Cat hindlimb motoneurons during locomotion. II. Normal activity patterns

Activity patterns were recorded from 51 motoneurons in the fifth lumbar ventral root of cats walking on a motorized treadmill at a range of speeds between 0.1 and 1.3 m/s. The muscle of destination of recorded motoneurons was identified by spike-triggered averaging of EMG recordings from each of the anterior thigh muscles. Forty-three motoneurons projected to one of the quadriceps (vastus medialis, vastus lateralis, vastus intermedius, or rectus femoris) or sartorius (anterior or medial) muscles of the anterior thigh. Anterior thigh motoneurons always discharged a single burst of action potentials per step cycle, even in multifunctional muscles (e.g., sartorius anterior) that exhibited more than one burst of EMG activity per step cycle. The instantaneous firing rates of most motoneurons were lowest upon recruitment and increased progressively during a burst, as long as the EMG was still increasing. Firing rates peaked midway through each burst and tended to decline toward the end of the burst. The initial, mean, and peak firing rates of single motoneurons typically increased for faster walking speeds. At any given walking speed, early recruited motoneurons typically reached higher firing rates than late recruited motoneurons. In contrast to decerebrated cats, initial doublets at the beginning of bursts were seen only rarely. In the 4/51 motoneurons that showed initial doublets, both the instantaneous frequency of the doublet and the probability of starting a burst with a doublet decreased for faster walking speeds. The modulations in firing rate of every motoneuron were found to be closely correlated to the smoothed electromyogram of its target muscle. For 32 identified motoneurons, the unit's instantaneous frequencygram was scaled linearly by computer to the rectified smoothed EMG recorded from each of the anterior thigh muscles. The covariance between unitary frequencygram and muscle EMG was computed for each muscle. Typically, the EMG profile of the target muscle accounted for 0.88-0.96 of the variance in unitary firing rate. The EMG profiles of the other anterior thigh muscles, when tested in the same way, usually accounted only for a significantly smaller fraction of the variance. Brief amplitude fluctuations observed in the EMG envelopes were usually also reflected in the individual motoneuron frequencygrams. To further demonstrate the relationship between unitary frequencygrams and EMG, EMG envelopes recorded during walking were used as templates to generate depolarizing currents that were applied intracellularly to lumbar motoneurons in an acute spinal preparation.(ABSTRACT TRUNCATED AT 400 WORDS)     

Cat hindlimb motoneurons during locomotion. I. Destination, axonal conduction velocity, and recruitment threshold

Fine flexible wire microelectrodes chronically implanted in the fifth lumbar ventral root (L5 VR) of 17 cats rendered stable records of the natural discharge patterns of 164 individual axons during locomotion on a treadmill. Fifty-one out of 164 axons were identified as motoneurons projecting to the anterior thigh muscle group. For these axons, the centrifugal propagation of action potentials was demonstrated by the technique of spike-triggered averaging using signals recorded from cuff electrodes implanted around the femoral nerve. The axonal conduction velocity was measured from the femoral nerve cuff records. For 43/51 motoneurons, the corresponding target muscle was identified by spike-triggered averaging of signals recorded from bipolar EMG electrodes implanted in each of the anterior thigh muscles: vastus intermedius, medialis and lateralis, sartorius anterior and medialis, and rectus femoris. For 32/51 motoneurons, the recruitment threshold during locomotion was determined from the mean value of the rectified digitally smoothed EMG of the target muscle measured at the time when the motoneuron fired its first spike for each step. The recruitment threshold of every motoneuron was relatively constant for a given speed of walking, but for some units there were small systematic variations as a function of treadmill speed (range: 0.1-1.3 m/s). Recruitment thresholds were standardized with respect to the mean value of peak EMG activity of the target muscle during 16 s of walking at 0.5 m/s. For 28/51 motoneurons recorded in nine cats, recruitment thresholds (range: 3-93% of peak target muscle EMG) were linearly correlated (r = 0.51, P less than 0.02) to axonal conduction velocities (range: 57-117 m/s). In addition, for seven recorded pairs of motoneurons that projected to the same muscle in the same cat, the recruitment thresholds were ordered by relative conduction velocities. Taken together, these results are consistent with the notion that, in normal cat locomotion up to a medium trot, anterior thigh motoneurons are progressively recruited in an orderly fashion.     

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

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

Proprioceptive input resets central locomotor rhythm in the spinal cat

Summary The reflex regulation of stepping is an important factor in adapting the step cycle to changes in the environment. The present experiments have examined the influence of muscle proprioceptors on centrally generated rhythmic locomotor activity in decerebrate unanesthetized cats with a spinal transection at Th12. Fictive locomotion, recorded as alternating activity in hindlimb flexor and extensor nerves, was induced by administration of nialamide (a monoamine oxidase inhibitor) and L-DOPA. Brief electrical stimulation of group I afferents from knee and ankle extensors were effective in resetting fictive locomotion in a coordinated fashion. An extensor group I volley delivered during a flexor burst would abruptly terminate the flexor activity and initiate an extensor burst. The same stimulus given during an extensor burst prolonged the extensor activity while delaying the appearance of the following flexor burst. Intracellular recordings from motoneurones revealed that these actions were mediated at premotoneuronal levels resulting from a distribution of inhibition to centres generating flexor bursts and excitation of centres generating extensor bursts. These results indicate that extensor group I afferents have access to central rhythm generators and suggest that this may be of importance in the reflex regulation of stepping. Experiments utilizing natural stimulation of muscle receptors demonstrate that the group I input to the rhythm generators arises mainly from Golgi tendon organ Ib afferents. Thus an increased load of limb extensors during the stance phase would enhance and prolong extensor activity while simultaneously delaying the transition to the swing phase of the step cycle.     

The role of Renshaw cells in locomotion: antagonism of their excitation from motor axon collaterals with intravenous mecamylamine

Summary The contribution of Renshaw cell (RC) activity to the production of fictive locomotion in the mesencephalic preparation was examined using the nicotinic antagonist mecamylamine (MEC). After the i.v. administration of 3 doses of MEC (1.0 mg/kg) the following observations were made: 1) ventral root (VR) evoked discharge of RCs was decreased by up to 87.7%, 2) recurrent inhibitory postsynaptic potentials recorded in alpha motoneurons were greatly reduced or abolished, and 3) the rhythmic firing of RCs during the fictive step cycle was abolished in 83% of the cells examined. Locomotor drive potentials (LDPs) in motoneurons persisted during the fictive step cycle after MEC administration. Bursts of motoneuron firing during each fictive step cycle were characterized by increased frequency and number of spikes after MEC, although the burst duration was unaltered for similar step cycle lengths. A greater number and frequency of spikes per burst was also observed in Ia inhibitory interneurons (IaINs), which remained rhythmically active after MEC administration. It is concluded that Renshaw cells are not an integral part of the spinal central pattern generator for locomotion, nor do they control the timing of the motoneuron or IaIN bursts of firing during fictive locomotion. The data are consistent with a role for RCs in limiting the firing rates of motoneurons and IaINs during each burst.     

Kinesiological studies of self- and cross-reinnervated FDL and soleus muscles in freely moving cats

The activity patterns in self- and cross-reinnervated flexor digitorum longus (FDL) and soleus (SOL) muscles were examined during natural movements in awake, unrestrained cats in which electromyographic (EMG) electrodes, tendon-force gauges, and muscle-length gauges had been chronically implanted under anesthesia and aseptic conditions. Kinesiological data were recorded between 13 and 22 mo after nerve surgery. Self-reinnervated FDL and SOL muscles (i.e., FDL----FDL and SOL----SOL, respectively) exhibited locomotor activity patterns that were the same as observed in normal, unoperated FDL and SOL muscles (26). FDL----FDL muscles exhibited primarily brief bursts of activity in early swing, just after the toes had left the ground, whereas SOL----SOL muscles showed bursts of activity just before and during stance. In contrast, the cross-reinnervated muscles (both SOL----FDL and FDL----SOL) that had little or no unwanted self-reinnervation showed the patterns of activity that are associated with the innervating foreign motoneurons. That is, cross-reinnervated SOL----FDL muscles were intensely active in quadrupedal standing and, during the stance phase of stepping, producing large force transients while actively lengthening. Conversely, cross-reinnervated FDL----SOL muscles were active mainly in short bursts at the onset of the swing phase of stepping, just after the foot had left the ground. There was considerable modulation of EMG and peak force output in FDL----SOL muscles with changing speed of locomotion, whereas little modulation was evident in SOL----FDL muscles. The activity patterns in self- and cross-reinnervated FDL and SOL muscles were also recorded during scratch and paw-shaking reflexes. As in locomotion, the observed patterns were in all cases consistent with those expected for the innervating motor pool rather than the innervated muscle. Muscles that had been dually reinnervated by both the original and foreign motor pools displayed activity patterns that were a mixture of the FDL and SOL activity patterns described above. The present results demonstrate that motoneuron activation patterns remain qualitatively unaltered when their motor axons reinnervate foreign muscles. In addition, the observations permit some quantitative estimates of the degree to which cross-reinnervated muscles are subjected to patterns of motoneuron activity and to conditions of mechanical loading that are markedly different from those in the self-reinnervated or normal conditions.     

Ia inhibitory interneurons and Renshaw cells as contributors to the spinal mechanisms of fictive locomotion

The activity of selected single alpha-motoneurons, Renshaw cells (RCs), and Ia inhibitory interneurons (IaINs) during fictive locomotion was recorded via microelectrodes in decerebrate (precollicular-postmammillary) cats in which fictive locomotion was induced by stimulation of the mesencephalic locomotor region. The interrelationships in the timing and frequency of discharge among these three interconnected cell types were determined by comparing their averaged step cycle firing histograms, which were normalized in reference to motoneuron activity recorded in ventral root filaments. Previous findings that RCs are rhythmically active during locomotion and discharge in phase with the motoneurons from which they are excited were confirmed, and further details of the phase relationships between RC and alpha-motoneuron activity during fictive locomotion were obtained. Flexor and extensor RCs became active after the onset of flexor and extensor motoneuron activity, respectively. Maximal activity in extensor RCs occurred at the end of the extension phase coincidental with the onset of hyperpolarization and a decrease in activity in extensor motoneurons. Maximal flexor RC activity occurred during middle to late flexion and was temporally related to the onset of reduced flexor motoneuron activity. The IaINs recorded in the present experiments were rhythmically active during fictive locomotion, as previously reported. The quadriceps IaINs were mainly active during the extension phase of the step cycle, along with extensor RCs. Thus the known inhibition of quadriceps IaINs by RCs coupled to quadriceps and other extensor motoneurons is obviously not sufficient to interfere with the appropriate phasing of IaIN activity and reciprocal inhibition during fictive locomotion, as had been speculated. Most of the quadriceps IaINs analyzed exhibited a decrease in discharge frequency at the end of the extension phase of the step cycle, which was coincidental with increased rates of firing in extensor RCs. These data are consistent with the possibility that extensor RCs contribute to the reduction in quadriceps IaIN discharge at the end of the extension phase of the step cycle. The possibility that IaIN rhythmicity during fictive locomotion arises from periodic inhibition, possibly from Renshaw cells, was tested by stimulating the reciprocal inhibitory pathway throughout the fictive step cycle. The amplitude of Ia inhibitory postsynaptic potentials (IPSPs) varied significantly throughout the fictive step cycle in 14 of the 17 motoneurons tested, and, in 11 of these 14 motoneurons, the Ia IPSPs were maximal during the phase of the step cycle in which the motoneuron was most     

Gait-related motor patterns and hindlimb kinetics for the cat trot and gallop

To assess speed- and gait-related changes in semitendinosus (ST) activity, EMG was recorded from three cats during treadmill locomotion. Selected step cycles were filmed, and hip and knee joint kinematics were synchronized with EMG records. Swing-phase kinetics for trot and gallop steps at 2.25 m/s were compared for gait-related differences. Also, swing kinetics for different gallop forms were compared. With few exceptions, ST-EMG was characterized by two bursts for each step cycle; the first preceded paw off (STpo), and the second preceded paw contact (STpc). The two-burst pattern for the walk was defined by a high-amplitude STpo burst and a brief, low-amplitude STpc burst; at the slowest walk speeds, the STpc burst was occasionally absent. For the trot, the STpo burst was biphasic, with a brief pause just after paw off. With increasing walk-trot speeds, the duration of both bursts (STpo, STpc) remained relatively constant, but recruitment increased. Also, the onset latency of the STpo burst shifted, and a greater proportion of the burst was coincident with knee flexion during early swing. At the trot-gallop transition, there was an abrupt change in the two-burst pattern, and galloping was characterized by a high-amplitude STpc burst and a brief, low-amplitude STpo burst. At the fastest gallop speeds, the STpo burst was often absent, and the reduction in or elimination of the burst was associated with a unique pattern of swing phase kinetics at the knee. Knee flexion during the gallop swing was sustained by two inertial torques related to hip linear acceleration (HLA) and leg angular acceleration (LAA); correspondingly, muscle contraction was unnecessary. Conversely, knee flexion at the onset of the trot swing relied on a flexor muscle torque at the knee acting with an inertial flexor torque (LAA). Rotatory and transverse gallops at 4.0 m/s had similar swing phase kinetics and ST-EMG. Gait-related changes in ST-EMG, particularly at the trot-gallop transition, are not congruent with neural models assuming that details of the ST motor pattern are produced by a spinal CPG. We suggest that motor patterns programed by the spinal CPG are modulated by input from supraspinal centers and/or motion-related feedback from the hindlimbs to provide appropriate gait-specific activation of the ST.     

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)     

Cat hindlimb motoneurons during locomotion. IV. Participation in cutaneous reflexes

The responses of 11 individual motoneurons, the muscle to which each projected, plus all other muscles in the anterior thigh of the cat, were recorded following single non-noxious electrical stimuli to cutaneous nerves while the intact animal walked on a treadmill. The various excitatory and/or inhibitory responses were qualitatively similar for stimuli within the range 1.1-10 times threshold for group I fibers in the stimulated nerve (usually saphenous). Monarticular knee extensor muscles in the vastus group and their motoneurons were usually inhibited in the period 10- to 25-ms poststimulus. The faster contracting vastus medialis and lateralis muscles tended to have an excitatory rebound at approximately 25- to 40-ms poststimulus that was confined to the stance phase of the step cycle when these muscles were normally active. Biarticular hip flexor muscles rectus femoris and both the anterior and medial parts of sartorius and their motoneurons all had similar bimodal excitatory responses, including an early period 3- to 18-ms poststimulus and a later period 20- to 35-ms poststimulus. The short-latency excitatory responses appeared to be proportional to the normal recruitment of the muscles in the step cycle, whereas the long-latency responses tended to be phase advanced with respect to normal recruitment. Motoneurons projecting to muscles with two excitatory peaks tended to have similar excitatory responses at both latencies and occasionally responded at both latencies to a single stimulus.     

Treadmill locomotion in normal mice—Step related multi-channel EMG profiles of thigh muscles

Mouse models are increasingly used in current research on motor disorders. In mice, the myoelectrical activation of thigh muscles during locomotion has not yet, however, been investigated in depth. Especially intramuscular coordination has hardly been clarified. Therefore, the aims of this study were to characterize myoelectrical activity in the vastus lateralis (VL) and the biceps femoris (BF) muscle of the healthy mouse for reference purposes. The VL and the BF muscles of 12 healthy mice performing a total of 1985 steps during treadmill locomotion were investigated with two subcutaneous arrays each incorporating four electrodes. Eight-channel EMG was recorded simultaneously with high-speed videography. The EMG curves of each step were rectified and smoothed by calculating root mean square (RMS) profiles and then time-normalized for comparisons within and between animals. The EMG-activity of both muscles increased during late swing phase. The VL activity rose steeply and peaked during mid-stance phase, while the biceps activity reached a plateau during early stance phase. With increasing gait velocity, stance time decreased. The increase in gait velocity was also associated with greater EMG amplitudes. The results suggest that the BF lifts the lower hind leg during swing phase and stabilizes the leg during stance, while the VL bears the weight of the body during the stance phase.     

Lateral turns in the Lamprey. I. Patterns of motoneuron activity.

The activity of motoneurons during lateral turns was studied in a lower vertebrate, the lamprey, to investigate how a supraspinal command for the change of direction during locomotion is transmitted from the brain stem and integrated with the activity of the spinal locomotor pattern generator. Three types of experiments were performed. 1) The muscular activity during lateral turns in freely swimming adult lampreys was recorded by electromyography (EMG). It was characterized by increased cycle duration and increased duration, intensity, and cycle proportion of the bursts on the side toward which the animal turns. 2) Electrical stimulation of the skin on one side of the head in a head-spinal cord preparation of the lamprey during fictive locomotion elicited asymmetric ventral root burst activity with similar characteristics as observed in the EMG of intact lampreys during lateral turns. The cycle duration and ventral root burst intensity, duration, and cycle proportion on the side of the spinal cord contralateral to the stimulus were increased; hence a fictive lateral turn away from the stimulus could be produced. The fictive turn propagated caudally with decreasing amplitude. The increase in burst duration during the turn correlated well with the increase in cycle duration, while changes in contralateral burst intensity and burst duration did not co-vary. Turning responses varied depending on the timing (phase) of the skin stimulation: stimuli in the first two-thirds of a cycle evoked a turn in the same cycle, whereas stimuli in the last third gave a turn in the following cycle. The largest turns were evoked by stimuli in the first third of a cycle. 3) Fictive turns were abolished after transection of the trigeminal nerve or a rhombencephalic midline split, but not in a rhombencephalic preparation with transected cerebellar commissure. High spinal hemisection was sufficient to block turning toward the lesioned side, while turns toward the intact side remained. Taken together these findings suggest that the reticulospinal turn command is essentially unilateral and generated in the rhombencephalon.     

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.     

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.     

Motor patterns in human walking and running

Despite distinct differences between walking and running, the two types of human locomotion are likely to be controlled by shared pattern-generating networks. However, the differences between their kinematics and kinetics imply that corresponding muscle activations may also be quite different. We examined the differences between walking and running by recording kinematics and electromyographic (EMG) activity in 32 ipsilateral limb and trunk muscles during human locomotion, and compared the effects of speed (3-12 km/h) and gait. We found that the timing of muscle activation was accounted for by five basic temporal activation components during running as we previously found for walking. Each component was loaded on similar sets of leg muscles in both gaits but generally on different sets of upper trunk and shoulder muscles. The major difference between walking and running was that one temporal component, occurring during stance, was shifted to an earlier phase in the step cycle during running. These muscle activation differences between gaits did not simply depend on locomotion speed as shown by recordings during each gait over the same range of speeds (5-9 km/h). The results are consistent with an organization of locomotion motor programs having two parts, one that organizes muscle activation during swing and another during stance and the transition to swing. The timing shift between walking and running reflects therefore the difference in the relative duration of the stance phase in the two gaits.     

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.     

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)     

Vibration-induced changes in EMG during human locomotion

The present study was set up to examine the contribution of Ia afferent input in the generation of electromyographic (EMG) activity. Subjects walked blindfolded along a walkway while tendon vibration was applied continuously to a leg muscle. The effects of vibration were measured on mean EMG activity in stance and swing phase. The results show that vibration of the quadriceps femoris (Q) at the knee and of biceps femoris (BF) at the knee enhanced the EMG activity of these muscles and this occurred mainly in the stance phase of walking. These results suggest involvement of Ia afferent input of Q and BF in EMG activation during stance. In contrast, vibration of muscles at the ankle and hip had no significant effect on burst amplitude. Additionally, the onset time of tibialis anterior was measured to look at timing of phase transitions. Only vibration of quadriceps femoris resulted in an earlier onset of tibialis anterior within the gait cycle, suggesting involvement of these Ia afferents in the triggering of phase transitions. In conclusion, the results of the present study suggest involvement of Ia afferent input in the control of muscle activity during locomotion in humans. A limited role in timing of phase transitions is proposed as well.     

Motoneuron input-resistance changes during fictive locomotion produced by stimulation of the mesencephalic locomotor region

Input-resistance changes during fictive locomotion were monitored in a variety of extensor and flexor hindlimb alpha-motoneurons in precollicular, postmammillary decerebrate cats induced to "walk" by electrical stimulation of the mesencephalic locomotor region (MLR). Using intracellular recording techniques and injected hyperpolarizing current pulses, the changes in the motoneuron input resistance recorded at the motoneuron soma were examined during nonlocomoting control periods as well as during the depolarized and hyperpolarized phases of the membrane potential oscillations (locomotor drive potentials, or LDPs) of fictive locomotion. In 28 of the 52 motoneurons examined, no change in the input resistance between the control and locomotor periods was observed. The remainder of the cells displayed a decrease (less than 20%) in input resistance when fictive stepping commenced. Over 80% of all the motoneurons depolarized (mean depolarization 4 mV), whereas only one LG motoneuron hyperpolarized (2 mV) with the onset of stimulation of the MLR. The remaining motoneurons did not display such changes. In 43 out of 52 motoneurons examined, no significant change in the input resistance could be observed between the depolarized and hyperpolarized phases of the step cycle. A decrease in the input resistance during the depolarized phase of the LDP was observed in four LG motoneurons, whereas five other motoneurons (2 LG, 1 TA, 1 PB, and 1 ST) displayed an increased input resistance during the depolarized phase compared with the hyperpolarized phase of locomotion. The data are consistent with the presence of an excitatory synaptic input alternating with an inhibitory input to the motoneuron during the fictive step cycle.(ABSTRACT TRUNCATED AT 250 WORDS)     

Effects of red nucleus microstimulation on the locomotor pattern and timing in the intact cat: a comparison with the motor cortex.

Effects of red nucleus microstimulation on the locomotor pattern and timing in the intact cat: a comparison with the motor cortex. To determine the extent to which the rubrospinal tract is capable of modifying locomotion in the intact cat, we applied microstimulation (cathodal current, 330 Hz; pulse duration 0.2 ms; maximal current, 25 microA) to the red nucleus during locomotion. The stimuli were applied either as short trains (33 ms) of impulses to determine the capacity of the rubrospinal tract to modify the level of electromyographic (EMG) activity in different flexors and extensors at different phases of the step cycle or as long trains (200 ms) of pulses to determine the effect of the red nucleus on cycle timing. Stimuli were also applied with the cat at rest (33-ms train). This latter stimulation evoked short-latency (average = 11.8-19.0 ms) facilitatory responses in all of the physiological flexor muscles of the forelimb that were recorded; facilitatory responses were also common in the elbow extensor, lateral head of triceps but were rare in the physiological wrist and digit extensor, palmaris longus. Responses were still evoked in most muscles when the current was decreased to near threshold (3-10 microA). Stimulation during locomotion with the short trains of stimuli evoked shorter-latency (average = 6.0-12.5 ms) facilitatory responses in flexor muscles during the swing phase of locomotion and, except in the case of the extensor digitorum communis, evoked substantially smaller responses in stance. The same stimuli also evoked facilitatory responses in the extensor muscles during swing and produced more complex effects involving both facilitation and suppression in stance. Increasing the duration of the train to 200 ms modified the amplitude and duration of the EMG activity of both flexors and extensors but had little significant effect on the cycle duration. In contrast, whereas stimulation of the motor cortex with short trains of stimuli during locomotion had very similar effects to that of the red nucleus, increasing the train duration to 200 ms frequently produced a marked reset of the step cycle by curtailing stance and initiating a new period of swing. The results suggest that whereas both the motor cortex and the red nucleus have access to the interneuronal circuits responsible for controlling the structure of the EMG activity in the step cycle, only the motor cortex has access to the circuits responsible for controlling cycle timing.