Critical points in the forelimb fictive locomotor cycle and motor coordination: evidence from the effects of tonic proprioceptive perturbations in the cat

During locomotion, the limbs of one girdle must remain coordinated in different conditions. To understand the neural mechanisms underlying such coordination, tonic protraction/ retraction of one shoulder or tonic flexion/extension of one elbow was applied during fictive locomotion in high decerebrate and paralyzed cats. We studied bilateral changes in the timing and amplitude characteristics of electroneurographic (ENG) muscle nerve bursts of cleidobrachialis (ClB, elbow flexor and shoulder protractor) and the two heads of triceps (long, TriLo, elbow extensor and shoulder retractor and lateral, TriLa, elbow extensor). Perturbations induced bilateral changes in amplitude and timing of ENG bursts that were anchored on certain critical points in the cycle. These critical points could correspond to morphological characteristics within the bursts or to bilateral onsets or offsets of ENG bursts. For instance, in response to shoulder and elbow perturbations, burst changes occur in relation to a fixed point, labeled point C, occurring at about mid-extensor burst and corresponding to a simultaneous abrupt increase in TriLa amplitude and a decrease in amplitude of contralateral ClB. At a point labeled B, corresponding to about mid-flexor burst, ClB amplitude increases above control with elbow extension or starts decreasing with shoulder protraction. Although cycle reorganization is specific for each type of tonic perturbation, a common feature is that the changes in burst duration are achieved through discrete shifts between consecutive critical points. It is postulated that coordination may be based on a discrete temporal cycle structure along which critical points delimiting burst components are shifted.     

Coordination between head and hindlimb motions during the cat scratch response

Coordination between motions of the head and the hindlimb paw ipsilateral to the stimulated pinna were assessed during the scratch cycle in freely moving cats. Motor patterns were determined by electromyographic (EMG) recordings made from epimysial-patch electrodes surgically implanted on the biventer cervicis (BC), complexus (CM), obliquus capitis inferior (OC), and splenius (SP) muscles and by fine-wire EMG electrodes implanted in two ankle muscles, medial gastrocnemius (MG), and tibialis anterior (TA). To assess head motions during the three phases of the scratch cycle (precontact, contact, postcontact), several responses were filmed, and in some cats an in vivo force transducer was implanted on an ankle extensor muscle (MG or plantaris, PL) to determine the tension profile during the scratch cycle. During the scratch cycle, the head's trajectory was usually characterized by a small oscillation in which the head was pushed away during paw contact (as hindlimb joints extended) and then repositioned during the noncontact phases (as hindlimb joints flexed). Neck muscle activity did not occur during all responses or during all cycles of a single multicycle scratch response, and when it occurred, neck muscle EMG was characterized as phasic (a single burst during the cycle) or tonic (low-level activity during the entire cycle). Neck muscles ipsilateral (i) to the scratching limb exhibited phasic bursts more than contralateral (c) muscles, and phasic activity was most frequently observed in the iBC, iSP, iOC, and cOC muscles. The cOC was reciprocally active with the ipsilateral muscles, and its burst coincided with the postcontact phase and the ankle flexor (TA) burst. The ipsilateral muscles (iOC, iSP, iBC) were active during paw contact, and the termination of all three bursts occurred synchronously just after peak tension of the ankle extensor was reached. The iBC was active before the onset of paw contact and may have been responsible for repositioning the head, along with the cOC, during the precontact phase. The iOC became active after the onset of paw contact (22 ms) and was recruited more often when the peak extensor tendon force was high (10–16 N). The iSP, in contrast, was active during the contact phase of most scratch cycles examined and its recruitment appeared to be unrelated to tendon forces. Our data suggest that phasic neck muscle activity is not obligatory during the cat scratch response, but is related to certain conditions such as a higher than average tendon force of an ankle extensor during contact and the need to reposition the head during the noncontact phases of the cycle. During contact it is possible that active muscle contraction helps to minimize head motion to provide a stable contact surface for the paw, thereby maximizing the possibility of the scratch stimulus being dislodged from the pinna. Possible neural mechanisms, both reflexes and central commands, responsible for coordinating motions of the head and hindlimb during the scratch cycle are discussed.     

Effects of motor cortex and single muscle stimulation on neurons of the lateral vestibular nucleus in the rat

The neuronal responses to stimulation of motor cortical sites and of forelimb single muscles were studied in the lateral vestibular nucleus of anaesthetized rats. Of the 228 neurons tested for response to stimulation of contralateral motor cortex, 63% responded to cortical sites controlling extensor muscles and 30% to those controlling flexors. The corresponding figures for responders to ipsilateral stimulation were 34 and 21%. Vestibulospinal units responded to cortical sites controlling extensor and flexor muscles whereas the remaining lateral vestibular nucleus neurons, very reactive to cortical sites controlling extensor muscles, responded little to contralateral and not at all to ipsilateral cortical sites controlling flexor muscles. The effects evoked by contralateral cortical sites controlling extensors varied, those induced by cortical sites controlling flexors were inhibitory in 77% of cases. The responses to ipsilateral motor cortex stimulation differed not so much by cortical sites controlling extensor or flexor muscles as by whether the neuron was in the dorsal or ventral zone of the lateral vestibular nucleus: mixed in the former, all inhibitory in the latter. Of the lateral vestibular nucleus units tested for response to stimulation of ipsilateral or contralateral forelimb distal muscles, only 11% responded. All the vestibulospinal units responsive to muscle stimulation lay in the dorsal zone of the nucleus. The remainder, dorsal or ventral, were not responsive to contralateral muscles. Single lateral vestibular nucleus cells influenced both by ipsilateral muscle and by contralateral motor cortex made up 24% of the pool, vestibulospinal and non-vestibulospinal. They fell into three groups: responsive to one or both structures but responding more strongly to combined stimulation; responsive to each of the two structures but showing a response to combined stimulation not significantly different from that evoked by the cortex alone; responsive only to combined stimulation. The lateral vestibular nucleus units included in these three groups accounted for 29% of those tested for response to extensor muscles and cortical sites controlling extensors and 15% of those tested for response to flexor muscles and cortical sites controlling flexors. Twenty-five per cent of the vestibulospinal neurons responded both to contralateral muscles and to ipsilateral motor cortex stimulation but none of the non-vestibulospinal neurons responded to both. All the responders to both were in the dorsal zone of the lateral vestibular nucleus and responded to extensor stimuli, always in the same way. These results indicate that motor cortex output exerts a major influence on lateral vestibular nucleus discharges, while the muscle afferents have a modulatory influence on the lateral vestibular nucleus responses to cortex.(ABSTRACT TRUNCATED AT 400 WORDS)     

Bilateral actions of the reticulospinal tract on arm and shoulder muscles in the monkey: stimulus triggered averaging

The motor output of the pontomedullary reticular formation (PMRF) was investigated to determine the reticulospinal system’s capacity for bilateral control of the upper limbs. Stimulus triggered electromyographic averages (StimulusTA) were constructed from muscles of both upper limbs while two awake monkeys (Macaca fascicularis) performed a reaching task using either arm. Extensor and flexor muscles were studied at the wrist, elbow, and shoulder; muscles acting on the scapula were also studied. Post-stimulus effects (PStEs) resulted from 435 (81%) of 535 sites tested. Of 1611 PStEs analyzed, 58% were post-stimulus suppression (PStS), and 42% were post-stimulus facilitation (PStF). Onset latency was earlier for PStF than PStS, duration was longer for PStS, and amplitude was larger for PStF. Ipsilateral and contralateral PStEs were equally prevalent; bilateral responses were typical. In the ipsilateral forelimb and shoulder, the prevalent pattern was flexor PStF and extensor PStS; the opposite pattern was prevalent contralaterally. Sites producing strong ipsilateral upper trapezius PStF were concentrated in a region caudal and ventral to abducens. The majority of muscles studied had no clear somatotopic organization. Overall, the results indicate the monkey PMRF has the capacity to support bilateral coordination of limb movements using reciprocal actions within a limb and between sides.     

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.     

Vestibulospinal and reticulospinal neuronal activity during locomotion in the intact cat. I. Walking on a level surface

To examine the function of descending brain stem pathways in the control of locomotion, we have characterized the discharge patterns of identified vestibulo- and reticulospinal neurons (VSNs and RSNs, respectively) recorded from the lateral vestibular nucleus (LVN) and the medullary reticular formation (MRF), during treadmill walking. Data during locomotion were obtained for 44 VSNs and for 63 RSNs. The discharge frequency of most VSNs (42/44) was phasically modulated in phase with the locomotor rhythm and the averaged peak discharge frequency ranged from 41 to 165 Hz (mean = 92.8 Hz). We identified three classes of VSNs based on their discharge pattern. Type A, or double peak, VSNs (20/44 neurons, 46%) showed two peaks and two troughs of activity in each step cycle. One of the peaks was time-locked to the activity of extensor muscles in the ipsilateral hindlimb while the other occurred anti-phase to this period of activity. Type B, or single pause, neurons (13/44 neurons, 30%) were characterized by a tonic or irregular discharge that was interrupted by a single pronounced and brief period of decreased activity that occurred just before the onset of swing in the ipsilateral hindlimb; some type B VSNs also exhibited a brief pulse of activity just preceding this decrease. Type C, or single peak, neurons (9/44 neurons, 23%) exhibited a single period of increased activity that, in most cells, was time-locked to the burst of activity of either extensor or flexor muscles of a single limb. The population of RSNs that we recorded included neurons that showed phasic activity related to the activity of flexor or extensor muscles [electromyographically (EMG) related, 26/63, 41%], those that were phasically active but whose activity was not time-locked to the activity of any of the recorded muscles (13/63, 21%) and those that were completely unrelated to locomotion (24/63, 38%). Most of the EMG-related RSNs showed one (15/26) or two (11/26) clear phasic bursts of activity that were temporally related to either flexor or extensor muscles. The discharge pattern of double-burst RSNs covaried with ipsilateral and contralateral flexor muscles. Peak averaged discharge activity in these EMG-related RSNs ranged from 4 to 98 Hz (mean = 35.2 Hz). We discuss the possibility that most VSNs regulate the overall activity of extensor muscles in the four limbs while RSNs provide a more specific signal that has the flexibility to modulate the activity of groups of flexor and extensor muscles, in either a single or in multiple limbs.     

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)     

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)     

Functional organization within the medullary reticular formation of the intact unanesthetized cat. III. Microstimulation during locomotion.

1. This article presents the results from stimulation in 21 loci within the medullary reticular formation (MRF; between 0.5 and 2.5 mm from the midline) and in 5 loci in the medial longitudinal fasciculus (MLF) of four intact, unanesthetized cats during locomotion. Stimulus trains (11 pulses, 0.2-ms duration, 330 Hz, stimulus strength 35 microA) were applied at those loci in each track at which the most widespread effects in each of the four limbs were obtained with the cat at rest. Electromyograms were recorded from flexor and extensor muscles of each limb. 2. As previously reported, stimulation with the cat at rest generally evoked brief, short-latency, twitch responses in both flexor and extensor muscles of more than one limb. In contrast, stimulation during locomotion evoked a more complex pattern of activity in which responses were normally evoked in one or other of the muscle pairs and incorporated into the locomotor pattern. 3. In the majority of sites, the stimulation evoked excitatory responses in the flexor muscles of each of the four limbs during that period of the step cycle in which each respective muscle was naturally active; stimulation in the stance phase of locomotion, although less effective, was also capable of producing responses in these muscles. All three ipsilateral extensor muscles studied [long and lateral heads of triceps and vastus lateralis (Tri, TriL, and VL, respectively)] were normally inhibited during their phase of muscle activity, although excitatory responses were occasionally seen. Responses in the contralateral (co) Tri were invariably excitatory and were largest during the period of muscle activity, whereas responses during the period of activity of the coVL were mixed, with both excitatory and inhibitory responses being seen from any one locus. 4. Excitatory responses were normally largest when stimulation was applied during the time that the muscle was active during the locomotor cycle. Responses evoked at times when the muscle was inactive were sometimes larger than those evoked with the animal at rest; such responses were most commonly seen in the hindlimb flexors and in the coVL. 5. In both flexors and extensors of each of the four limbs, the latency of the responses was greatest when the cat was at rest and least for stimuli given during the period of activity of the respective muscle. Average latencies during the period of muscle activity ranged from a minimum of 9.0 +/- 2.6 (SD) ms for inhibitory responses in the ipsilateral Tri and TriL to a maximum of 17.1 +/- 3.0 ms for the responses evoked in the ipsilateral semitendinosus.(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)     

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.     

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.     

Supraspinal and segmental signals can be transmitted through separate spinal cord pathways to enhance locomotor activity in extensor muscles in the cat

 The fine control of locomotion results from a complex interaction between descending signals from supraspinal structures and sensory feedback from the limbs. In this report, we studied the interaction between vestibulospinal volleys descending from Deiters’ nucleus and group I afferent input from extensor muscles. It has been shown that both pathways can exert powerful control over the amplitude and the timing of muscle bursting activity in the different phases of the step cycle. The effects of stimulating these pathways on the fictive locomotor rhythm were compared in decerebrate, partially spinal cats (ipsilateral ventral quadrant intact) injected with nialamide and l-dopa. As reported before, stimulation of both Deiters’ nucleus and group I fibres from ankle extensor muscles, when given during the flexor phase, stopped the flexor activity and initiated activity in extensors. When applied during the extensor phase, the same stimulation prolonged the extensor activity and therefore delayed the onset of flexor activity. This similarity suggests that the two pathways might converge on common spinal interneurones. This possibility was tested with the spatial facilitation technique in lumbosacral motoneurones. Deiters’ nucleus and group I fibres from extensor muscles were stimulated with different intensities and with several different coupling intervals. Motoneurones showing clear di- and/or polysynaptic excitation from both pathways were retained for analysis. Surprisingly, in all cases, there were no signs of spatial facilitation, but a simple algebraic sum of the two excitatory postsynaptic potentials. This result indicates that each input acts on the rhythm generator through separate interneuronal pathways. Received: 20 August 1996 / Accepted: 14 November 1996     

Reflexes induced by nerve stimulation in walking cats with implanted cuff electrodes

Summary Neural cuffs, implanted around various hindlimb nerves (sural, common peroneal, posterior tibial), were used to deliver brief stimulus trains to unrestrained cats walking on a treadmill. The resulting perturbations of the step cycle were evaluated by analyzing the EMG bursts recorded from the ankle extensors and by high speed cinematography. It was found that relatively weak stimulation (1.4 to 2 X T) of the posterior tibial nerve was very effective in eliciting a prolongation of the flexion phase provided the stimuli were applied just prior to the expected onset of the ankle extensor EMG burst. This ipsilateral hyperflexion was correlated with a prolongation of the contralateral extension. The same stimuli applied during stance usually evoked a yielding of the stimulated leg and a prolongation of the ongoing contralateral stance. In addition to these flexor and extensor reflex effects, it was found that low threshold stimulation of the sural and common peroneal nerves resulted in a powerful reflex activation of the ankle extensors. In contrast, stimulation of the gastrocnemius-soleus nerve (a muscle nerve) produced no discernible behavioral effects, even for stimuli at 3 X T, indicating that the observed reflexes are probably mediated by cutaneous afferents. The results were largely confirmed in experiments using the same cuffs implanted in spontaneously walking premammillary cats.     

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.     

Effect of hindlimb unloading on two hindlimb muscles during treadmill locomotion in rats

The purpose of the study was to examine the pattern of electromyographic (EMG) activity of the rat soleus (SOL) and tibialis anterior (TA) muscles during treadmill locomotion at various speeds after 7 days of hindlimb unloading (HU). Raw EMG signals were processed to determine cycle duration, burst duration and mean EMG (burst surface divided by its duration). Cycle duration and SOL burst duration increased after HU (+7% and +5%, respectively) while TA burst duration decreased (?16%). After HU, the alternating pattern of activity between extensor and flexor muscles was maintained. Nevertheless, a co-activation of the two muscles was sometimes observed. The EMG pattern revealed no difference in the timing of the coordination between flexor and extensor muscles after HU. The delay between TA offset and SOL onset was increased (+12 ms), but this increase could be explained by the decrease in TA burst duration. Neither TA burst duration nor TA mean EMG were changed with increased treadmill speed, so that the flexor muscle activity was not related to speed of locomotion. These results would suggest that SOL activity is centrally programmed. Moreover, it is proposed that a decline in afferent feedback from SOL in rats which are suspended has an effect upon the locomotor pattern, leading to an hyperexcitability of SOL motoneurons and, via reciprocal inhibition, to a reduction in TA activity.     

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.     

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)     

Short-latency crossed inhibitory responses in extensor muscles during locomotion in the cat

During locomotion, contacting an obstacle generates a coordinated response involving flexion of the stimulated leg and activation of extensors contralaterally to ensure adequate support and forward progression. Activation of motoneurons innervating contralateral muscles (i.e., crossed extensor reflex) has always been described as an excitation, but the present paper shows that excitatory responses during locomotion are almost always preceded by a short period of inhibition. Data from seven cats chronically implanted with bipolar electrodes to record electromyography (EMG) of several hindlimb muscles bilaterally were used. A stimulating cuff electrode placed around the left tibial and left superficial peroneal nerves at the level of the ankle in five and two cats, respectively, evoked cutaneous reflexes during locomotion. During locomotion, short-latency ( approximately 13 ms) inhibitory responses were frequently observed in extensors of the right leg (i.e., contralateral to the stimulation), such as gluteus medius and triceps surae muscles, which were followed by excitatory responses ( approximately 25 ms). Burst durations of the left sartorius (Srt), a hip flexor, and ankle extensors of the right leg increased concomitantly in the mid- to late-flexion phases of locomotion with nerve stimulation. Moreover, the onset and offset of Srt and ankle extensor bursts bilaterally were altered in specific phases of the step cycle. Short-latency crossed inhibition in ankle extensors appears to be an integral component of cutaneous reflex pathways in intact cats during locomotion, which could be important in synchronizing EMG bursts in muscles of both legs.     

Entrainment of the locomotor rhythm by group Ib afferents from ankle extensor muscles in spinal cats

Summary 1. Previous studies have concluded that the timing of the locomotor rhythm can be strongly influenced by input from group Ib afferents from leg extensor muscles (Duysens and Pearson 1980; Conway et al. 1987). The main objective of the present study was to obtain additional evidence for this conclusion by examining the characteristics of entrainment of the locomotor rhythm by rhythmic stimulation of group I afferents and by rhythmic force pulses in the ankle extensor muscles. 2. A reduced, non-immobilized preparation was developed in spinal cats that allowed isometric contractions of ankle extensor muscles to be elicited by ventral root stimulation during the expression of locomotor activity. The same preparation was used to examine the influence of electrically stimulating group I afferents from the ankle extensors and the effect of rhythmically stretching these muscles. The locomotor rhythm was initiated by sustained mechanical stimulation of the perineum following the administration of Clonidine and, in some preparations, Naloxone. 3. The timing of the onset of flexor burst activity was examined during entrainment with saw-tooth and ramp-and-hold stretches of the ankle extensor muscles. Flexor bursts were initiated about 200 ms following the release from the stretch, and this latency was independent of the entrainment frequency. 4. The locomotor rhythm was readily entrained by rhythmic contractions of the ankle extensor muscles produced by ventral root stimulation provided the magnitude of the contractions was greater than about 10N. Repetitive stimulation of group I muscle afferents from the ankle extensors also entrained the locomotor rhythm, with the timing of motor activity being similar to that during entrainment with rhythmic muscle contractions. Burst activity in the ipsilateral extensors was coincident with the stimulus trains in both cases. This similarity argues for entrainment being produced mainly by input from group Ib afferents. 5. The functional implication of the results of this and previous studies is that input from group Ib afferents during the stance phase of walking acts to inhibit generation of flexor burst activity and to promote extensor activity. The proposal that a decline in Ib activity near the end of the stance phase is involved in regulating the stance to swing transition is discussed.CRSN, Physiologie, Faculte de Medecine, C.P. 6128, succursal A, Montréal, Quebec, Canada, H3C 3J7