Speed-related changes in hindlimb intersegmental dynamics during the swing phase of cat locomotion

Summary To determine speed-related changes in hindlimb motion that might account for the mutability of bifunctional (hip extensor/knee flexor) muscle activity during the E1 phase of swing, we studied hip and knee joint kinematics and kinetics during swing over a ten-fold increase in locomotor speed (0.35 to 3.5 m/s). Three cats were filmed (100 frames/s) while locomoting on a motorized treadmill; kinematics were analyzed for the entire step cycle and kinetics for the swing phase. During swing, angular excursions at the hip and knee joints were similar for walking and trotting, but hip flexion and extension were significantly less after the transition from trot to gallop, while knee-angle range of motion increased during gallop phases E1, E2, and E3. During swing, knee-extension velocity peaked early in E1 and increased linearly with speed, while hip-flexion velocity peaked late in the flexion (F) phase and also increased linearly, but decreased precipitously at the trotgallop transition and remained constant as speed of galloping increased. Muscle torque directions during E1, flexor at the knee and extensor at the hip, were consistent with the proposed role of bifunctional posterior thigh muscles to decelerate thigh and leg segments for paw contact. At the knee joint, muscle torque during E1 counteracted a large interactive torque due to leg angular acceleration; the magnitudes of both torques were speed related with maximal values at the fastest speed tested (3.5 m/s). At the hip joint, muscle torque during E1 also counteracted a large interactive torque due to leg angular acceleration; the magnitudes of these two torques were speed related during the walk and trot, and like hip flexion velocity, decreased at the trot-gallop transition. Our data on speed-related changes in hindlimb dynamics suggest that the E1 burst amplitude (and perhaps duration) of posterior thigh muscles will be speed related during the walk and trot. After the trot-gallop transition at about 2.5 m/s, the recruitment of these bifunctional muscles may decline due to the changes in hindlimb dynamics. Because activity of these muscles counteracts interactive torques primarily related to leg angular acceleration, we suggest that motion-related feedback decoding this action may be important for regulating recruitment during E1.     

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)     

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)     

Recruitment of gastrocnemius muscles during the swing phase of stepping following partial denervation of knee flexor muscles in the cat

In walking cats, the biarticular medial and lateral gastrocnemius (MG–LG) muscles act to produce extension and flexion torques at the ankle and knee, respectively, and they usually display only one burst of activity beginning just before ground contact and ending near the end of the stance phase. Currently, the MG–LG muscles are considered to function primarily to control extension movements around the ankle joint during the stance phase. However, their flexion action at the knee means that they have the capacity to regulate rotations at the knee, but this role has not yet been clearly defined. Following partial denervation of the other muscles that normally act to flex the knee during swing, we observed that the MG–LG muscles, but not the Soleus muscle (a pure ankle extensor), often generated strong bursts of activity during early swing. These bursts were enhanced following mechanical stimulation of the paw, and they were especially prominent when the leg trailed over an object. They were absent when the leg led over an object. During treadmill walking the swing-related bursts in MG and LG had little influence on ankle flexion at the beginning of swing, but they were associated with slowing of ankle flexion when the leg trailed over an object. We hypothesized that the recruitment of these bursts functions to partially compensate for the reduction in knee torque resulting from the denervation of other knee flexors. Consistent with this hypothesis was our finding that the magnitude of the swing-related activity in the MG–LG muscles was linearly correlated to the extent of the knee flexion and to the peak angular velocity of knee flexion, and that the timing of the bursts was similar to that in the denervated muscles prior to denervation. Our findings suggest that an excitatory pathway exists from the flexor half-center of the central pattern-generating network to MG–LG motoneurons, and that this pathway is strongly regulated by central and/or peripheral signals.     

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.     

Simultaneous control of two rhythmical behaviors. I. Locomotion with paw-shake response in normal cat

We investigated the ability of normal cats, trained to maintain a constant position while walking on a treadmill, to combine the paw-shake response with quadrupedal locomotion. Hindlimb paw-shake responses were elicited during walking after the right hindpaw was wrapped with tape. To assess intralimb and interlimb coordination of the combined behaviors, electromyographic (EMG) recordings from forelimb extensor muscles and from selected flexor and extensor muscles at the three major hindlimb joints were correlated with joint motion by using high-speed, cinefilm analysis. When paw shaking was combined with walking, the response occurred during the swing phase of the taped hindlimb. To accommodate the paw-shake response, swing duration of the shaking hindlimb and of the homolateral forelimb increased and was followed by a brief recovery step. Concurrently, to compensate for the response, stance durations of the contralateral forelimb and hindlimb increased. The magnitude of these adjustments in interlimb coordination was influenced by the number of paw-shake cycles, which ranged from one to four oscillations. Transitions between the muscle synergies for the paw-shake response and swing were smooth in the shaking limb. Early in the swing phase, when the flexor muscles were still active (F phase), the paw shake was initiated by an early onset of knee extensor activity, which preceded extensor activity at the hip and ankle. This action provided a transition from the general reciprocal synergy between flexor and extensor muscles of locomotion to the mixed synergy that is typical of the paw shake (30). Following the last paw-shake cycle, an extensor synergy initiated the E-1 phase of swing, and the resultant joint motion was in-phase extension of the hip, knee, and ankle to lower the paw for stance. Average cycle period and burst duration for muscles participating in the paw-shake response were similar to those reported for normal cats assuming a standing posture (28, 30). The average number of paw-shake cycles, however, decreased from eight to three when the response occurred during walking, suggesting that the response was truncated to provide for continued locomotion. Further, hip motion was variable when the paw shake was combined with swing, and sometimes the hip failed to oscillate and its trajectory was similar to that of an unperturbed swing phase. When hip joint oscillations occurred during the paw-shake response, they were in-phase with ankle motions.(ABSTRACT TRUNCATED AT 400 WORDS)     

Coordination of two-joint rectus femoris and hamstrings during the swing phase of human walking and running

It has been hypothesized previously that because a strong correlation was found between the difference in electromyographic activity (EMG) of rectus femoris (RF) and hamstrings (HA; EMG_RF–EMG_HA) and the difference in the resultant moments at the knee and hip (M_k–M_h) during exertion of external forces on the ground by the leg, input from skin receptors of the foot may play an important role in the control of the distribution of the resultant moments between the knee and hip by modulating activation of the two-joint RF and HA. In the present study, we examined the coordination of RF and HA during the swing phase of walking and running at different speeds, where activity of foot mechanoreceptors is not modulated by an external force. Four subjects walked at speeds of 1.8 m/s and 2.7 m/s and ran at speeds of 2.7 m/s and 3.6 m/s on a motor-driven treadmill. Surface EMG of RF, semimembranosus (SM), and long head of biceps femoris (BF) and coordinates of the four leg joints were recorded. An inverse dynamics analysis was used to calculate the resultant moments at the ankle, knee, and hip during the swing phase. EMG signals were rectified and low-pass filtered to obtain linear envelopes and then shifted in time to account for electromechanical delay between EMG and joint moments. During walking and running at all studied speeds, mean EMG envelope values of RF were statistically (P<0.05) higher in the first half of the swing (or at hip flexion/knee extension combinations of joint moments) than in the second half (or at hip extension/knee flexion combinations of joint moments). Mean EMG values of BF and SM were higher (P<0.05) in the second half of the swing than in the first half. EMG and joint moment peaks were substantially higher (P<0.05) in the swing phase of walking at 2.7 m/s than during the swing phase of running at the same speed. Correlation coefficients calculated between the differences (EMG_RF–EMG_HA) and (M_k–M_h), taken every 1% of the swing phase, were higher than 0.90 for all speeds of walking and running. Since the close relationship between EMG and joint moments was obtained in the absence of an external force applied to the foot, it was suggested that the observed coordination of RF and HA can be regulated without a stance-specific modulation of cutaneous afferent input from the foot. The functional role of the observed coordination of RF and HA was suggested to reduce muscle fatigue. Received: 7 August 1997 / Accepted: 9 February 1998     

Modelling the optimal control of cyclical leg movements induced by functional electrical stimulation.

An optimal control strategy for FES-induced cyclical leg movements in paraplegics is proposed. The control of the cyclical movement of a freely swinging leg is considered as an example. Quadriceps and the flexion withdrawal reflex are stimulated in order to generate a cyclical movement, of which the forward swing resembles the swing phase of gait. Optimal stimulation patterns are determined on the basis of an optimization criterion and a dynamic model of the system. The criterion is based on desired movement parameters and a minimal duration of the stimulation bursts. The movement parameters should ensure the generation of the desired cyclical movement: a desired hip angle range, sufficient foot clearance during the forward swing and knee extension at the beginning of the backward swing. Minimal duration of the stimulation bursts is assumed to yield minimal fatigue. A dynamic model, describing the dynamics of the neural system, the muscles and the leg, was constructed and its parameters identified on the basis of preliminary experiments and literature. Optimal timing of the quadriceps and flexion reflex stimulation bursts was determined by means of computer simulation. These simulations predicted that the flexion reflex should be stimulated in a short burst approximately 150 ms before the start of the forward swing. The quadriceps should be stimulated approximately starting 200 ms before the end of the forward swing in order to ensure knee extension at the beginning of the backward swing. The duration of one cycle of the movement was between 1300 and 1500 ms in these simulations. These results predict that the movement specified by the functional objectives can be realised using only two channels of stimulation. On the basis of the optimal timing, an adaptive control strategy can be designed, which varies the stimulation burst width when muscles fatigue.     

Contrasting roles of inertial and muscle moments at knee and ankle during paw-shake response

Intralimb kinetics of the paw-shake response (PSR) were studied in four spinal, adult cats. Using rigid body equations of motion to determine the dynamic interactions between limb segments, knee and ankle joint kinetics were calculated for the steady-state cycles as defined in the preceding paper. Hindlimb motion was filmed (200 frames/s) to obtain knee and ankle kinematics. Responses of flexors and extensors at both joints were recorded synchronously with cinefilm. Ankle and knee joint kinematics were determined from 51 steady-state cycles of 16 PSRs. Average maximum displacements, velocities, and accelerations were substantially greater for the ankle than for the knee joint. Knee and ankle motions were out of phase in the first part of the cycle; knee extension occurred simultaneously with ankle flexion. In the second part of the cycle, motions at the two joints were sequential; rapid knee flexion, accompanied by negligible ankle displacement, preceded rapid ankle extension with minimal knee displacement. At the ankle joint, peak net moments tending to cause flexion and extension were similar in magnitude and determined primarily by muscle moments. Moments due to leg angular acceleration contributed significantly to an extensor peak in the net moment near the end of the cycle. Other inertial and gravitational moments were small. At the knee joint, net moments tending to cause flexion and extension were also similar, but smaller than those at the ankle. The knee muscle moments, however, were large and counteracted large inertial moments due to paw angular acceleration. Also, moments due to leg angular acceleration and knee linear acceleration were substantial and opposite in effect. Other inertial and the gravitational moments were negligible. Muscle moments slowed and reversed joint motions, and active muscle force components of muscle moments were derived from lengthening of active musculotendinous units. Segmental interactions, in which proximal segment motion augmented distal segment velocity, increased the effectiveness of PSR steady-state cycles by facilitating the generation of extremely large paw linear accelerations. Limb oscillations during PSR steady-state result from interactions between muscle synergies and motion-dependent limb dynamics. At the ankle, muscle activity functioned to control paw acceleration, whereas at the knee, muscle activity functioned to control leg and paw inertial interactions.(ABSTRACT TRUNCATED AT 400 WORDS)     

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.     

Stumbling corrective reaction: a phase-dependent compensatory reaction during locomotion.

1. Tactile stimuli to the paw consisting of a stick making contact or an air puff aimed at the dorsum were used to study the phasic influence of locomotor activity on the reflex pattern elicited in extensor and flexor muscles and on the induced compensatory movements in intact cats. The resulting movements and reflex pattern are called "stumbling corrective reactions." 2. The basic reflex pattern and movements of the stumbling corrective reaction are: a) if the stimulus occurs during the swing phase, a short-latency activation of the flexor muscles, which induces an additional flexion of the limb lifting the paw over the obstacle; b) if the stimulus occurs during the support phase, an inhibition followed by an excitation of the extensor muscles, which neither increase nor decrease the extension. However, the stimulus evokes an increased flexor activity in the succeeding swing phase, which induces a brisker flexion. 3. Tactile stimuli to the proximal part of the limb or to the belly in front of the knee evoked the same type of phase-dependent compensatory reactions. Such reactions would presumably be beneficial for the animal to avoid high obstacles that impede movement. 4. A nonnoxious electrical stimulus (typically 2 mA; 1 ms) applied to the dorsum of the paw was used to study systematically the reflex pattern of the stumbling corrective reaction. Two pathways were defined to the knee flexor (semitendinosus). One early burst was evoked at about 10 ms and one later at about 25 ms after the stimulus. Short inhibitory pathways and longer excitatory pathways (20-50 ms) projected to the extensor nuclei. A short-latency (10 ms) excitatory pathway to the ankle extensor (lateral gastrocnemius) was also activated. 5. A painful electrical stimulus applied to the dorsum of the paw evoked large flexor responses during the whole step cycle. During the support phase the locomotion was disrupted as the supporting limb was withdrawn. 6. The results demonstrate that intact cats are able to compensate rapidly for unpredicted perturbations and that the reflex pattern and the induced corrective movements are adapted to the locomotor activity so that functionally meaningful movements are evoked in each phase of the step cycle. 7. The evoked reflexes and their modulation are consistent with those previously found in chronic spinal cats during walking and in paralyzed spinal cats performing "fictive locomotion." It is suggested that the same spinal pathways are used, and that they are controlled by the spinal "locomotor generator."     

Incomplete spinal cord injury promotes durable functional changes within the spinal locomotor circuitry

While walking in a straight path, changes in speed result mainly from adjustments in the duration of the stance phase while the swing phase remains relatively invariant, a basic feature of the spinal central pattern generator (CPG). To produce a broad range of locomotor behaviors, the CPG has to integrate modulatory inputs from the brain and the periphery and alter these swing/stance characteristics. In the present work we raise the issue as to whether the CPG can adapt or reorganize in response to a chronic change of supraspinal inputs, as is the case after spinal cord injury (SCI). Kinematic data obtained from six adult cats walking at different treadmill speeds were collected to calculate the cycle and subphase duration at different stages after a first spinal hemisection at T(10) and after a subsequent complete SCI at T(13) respectively aimed at disconnecting unilaterally and then totally the spinal cord from its supraspinal inputs. The results show, first, that the neural control of locomotion is flexible and responsive to a partial or total loss of supraspinal inputs. Second, we demonstrate that a hemisection induces durable plastic changes within the spinal locomotor circuitry below the lesion. In addition, this study gives new insights into the organization of the spinal CPG for locomotion such that phases of the step cycle (swing, stance) can be independently regulated for adapting to speed and also that the CPGs controlling the left and right hindlimbs can, up to a point, be regulated independently.     

Step, swim, and scratch motor patterns in the turtle

The turtle generates a variety of coordinated hindlimb movements, including different forms of locomotion and scratching. The intact turtle produces forward step, forward swim, and backpaddle. Following spinal cord transection, rostral, pocket, and caudal scratches can be evoked by mechanical stimulation of the shell. Comparisons of the kinematics and motor patterns of these six behaviors provide insights regarding neuronal mechanisms underlying their production. All six behaviors were characterized by alternating hip flexion and extension and by an event during which force was exerted against a substrate. The portion of the cycle occupied by hip flexion or extension movement varied across behaviors. Hip extension occupied well over half the cycle period in the forward step and the caudal scratch. The cycle was split into approximately half hip flexion and half hip extension for the forward swim, the backpaddle, and the rostral scratch. Hip flexion occupied over half the cycle in the pocket scratch. The swim and scratch forms had curvilinear, crescent-shaped toe trajectories and a single burst of monoarticular knee extensor activity during each cycle. The forward step had a linear toe trajectory and two bursts of knee extensor activity during each cycle, one during swing and one during stance. Timing of monoarticular knee extensor onset was similar for: the forward swim, the rostral scratch, and the swing phase burst of forward step; the pocket scratch and the stance phase burst of forward step; and the backpaddle and the caudal scratch. Amplitudes of muscle activity varied among the six behaviors; high amplitudes of activity were associated with events during which force was exerted against a substrate. These times of force exertion were: stance phase in the forward step, powerstroke in the forward swim and the backpaddle, and rubs of the limb against the shell in the scratch forms. The six behaviors studied represent a range of parameter values, as evidenced by relative durations of hip flexion to hip extension, knee extensor phasing, and electromyogram (EMG) amplitudes. This range of behaviors could be produced by assembling different combinations of neurons from a common pool, with all six behaviors likely sharing some basic circuitry. The extent of shared circuitry may be greater between behaviors with similar timing, e.g., backpaddle and caudal scratch.     

The locomotion of the low spinal cat. II. Interlimb coordination

The interaction of the two hindlimbs were investigated by an analysis of the muscular activity and the movements in 14 chronic spinal kittens during treadmill locomotion (i.e. in kittens subjected to a transection of the spinal cord (Th_10–12) one or two weeks after birth). At low speed the limbs were alternating (walk or trot). At higher they were activated more simultaneous, as during gallop. The two limbs could walk at different velocities, as during walking in a circle, when the two belts of the treadmill were driven at different speeds. The duration of the support phases was mainly influenced by the speed of the belt on which the limb was walking. The limbs could still maintain a common rhythm up to a two or three fold speed difference, as the flexion or the first extension phase of the limb walking on the “fast” belt was prolonged and the flexion phase of the “slow limb” was shortened. At extreme speed differences the limb on the “fast belt” performed 2, 3 and even 4 steps during one stepcycle of the “slow limb”. The placement of the feet was found to maintain the most stable relationship during alternating gaits at different speed differences. It is concluded that all phases of the step cycle are modifiable and that there are several mechanisms coordinating the limbs within the spinal cord.     

Knee kinematics during walking at different speeds in people who have undergone total knee replacement

People who have undergone total knee replacement (TKR) experience difficulties in some daily activities including walking. Walking at faster speeds requires more knee flexion and may therefore present a greater challenge following TKR. The aim of this study was to compare the knee kinematics of patients following TKR and unimpaired controls during comfortable and fast walking speeds. Forty patients (22 women, 18 men) 12 months following TKR and 40 control participants (matched for age and sex) were assessed during walking at self-selected comfortable and fast speeds using three dimensional motion analysis. The group averages of spatiotemporal and peak kinematic characteristics in the sagittal, coronal and transverse movement planes were compared using univariate analysis of variance with walking speed as a co-variate. The TKR group walked with significantly reduced cadence (p < 0.001 at both speeds) and reduced stride length (p < 0.001 at both speeds), less knee flexion during stance and swing phases (p < 0.001 for both speeds) and less knee extension during stance phase (p < 0.024 for comfortable speed; p < 0.042 for fast speed). The TKR group also walked with less peak knee external rotation than controls at both speeds (p < 0.001 for both speeds). Both groups increased their velocity, cadence and stride length by a similar proportion when walking at fast speed. When walking at a faster speed, spatiotemporal gait parameters and knee motion are altered in a similar manner for both TKR patients and controls. However, at both walking speeds, TKR patients exhibit residual deficits 12 months following surgery.     

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)     

Coupling of upper and lower limb pattern generators during human crawling at different arm/leg speed combinations

A crawling paradigm was performed by healthy adults to examine inter-limb coupling patterns and to understand how central pattern generators (CPGs) for the upper and lower limbs are coordinated. Ten participants performed hands-and-feet crawling on two separate treadmills, one for the upper limbs and another one for the lower limbs, the speed of each of them being changed independently. A 1:1 frequency relationship was often maintained even when the treadmill speed was not matched between the upper and lower limbs. However, relative stance durations in the upper limbs were only affected by changes of the upper limb treadmill speed, suggesting that although absolute times are adjusted, the relative proportions of stances and swing do not adapt to changes in lower limb treadmill speeds. With large differences between treadmill speeds, changes in upper and lower limb coupling ratio tended to occur when the upper limbs stepped at slower speeds than the lower limbs, but more rarely the other way around. These findings are in sharp contrast with those in the cat, where forelimbs always follow the rhythm of the faster moving hindlimbs. However, the fact that an integer frequency ratio is often maintained between the upper and lower limbs supports evidence of coupled CPG control. We speculate that the preference for the upper limb to decrease step frequency at lower speeds in humans may be due to weaker ascending propriospinal connections and/or a larger influence of cortical control on the upper limbs which allows for an overriding of spinal CPG control.     

A kinematic and electromyographic study of cutaneous reflexes evoked from the forelimb of unrestrained walking cats

A kinematic and electromyographic (EMG) analysis was undertaken of the responses evoked in the forelimb of the cat by either mechanical obstruction of the forelimb during the swing phase of locomotion or by electrical stimulation of low-threshold cutaneous afferents during both swing and stance. Mechanical obstruction of the forelimb with a stiff metal rod evoked a complex response that allowed the cat to smoothly negotiate the obstacle without undue disruption of the overall locomotor rhythm. The initial movements were a flexion of the shoulder, together with a locking of the elbow joint, and a dorsiflexion of the wrist, which caused the limb to withdraw from the obstacle. They were followed by an extension of the shoulder, a flexion of the elbow, and a ventroflexion of the wrist, which together brought the limb forward and above the obstacle. The associated and complex pattern of short- and long-latency EMG responses was shown to be related to different aspects of the movement. At the shoulder there was a strong activation of flexor muscles; these responses were of long duration (greater than or equal to 100 ms) and generally lasted throughout the period of shoulder flexion. At the elbow, both flexor and extensor muscles were activated at short latency (9-13 ms). In flexors, this was followed by a cessation and subsequently an augmentation and prolongation of their activity. Dorsiflexors of both the wrist and digits were activated at short latency (10-12 ms) and remained active throughout the period of dorsiflexion of these joints. An injection of a local anesthetic into the area of skin contacted by the metal rod reduced or abolished all of the reflex responses, which suggests that the integrity of cutaneous reflex pathways is essential for the elaboration of these responses. Electrical stimulation of a cutaneous nerve innervating the distal forelimb (the superficial radial nerve) resulted in qualitatively similar, although weaker, responses to those obtained with the mechanical stimulation. Terminal experiments confirmed that these responses were mediated by low-threshold cutaneous afferents. Electrical stimulation also evoked short-latency excitatory responses (10-12 ms) in extensor muscles of the elbow. Generally, the largest reflex effects were obtained during the period of swing for flexor, extensor, and bifunctional muscles. During stance the stimulus was normally ineffective in exciting flexor muscles and in extensors evoked a short-latency inhibition, which was frequently followed by an increase in activity.(ABSTRACT TRUNCATED AT 400 WORDS)     

The role of cutaneous afferents in controlling locomotion evoked by epidural stimulation of the spinal cord in decerebrate cats

The effects of the cutaneous input on the formation of the locomotor pattern in conditions of epidural stimulation of the spinal cord in decerebrate cats were studied. Locomotor activity was induced by rhythmic stimulation of the dorsal surface of spinal cord segments L4-L5 at a frequency of 3-5 Hz. Electromyograms (EMG) recorded from the antagonist muscles quadriceps, semitendinosus, tibialis anterior, and gastrocnemius lateralis were recorded, along with the kinematics of stepping movements during locomotion on a moving treadmill and reflex responses to single stimuli. Changes in the pattern of reactions observed before and after exclusion of cutaneous receptors (infiltration of lidocaine solution at the base of the paw or irrigation of the paw pads with chlorothane solution) were assessed. This treatment led to impairment of the locomotor cycle: the paw was placed with the rear surface downward and was dragged along in the swing phase, and the duration of the stance phase decreased. Exclusion of cutaneous afferents suppressed the polysynaptic activity of the extensor muscles and the distal flexor muscle of the ipsilateral hindlimb during locomotion evoked by epidural stimulation of the spinal cord. The effects of exclusion of cutaneous afferents on the monosynaptic component of the EMG response were insignificant.     

Peripheral control of the cat's step cycle

Acute low spinal and curarized cats injected with noradrenergic agonists i.v. can elicit an efferent burst pattern which can be recorded in muscle nerve filaments and can be referred to as “fictive locomotion”. This study investigates the effect that feedback, arising from movements in the hip joint, can exert on the central network generating fictive locomotion. The central network is uncoupled from generating any active movements by curarization. The motor pattern could be entrained by applying sinusoidal hip movements, even when a very extensive denervation of the leg had been performed leaving only some of the muscles around the hip and the hip joint innervated. During flexion movements, efferents to different flexor muscles became active and during movements in the reverse direction (extension), efferents to extensors were active. With an increasing movement frequency the onsets of both flexor and extensor bursts were delayed in the movement cycle. The duration of the extensor bursts varied markedly with the movement cycle, whereas pure flexors changed less in burst duration. The frequency range within which the efferent burst activity was entrained in a strict 1:1 relation to the movement varied between 5 to 70% above and below the resting burst period. In preparations with a narrow 1:1 range, a “relative coordination” was encountered outside this range. The flexor burst duration was in these cases dependent on where in the hip movement cycle the bursts appeared.