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)     

Paw-shake responses with joint immobilization: EMG changes with atypical feedback

Summary To determine the effects of atypical motion-related feedback on motor patterns of the paw shake, EMG patterns of selected flexor and extensor muscles were recorded under four conditions of joint immobilization (hip and ankle alone, hip-knee, hip-knee-ankle) and compared to responses evoked in the freely-moving hindlimb of the chronic-spinal cat. With only the ankle joint casted, paw shaking was easily evoked by applying tape to the paw, and cyclic characteristics were not altered. However, under the three conditions with hip-joint immobilization (hip alone, hip-knee, hip-knee-ankle), responses were difficult to obtain, and if elicited, the number of cycles within a response decreased and cycle periods were prolonged. The temporal organization of consecutive cycles, however, was not altered by immobilization of any joint(s). Ankle (LG) and hip (GM) extensor activity was relatively unaffected by conditions of joint immobilization. In contrast, hip flexor (IP) and knee extensor (VL) bursts were often absent under all three conditions of hip-joint immoblization, and if present, VL burst durations decreased under the casted hip-knee-ankle condition, while the onset of IP activity occurred early in the cycle with prolonged bursts under casted ankle and casted hip-knee-ankle conditions. The coactivity of the knee extensor (VL) and ankle flexor (TA) was disrupted by conditions of hip-joint immobilization: VL onset was dissociated from TA onset and coincident with LG onset. These results suggest that motion-related feedback from the hip joint is particularly important in the initiation, cycle frequency, and the number of cycles of paw-shake responses. The presence of atypical motion-dependent feedback from the hip joint altered activity of knee and ankle anterior muscles, while motion-dependent feedback from the ankle joint changed activity of the anterior hip muscle. Moreover, the results suggest a differential control of posterior and anterior muscles of the hindlimb, consistent with paw-shake limb dynamics.     

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)     

The distal hindlimb musculature of the cat

Summary Chronic recordings were made of electromyographic (EMG) activity, tension, and length of distal hindlimb muscles in six cats performing a variety of normal motor tasks. Muscles studied thoroughly or in part were medial gastrocnemius, lateral gastrocnemius, plantaris, soleus, flexor digitorum brevis, flexor digitorum longus, flexor hallucis longus, tibialis posterior, tibialis anterior, extensor digitorum longus, peroneus longus, and peroneus brevis. Postural and locomotor activities were examined, as well as jumping, landing, scratching, and paw shaking. In general, muscles could be assigned to traditional groupings (e.g. extensor, flexor) related to the demands of the motor task. Patterns of muscle activity were most often consistent with current understanding of muscle mechanics and neural coordination. However, purely functional distinctions between flexor digitorum longus and flexor hallucis longus (anatomical synergists) were made on the basis of activity patterns. Likewise, the activity of plantaris and flexor digitorum brevis, which are attached in series, was differentiated in certain tasks. The rhythmical oscillatory patterns of scratching and paw shaking were found to differ temporally in a manner consistent with the limb mechanics. In several cases, mechanical explanations of specific muscle activity required length and force records, as well as EMG patterns. Future efforts to study motor patterns should incorporate information about the relationships between muscle activation, tension, length and velocity.Abbreviations EDL extensor digitorum longus - FDB flexor digitorum brevis - FDL flexor digitorum longus - FHL flexor hallucis longus - LG lateral gastrocnemius - MG medial gastrocnemius - PB peroneus brevis - PL peroneus longus - PLT plantaris - SOL soleus - TA tibialis anterior - TP tibialis posterior Limbs A ankle - K knee - LF left forelimb - LH left hindlimb - RF right forelimb - RH right hindlimb Step Cycle Phases E₁ first extension, late swing phase prior to footfall - E₂ second extension, early stance phase - E₃ third extension, late stance phase - F flexion, early swing phase     

Proprioceptive modulation of hip flexor activity during the swing phase of locomotion in decerebrate cats

This study examined the influence of proprioceptive input from hip flexor muscles on the activity in hip flexors during the swing phase of walking in the decerebrate cat. One hindlimb was partially denervated to remove cutaneous input and afferent input from most other hindlimb muscles. Perturbations to hip movement were applied either by 1) manual resistance or assistance to swing or by 2) resistance to hip flexion using a device that blocked hip flexion but allowed leg extension. Electromyographic recordings were made from the iliopsoas (IP), sartorius, and medial gastrocnemius muscles. When the hip was manually assisted into flexion, there was a reduction in hip flexor burst activity. Conversely, when hip flexion was manually resisted or mechanically blocked during swing, the duration and amplitude of hip flexor activity was increased. We also found some specificity in the role of afferents from individual hip flexor muscles in the modulation of flexor burst activity. If the IP muscle was detached from its insertion, little change in the response to blocking flexion was observed. Specific activation of IP afferent fibers by stretching the muscle also did not greatly affect flexor activity. On the other hand, if conduction in the sartorius nerves was blocked, there was a diminished response to blocking hip flexion. The increase in duration of the flexor bursts still occurred, but this increase was consistently lower than that observed when the sartorius nerves were intact. From these results we propose that during swing, feedback from hip flexor muscle afferents, particularly those from the sartorius muscles, enhances flexor activity. In addition, if we delayed the onset of flexor activity in the contralateral hindlimb, blocking hip flexion often resulted in the prolongation of ipsilateral flexor activity for long periods of time, further revealing the reinforcing effects of flexor afferent feedback on flexor activity. This effect was not seen if conduction in the sartorius nerves was blocked. In conclusion, we have found that hip flexor activity during locomotion can be strongly modulated by modifying proprioceptive feedback from the hip flexor muscles.     

Stumbling corrective reaction during fictive locomotion in the cat

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

Intralimb coordination of the paw-shake response: a novel mixed synergy

Intralimb coordination of the paw-shake response (PSR) was studied in five normal and eleven spinal adult cats. Representative extensor and flexor muscles that function at the hip, knee, and ankle joints were recorded, and in six spinal cats the kinematics of these joints were determined from high-speed cinefilm. The PSR was characterized uniquely by mixed (flexor-extensor) synergies. Knee extensor (VL) and ankle flexor (TA) coactivity constituted one synergy, while the second synergy included hip extensors (GM, BF), knee flexors (BF, LG), and ankle extensor (LG). Joint displacements reflected the mixed synergy. Motions at the knee and ankle were out of phase, while motions at the hip were in phase with movements of the knee. Electromyographic burst durations and onset latencies were similar for normal and spinal cats, and in all cycles of a given PSR, the recruitment pattern was consistent for all muscles, except VL. High variability and missing bursts marked the activity of VL in some spinal cats. In PSRs with missing VL bursts, oscillations at the knee joint were not coordinated with cyclic actions at the hip and ankle. From the kinematic records three distinct phases of the PSR were identified: start-up consisted of the initial four to six cycles during which hip, knee, and ankle actions progressively became organized; steady-state included the middle three to five cycles that were characterized by consistent displacement at all three joints; and slow-down comprised the last three to four cycles during which the rate of oscillations slowed, and joint excursions decreased. During steady-state cycles, muscle contractions acted to reverse joint motions at the knee and ankle joints. Thus, knee and ankle extensor recruitment coincided with joint flexion, while joint flexors were recruited during joint extension. Muscle activity at the hip, however, was in phase with displacement. While neural input to muscle is consistent throughout the three phases of the PSR, segment motions can become progressively organized during start-up to achieve stable oscillations. Whether the PSR attains steady-state or not may hinge on the sensitive interplay that occurs between muscle activities and intersegmental mechanical interactions. That kinetic interplay is detailed in the following paper.     

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."     

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.     

Locomotor deficits in the mutant mouse,Lurcher

Summary The effect of total Purkinje cell degeneration on treadmill locomotion was studied in the cerebellar mutant mouse Lurcher. Other movements such as swimming and scratchting were also studied in order to evaluate the cerebellar control of rhythmic actions. Cinematographic and electromyographic recordings were taken from normal and Lurcher mice that were subsequently perfused to obtain a Purkinje cell count. Walking deteriorated progressively and was clearly abnormal in 30 day old Lurchers with 90% Purkinje cell degeneration. In adult Lurcher mice in which Purkinje cells were totally absent, walking was characterized by short steps with exaggerated hindlimb flexion in the swing phase. Also, both the interlimb step ratio, defined as the step length of the reference limb divided by the step length of the opposite limb, and the interlimb coupling, defined as the temporal relation of one footfall with respect to the footfall of another limb, varied more than in normal mice. Furthermore, the locomotion of Lurcher mice displayed increased vertical displacement of the hip and an inability to produce continuous step cycles without stumbling. Both the EMG onset relative to foot contact and the EMG burst duration were highly variable, and a greater overlap in the activities of antagonist muscles at the transition from ankle extension to flexion was evident. Although both walking and swimming involve cyclical limb movements, the disorganization of the cycle and the irregular EMG pattern seen in the Lurcher during walking were not observed during swimming. Furthermore, scratching was well executed in the Lurcher mice. However, a consistently higher tonic extensor activity at the ankle appeared during walking, swimming and scratching. These results suggest that, in contrast to swimming and scratching, the requirements of walking depend to a greater degree on a functional cerebellar cortex for successful performance.     

Contribution of the motor cortex to the structure and the timing of hindlimb locomotion in the cat: a microstimulation study

We used microstimulation to examine the contribution of the motor cortex to the structure and timing of the hindlimb step cycle during locomotion in the intact cat. Stimulation was applied to the hindlimb representation of the motor cortex in 34 sites in three cats using either standard glass-insulated microelectrodes (16 sites in 1 cat) or chronically implanted microwire electrodes (18 sites in 2 cats). Stimulation at just suprathreshold intensities with the cat at rest produced multi-joint movements at a majority of sites (21/34, 62%) but evoked responses restricted to a single joint, normally the ankle, at the other 13/34 (38%) sites. Stimulation during locomotion generally evoked larger responses than the same stimulation at rest and frequently activated additional muscles. Stimulation at all 34 sites evoked phase-dependent responses in which stimulation in swing produced transient increases in activity in flexor muscles while stimulation during stance produced transient decreases in activity in extensors. Stimulation with long (200 ms) trains of stimuli in swing produced an increased level of activity and duration of flexor muscles without producing changes in cycle duration. In contrast, stimulation during stance decreased the duration of the extensor muscle activity and initiated a new and premature period of swing, resetting the step cycle. Stimulation of the pyramidal tract in two of these three cats as well as in two additional ones produced similar effects. The results show that the motor cortex is capable of influencing hindlimb activity during locomotion in a similar manner to that seen for the forelimb.     

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)     

Interlimb coordination during fictive locomotion in the thalamic cat

Summary Efferent discharges in muscle nerves of the four limbs were recorded simultaneously during spontaneous fictive locomotion in thalamic cats with the goal of understanding how the central nervous system controls interlimb coordination during stepping. The onset of the bursts of activity in the nerve of a selected flexor muscle in each limb allowed the temporal and the phase relationships between the fictive step cycle of a pair of limbs to be determined. Our main results are the following: 1) the fictive step cycles of the two forelimbs are always strictly alternated whereas the phasing of the step cycles of either the two hindlimbs or pairs of homolateral or diagonal limbs is more variable; 2) the time interval between the onsets of the flexor bursts of one of the two pairs of diagonal limbs is independent of the step cycle duration; 3) distinct patterns of interlimb coordination exist during fictive locomotion; a small number of patterns of coordination involving all four limbs, which correspond to the walking and the trotting gaits in the intact cat, occur very frequently. The results demonstrate that the central nervous system deprived of phasic afferent inputs from the periphery has the capacity to generate most of the patterns of interlimb coordination which occur during real locomotion. They further support the view that the central pattern of interlimb coordination essentially results from diagonal interactions between a forelimb generator for locomotion and a hindlimb one.Abbreviations CD step cycle duration - LF left forelimb - LH left hindlimb - m slope of correlation curve - N number of step cycles - r correlation coefficient - RF right forelimb - RH right hindlimb - Ti time interval     

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.     

Motor functions in rat hindlimb muscles following neonatal sciatic nerve crush.

The sciatic nerve was crushed in the right hindlimb in newborn (3-8 h old) rats. Two to four months later, electromyographic activity was recorded from both the control and reinnervated ankle extensor muscles soleus or lateral gastrocnemius and from the ankle flexor muscle tibialis anterior. Tonic postural activity was present in the extensor muscles on both sides during quiet stance. The control flexor muscles were usually silent in this situation, but the reinnervated flexors exhibited abnormal sustained activity. During locomotion, the control extensors were activated during the stance phase and their mean burst made up 61.5% of the step cycle. The control tibialis anterior muscle fired only during the swing phase, with the burst lasting 18.1% of the step cycle. In the reinnervated extensor muscles, the mean burst duration was decreased (46% of the cycle) but the basic locomotor pattern was not impaired. The reinnervated tibialis muscle, however, was activated abnormally, with one appropriate flexor burst during the swing phase and an "extensor-like" burst during the stance phase of the step. Reflex responses to stretch were weak or absent on the operated side. Histological examination showed that the reinnervated soleus and tibialis muscles were almost devoid of muscle spindles. The motor unit mean firing rates in the reinnervated soleus (22 imp/s) and lateral gastrocnemius (45 imp/s) matched those of the control muscles (25 and 42 imp/s, respectively). In contrast to the phasic, high-frequency firing (52-80 imp/s) in the control tibialis, the reinnervated tibialis motor units fired at significantly lower rates (22-56 imp/s).(ABSTRACT TRUNCATED AT 250 WORDS)     

Study of cutaneous reflex compensation during locomotion after nerve section in the cat

In the cat, section of all cutaneous nerves of the hindfeet except the tibial (Tib) nerve supplying the plantar surface results in a long-lasting decrease in the intensity of Tib stimulation needed for a threshold response in flexor muscles and an increase in the amplitude of the phase-dependent responses recorded in various muscles during locomotion. Stimulating through chronically implanted nerve cuffs ensured a stable stimulation over time. The increase in reflex amplitude was well above the small increase in the amplitude of the locomotor bursts themselves that results from the denervation. Short latency responses (P1) were seen in flexor muscles, especially at the knee (semitendinosus) and ankle (tibialis anterior and extensor digitorum longus), with stimuli applied in the swing phase and also to a lesser degree in the later part of the cycle. Longer latency responses (P2) were increased in hip, knee, and ankle flexors, as well as in a contralateral extensor (vastus lateralis) when applied in late stance. Responses evoked from stimulating the proximal end of sectioned nerves were not larger than before neurectomy. This suggests that the increased responsiveness to Tib stimulation is not simply caused by an increase in motoneuron excitability, because this would have resulted in a nonspecific increase of responses to stimulation of any nerve. It is concluded that the adult locomotor system is capable of central reorganization to enhance specific remaining cutaneous reflex pathways after a partial cutaneous denervation of the paw.     

The role of cutaneous afferents from the distal hindlimb in the regulation of the step cycle of thalamic cats

Summary The pad and the plantar surface of the foot were stimulated electrically in thalamic cats. Weak stimulation evoked an extensor reflex in the animal at rest. The same stimuli in a spontaneously walking animal applied during the stance phase produced an increase both in amplitude and duration of the ongoing extensor activity. When given during the swing phase, the stimuli either prolonged the ongoing flexor activity and/or shortened the following extensor burst. These changes in flexor and extensor burst duration were reflected in changes in the step cycle duration.Similar results were seen with direct stimulation of the sural nerve. For the latter experiments the ipsilateral hindlimb was fixed and denervated except for the ankle extensors and flexors, which showed rhythmic contractions correlated normally with the walking movements of the three remaining limbs. At rest, threshold stimulation of the sural nerve evoked a reflex contraction in the triceps surae of the fixed leg. The same stimuli applied during the contraction phase of the fixed triceps surae during walking resulted in a larger and longer extensor contraction and a delay of the following flexion. Stimulation during the relaxation phase of the fixed triceps surae reduced the duration of the following contraction phase. The findings are discussed in relation to the possible role of cutaneous input during locomotion.     

Interlimb coordination in cat locomotion investigated with perturbation

Summary During locomotion of decerebrate and awake walking cats, perturbation (mechanical tap) was applied to the paw dorsum of the left forelimb (LF), and the responses of both forelimbs were recorded cinematographically and electromyographically (EMG). When the tap was applied during the LF stance phase, the duration of the ongoing LF stance was shortened by 10%; in the right forelimb (RF), the duration of the concomitant swing was shortened by 32%. A tap during the LF swing phase prolonged the duration of the ongoing LF swing phase and the concomitant RF stance phase by 55 and 15%, respectively. Analysis of RF joint angle excursions showed that the shortening of the RF swing phase was related mainly to acceleration of extension movement in the late swing phase; the prolongation of the RF stance phase was related to prolonged extension movement in the late stance phase. While EMG activities were relevant to these limb movements, a notable observation was that, by tapping the LF during the LF stance phase, EMG activity in the RF extensor started well before onset of the elbow extension movement to place down the limb; without the tap, the extensor activity started shortly after onset of the extension. Closely related to changes in phase durations of each forelimb, the period of bisupport phase where both forelimbs were in stance, was retained for more than 40% of that of unperturbed steps, even when the RF or LF made the first touchdown after the tap. The rostrocaudal level at RF touchdown after the tap was comparable to unperturbed steps. These findings on interlimb relation suggest that neural control ensures coordinated movements between symmetric limbs during locomotion.Supported by grant no. 557033 from the Japanese Ministry of Education, Science, and Culture     

Scratch responses in normal cats: hindlimb kinematics and muscle synergies

1. Scratch responses evoked by a tactile stimulus applied to the outer ear canal were characterized in nine adult cats. Chronic electromyographic (EMG) electrodes were surgically implanted in selected flexor and extensor muscles of the hip, knee, and ankle joints to determine patterns of muscle activity during scratching. In some trials EMG records were synchronized with kinematic data obtained by digitizing high-speed ciné film, and in one cat, medial gastrocnemius (MG) tendon forces were recorded along with EMG. For analysis the response was divided into three components: the approach, cyclic, and return periods. Usually scratch responses were initiated with the cat in a sitting position, but in some trials the animal initiated the response from a standing or lying posture. 2. During the approach period the hindlimb ipsilateral to the stimulated ear was lifted diagonally toward the head by a combination of hip and ankle flexion with knee extension. Hindlimb motions during the approach period were associated with sustained EMG activity in hip-flexor, knee-extensor (occasionally), and ankle-flexor muscles. Initial hindlimb motions were typically preceded by head movements toward the hindpaw, and at the end of the approach period, the head was tilted downward with the stimulated pinna lower than the contralateral ear. During the return period movements were basically the reverse of the approach period, with the hindpaw returning to the ground and the head moving away from the hindlimb. 3. During the cyclic period the number of cycles per response varied widely from 1 to 60 cycles with an average of 13 cycles, and cycle frequency ranged from 4 to 8 cycles/s, with a mean of 5.6 cycles/s. During each cycle the paw trajectory followed a fairly circular path, and the cycle was defined by three phases: precontact, contact, and postcontact. On average the contact phase occupied approximately 50% of the cycle and was characterized by extensor muscle activity and extension at the hip, knee, and ankle joints. The hindpaw contacted the pinna or neck at the base of the pinna throughout the contact phase, and paw contact typically resulted in a rostral motion of the head as the hindlimb extended. 4. The postcontact phase constituted approximately 24% of scratch cycle and was usually initiated by the onset of knee flexion. Ankle and then hip flexion followed knee flexion, and flexor muscles were active during the postcontact phase as the paw was withdrawn from the head. The precontact phase constituted approximately 26% of scratch cycle and was initiated by knee joint extension and knee-extensor 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.