Reaction time and spinal excitability in a simple reaction time task

H and Tendon reflexes were elicited from subjects at varying intervals after the reaction signal (RS) of a simple reaction time (RT) task when the response involved a rapid plantar flexion of the left foot. It was found that a reflex stimulus presented in close temporal proximity to the RS significantly shortened RT in comparison with the RT when no reflex stimulus was presented. This effect was discussed in terms of the intersensory facilitation of RT. No significant evidence was found in support of previous evidence that RT is delayed when a reflex stimulus occurs about 100 msec after the RS. A large facilitation of reflex amplitude was observed commencing 90–120 msec before the onset of the voluntary movement. This was discussed in terms of the hypothesised stages of information processing involved in the latent period of simple RT.     

Intracellular analysis of reflex pathways underlying the stumbling corrective reaction during fictive locomotion in the cat

In cat and humans, contact between an obstacle and the dorsum of the foot evokes the stumbling corrective reaction (reflex) that lifts the foot to avoid falling. This reflex can also be evoked by short trains of stimuli to the cutaneous superficial peroneal (SP) nerve in decerebrate cats during the flexion phase of fictive locomotion. Here we examine intracellular events in hindlimb motoneurons accompanying stumbling correction. SP stimulation delivered during the flexion phase excites knee flexor motoneurons at short latency [minimum excitatory postsynaptic potential (EPSP) latency 1.8 ms; mean 2.7 ms]. Although a similar short latency excitation occurs in ankle extensors (mean latency, 2.8 ms), recruitment is delayed until successive shocks in the stimulus train overcome the locomotor-related hyperpolarization of ankle extensors. In ankle flexor motoneurons, SP stimulation evokes an inhibition (mean latency, 2.7 ms) that briefly reduces or stops their firing during the flexion phase. There is a phase-dependent modulation of SP-evoked EPSP amplitude as well as latency during locomotion. However, the more obvious change in SP reflex pathways with the onset of fictive locomotion is the reduced inhibition of ankle extensor motoneurons and the increased inhibition of ankle flexors. These results show that the characteristic pattern of hindlimb motoneuron activation during SP nerve-evoked stumbling correction results from 1) di- and trisynaptic excitation of knee flexor and ankle extensor motoneurons; 2) increased inhibitory postsynaptic potentials in ankle flexors and a suppression of inhibition in extensors, 3) sculpting of the short-latency SP postsynaptic effects by motoneuron membrane potential, and 4) longer latency excitatory effects that are likely evoked by lumbar interneurons involved in the generation of fictive locomotion.     

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.     

Riluzole decreases flexion withdrawal reflex but not voluntary ankle torque in human chronic spinal cord injury

The objectives of this study were to probe the contribution of spinal neuron persistent sodium conductances to reflex hyperexcitability in human chronic spinal cord injury. The intrinsic excitability of spinal neurons provides a novel target for medical intervention. Studies in animal models have shown that persistent inward currents, such as persistent sodium currents, profoundly influence neuronal excitability, and recovery of persistent inward currents in spinal neurons of animals with spinal cord injury routinely coincides with the appearance of spastic reflexes. Pharmacologically, this neuronal excitability can be decreased by agents that reduce persistent inward currents, such as the selective persistent sodium current inhibitor riluzole. We were able to recruit seven subjects with chronic incomplete spinal cord injury who were not concurrently taking antispasticity medications into the study. Reflex responses (flexion withdrawal and H-reflexes) and volitional strength (isometric maximum voluntary contractions) were tested at the ankle before and after placebo-controlled, double-blinded oral administration of riluzole (50 mg). Riluzole significantly decreased the peak ankle dorsiflexion torque component of the flexion withdrawal reflex. Peak maximum voluntary torque in both dorsiflexion and plantarflexion directions was not significantly changed. Average dorsiflexion torque sustained during the 5-s isometric maximum voluntary contraction, however, increased significantly. There was no effect, however, on the monosynaptic plantar and dorsiflexor H-reflex responses. Overall, these results demonstrate a contribution of persistent sodium conductances to polysynaptic reflex excitability in human chronic spinal cord injury without a significant role in maximum strength production. These results suggest that intrinsic spinal cellular excitability could be a target for managing chronic spinal cord injury hyperreflexia impairments without causing a significant loss in volitional strength.     

Stumbling corrective reaction during fictive locomotion in the cat

An obstacle contacting the dorsal surface of a cat's hind foot during the swing phase of locomotion evokes a reflex (the stumbling corrective reaction) that lifts the foot and extends the ankle to avoid falling. We show that the same sequence of ipsilateral hindlimb motoneuron activity can be evoked in decerebrate cats during fictive locomotion. As recorded in the peripheral nerves, twice threshold intensity stimulation of the cutaneous superficial peroneal (SP) nerve during the flexion phase produced a very brief excitation of ankle flexors (e.g., tibialis anterior and peroneus longus) that was followed by an inhibition for the duration of the stimulus train (10-25 shocks, 200 Hz). Extensor digitorum longus was always, and hip flexor (sartorius) activity was sometimes, inhibited during SP stimulation. At the same time, knee flexor and the normally quiescent ankle extensor motoneurons were recruited (mean latencies 4 and 16 ms) with SP stimulation during fictive stumbling correction. After the stimulus train, ankle extensor activity fell silent, and there was an excitation of hip, knee, and ankle flexors. The ongoing flexion phase was often prolonged. Hip extensors were also recruited in some fictive stumbling trials. Only the SP nerve was effective in evoking stumbling correction. Delivered during extension, SP stimulus trains increased ongoing extensor motoneuron activity as well as increasing ipsilateral hip, knee, and ankle hindlimb flexor activity in the subsequent step cycle. The fictive stumbling corrective reflex seems functionally similar to that evoked in intact, awake animals and involves a fixed pattern of short-latency reflexes as well as actions evoked through the lumbar circuitry responsible for the generation of rhythmic alternating locomotion.     

In vivo muscle fibre behaviour during counter-movement exercise in humans reveals a significant role for tendon elasticity

Six men performed a single ankle plantar flexion exercise in the supine position with the maximal effort with counter movement (CM, plantar flexion preceded by dorsiflexion) and without counter movement (NoCM, plantar flexion only) produced by a sliding table that controlled applied load to the ankle (40 % of the maximal voluntary force). The reaction force at the foot and ankle joint angle were measured using a force plate and a goniometer, respectively. From real-time ultrasonography of the gastrocnemius medialis muscle during the movement, the fascicle length was determined. The estimated peak force, average power, and work at the Achilles' tendon during the plantar flexion phase in CM were significantly greater than those in NoCM. In CM, in the dorsiflexion phase, fascicle length initially increased with little electromyographic activity, then remained constant while the whole muscle-tendon unit lengthened, before decreasing in the final plantar flexion phase. In NoCM, fascicle length decreased throughout the movement and the fascicle length at the onset of movement was longer than that of the corresponding phase in CM. It was concluded that during CM muscle fibres optimally work almost isometrically, by leaving the task of storing and releasing elastic energy for enhancing exercise performance to the tendon.     

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.     

Contribution of muscle afferents to prolonged flexion withdrawal reflexes in human spinal cord injury

The contribution of force-sensitive muscular afferents to prolonged flexion withdrawal reflexes, or flexor spasms, after human spinal cord injury (SCI) was investigated. In three separate experimental conditions, flexion reflexes were triggered in subjects with SCI using trains of electrocutaneous stimuli delivered at the foot and lower leg and compared with reflexes elicited via intramuscular (i.m.) electrical stimuli. In the first experiment, flexion reflexes were elicited using i.m. stimuli to the tibialis anterior (TA) in the majority of subjects tested. The ratio of peak isometric ankle to hip torques during i.m.-triggered reflexes were proportionally similar to those evoked by electrocutaneous foot or shank stimulation, although the latency to onset and peak flexion torques were significantly longer with i.m. stimulation. In the second experiments, the amplitude and frequency of i.m. TA stimulation were varied to alter the stimulus-induced muscle torque. Peak ankle and hip torques generated during the flexion reflex responses were correlated to a greater extent with stimulus-induced muscle torques as compared with the modulated stimulus parameters. In the third experimental series, i.m. stimuli delivered to the gastrocnemius (GS) elicited flexion reflexes in approximately half of the subjects tested. The combined data indicate a potentially prominent role of the stimulus-induced muscle contraction to the magnitude and latency of flexor reflex behaviors after i.m. TA stimulation. Results after i.m. GS stimulation indicate multi-joint flexion reflexes can also be elicited, although to a lesser extent than i.m. TA stimulation.     

Cyclic modulation of the H-reflex in a wrist flexor during rhythmic flexion-extension movements of the ipsilateral foot

 In 12 subjects, each sitting on an armchair with the right forearm prone, the H-reflex elicited in the resting flexor carpi radialis muscle underwent cyclic excitability changes correlated with rhythmic flexion-extension movements of the ipsilateral foot (frequency of oscillations between 1.5 and 2.5 Hz). During foot plantar flexion, the H-reflex underwent a clear-cut increase, the maximum facilitation falling, in most subjects, within the second half of that phase; then, a gradual reduction in size led the reflex amplitude back to the initial value at the end of foot dorsal extension. If present also when the wrist and the ankle are moved together, this facilitation should favour the in-phase (isodirectional) association between movements and, conversely, hinder the anti-phase coupling. Received: 16 June 1997 / Accepted: 10 July 1997     

Role of proprioceptive information in the temporal coordination between joints

Ten subjects made rapid, simultaneous movements of jaw (elevation or lowering) and right foot (ankle flexion or extension) in two experimental situations: (1) in response to an external signal (reaction-time situation), and (2) in a self-paced situation. We calculated the mean time intervals between the onsets of electromyographic (EMG) activity of agonist muscles (tibialis anterior or gastrocnemius lateralis compared with masseter or digastricus pars anterior) and those between the onsets of movement acceleration at each joint. Despite the fact that subjects reported simultaneous jaw-foot movements, there was always a short time interval between the two movements as between the agonist EMG activities. When the subjects were asked to perform a jaw elevation movement simultaneously with an ankle movement (flexion or extension), the sign of the time interval was dependent on the situation of movement initiation. In the reaction-time situation, the jaw motor activity preceded that of the ankle, whereas the reverse temporal order was observed in the self-paced situation. This is consistent with a previous hypothesis suggesting that the simultaneity of two motor actions is centrally established through two separate central processes: reactive or predictive. When subjects tried to perform simultaneous jaw lowering and foot flexion or extension movements, the strict temporal order observed when considering jaw elevation and ankle movements disappeared. The jaw motor activity generally preceded that of ankle in the reactive situation, but, depending on the subjects, it preceded or followed the ankle motor activity during self-paced movements. It is likely that the specific spindle supply of jaw muscles accounts for these results. Indeed, the jaw depressor muscles, in contrast to the elevators, lack muscle spindles. Our results suggest that the kinesthetic inputs used by the upper central nervous system to synchronize two rapid voluntary movements are mainly those from spindles located in the muscles that accelerate the movement, suggesting a strong α-γ linkage. Received: 31 October 1996 / Accepted: 1 September 1997     

Increases in reflex excitability of monkey masseter motoneurons before a jaw-bite reaction-time response.

Reflex excitability of the motoneurons innervating the masseter muscle of monkeys was tested before a phasic voluntary activation of the jaw-closing muscles (a RT bite response). Single test shocks were delivered to the Mes V which supplies a monosynaptic excitatory input to the jaw-muscle motoneurons. Changes in reflex excitability were assessed by measuring the amplitude of the synchronous muscle potential evoked by the test shock. Amplitudes of the muscle potentials evoked by shocks which occurred just before the beginning of the voluntary EMG response, as judged by the onset of EMG activity of the masseter muscle contralateral to the test shock were many times larger than potentials evoked immediately following the visual RT stimulus. Curves relating the average amplitude of the evoked response to its time before the beginning of the voluntary response suggest that the reflex excitability of the motoneuron pool begins to increase 25-45 ms before the first detectable EMG activity occurs. These results suggest that inputs arrive at the motoneurons of agonist muscles used in rapid RT tasks substantially before changes in the EMG of the muscle are noted. These results, in part, would account for the time interval noted between the beginning of neural activity in suprasegmental structures which presumably excites spinal motoneurons, and the first EMG activity of muscles which are innervated by these motoneurons.     

Facilitation of transmission via inhibitory pathway 1a to spinal extensor motoneurons in response to stimulation of the forelimb nerves in cats

Activation of forelimb flexor reflex afferent in cats anesthetized with a mixture of Chloralose and Nembutal evoked temporospatial facilitation in the reciprocal inhibitory pathways to spinal extensor motoneurons. The amplitude of disynaptic reciprocal 1a inhibitory postsynaptic potentials (IPSP) evoked in the extensor motoneurons by activation of the most excitable fibers of the nerve supplying the antagonist muscle increased several-fold using conditioning stimulation of the forelimb nerve. Facilitation of 1a IPSP occurred on a background of IPSP evoked by descending inter-limb discharges. Facilitation of 1a IPSP had a latent period of 18–20 msec and could last to 60 msec. The possible role of inhibitory 1a interneurons in interlimb coordination is discussed. Translated from Rossiiskii Fiziologicheskii Zhurnal imeni I. M. Sechenova, Vol. 85, No. 3, pp. 430–435, March, 1999.     

Withdrawal reflex organisation to electrical stimulation of the dorsal foot in humans

The present study investigated excitatory reflex receptive fields for various muscle reflex responses and reflex mediated ankle joint movements using randomised electrical stimulation of the dorsal and plantar surface of the foot in 12 healthy subjects. Eleven electrodes (0.5-cm2 cathodes) were mounted on the dorsal side and three on the plantar side of the foot. A low (1.5 times pain threshold) and a high (2.3 times pain threshold) stimulus intensity were used to elicit the reflexes. EMG signals were recorded from tibialis anterior (TA), gastrocnemius medialis (GM), soleus (SO), biceps femoris (BF), and rectus femoris (RF) muscles together with the ankle movement measured by a goniometer. The withdrawal pattern evoked from the dorsal side consisted of two separate responses with different receptive fields: (1) early EMG responses in GM and BF (50-120 ms) evoking knee flexion, probably of purely spinal origin, and (2) a late response in GM and SO (120-200 ms) that may be under supraspinal control. The ankle flexor TA was significantly activated in both time windows, but in 11 of 12 subjects its contraction was too small to cause significant dorsal flexion. In the ankle joint inversion was the most dominant movement. Stimulation of the plantar side resulted in activation of TA when stimulating the forefoot and in activation of triceps surae when stimulating the heel. These observations show that painful stimuli activate appropriate muscles depending on stimulus location to initiate the adequate withdrawal. For proximal muscles (e.g. knee flexors) the receptive field covers almost the entire foot (dorsal and plantar sides) while more distal muscles have a smaller receptive field covering only a part of the foot. This adequate withdrawal movement suggests a more refined withdrawal reflex organisation than a stereotyped flexion of all joints to avoid tissue damage.     

Effects of changes in hip joint angle on H-reflex excitability in humans

We examined the amplitude modulation of the soleus (Sol) H-reflex during controlled variations of the hip joint angle in 21 healthy adult human subjects. Hip angle variations were imposed separately, or in combination either with stimulation of the plantar skin or with electrical activation of muscle afferents from the medial gastrocnemius (MG) nerve. We found that with subjects in the supine position, flexion of the hip significantly depressed Sol H-reflex excitability, by as much as 50% of control reflex values (Ho) recorded at 10 degrees of hip flexion. Conversely, significant facilitation of the H-reflex was observed when the hip joint was extended (10 degrees), with amplitudes reaching 200+/-15.3% of Ho. Changes in H-reflex amplitude were also observed during electrical stimulation of either the foot sole or the MG nerve, when stimuli were delivered at different hip angles. Foot sole stimulation resulted in facilitation of the H-reflex with the hip extended while depression of the reflex was recorded with the hip flexed. In contrast, MG nerve stimulation at group-I muscle afferent strength resulted in a significant increase in the Sol H-reflex magnitude with the hip flexed, while during hip extension, no significant effects were observed [corrected]. This study provides evidence for the existence of a spinal mechanism, determined principally by the hip joint angle, which promotes switching between inhibitory and facilitatory pathways during hip flexion and extension. The origins of such a spinal mechanism are discussed.     

Electromyographic behaviour of the gastrocnemius muscle during knee extension and flexion performed on the leg press.

The gastrocnemius was analysed in 10 male volunteers during knee flexion and extension with the foot in normal, plantar flexion and dorsal flexion positions. Hewlett-Packard surface electrodes, an electromyographic signal amplifier, a computer equipped with an A/D conversion plaque (Model CAD 10/26), a software specially designed to record and analyse the signals, a horizontal leg press, and electrogoniometers were used. The gastrocnemius muscle showed strong potentials at the end of knee extension and beginning of knee flexion. The muscle presented a similar activity both in the upper and lower platforms. As to bilateral action, the right gastrocnemius presented stronger potentials on the upper platforms, whereas the potentials were bilaterally similar on the lower platforms. As for foot position, the gastrocnemius presented strong potentials when the foot was in plantar flexion. The remaining positions had no effect on the work of the muscle.     

Stretch reflex modulation during imposed static and dynamic hip movements in standing humans

The purpose of this study was to investigate the effects of hip proprioceptors on soleus stretch reflex excitability in standing humans. A custom-made device to stretch the ankle extensors was mounted on the lower leg portion of a gait orthosis and was used to elicit stretch reflex responses while standing. Six subjects with motor complete spinal cord injury (SCI) and six spinal intact subjects were placed in the orthosis, and stretch reflex responses were elicited when static and/or dynamic hip joint angle changes were imposed. We found that static hip extension significantly enhanced the stretch reflex responses as compared to the neutral position and the hip flexion position only in the SCI group. The EMG magnitude induced by hip extension was 142 ± 16.6% greater than that induced by the neutral position. When the leg was dynamically swung, the reflex responses also changed with the phase of the hip angle in the SCI group; in particular, the reflex amplitude was enhanced with hip extension and in the transition phase from flexion to extension. Although the magnitude of the changes was less than that in the SCI group, a similar type of modulation was found in the normal group. Given the fact that the persons with SCI had lost the neural connection between higher nervous center and the paralyzed lower limb muscles, the mechanism underlying the present results can be attributed to the peripheral afferent input due to the hip angle changes. We concluded that hip mediated afferent input has a significant influence on the excitability modulation of the soleus stretch reflex pathway. Such neural modulation may play a role in the mechanism responsible for the phase-dependent modulation of the stretch reflex while walking.     

Acute inflammation of the knee joint in the cat alters responses of flexor motoneurons to leg movements

1. This is a report of changes in reflex excitability of flexor motoneurons in response to innocuous mechanical stimuli following initiation of an acute experimental inflammation of the knee joint in the chloralose-anaesthetized cat spinalized at level T12. 2. Activity of functionally single alpha- or gamma- motoneurons, identified by ventral root stimulation, was recorded in filaments of the nerve to biceps semitendinosus before and after onset of an acute inflammation. The inflammation was evoked by injection into the knee cavity of the compounds kaolin and carrageenan. 3. Measurements were made of resting activity, responses to local pressure applied to parts of the upper or lower limb, and to flexion or extension movements at the knee joint before and after onset of the inflammation. In two experiments, one in which no inflammation was initiated, and another in which recordings were made only after the inflammation had fully developed, a survey of large numbers of neurons (44 alpha and 84 gamma) showed that under control conditions only 14% of alpha-motoneurons showed a response to mechanical stimulation compared with 41% of gamma-motoneurons. In the presence of an inflammation 41% of alpha-cells were responsive compared with 64% of gamma-cells. 4. Inflammation-induced changes in activity of motoneurons including both excitatory and inhibitory effects, took 1-2 h to fully develop. Excitatory effects included a rise in resting discharge, an increase in the response to local pressure, and an increased response to flexion and/or to extension of the knee. Inhibitory effects included falls in resting discharge and/or in the responses to leg movements. 5. For 35 cells identified as alpha-motoneurons and tested both before and after initiation of an inflammation, 20 remained unresponsive throughout the experiment, whereas 4 that had not responded during the control period began after inflammation to respond to local pressure and/or flexion/extension movements. Of 11 units that exhibited some response before inflammation, 8 showed an increase with inflammation, whereas 2 became unresponsive. 6. The inflammation had rather more dramatic effects on gamma-neurons. Many showed control responses to leg movements, and these were measurably modified by inflammation. Of 56 gamma-cells tested under control conditions, 26 were unresponsive to all stimuli, whereas 30 showed some form of response including activity during flexion and extension movements of the leg. Where responses were tested both before and after onset of an inflammation, 11 of 13 unresponsive units remained unresponsive.(ABSTRACT TRUNCATED AT 400 WORDS)     

Windup of flexion reflexes in chronic human spinal cord injury: a marker for neuronal plateau potentials?

The physiological basis of flexion spasms in individuals after spinal cord injury (SCI) may involve alterations in the properties of spinal neurons in the flexion reflex pathways. We hypothesize that these changes would be manifested as progressive increases in reflex response with repetitive stimulus application (i.e., "windup") of the flexion reflexes. We investigated the windup of flexion reflex responses in 12 individuals with complete chronic SCI. Flexion reflexes were triggered using trains of electrical stimulation of plantar skin at variable intensities and inter-stimulus intervals. For threshold and suprathreshold stimulation, windup of both peak ankle and hip flexion torques and of integrated tibialis anterior electromyographic activity was observed consistently in all patients at inter-stimulus intervals < or =3 s. For subthreshold stimuli, facilitation of reflexes occurred only at intervals < or =1 s. Similarly, the latency of flexion reflexes decreased significantly at intervals < or =1 s. Patients that were receiving anti-spasticity medications (e.g., baclofen) had surprisingly larger windup of reflex responses than those who did not take such medications, although this difference may be related to differences of spasm frequency between the groups of subjects. The results indicate that the increase in spinal neuronal excitability following a train of electrical stimuli lasts for < or =3 s, similar to previous studies of nociceptive processing. Such long-lasting increases in flexion reflex responses suggest that cellular mechanisms such as plateau potentials in spinal motoneurons, interneurons, or both, may partially mediate spinal cord hyperexcitability in the absence of descending modulatory input.     

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.     

Phase-dependent reversal of reflexly induced movements during human gait

Summary To investigate whether phase-dependent reversals in reflex responses on electromyography (EMG) are accompanied by movement reversals, a series of human volunteers were studied for their behavioural responses to sural nerve stimulation during running or walking on a treadmill. Low-intensity stimulation (< 2.5 x perception threshold, T) of the sural nerve yielded facilitatory responses in the tibialis anterior muscle (TA), correlated with an induced ankle dorsiflexion (mean maximum 4°) in early swing. The same stimuli yielded primarily TA suppression and weak ankle plantar flexion (mean maximum 1°) at end swing. The correlated induced knee angle changes did not precede the ankle changes, and they were relatively small. Mean maximum flexion in early swing was 6.2°, while mean maximum extension was 3.7°. High-intensity stimulation of the sural nerve (> 2.5 x T) always gave rise to suppression of the ongoing activity. This resulted in a second type of movement reversal. During late stance and early swing the responses in TA were suppressive (i.e. below background activity) and related to ankle plantar flexion. In contrast, the responses during early and middle stance consisted of suppression in extensor activity (gastrocnemius medialis and soleus) and ankle dorsiflexion.The data are discussed in terms of a new hypothesis, which states that the responses to electrical stimulation of cutaneous nerves during locomotion do not correspond directly to corrections for stumbling following mechanical perturbations during the step cycle. Instead, the data invite a reinterpretation in terms of the opening and closing of reflex pathways, presumably by a central pattern generator for locomotion.