The effects of self-reinnervation of cat medial and lateral gastrocnemius muscles on hindlimb kinematics in slope walking

The aim of this study was to investigate the effects of self-reinnervation of the medial (MG) and lateral gastrocnemius (LG) muscles on joint kinematics of the whole hindlimb during overground walking on surfaces of varying slope in the cat. Hindlimb kinematics were assessed (1) with little or no activity in MG and LG (short-term effects of self-reinnervation), and (2) after motor function of these muscles was presumably recovered but their proprioceptive feedback permanently disrupted (long-term effects of self-reinnervation). The stance phase was examined in three walking conditions: downslope (−50%, i.e. −26.6°), level (0%) and upslope (+50%, +26.6°). Measurements were performed prior to and at consecutive time points (between 1 and 57 weeks) after transecting and immediately suturing MG and LG nerves. It was found that MG-LG self-reinnervation did not significantly change hip height and hindlimb orientation in any of the three walking conditions. Substantial short-term effects were observed in the ankle joint (e.g., increased flexion in early stance) as well as in metatarsophalangeal and knee joints, leading to altered interjoint coordination. Hindlimb kinematics in level and upslope walking progressed back towards baseline within 14–19 weeks. Thus in these two conditions the cats were walking without any detectable kinematic deficits, despite the absence of length feedback from two major ankle extensors. This was verified in a decerebrate preparation for four of the five cats. In contrast, ankle joint kinematics as well as interjoint coordination in downslope walking gradually progressed towards, but never reached their baseline patterns. The short-term effects can be explained by both mechanical and neural factors that are affected by the functional elimination of MG and LG. Permanent changes in kinematics during downslope walking indicate the importance of proprioceptive feedback from the MG and LG muscles in regulating locomotor activity of ankle extensors. Full recovery of hindlimb kinematics during level and upslope walking suggests that the proprioceptive loss is compensated by other sensory sources (e.g. cutaneous receptors) or altered central drive.     

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     

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.     

Adaptive control for backward quadrupedal walking. II. Hindlimb muscle synergies

1. To compare the basic hindlimb synergies for backward (BWD) and forward (FWD) walking, electromyograms (EMG) were recorded from selected flexor and extensor muscles of the hip, knee, and ankle joints from four cats trained to perform both forms of walking at a moderate walking speed (0.6 m/s). For each muscle, EMG measurements included burst duration, burst latencies referenced to the time of paw contact or paw off, and integrated burst amplitudes. To relate patterns of muscle activity to various phases of the step cycle, EMG records were synchronized with kinematic data obtained by digitizing high-speed ciné film. 2. Hindlimb EMG data indicate that BWD walking in the cat was characterized by reciprocal flexor and extensor synergies similar to those for FWD walking, with flexors active during swing and extensors active during stance. Although the underlying synergies were similar, temporal parameters (burst latencies and durations) and amplitude levels for specific muscles were different for BWD and FWD walking. 3. For both directions, iliopsoas (IP) and semitendinosus (ST) were active as the hip and knee joints flexed at the onset of swing. For BWD walking, IP activity decreased early, and ST activity continued as the hip extended and the knee flexed. For FWD walking, in contrast, ST activity ceased early, and IP activity continued as the hip flexed and the knee extended. For both directions, tibialis anterior (TA) was active throughout swing as the ankle flexed and then extended. A second ST burst occurred at the end of swing for FWD walking as hip flexion and knee extension slowed for paw contact. 4. For both directions, knee extensor (vastus lateralis, VL) activity began at paw contact. Ankle extensor (lateral gastrocnemius, LG) activity began during midswing for BWD walking but just before paw contact for FWD walking. At the ankle joint, flexion during the E2 phase (yield) of stance was minimal or absent for BWD walking, and ankle extension during BWD stance was accompanied by a ramp increase in LG-EMG activity. At the knee joint, the yield was also small (or absent) for BWD walking, and increased VL-EMG amplitudes were associated with the increased range of knee extension for BWD stance. 5. Although the uniarticular hip extensor (anterior biceps femoris, ABF) was active during stance for both directions, the hip flexed during BWD stance and extended during FWD stance.(ABSTRACT TRUNCATED AT 400 WORDS)     

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.     

Adaptive changes in motor activity associated with functional recovery following muscle denervation in walking cats.

In this investigation we examined the changes in the pattern of activity in the medial gastrocnemius (MG) muscle in walking cats following transection of the nerves innervating synergist muscles (lateral gastrocnemius, soleus, and plantaris). Immediately following the nerve transections, there was a large increase in ankle flexion during early stance (from approximately 10 to approximately 30 degrees ) and a marked increase in the magnitude of the MG bursts during stance. We attribute this increase in the magnitude of the MG bursts to an increase in afferent feedback from the abnormally stretched MG muscle. During the week after the nerve transections, there was a progressive decrease in ankle yield. This improvement in ankle function was correlated with an increase in magnitude of two components of the MG bursts; the initial component starting during late swing and ending approximately 40 ms after ground contact, and a late component associated with stance. The time courses of the increases in the initial and late components of the MG bursts were different. Large and significant increases in the late component occurred the day after the nerve transections, whereas increases in the initial component occurred more gradually. This difference in time course was reflected in the kinematics of ankle movement. Over the first few days after the nerve transections, improvement in ankle movement occurred primarily late in the stance phase, and there was little change in ankle yield during early stance. At 1 wk, however, there was a significant reduction in ankle yield during early stance. This decreased yield was most likely due to an increase in stiffness of the MG muscle at the time of ground contact resulting from the increase in magnitude of the initial component of the MG bursts. The increases in the magnitude of the initial and late components of the MG bursts, as well as the improvement in ankle function, depended on use of the leg. All these changes were delayed by immobilizing the leg for 6 days in an extended position. We discuss possible mechanisms underlying the increase in the magnitude of the MG bursts and propose that proprioceptive signals from the stretched MG muscles provide an error signal for rescaling the magnitude of the centrally generated initial component. Our data support the concept that proprioceptive feedback functions to scale the magnitude of feed-forward motor commands to ensure they are appropriate for the biomechanical properties of the musculoskeletal system.     

How Crouch Gait Can Dynamically Induce Stiff-Knee Gait

Children with cerebral palsy frequently experience foot dragging and tripping during walking due to a lack of adequate knee flexion in swing (stiff-knee gait). Stiff-knee gait is often accompanied by an overly flexed knee during stance (crouch gait). Studies on stiff-knee gait have mostly focused on excessive knee muscle activity during (pre)swing, but the passive dynamics of the limbs may also have an important effect. To examine the effects of a crouched posture on swing knee flexion, we developed a forward-dynamic model of human walking with a passive swing knee, capable of stable cyclic walking for a range of stance knee crouch angles. As crouch angle during stance was increased, the knee naturally flexed much less during swing, resulting in a ‘stiff-knee’ gait pattern and reduced foot clearance. Reduced swing knee flexion was primarily due to altered gravitational moments around the joints during initial swing. We also considered the effects of increased push-off strength and swing hip flexion torque, which both increased swing knee flexion, but the effect of crouch angle was dominant. These findings demonstrate that decreased knee flexion during swing can occur purely as the dynamical result of crouch, rather than from altered muscle function or pathoneurological control alone.     

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.     

Adaptive changes in locomotor activity following botulinum toxin injection in ankle extensor muscles of cats.

The present study investigated the adaptations made in motor behavior following a temporary reduction in ankle extensor activity in the walking cat. Temporary muscle weakness was induced by injecting botulinum toxin into the lateral gastrocnemius (LG), plantaris (PL), and soleus (SOL) muscles, or SOL alone. The medial gastrocnemius (MG) muscle was not injected. Adaptations in the level of muscle activity were recorded using chronically implanted electromyographic (EMG) electrodes. Serial recordings were made prior to botulinum toxin injections and for several days following the injections. Kinematic analysis of ankle joint movements was made from video records to assess the impact of the botulinum toxin injections on the function of the ankle joint during walking. Following injection of the LG, PL, and SOL muscles with botulinum toxin, the amplitude of the MG burst increased over a period of a few days to a week. This increase was similar to the previously reported changes produced in MG following transection of the nerves serving LG, PL, and SOL. Following the weakening of the ankle extensor muscles, there was a temporary deficit in ankle function during walking as evidenced by a marked increase in the amount of ankle flexion that occurred at stance onset. This functional deficit recovered relatively quickly and was not associated with a return of the EMG pattern to the preinjection pattern. After recovery from the initial injections, a second injection of botulinum toxin into SOL alone was performed. No functional deficits were observed in the ankle movements during walking following this second injection. However, weakening SOL produced increases in the burst amplitudes of the MG, LG, and PL muscles over a period of a few days. This suggests that normal movements at the ankle during walking can be generated with more than one pattern of ankle extensor activity and that there is flexibility in how the necessary torque is produced. A final procedure, transection of the nerves serving LG, PL, and SOL, failed to produce any functional deficits in ankle movements. The implication is that adaptations to the neural control of ankle extensor activity that were induced by the initial procedure persisted after the recovery of the injected muscles and were sufficient to compensate for the subsequent challenges.     

A knee and ankle flexing hybrid orthosis for paraplegic ambulation

Limitations of mechanical walking orthoses for paraplegics are high energy consumption and upper limb loading. Flexing of the knee during swing phase has been used as a means of attempting to reduce these. It has been found that this has little effect because using knee flexion results in no change in the compensatory mechanisms required for swing foot clearance. This is because knee flexion can result in an increase in effective leg length, i.e. hip to toe distance. A combination of knee flexion and ankle dorsiflexion during swing phase is suggested as a means of reducing compensatory mechanisms. To examine this hypothesis, an orthosis incorporating knee and ankle flexion was constructed. The design used a novel mechanism to link the motion of the knee to that of the ankle, and also used functional electrical stimulation. Two spinal cord-injured subjects were trained to use the orthosis in two configurations. The first configuration used knee flexion and ankle dorsiflexion and the second configuration used knee flexion alone. Kinematic data were obtained to measure the compensatory mechanisms used during gait. The results showed that a combination of knee flexion and ankle dorsiflexion during swing phase resulted in a reduction in compensatory mechanisms when compared with knee flexion alone.     

Adaptive control for backward quadrupedal walking. I. Posture and hindlimb kinematics

1. To gain new perspectives on the neural control of different forms of quadruped locomotion, we studied adaptations in posture and hindlimb kinematics for backward (BWD) walking in normal cats. Data from four animals were obtained from high-speed (100 fr/s) ciné film of BWD treadmill walking over a range of slow walking speeds (0.3-0.6 m/s) and forward (FWD) treadmill walking at 0.6 m/s. 2. Postural adaptations during BWD walking included flexion of the lumbar spine, compared to a relatively straight spine during FWD walking. The usual paw-contact sequence for FWD walking [right hindlimb (RH), right forelimb (RF), left hindlimb (LH), left forelimb (LF)] was typically reversed for BWD walking (RH, LF, LH, RF). The hindlimbs alternated consistently with a phase difference averaging 0.5 for both forms of walking, but the phasing of the forelimbs was variable during BWD walking. 3. As BWD walking speed increased from 0.3 to 0.6 m/s, average hindlimb cycle period decreased 21%, stance-phase duration decreased 29%, and stride length increased 38%. Compared to FWD walking at 0.6 m/s, stride length was 30% shorter, whereas cycle period and stance-phase duration were 17% shorter for BWD walking. For both directions, stance occupied 64 +/- 4% (mean +/- SD) of the step cycle. 4. During swing for both forms of walking, the hip, knee, and ankle joints had flexion (F) and extension (E1) phases; however, the F-E1 reversals occurred earlier at the hip and later at the knee for BWD than for FWD walking. At the ankle joint, the ranges of motion during the F and E1 phases were similar for both directions. During BWD walking, however, the knee flexed more and extended less, whereas the hip flexed less and extended more. Thus horizontal displacement of the limb resulted primarily from hip extension and knee flexion during BWD swing, but hip flexion and knee extension during FWD swing. 5. At the knee and ankle joints, there were yield (E2) and extension (E3) phases during stance for both forms of walking; however, yields at the knee and ankle joints were reduced during BWD walking. At the hip, angular motion was unidirectional, as the hip flexed during BWD stance but extended during FWD stance. Knee extension was the prime contributor to horizontal displacement of the body during BWD stance, but hip extension was the prime contributor to horizontal displacement during FWD stance. 6. Our kinematic data revealed two discriminators between BWD and FWD walking.(ABSTRACT TRUNCATED AT 400 WORDS)     

Simultaneous control of two rhythmical behaviors. II. Hindlimb walking with paw-shake response in spinal cat

The simultaneous control of the hindlimb paw-shake response and hindlimb walking at slow treadmill speeds (0.2-0.4 m/s) was examined in adult cats spinalized at the T12 level, 3-6 mo earlier. Paw shaking was elicited by either 1) application of adhesive tape or 2) water to the right hindpaw. To assess intralimb and interlimb coordination of the combined behaviors, activity from selected flexor and extensor muscles at the hip, knee, and ankle was recorded, and the kinematics of these joints were determined from high-speed cinefilm. When paw shaking was combined with hindlimb walking, the response in the stimulated limb was initiated during swing (F phase) of the step cycle. The onset of knee extensor activity provided the transition from the flexor synergy of swing to the mixed synergy of paw shake. At the end of the paw shake, an extensor synergy initiated the E-1 phase of swing, and the resultant joint motion was in-phase extension at the hip, knee, and ankle to lower the paw for contact with the treadmill belt. During the rapid (81 ms) paw-shake cycles, knee extensor and ankle flexor muscles exhibited single, coactive bursts that were reciprocal with coactive hip and ankle extensor bursts. This mixed synergy was reflected in the limb coordination, as knee flexion coincided with ankle extension and knee flexion coincided with ankle extension. Phasing of hip motions was variable, reflecting the role of the proximal in stabilization during paw shake (16). Although the number of paw-shake cycles combined during swing varied greatly from 2 to 14, average cycle periods, burst durations, and intralimb synergies were similar to those previously reported for spinal cats tested under conditions in which the trunk was suspended and hindlimbs were pendent (23, 27). For step cycles during which a long paw-shake response of 8-14 cycles occurred, swing duration of the shaking limb increased by 1 s, and during this prolonged interval, the contralateral hindlimb completed two support steps. Stance duration of the support steps was also prolonged. This adjustment maximized the duration of paw-contact and minimized any period of nonsupport by the contralateral hindlimb during paw shake. Completion of the paw-shake response was followed by either an alternating, or a nonalternating, gait pattern on the recovery steps. One spinal cat combined locomotion with short two-cycle paw-shake responses, and because the shortened response was limited primarily to the time ordinarily devoted to swing, interlimb adjustments were slight.(ABSTRACT TRUNCATED AT 400 WORDS)     

Chemical ablation of sensory afferents in the walking system of the cat abolishes the capacity for functional recovery after peripheral nerve lesions

Weakening the ankle extensor muscles of cats by denervation of the synergists of the medial gastrocnemius (MG) muscle results in transient increase in yield at the ankle during early stance. Recovery of ankle function occurs over a period of 1–2 weeks, is use-dependent, and is associated with increases in the strength of reflexes from MG group I muscle afferents and an increase in the magnitude of bursts in the MG muscles during stance. These observations have led to the hypothesis that feedback from large muscle afferents is necessary for functional recovery. In this investigation we have tested this hypothesis by examining functional recovery in animals treated with pyridoxine, a drug known to destroy large muscle afferents. In four adult animals we confirmed that pyridoxine abolished the group I-mediated tendon-tap reflex in the ankle extensor muscle, and subsequently found that group I afferents from MG were either destroyed or non-conducting. Immediately after pyridoxine treatment the animals showed severe locomotor dysfunction but all recovered significantly over a period of 1 or 2 months and showed only minor kinematics deficits at the time of the muscle denervations. In all four pyridoxine-treated animals, weakening of the ankle extensors by denervation of the synergists of the MG muscle resulted in a large increase in yield at the ankle that persisted almost unchanged for a month after the operation. The magnitude of burst activity in the MG muscle during early stance of the pyridoxine-treated animals either did not increase or increased only slightly after the denervation of synergists. These observations are consistent with the hypothesis that feedback from group I afferents is necessary for functional recovery in untreated animals. Electronic Publication     

Coherence analysis of muscle activity during quiet stance

Studies of muscle activation during perturbed standing have demonstrated that the typical patterns of coordination (“ankle strategy” and “hip strategy”) are controlled through multiple muscles activated in a distal-to-proximal or proximal-to-distal temporal pattern. In contrast, quiet stance is thought to be maintained primarily through the ankle musculature. Recently, spectral analysis of inter-segment body motion revealed the coexistence of both ankle and hip patterns of coordination during quiet stance, with the predominating pattern dependent on the frequency of body sway. Here we use frequency domain techniques to determine if these patterns are associated with the same muscular patterns as observed during perturbed stance. Six of the seven muscles measured showed a linear relationship to the sway of at least one body segment, all being leg muscles. Muscle–segment phases were consistent with that required to resist gravity at low frequencies, with increasing phase lag as frequency increased. Visual information had effects only at frequencies below 0.5 Hz, where the shift from in-phase to anti-phase trunk–leg co-phase was observed. These results indicate that co-existence of the ankle and hip pattern during quiet stance involves only leg musculature. Anti-phase movement of the trunk relative to the legs at higher frequencies arises from indirect biomechanical control from posterior leg muscles.     

A role for hip position in initiating the swing-to-stance transition in walking cats

In this investigation, we obtained data that support the hypothesis that afferent signals associated with hip flexion play a role in initiating the swing-to-stance transition of the hind legs in walking cats. Direct evidence came from observations in walking decerebrate cats. Assisting the flexion of the hip joint during swing advanced the onset of activity in ankle extensor muscles, and this advance was strongly correlated with a reduction in the duration of hip flexor muscle activity. The hip angle at the time of onset of the flexion to extension transition was similar during assisted and unassisted steps. Additional evidence for the hypothesis that sensory signals related to hip flexion are important in regulating the swing-to-stance transition came from four normal animals trained to walk in a variety of situations designed to alter the coordination of movements at the hip, knee, and ankle joints during the swing phase. Although there were exceptions in some tasks and preparations, the angle of the hip joint at the time of onset of extensor activity was generally less variable than that of the knee and ankle joints. We also found no clear relationships between the angle of the limb and body axes, or the length of the limb axis, and the time of onset of extensor activity. Finally, there were no indications that the stretching of ankle extensor muscles during swing was a factor in regulating the transition from swing-to-stance.     

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.     

Kinetic and energetic patterns for hindlimb obstacle avoidance during cat locomotion

 The safe control of walking over different terrains requires appropriate adaptations in the dynamic and kinematic limb patterns. To date, the study of locomotor dynamics in the cat has been confined to level, unobstructed walking. The present study extends the work of Lavoie et al. by applying linked segment analyses to estimate muscle contributions to torque and mechanical power at the hindlimb joints of two female cats during both unobstructed walking and obstacle avoidance. Data during obstacle avoidance were analyzed both when the hindlimb led in clearance and was farthest from the obstacle, and when it trailed in clearance and was closest or near to the obstacle. It was found that, in both the Far and Near obstructed conditions, the cats cleared the obstacles primarily by increasing the knee flexor torque already used during unobstructed gait. Contributions from the hip and ankle muscle groups were more variable. There was more emphasis on the hip extensors in mid to late stance, and the hip flexors generated a small amount of energy at paw-lift in the Far condition. In the Near condition, the hip extensors were employed to control hip flexion. We suggest that hip flexor generation power in mid-swing contributes to the clearance of the upcoming obstacle in the Far condition while, in the Near condition, hip flexion advances the already extended limb ahead of the obstacle. The ankle was actively dorsiflexed in the Near condition but was maintained in extension in the Far condition. The emphasis on active knee flexor control by the cat to avoid obstacles, as well as the dependence of ankle control on obstacle proximity, is similar to strategies seen for humans. However, the knee flexor strategy is innate to the cat’s normal level walking control, whereas in humans active knee flexion at toe-off requires a reorganization from level, non-obstructed gait. Received: 2 February 1998 / Accepted: 21 September 1998     

Activity of spindle afferents from cat anterior thigh muscles. II. Effects of fusimotor blockade

Chronically implanted electrodes and nerve cuff catheters were used to record the activity of individual muscle spindle afferents during treadmill walking as low doses of lidocaine were infused around the femoral nerve to progressively block gamma motoneuron activity. Both primary and secondary endings from both the monarticular knee extensors and the biarticular hip/knee muscles of the anterior thigh showed large decreases in afferent activity, usually well before changes in the electromyographic activity, force output, or length and velocity were seen in the parent muscles. This decline in the proprioceptive signal feeding back onto the spinal cord, which we presume to have involved most of the spindles supplied by the femoral nerve, did not cause noticeable irregularities or instability of the walking gait. At the peak of the fusimotor blockade, spindle afferents from knee extensor muscles lost about half of their usually brisk activity during the near-isometric contraction of the stance phase. Significant decreases in the response to passive stretch during the flexion phase also occurred. At the peak of the fusimotor blockade, spindle afferents from the biarticular muscles lost all of their activity during the rapidly shortening swing phase and about half of their activity during the rapidly lengthening stance phase. For both monarticular and biarticular muscle spindles, the activity decreases in stance and swing phase often occurred at distinctly different stages of the progressive fusimotor blockade, indicating several different sources of fusimotor control. From these data, we infer that the sensitivity of most spindle afferents is substantially influenced by fusimotor activity during phases of both extrafusal activity and extrafusal silence. At least some of this influence appears to come from fusimotor neurons whose recruitment is independent of the extrafusal recruitment. The extent and type of fusimotor effects on spindle afferent sensitivity (dynamic or static) appear to be specialized for the mechanical events that tend to occur during those phases.     

Ageing effects on knee and ankle joint angles at key events and phases of the gait cycle

The objective of this research was to determine whether joint angles at critical gait events and during major energy generation/absorption phases of the gait cycle would reliably discriminate age-related degeneration during unobstructed walking. The gaits of 24 healthy adults (12 young and 12 elderly) were analysed using the PEAK Motus motion analysis system. The elderly participants showed significantly greater single (60.3% versus 62.3%, p < 0.01) and double ( p < 0.05) support times, reduced knee flexion (47.7 degrees versus 43.0 degrees , p < 0.05) and ankle plantarflexion (16.8 degrees compared to 3.3 degrees , p = 0.053) at toe off, reduced knee flexion during push-off and reduced ankle dorsiflexion (16.8 degrees compared to 22.0 degrees , p < 0.05) during the swing phase. The plantarflexing ankle joint motion during the stance to swing phase transition (A2) for the young group (31.3 degrees ) was about twice ( p < 0.05) that of the elderly (16.9 degrees ). Reduced knee extension range of motion suggests that the elderly favoured a flexed-knee gait to assist in weight acceptance. Reduced dorsiflexion by the elderly in the swing phase implies greater risk of toe contact with obstacles. Overall, the results suggest that joint angle measures at critical events/phases in the gait cycle provide a useful indication of age-related degeneration in the control of lower limb trajectories during unobstructed walking.     

Contribution of cutaneous inputs from the hindpaw to the control of locomotion. I. Intact cats

The goal of this study was to evaluate the role of hindpaw cutaneous feedback in the control of locomotion, by cutting some (in one cat) or all (in 2 cats) cutaneous nerves bilaterally at ankle level. Kinematic and electromyographic (EMG) recordings were obtained before and for several weeks after denervation during level and incline (15 degrees up and down) treadmill walking. Ladder walking and ground reaction forces were also documented sporadically. Early after the denervation (1-3 days), cats could not walk across a ladder, although deficits were small during level treadmill walking. Increased knee flexion velocity caused a 14% reduction in swing phase duration. EMG activity was consistently increased in knee, ankle, and toe flexors, and in at least one knee or ankle extensor. The adaptive changes during walking on the incline were much reduced after denervation. Ladder walking gradually recovered within 3-7 wk. By this time, level treadmill walking kinematics had completely returned to normal, but EMG activity in flexors remained above control. Incline walking improved but did not return to normal. Mediolateral ground reaction forces during overground walking were increased by 200%. It is concluded that in intact cats, cutaneous inputs contribute more to demanding situations such as walking on a ladder or on inclines than to level walking. Active adaptive mechanisms are likely involved given that the EMG locomotor pattern never returned to control level. The companion paper shows on the other hand that when the same cats are spinalized, these cutaneous inputs become critical for foot placement during locomotion.