Contribution of afferent feedback to the soleus muscle activity during human locomotion

During the stance phase of the human step cycle, the ankle undergoes a natural dorsiflexion that stretches the soleus muscle. The afferent feedback resulting from this stretch enhances the locomotor drive. In this study a robotic actuator was used to slightly enhance or reduce the natural ankle dorsiflexion, in essence, mimicking the small variations in the ankle dorsiflexion movement that take place during the stance phase of the step cycle. The soleus (SOL) and tibialis anterior EMG were analyzed in response to the ankle trajectory modifications. The dorsiflexion enhancements and reductions generated gradual increments and decrements, respectively, in the ongoing SOL EMG. We exercised care to ensure that the imposed ankle movements were too slow to elicit distinct burst-like stretch reflex responses that have been investigated previously. The increased SOL EMG after the dorsiflexion enhancements was reduced when the group Ia afferents were blocked with peripheral ischemia at the thigh, and during high-frequency Achilles tendon vibration. However, neither ischemia nor tendon vibration affected the decrements in the SOL EMG during the dorsiflexion reductions. These findings give evidence of the contribution of afferent feedback to the SOL activity in an ongoing basis during the stance phase. The results suggest that mainly feedback from the group Ia pathways is responsible for the increments in the SOL EMG during the dorsiflexion enhancements. However, the decrements in the SOL activity might be mediated by different afferent mechanisms.     

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.     

Locomotor changes in length and EMG activity of feline medial gastrocnemius muscle following paralysis of two synergists

The mechanism of the compensatory increase in electromyographic activity (EMG) of a cat ankle extensor during walking shortly after paralysis of its synergists is not fully understood. It is possible that due to greater ankle flexion in stance in this situation, muscle spindles are stretched to a greater extent and, thus, contribute to the EMG enhancement. However, also changes in force feedback and central drive may play a role. The aim of the present study was to investigate the short-term (1- to 2-week post-op) effects of lateral gastrocnemius (LG) and soleus (SO) denervation on muscle fascicle and muscle–tendon unit (MTU) length changes, as well as EMG activity of the intact medial gastrocnemius (MG) muscle in stance during overground walking on level (0%), downslope (−50%, presumably enhancing stretch of ankle extensors in stance) and upslope (+50%, enhancing load on ankle extensors) surfaces. Fascicle length was measured directly using sonomicrometry, and MTU length was calculated from joint kinematics. For each slope condition, LG-SO denervation resulted in an increase in MTU stretch and peak stretch velocity of the intact MG in early stance. MG muscle fascicle stretch and peak stretch velocity were also higher than before denervation in downslope walking. Denervation significantly decreased the magnitude of MG fascicle shortening and peak shortening velocity during early stance in level and upslope walking. MG EMG magnitude in the swing and stance phases was substantially greater after denervation, with a relatively greater increase during stance of level and upslope walking. These results suggest that the fascicle length patterns of MG muscle are significantly altered when two of its synergists are in a state of paralysis. Further, the compensatory increase in MG EMG is likely mediated by enhanced MG length feedback during downslope walking, enhanced feedback from load-sensitive receptors during upslope walking and enhanced central drive in all walking conditions.     

Afferent-mediated modulation of the soleus muscle activity during the stance phase of human walking

The aim of this study was to investigate the contribution of proprioceptive feedback to the amplitude modulation of the soleus muscle activity during human walking. We have previously shown that slow-velocity, small-amplitude ankle dorsiflexion enhancements and reductions applied during the stance phase of the step cycle generate, respectively, increments and decrements on the ongoing soleus activity. We have also shown that the increments in soleus activity are at least partially mediated by feedback from group Ia fibres. In the present study, we further investigated the afferent-mediated contribution from muscle group II afferents, cutaneous and proprioceptive afferents from the foot, and load-sensitive afferents to the soleus EMG. Slow-velocity, small-amplitude ankle trajectory modifications were combined with the pharmaceutical depression of group II polysynaptic pathways with tizanidine hydrochloride, anaesthetic blocking of sensory information from the foot with injections of lidocaine hydrochloride, and modulation of load feedback by increasing and decreasing the body load. The depression of the group II afferents significantly reduced the soleus response to the ankle trajectory modifications. Blocking sensory feedback from the foot did not have an effect on the soleus muscle activity. Changes in body load affected the ongoing soleus activity level; however, it did not affect the amplitude of the soleus EMG responses to the ankle trajectory modifications. These results suggest that the feedback from group II afferents, and possibly from load-sensitive afferents, contribute to the amplitude modulation of the soleus muscle activity during the stance phase of the step cycle. However, feedback from cutaneous afferents and instrinsic proprioceptive afferents from the foot does not seem to contribute to this muscle activation.     

Lack of on-going adaptations in the soleus muscle activity during walking in patients affected by large-fiber neuropathy

The aim of this study was to investigate the contribution of feedback from large-diameter sensory fibers to the adaptation of soleus muscle activity after small ankle trajectory modifications during human walking. Small-amplitude and slow-velocity ankle dorsiflexion enhancements and reductions were applied during the stance phase of the gait cycle to mimic the normal variability of the ankle trajectory during walking. Patients with demyelination of large sensory fibers (Charcot-Marie-Tooth type 1A and antibodies to myelin-associated glycoprotein neuropathy) and age-matched controls participated in this study. The patients had absent light-touch sense in the toes and feet and absent quadriceps and Achilles tendon reflexes, indicating functional loss of large sensory fibers. Moreover, their soleus stretch reflex response consisted of a single electromyographic (EMG) burst with delayed onset and longer duration (P < 0.01) than the short- and medium-latency reflex responses observed in healthy subjects. In healthy subjects, the soleus EMG gradually increased or decreased when the ankle dorsiflexion was, respectively, enhanced or reduced. In the patients, the soleus EMG increased during the dorsiflexion enhancements; however, the velocity sensitivity of this response was decreased compared with the healthy volunteers. When the dorsiflexion was reduced, the soleus EMG was unchanged. These results indicate that the enhancement of the soleus EMG is mainly sensitive to feedback from primary and secondary muscle spindle afferents and that the reduction may be mediated by feedback from the group Ib pathways. This study provides evidence for the role of sensory feedback in the continuous adaptation of the soleus activity during the stance phase of human walking.     

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.     

The rat lumbosacral spinal cord adapts to robotic loading applied during stance

Load-related afferent information modifies the magnitude and timing of hindlimb muscle activity during stepping in decerebrate animals and spinal cord-injured humans and animals, suggesting that the spinal cord mediates load-related locomotor responses. In this study, we found that stepping on a treadmill by adult rats that received complete, midthoracic spinal cord transections as neonates could be altered by loading the hindlimbs using a pair of small robotic arms. The robotic arms applied a downward force to the lower shanks of the hindlimbs during the stance phase and measured the position of the lower shank during stepping. No external force was applied during the swing phase of the step. When applied bilaterally, this stance force field perturbed the hindlimb trajectories so that the ankle position was shifted downward during stance. In response to this perturbation, both the stance and step cycle durations decreased. During swing, the hindlimb initially accelerated toward the normal, unperturbed swing trajectory and then tracked the normal trajectory. Bilateral loading increased the magnitude of the medial gastrocnemius electromyographic (EMG) burst during stance and increased the amplitude of the semitendinosus and rectus femoris EMG bursts. When the force field was applied unilaterally, stance duration decreased in the loaded hindlimb, while swing duration was decreased in the contralateral hindlimb, thereby preserving interlimb coordination. These results demonstrate the feasibility of using robotic devices to mechanically modulate afferent input to the injured spinal cord during weight-supported locomotion. In addition, these results indicate that the lumbosacral spinal cord responds to load-related input applied to the lower shank during stance by modifying step timing and muscle activation patterns, while preserving normal swing kinematics and interlimb coordination.     

Adaptation of the walking pattern to uphill walking in normal and spinal-cord injured subjects

 Lower-limb movements and muscle-activity patterns were assessed from seven normal and seven ambulatory subjects with incomplete spinal-cord injury (SCI) during level and uphill treadmill walking (5, 10 and 15°). Increasing the treadmill grade from 0° to 15° induced an increasingly flexed posture of the hip, knee and ankle during initial contact in all normal subjects, resulting in a larger excursion throughout stance. This adaptation process actually began in mid-swing with a graded increase in hip flexion and ankle dorsiflexion as well as a gradual decrease in knee extension. In SCI subjects, a similar trend was found at the hip joint for both swing and stance phases, whereas the knee angle showed very limited changes and the ankle angle showed large variations with grade throughout the walking cycle. A distinct coordination pattern between the hip and knee was observed in normal subjects, but not in SCI subjects during level walking. The same coordination pattern was preserved in all normal subjects and in five of seven SCI subjects during uphill walking. The duration of electromyographic (EMG) activity of thigh muscles was progressively increased during uphill walking, whereas no significant changes occurred in leg muscles. In SCI subjects, EMG durations of both thigh and leg muscles, which were already active throughout stance during level walking, were not significantly affected by uphill walking. The peak amplitude of EMG activity of the vastus lateralis, medial hamstrings, soleus, medial gastrocnemius and tibialis anterior was progressively increased during uphill walking in normal subjects. In SCI subjects, the peak amplitude of EMG activity of the medial hamstrings was adapted in a similar fashion, whereas the vastus lateralis, soleus and medial gastrocnemius showed very limited adaptation during uphill walking. We conclude that SCI subjects can adapt to uphill treadmill walking within certain limits, but they use different strategies to adapt to the changing locomotor demands. Received: 10 March 1998 / Accepted: 29 December 1998     

Loading during the stance phase of walking in humans increases the extensor EMG amplitude but does not change the duration of the step cycle

 Prior work from mammals suggests that load experienced by extensor muscles of the hindlimbs (i.e. Duysens and Pearson 1980; Pearson and Collins 1993; Fouad and Pearson 1997) or cutaneous afferents from the plantar surface of the foot (Duysens and Pearson 1976; Guertin et al. 1995) enhances activity in extensor muscles during the stance phase, and delays the onset of flexor activity associated with the swing phase. The presumed functional significance of this phenomenon is that extensor activity of the supporting limb during walking can: (a) reinforce the supporting function in proportion to the load experienced, and (b) prolong the stance phase until unloading of the limb has occurred. Whether a similar functional role exists for load-sensitive afferents during walking in the human is unknown. In this study, the effect of adding or removing a substantial load (30% of body weight) at the centre of mass was studied in healthy adult human subjects. Loads were applied near the centre of mass to avoid the need for postural adjustments which might confound the interpretation of the results. Subjects walked on a treadmill with either: (a) a sustained increase or decrease in load, or (b) a sudden unexpected increase or decrease in load. In general, subjects responded to the changes in load by changing the amplitude of the extensor electromyographic (EMG) bursts. For example, with sudden unexpected additions in load, the average increase in amplitude was 40% for the soleus across the stance phase, and 134% for the quadriceps during the early part of the stance phase. Extensor EMGs increased with both sustained and sudden increases in load. Extensor EMG durations also increased (average increase in duration of 4% for soleus with sudden loading, and 7% for sustained loading). Cycle duration hardly changed (average increase of 0.5% with both sudden and sustained loading). These results differ from those of infants subjected to a similar perturbation during supported walking. A large change in timing (i.e. an increase in the duration of the stance phase by 30% and the step cycle by 28%) was seen in the infants, with no change in the amplitude of the EMG burst (Yang et al. 1998). These results suggest that the central nervous system can control the timing and amplitude of extensor EMG activity in response to loading independently. Maturation of the two components most likely occurs independently. In the adult, independent control of the two components may provide greater flexibility of the response. Received: 28 April 1998 / Accepted: 3 September 1998     

Modulation of stretch reflexes during imposed walking movements of the human ankle.

Our overall objectives were to examine the role of peripheral afferents from the ankle in modulating stretch reflexes during imposed walking movements and to assess the mechanical consequences of this reflex activity. Specifically we sought to define the changes in the electromyographic (EMG) and mechanical responses to a stretch as a function of the phase of the step cycle. We recorded the ankle position of a normal subject walking on a treadmill at 3 km/h and used a hydraulic actuator to impose the same movements on supine subjects generating a constant level of ankle torque. Small pulse displacements, superimposed on the simulated walking movement, evoked stretch reflexes at different phases of the cycle. Three major findings resulted: 1) soleus reflex EMG responses were influenced strongly by imposed walking movements. The response amplitude was substantially smaller than that observed during steady-state conditions and was modulated throughout the step cycle. This modulation was qualitatively similar to that observed during active walking. Because central factors were held constant during the imposed walking experiments, we conclude that peripheral mechanisms were capable of both reducing the amplitude of the reflex EMG and producing its modulation throughout the movement. 2) Pulse disturbances applied from early to midstance of the imposed walking cycle generated large reflex torques, suggesting that the stretch reflex could help to resist unexpected perturbations during this phase of walking. In contrast, pulses applied during late stance and swing phase generated little reflex torque. 3) Reflex EMG and reflex torque were modulated differently throughout the imposed walking cycle. In fact, at the time when the reflex EMG response was largest, the corresponding reflex torque was negligible. Thus movement not only changes the reflex EMG but greatly modifies the mechanical output that results.     

Flexion reflex modulation during stepping in human spinal cord injury

The flexion reflex modulation pattern was investigated in nine people with a chronic spinal cord injury during stepping using body weight support on a treadmill and manual assistance by therapists. Body weight support was provided by an upper body harness and was adjusted for each subject to promote the best stepping pattern with the least manual assistance required by the therapists. The flexion reflex was elicited by sural nerve stimulation with a 30 ms pulse train at 1.2–2 times the tibialis anterior reflex threshold. During stepping, stimuli were randomly dispersed across the gait cycle which was divided into 16 equal bins. A long latency (>110 ms) flexion reflex was present in all subjects, while a short (>30 ms) and a medium latency (>70 ms) flexion reflex were present only in three subjects. For each response, the non-stimulated EMG was subtracted from the stimulated EMG at identical time windows and bins, normalized to the maximal corresponding EMG, and significant differences were established with a Wilcoxon rank-sum test. The long latency flexion reflex was facilitated at late stance and during the swing-to-stance transition phase. A reflex depression was present from heel strike until mid-stance and during the swing-to-stance transition phase. The short and medium latency flexion reflexes were depressed during mid-stance followed by facilitation during the stance-to-swing transition phase. Regardless of the latency, facilitatory flexion responses during the swing phase coincided with decreased activity of ipsilateral ankle extensors. The flexion reflex was modulated in a phase dependent manner, a behavior that was absent for the soleus H-reflex in most of these patients (Knikou et al. in Exp Brain Res 193:397–407, 2009). We propose that training should selectively target spinal reflex circuits in which extensor muscles and reflexes are involved in order to maximize sensorimotor recovery in these patients.     

Segmental reflexes and ankle joint stiffness during co-contraction of antagonistic ankle muscles in man

The size of soleus H-reflexes and short-latency stretch reflexes was measured at different levels of plantar flexion or co-contraction (simultaneous activation of dorsi- and plantar flexors) in seven healthy subjects. In four of seven subjects the short-latency stretc reflex was smaller during weak co-contraction than during isolated plantar flexion at matched background electromyogram (EMG) levels in the soleus muscle. In three of these four subjects the stretch reflex was larger during strong co-contraction than during plantar flexion, whereas it had the same size during the two tasks in the last subject. In the remaining subjects the stretch reflex either had the same size or was larger at all levels of co-contraction than at similar levels of plantar flexion. In contrast, the H-reflex was found to decrease with co-contraction at all contraction levels in all subjects. The decrease in the reflexes during weak co-contraction might be caused by presynaptic inhibition of Ia afferents. It is unclear why only the H-reflex decreased during strong co-contraction. The stiffness of the ankle joint was measured from the torque increment following the stretch of the plantar flexors divided by the stretch amplitude. In all subjects the total stiffness of the ankle joint was larger during strong co-contraction than during plantar flexion of similar strength. The stiffness was smaller during weak co-contraction than during weak plantar flexion in three out of seven subjects. The medial gastrocnemius muscle was more active at a given level of soleus activity during the co-contraction task than during the isolated plantar flexion task. It is suggested that the increase in the stiffness during co-contraction as compared to isolated plantar flexion was mainly due to the mechanical contribution of the activity in the tibialis anterior and medial gastrocnemius muscles. The decrease in stiffness during weak co-contraction was, in contrast, most likely mainly caused by modulation of reflex stiffness.     

Interindividual differences in H reflex modulation during normal walking

Based on previous studies, at least two different types of soleus Hoffmann (H) reflex modulation were likely to be found during normal human walking. Accordingly, the aim of the present study was to identify different patterns of modulation of the soleus H reflex and to examine whether or not subjects with different H reflex modulation would exhibit different walking mechanics and different EMG activity. Fifteen subjects walked across two force platforms at 4.5 km/h (+/-10%) while the movements were recorded on video. The soleus H reflex and EMG activity were recorded separately during treadmill walking at 4.5 km/h. Using a two-dimensional analysis joint angles, angular velocities, accelerations, linear velocities and accelerations were calculated, and net joint moments about the ankle, knee and hip joint were computed by inverse dynamics from the video and force plate data. Six subjects (group S) showed a suppressed H reflex during the swing phase, and 9 subjects (group LS) showed increasing reflex excitability during the swing phase. The plantar flexor dominated moment about the ankle joint was greater for group LS. In contrast, the extensor dominated moment about the knee joint was greater for the S group. The hip joint moment was similar for the groups. The EMG activity in the vastus lateralis and anterior tibial muscles was greater prior to heel strike for the S group. These data indicate that human walking exhibits at least two different motor patterns as evaluated by gating of afferent input to the spinal cord, by EMG activity and by walking mechanics. Increasing H reflex excitability during the swing phase appears to protect the subject against unexpected perturbations around heel strike by a facilitated stretch reflex in the triceps surae muscle. Alternatively, in subjects with a suppressed H reflex in the swing phase the knee joint extensors seem to form the primary protection around heel strike.     

Excitability of the soleus H reflex during graded walking in humans

The excitability of the soleus Hoffmann (H) reflex was measured in five healthy male subjects during graded treadmill walking. Uphill and downhill walking at an 8% grade as well as level walking were used to vary the demands for lengthening and shortening contractions of the soleus muscle. These changes were assumed to cause differences in control of the afferent input in the spinal cord and the voluntary output to the soleus muscle. The H reflex was strongly modulated in all three walking conditions, high during the stance phase and low or absent during the swing phase. The shape of the modulations was, however, different. At uphill walking the reflex increased gradually during the whole stance phase and seemed to follow the soleus electromyogram (EMG) pattern closely. In the downhill condition the reflex excitability increased rapidly at heel strike like the soleus EMG and co-contraction of the anterior tibial muscle was observed. At level walking a fast rise in reflex excitability was seen just after heel strike with low or absent soleus EMG. Mean soleus EMG was lower during downhill than during uphill or level walking, but the mean H reflex amplitude was similar in all three conditions. However, when the H reflex was related directly to the EMG activity by linear regression the reflex gain was lower during uphill walking than in the two other conditions. Furthermore, the ratio between H reflex and EMG amplitude was high during the first half of the stance phase at level walking indicating an elevated reflex excitability independent of the voluntary motor output. It is therefore concluded that the modulation of reflexes during walking cannot be interpreted in terms of the idea of automatic gain compensation. The reflexes must be controlled specifically and independently during the different phases of the motor output to meet the mechanical requirements of the movement task. Most explicitly this was seen during downhill walking, where an elevated reflex excitability together with co-contraction at the ankle joint seem to provide increased joint stiffness and security, when the kinetic energy of the body has to be brought under control at heel strike.     

On the origin of the soleus H-reflex modulation pattern during human walking and its task-dependent differences

Recently, Brooke and colleagues have suggested "that the strong inhibition arising from passive movement about the knee and hip joints, lays down the base for the soleus H-reflex gain modulation seen during human gait." In particular stretch-evoked afferent activity from the quadriceps muscle was emphasized as the most important source of movement-induced inhibition of the H-reflex. To test this hypothesis we examined the kinematics and electromyographic (EMG) activity of the leg during human walking and correlated these with the modulation pattern of the soleus H-reflex. To further test the possible contribution of stretch-evoked quadriceps afferent activity to the soleus H-reflex modulation pattern during walking different walking gaits were studied. In one condition subjects were asked to walk with their knee locked in full extension by a rigid knee brace. In a second condition subjects were asked to walk backwards. During normal walking, the soleus H-reflex modulation pattern is strongly correlated with the EMG events of the soleus and tibialis anterior (TA), but not with hip, knee, or ankle angular displacement or velocity. When subjects walked with the knee locked in full extension, the amplitude of the H-reflex, its modulation pattern, and the task-dependent changes of its amplitude were the same as during normal walking. During backward walking, the H-reflex increases in late swing before activity of the soleus has begun and while the knee is flexing, an observation that highlights central control of the H-reflex amplitude. The effects of imposed flexion of the knee in passive subjects were also reexamined. The knee flexion imposed by the experimenter followed the same trajectory as that which occurred during the swing phase of the subject's step cycle. It was found that imposed knee flexions elicited a burst of TA EMG activity with an average latency of 81.6 ms (SD = 21 ms) in six out of eight subjects. Inhibition of the H-reflex, when it occurred, was associated with the occurrence of this burst. When subjects voluntarily flexed their right knee from an initial quiet standing posture, the inhibition of the soleus H-reflex began before flexion of the knee or that of any other leg segment. Once again the onset of inhibition was closely associated with the onset of activity in the TA. In the discussion section the present observations are examined in light of the predictions made by the movement-induced inhibition hypothesis of Brooke et al. It will be concluded that none of the predictions of this hypothesis were corroborated by present tests done during human walking. In consequence, we suggest that the modulation pattern of the H-reflex observed during normal human walking is centrally determined, as are the task-dependent differences of its amplitude (e.g., standing versus the stance phase of human walking).     

Post-effect of forward and backward locomotion on body orientation in space during quiet stance

Neural circuits responsible for stance control serve other motor tasks as well. We investigated the effect of prior locomotor tasks on stance, hypothesizing that postural post-effects of walking are dependent on walking direction. Subjects walked forward (WF) and backward (WB) on a treadmill. Prior to and after walking they maintained quiet stance. Ground reaction forces and centre of foot pressure (CoP), ankle and hip angles, and trunk inclination were measured during locomotion and stance. In WF compared to WB, joint angle changes were reversed, trunk was more flexed, and movement of CoP along the foot sole during the support phase of walking was opposite. During subsequent standing tasks, WB induced ankle extension, hip flexion, trunk backward leaning; WF induced ankle flexion and hip extension. The body CoP was displaced backward post-WB and forward post-WF. The post-effects are walking-direction dependent, and possibly related to foot-sole stimulation pattern and trunk inclination during walking.     

Monosynaptic and dorsal root reflexes during locomotion in normal and thalamic cats

1. In normal and thalamic walking cats electrical stimulation of muscle nerves via chronically implanted electrodes produced electromyographic (EMG) and neurographic responses that were modulated in amplitude depending on the phase of the step cycle. These responses were examined for possible indications of effects of primary afferent depolarization (PAD) during stepping. 2. Monosynaptic reflexes (MSRs) produced by stimulating the lateral gastrocnemius (LG) and medial gastrocnemius (MG) nerves were recorded as EMGs in MG or LG muscles during treadmill locomotion in normal cats. These heteronymous MSR responses were greatest during the stance (extensor) phase. 3. In the same animals, after decerebration, similar modulation of the heteronymous ankle extensor MSRs occurred during spontaneous locomotion with the use of the same stimulus and recording sites. 4. In both normal and thalamic cats the amplitude of neurogram responses recorded from LG or MG nerve after stimulation of the other muscle nerve varied with phase of stepping but did not parallel the variations of the MSR measured as EMG amplitude in the same muscle. The nerve responses were largest during the flexion phase of the step cycle and had a calculated central latency of 0.6-1.0 ms. These are interpreted as arising from antidromic activity in large-caliber afferent nerve fibers (i.e., dorsal root reflexes). 5. Spontaneous antidromic activity in severed L7 dorsal rootlet fibers to triceps surae was observed in the thalamic cats during episodes of locomotion and was closely correlated with flexion phase EMG activity in semitendinosus, a bifunctional muscle. 6. In decerebrate cats, dorsal root reflexes (DRRs) in severed filaments of L4-L7 dorsal roots were produced by stimulation of saphenous and posterior tibial nerves. These DRRs were always smaller during locomotion than during rest and were smallest during the flexion phase. 7. The short-latency antidromic activity produced in muscle nerves by stimulating heteronymous muscle nerves thus appears to be a DRR produced in Group I terminal arborizations that are depolarized close to threshold during the flexion phase. Such PAD could account for changes in the MSR that do not always parallel the levels of recruitment of the motor pools as manifest by background EMG amplitude.     

Short-term motor compensations to denervation of feline soleus and lateral gastrocnemius result in preservation of ankle mechanical output during locomotion

Denervation of selected ankle extensors in animals results in locomotor changes. These changes have been suggested to permit preservation of global kinematic characteristics of the hindlimb during stance. The peak ankle joint moment is also preserved immediately after denervation of several ankle extensors in the cat, suggesting that the animal's response to peripheral nerve injury may also be aimed at preserving ankle mechanical output. We tested this hypothesis by comparing joint moments and power patterns during walking before and after denervation of soleus and lateral gastrocnemius muscles. Hindlimb kinematics, ground reaction forces and electromyographic activity of selected muscles were recorded during level, downslope (-50%) and upslope (50%) walking before and 1-3 weeks after nerve denervation. Denervation resulted in increased activity of the intact medial gastrocnemius and plantaris muscles, greater ankle dorsiflexion, smaller knee flexion, and the preservation of the peak ankle moment during stance. Surprisingly, ankle positive power generated in the propulsion phase of stance was increased (up to 50%) after denervation in all walking conditions (p < 0.05). The obtained results suggest that the short-term motor compensation to denervation of lateral gastrocnemius and soleus muscles may allow for preservation of mechanical output at the ankle. The additional mechanical energy generated at the ankle during propulsion can result, in part, from increased activity of intact synergists, the use of passive tissues around the ankle and by the tendon action of ankle two-joint muscles and crural fascia.     

Early corrective reactions of the leg to perturbations at the torso during walking in humans

The contribution of afferent feedback to the regulation of locomotion in humans is not well understood. Animal experiments have suggested that loading of the leg during the stance phase may enhance the magnitude of extensor burst activity and delay the onset of swing phase. The aim of the present study was to determine whether transient loading of the leg at the end of stance would enhance extensor-muscle activity and delay the onset of swing in walking humans. To test this hypothesis, we applied loads to the hips of subjects so that the load was applied along the long axis of the leg at the end of stance (down-back unsupported, DBU). This resulted in an unexpectedly complex reaction characterised by rapid co-contraction of antagonist pairs of muscles around the ankle and knee and a prolongation of the stance phase. We speculated that the complexity of the reaction was, in part, due to a disturbance in equilibrium. To address this possibility, two additional perturbation paradigms were tested: (1) subjects held a rail during the loading paradigm (down-back supported, DBS), or (2) subjects received only a posteriorly directed perturbation of the hips, which added no additional load to the leg (backward unsupported, BU). As predicted, the DBS perturbation resulted in an enhancement of the ongoing soleus-muscle activity, and the unexpected tibialis anterior burst that was observed during the DBU paradigm was absent. Allowing the subjects to hold a rail substantially reduced the change in the timing of the step cycle observed in the DBU paradigm. The BU perturbation prolonged the stance phase duration and, as expected, resulted in a burst of activity in tibialis activity. This was usually accompanied by a reduction in the ongoing soleus activity. Two important conclusions are drawn from the present study. First, loading of the leg at the end of stance phase enhances the ongoing extensor-muscle activity. We suggest that afferent feedback responding to the increase load supported by the leg leads to rapid enhancement of the active extensor muscles to compensate for the increased load and prevent collapse of the leg. Interestingly, the duration of the stance phase was only marginally increased when loading was applied without a postural disturbance (DBS). Second, posterior perturbation of the centre of mass at the end of stance phase evokes an "automatic postural response" in tibialis anterior. Of particular interest, this evoked postural response can occur simultaneously with an enhanced activation of soleus. This indicates that the DBU perturbation employed in this study elicited two responses, one to prevent the collapse of the leg and the other to stabilise the centre of mass.     

Motor compensatory reactions following a forward fall in subjects with unilateral abolition of the triceps-surae H reflex

The electromyograms of the right and left soleus and tibialis anterior muscles of 6 subjects with unilateral abolition of Achilles tendon reflex due to S1 radiculitis were recorded during a forward fall involving stepping to recover balance. Each subject took part in two series of experiments, one in which the step was performed with the unaffected leg and a second in which the affected leg was used. A unilateral deficiency of peripheral proprioceptive afferents affected ankle muscles EMG activities bilaterally, except for the EMG activity of the soleus of the starting foot. The tibialis anterior of both the affected and the unaffected side, showed either a normal pattern (i.e. a phasic contraction after soleus contraction stopped), or an early contraction. On the stance side, the early contraction was associated with a depressed soleus EMG activity. Some abnormal motor patterns could be due to the ipsilateral deficiency of the Ia inhibitory projection from soleus to tibialis anterior. The presence of abnormal patterns on the unaffected side indicates that the motor activity in one lower limb can be modified by a loss of peripheral afference in the contralateral limb. This suggests that crossed pathways between lower limbs are involved in balance recovery movement.