities below pain threshold. In all subjects, MED stimulation induced bilaterally a marked depression of masseter EMG activity, which occurred at a latency of 23.6 +/- 2.1 ms and lasted 27.8 +/- 6.6 ms. RAD stimulation also induced a marked reduction in masseter EMG activity, but this effect was clearly observed in 9 out of 18 subjects, and it showed latency (30.2 +/- 7.5 ms) and duration (44.9 +/- 5.4 ms) significantly longer in comparison with the MED-induced effect. All subjects exhibited the inhibitory period in masseter EMG following the simultaneous stimulation of both nerves; this one appeared at a latency not significantly different (25.3 +/- 5.9 ms) and lasted much more (37.4 +/ - 8.2 ms) than EMG depression evoked by MED stimulation. The duration of masseter muscle inhibition, induced by MED and/or RAD stimulation, was inversely related to the level of EMG activity, while latency was not related to it. Significant increases in the inhibitory period duration were also observed by increasing stimulus intensity, within a subthreshold range for the activation of nociceptive fibres. In all cases, the inhibitory period was followed by a later excitatory rebound activity, whose latency and duration depended on the duration of the preceding EMG inhibition and on the background level of masseter activation. In conclusion, results evidenced that the activation of arm somatosensory fibres modulates masseter muscle activity in normal man. This might lead to a coordination between limb and masticatory muscle activity, which is required in several complex motor acts.     

Synaptic linkage of masseter muscle afferents to masseteric motoneurones in the cat

The synaptic linkage of masseter muscle afferents to masseteric motoneurones was investigated under blockage of soma-dendritic invasion of antidromic spikes by passing constant inward current across the cell membrane. A monosynaptic latency for excitatory postsynaptic potentials produced by group Ia afferents was measured as 1.3 ms and no group lb component was obtained. Inhibitory postsynaptic potentials with latencies of 5.5 ms were produced at a stimulus strength of 4.5 times the threshold of group Ia fibers. On the basis of stimulus strength, muscle afferents activated at 4.5 times the threshold and producing inhibitory postsynaptic potentials in masseteric motoneurones are probably group II afferents. The same reversal point was obtained in both the lingually induced and the group II IPSPs, indicating that the group II inhibitory postsynaptic potential is dependent on an increased permeability to Cl ions. The inhibitory postsynaptic potentials produced by stimulation of the high threshold muscle afferents were the composite of a strychnine-sensitive and strychnine-insensitive inhibitory postsynaptic potential. The latency of the inhibitory postsynaptic potentials caused by the high threshold muscle afferents was about 10 ms.     

Stimulation of the group I extensor afferents prolongs the stance phase in walking cats

Group I afferents in nerves innervating the lateral gastrocnemius-soleus (LG-Sol), plantaris (P1), and vastus lateralis/intermedius (VL/VI) muscles were stimulated during walking in decerebrate cats. The stimulus trains were triggered at a fixed delay following the onset of bursts in the medial gastrocnemius muscle. Stimulation of all three nerves with long stimulus trains (>600 ms) prolonged the extensor bursts and delayed the onset of flexor burst activity. LG-Sol nerve stimulation had the strongest effect; often delaying the onset of flexor burst activity until the stimulus train was ended. By contrast, flexor bursts were usually initiated before the end of the stimulus train to the P1 and VL/VI nerves. The minimum stimulus strength required to increase the cycle period was between 1.3×threshold and 1.6×threshold for all three nerves. Simultaneous stimulation of the P1 and VL/VI nerves produced a larger effect on the cycle period than stimulation of either nerve alone. The spatial summation of inputs from knee and ankle muscles suggests that the excitatory action of the group I afferents during the stance phase is distributed to all leg extensor muscles. Stimulation of the group I afferents in extensor nerves generally produced an increase in the amplitude of the heteronymous extensor EMG towards the end of the stance phase. This increase in amplitude occurred even though there were only weak monosynaptic connections between the stimulated afferents and the motoneurones that innervated these heteronymous muscles. This suggests that the excitation was produced via oligosynaptic projections onto the extensor motoneuronal pool. Stimulation with 300 ms trains during the early part of flexion resulted in abrupt termination of the swing phase and reinitiation of the stance phase of the step cycle. The swing phase resumed coincidently with the stimulus offset. Usually, stimulation of two extensor nerves at group I strengths was required to elicit this effect. We were unable to establish the relative contributions of input from the group 1a and group 1b afferents to prolonging the stance phase. However, we consider it likely that group Ib afferents contribute significantly, since their activation has been shown to prolong extensor burst activity in reduced spinal preparations. Thus, our results add support to the hypothesis that unloading of the hindlimb during late stance is a necessary condition for the initiation of the swing phase in walking animals.     

Neuromuscular and biomechanical coupling in human cycling: modulation of cutaneous reflex responses to sural nerve stimulation

This study tested the hypothesis that the modulation of cutaneous reflexes during human cycling would be dependent on muscle biomechanical function and phase of leg movement. The coupling between neuromuscular (electromyographic, EMG), kinetic and kinematic responses to brief innocuous (75% of the pain threshold PnT) and noxious (125% PnT) sural nerve stimulation were studied. Stimuli were delivered pseudorandomly at eight equidistant (45°) positions of the crank cycle. Peak ipsilateral middle latency EMG reflex responses were calculated between 70 and 130 ms post stimulus in Biceps Femoris (BF), Rectus Femoris (RF), Tibialis Anterior (TA) and Soleus (SOL). Peak torque, knee and ankle joint angle changes were calculated between 140 and 220 ms post stimulus to quantify net kinetic and kinematic reflex modulation. Reflex responses were predominately suppressive during early activation of all muscles and facilitatory during BF and TA muscle inactivation. EMG reflex responses in monoarticular lower leg muscles TA and SOL were well correlated with ankle angle in dorsi/plantaflexion, whereas the correlation between reflex modulation in biarticular upper leg muscles (BF and RF) and knee angle changes in flexion/extension was weaker. Stimulation provoked significant ankle eversion over the whole crank cycle for both stimulus intensities, which was correlated with TA and BF EMG reflex responses. Torque modulation followed EMG and kinematic changes in a movement phase-dependent manner. Reflex magnitude was stimulation intensity-dependent. Supplementary nociceptive activation may contribute for this increase. We conclude that sural nerve stimulation during human cycling evokes distinct reflex responses in muscles operating around the knee (BF and RF) and the ankle (TA and SOL). These reflexes are modulated in a phase-dependent manner depending on muscle biomechanical function to generate energy for limb and crank propulsion during a specific region in the cycle. This modulation contributed to a specific adaptation of joint motion and force production in order to maintain task performance.     

Reflex responses in active muscles elicited by stimulation of low-threshold afferents from the human foot.

1. Reflex responses were elicited in muscles that act at the ankle by electrical stimulation of low-threshold afferents from the foot in human subjects who were reclining supine. During steady voluntary contractions, stimulus trains (5 pulses at 300 Hz) were delivered at two intensities to the sural nerve (1.2-4.0 times sensory threshold) or to the posterior tibial nerve (1.1-3.0 times motor threshold for the intrinsic muscles of the foot). Electromyographic (EMG) recordings were made from tibialis anterior (TA), peroneus longus (PL), soleus (SOL), medial gastrocnemius (MG), and lateral gastrocnemius (LG) muscles by the use of intramuscular wire electrodes. 2. As assessed by averages of rectified EMG, stimulation of the sural or posterior tibial nerves at nonpainful levels evoked a complex oscillation with onset latencies as early as 40 ms and lasting up to 200 ms in each muscle. The most common initial responses in TA were a decrease in EMG activity at an onset latency of 54 ms for sural stimuli, and an increase at an onset latency of 49 ms for posterior tibial stimuli. The response of PL to stimulation of the two nerves began with a strong facilitation of 44 ms (sural) and 49 ms (posterior tibial). With SOL, stimulation of both nerves produced early inhibition beginning at 45 and 50 ms, respectively. With both LG and MG, sural stimuli produced an early facilitation at 52-53 ms. However, posterior tibial stimuli produced different initial responses in these two muscles: facilitation in LG at 50 ms and inhibition in MG at 51 ms. 3. Perstimulus time histograms of the discharge of 61 single motor units revealed generally similar reflex responses as in multiunit EMG. However, different reflex components were not equally apparent in the responses of different single motor units: an individual motor unit could respond slightly differently with a change in stimulus intensity or background contraction level. The multiunit EMG record represents a global average that does not necessarily depict the precise pattern of all motor units contributing to the average. 4. When subjects stood erect without support and with eyes closed, reflex patterns were seen only in active muscles, and the patterns were similar to those in the reclining posture. 5. It is concluded that afferents from mechanoreceptors in the sole of the foot have multisynaptic reflex connections with the motoneuron pools innervating the muscles that act at the ankle. When the muscles are active in standing or walking, cutaneous feedback may play a role in modulating motoneuron output and thereby contribute to stabilization of stance and gait.     

Posture-related changes in heteronymous recurrent inhibition from quadriceps to ankle muscles in humans

The possibility was investigated that changes in heteronymous recurrent inhibition (RI) from quadriceps (Q) to soleus (Sol) and tibialis anterior (TA) motoneurons (MNs) occur during postural tasks requiring cocontraction of Q with one of these muscles. Stimulation of the femoral nerve (FN), which elicited a Q H-reflex discharge, was used to activate Renshaw cells. The resulting inhibition of TA and Sol MNs was assessed using three test responses: (1) the rectified and averaged ongoing electromyogram (EMG) activity in TA or Sol; (2) the motor-evoked potential (MEP) elicited by cortical stimulation in these muscles; and (3) the Sol H reflex. The characteristics of the depression (appearance and increase with the conditioning reflex discharge, short central delay and long duration) are consistent with a Renshaw origin. In addition, results obtained in control experiments (no change in the EMG suppression after an ischaemic blockade of group-I afferents from the leg, time course of the FN-induced depression of the MEP similar to that of the ongoing EMG) made a significant contribution from other pathways activated by FN stimulation unlikely. Posture-related heteronymous RI from Q was compared in different postural tasks at matched levels of background EMG activity: voluntary co-contraction of Q and of the relevant ankle muscle while sitting (control situation), postural co-contraction of Q and TA (while leaning backwards during stance), or contraction of Sol with (preparation for hopping) and without (standing on tip of toes and leaning forwards during stance) associated contraction of the Q. During stance, heteronymous RI from Q was reduced to TA (but not to Sol) while leaning backwards and to Sol in preparation for hopping, but not in the other situations. Thus, RI from Q to TA or Sol was specifically decreased when a co-contraction of the Q and of the relevant muscle operating at the ankle was required to maintain bipedal stance. It is argued that this control of Renshaw cells is descending in origin and contributes to selection of the appropriate synergism in various postural tasks.     

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.     

Do muscle afferents contribute to the cervical response evoked by electrical stimulation of the median nerve in man?

Summary The possible contribution of low threshold muscle afferents to the postsynaptic component (N₁₃) of the cervical response evoked by electrical stimulation of the median nerve (MN) was investigated in normal subjects. Electroneurographic (ENG) and electromyographic (EMG) correlates of the reflex motoneuronal discharge (RMND) were recorded simultaneously. A. No reflex activity could be elicited by stimulation of the MN at the wrist, at least in the resting subjects, while well developed ENG (P₂ efferent volley) and EMG (H reflex) monosynaptic responses occurred following stimulation of the MN at the elbow at suitable strengths. In neither case could a surface correlate of interneuronal activity evoked by muscle afferents be demonstrated. B. Recruitment curves showed that at stimulus intensities above maximal for the H reflex both P₂ and H responses started to decrease until they completely disappeared, while N₁₃ showed further enhancement. C. Subthreshold conditioning stimulation of the MN enhanced both P₂ and H responses, while vibratory muscle stimuli provoked a clearcut suppression of these two responses. In contrast, N₁₃ was completely unaffected by either manuvre. D. No cervical evoked activity could be detected following tendon tapping of the anterior forearm muscles in spite of the appearance of well developed cortical responses and the ENG and EMG correlates of the T reflex. E. Conditioning volleys elicited by tendon taps of the anterior forearm muscles suppressed both P₂ and H responses following stimulation of the MN at the elbow without affecting the related N₁₃ component. F. Conditioning supramaximal stimulation of the MN at the wrist suppressed the N₁₃ component of the cervical response evoked by stimulation of the MN at the elbow without affecting the related reflex responses. No component chronologically related to the RMND could be recorded at the posterior neck region during suppression of N₁₃, thus ruling out the possibility that failure to detect the RMND (as well as its interneuronal concomitants) with cervical electrodes is due to a masking effect of the N₁₃ component. G. Conditioning tendon taps of anterior forearm muscles provoked a clearcut reduction of the primary cortical response to finger stimulation without affecting the postsynaptic component of the related cervical response. It is concluded that neither segmental (motoneuronal or interneuronal in origin) nor ascending postsynaptic impulses generated in the spinal cord by stimulation of low threshold muscle afferents contribute to N₁₃, the latter being probably due to activation of both short and long axoned spinal neurons by cutaneous afferents.     

Absence of effects of contralateral group I muscle afferents on presynaptic inhibition of Ia terminals in humans and cats

Crossed effects from group I afferents on reflex excitability and their mechanisms of action are not yet well understood. The current view is that the influence is weak and takes place indirectly via oligosynaptic pathways. We examined possible contralateral effects from group I afferents on presynaptic inhibition of Ia terminals in humans and cats. In resting and seated human subjects the soleus (SO) H-reflex was conditioned by an electrical stimulus to the ipsilateral common peroneal nerve (CPN) to assess the level of presynaptic inhibition (PSI_control). A brief conditioning vibratory stimulus was applied to the triceps surae tendon at the contralateral side (to activate preferentially Ia muscle afferents). The amplitude of the resulting H-reflex response (PSI_conditioned) was compared to the H-reflex under PSI_control, i.e., without the vibration. The interstimulus interval between the brief vibratory stimulus and the electrical shock to the CPN was -60 to 60 ms. The H-reflex conditioned by both stimuli did not differ from that conditioned exclusively by the ipsilateral CPN stimulation. In anesthetized cats, bilateral monosynaptic reflexes (MSRs) in the left and right L(7) ventral roots were recorded simultaneously. Conditioning stimulation applied to the contralateral group I posterior biceps and semitendinosus (PBSt) afferents at different time intervals (0-120 ms) did not have an effect on the ipsilateral gastrocnemius/soleus (GS) MSR. An additional experimental paradigm in the cat using contralateral tendon vibration, similar to that conducted in humans, was also performed. No significant differences between GS-MSRs conditioned by ipsilateral PBSt stimulus alone and those conditioned by both ipsilateral PBSt stimulus and contralateral tendon vibration were detected. The present results strongly suggest an absence of effects from contralateral group I fibers on the presynaptic mechanism of MSR modulation in relaxed humans and anesthetized cats.